UNIT 9 NUTRITION, EXCRETION AND

OSMOREGULATION
Structure
9.1 Introduction

_ 9.2
Objectives
Nutrition in Non-Chordates
Feeding and Digestion in Sponges, Coelenterates and Flatworms
Feeding and Digestion in Annelids
Feeding and Digestion in Molluscs
Feeding and Digestion in Echinoderms
Feeding and Digestion in Arthropods
9.3 Excretion in Non-Chordates
Protonephridia and Metanephridia
Malpighian Tubules
Goelornoducts of Molluscs
9.4 Osmoregulation in Non-Chordates
Os~~~oregulationin Fieshwater Non-Chordates
Osrnoregulation in Marine Non-Chordates
Water Conservation in Ttrrestrial Non-Chordates
9.5 Summary
9.6 Terminal Questions
9.7 Answers

INTRODUCTIOP'
In Unit 2 of this course, you have learnt about mechanisms of feeding and digestion as
well as the r,lGchanismsof excretion and osmoregulation in protozoans. In this unit you
will study aspects of nutrition, excretion and osmoregulation in multicellular non -
chordate animals. These animals exhibit a variety of feeding habits. The success with
1 which the animals exploit the food resources could be compared only with the ~legance
and complexity of their structural and functional adaptations. In this unit you will first
study nutrition, that is, the feeding habits, and the various adaptations for feeding and
digestion in the multicellular non-chordates. The second aspect that will be covered in
this unit is excretion.

Excretion is concerned with the removal of metabolic wastes that arise as a result of
oxidation of energy rich compounds and metabolism of proteins and nucleic acids. This
is carried out in non-chordate metazoan animais by two distinct sets of tubular structures
- the nephridia and coelomoducts. The ne:,hridia and coe~omoductsare variously named
in different groups of animals depending on their complexity and location. Terrestrial
arthropods have evolved another type of excretory organ known as Malpighian tubules
st~qcturallydifferent form nephridia and coelomoducts. In this unit you will study in
detail these organs and how they carry out excretion.

A thiid aspect of study covered in this unit relates to the regulhtion of water and ionic
content of the body of non-chordate metazoans. These may be osmoconformers or
osmoregulators. Osmoconformers maintain their internal body fluid in osmotic
equilibrium with the aqueous environment in which they live. In other words, the salt
concentration of the internal and external media is more or less same. Other organisms
.,are osmoregulators and they maintain the~oncentration of their internal body fluids
relatively constant. This may be often at a different osmotic and ionic level from that of
the environment. In this unit, you will also study the mechanisms of regulation of water
and ionic content of body fluids in these animals.
Objectives
After studying this unit, you should be able to:
describe the structures associated with feeding and the mode dffeeding in different
non-chor te metazoan phyla,
list the excretory organs found among the various groups of non-chordate
metazoan invertebrates and describe their function@g,and
Comparative Forms and outline the mechanisms of water and ion regulation in organisms occupying different
Functions . habitats.

9.2 NUTRITION IN NON-CHORDATE METAZOANS

The vast majority of invertebrates feed on particulate food material of very small size.
They can be broadly classified as microphagous organisms. 1.. contrast, feeding on large
masses of food is called macrophagy. Generally large invert~~l-ates are macrophagou?.
Macrophagous feeders could be active predators and may feed on live material.
Cephalopod n~olluscs,for example, are completely predaceous. Large crustaceans ,nd in
general all the living arthropods are macrophagous. Before you proceed further, it is
suggested that you go back to the Unit 2, Block 1 of this course 'Protozoa' and revise the
concepts on various tlzpesof animal nutrition such as 'autotrophy, heterotrophy, holozoic
and saprozoic'.

Microphagous animals make effective use of their cilia or setae for feeding . They are
therefore commonly known as ciliary feeders. Ciliary feeders fall into two categories.
One type known as .~spensionfeeders feed on minute organisms and other particulate
matter suspended in water. In this type of feeding food particles are extracted by filtering
water. Hence the orgallisms are known as filter feeders.

Filter feeders set up a current in surrounding water with their cilia or setae. The food is
collected by filtration as well as by trapping food in mucous. Much of the filtered
material may be inedible or harmful or larger than the size of the particle that the
organism can feed on. Such materials are usually discarded by a w.ell developed sorting
cum rejecting mechanism. The chosen food particles are then directed towards the
mouth. Both sessile and free swimming organisms have evolved filter feeding
mechanisms. Free swimming animals move in water containing food particles. Sessile
animals depend on natural currents in water as well as those created by cilia and other
appendages. The"sessile organisms also feed on deposits of organic material that
accumulate on substratum as well as in the sand or mud. The deposit feeders, like
suspension feeders, depend on cilia for feeding. In fact some organisms may feed on
both suspended and deposited food materials. When the material is deposited on sand or
mud, the substratum itsqlf is swallowed. Other feeders depend on encrusted organisms
like algae, polyzoans and sponges as food organisms. The mouth parts of such feeders
are modified for rasping and browsing food. Sedentary polychaets, molluscs - more
particularly lamellibranchs, sponges, pterobranchs, small crustaceans as well as a number
of other groups of small animals are all microphagous filter feeding organisms. Rasping
feeding is very characteristic of many gastropod molluscs and they use their radula for
such a purpose. You will now study feeding and digestion in chosen groups of
organisms. But before doing so, attempt the following questions

SAQ 1
Fill in the blanks with suitable words:
i) Feeding on large masses of food is termed ...............................
ii) Microphagous animals make use of ............................ for effective feeding.
iii) The .............................cum .......................... mechanism discards too large
or otherwise unsuited particles.
iv) Rasping feeders such as gastropods make use of ........................... for feeding.

9.2.1 FEEDING AND DIGESTION IN SPONGES, COELENTE-

RATES AND FLATWORMS
Sponges
Sponges make use of their canal system for feeding, respiration and excretion. For a
detailed account of canal system in sponges, you may refer to unit 5 of Block I1 of this
course.

Sponges feed by filtering particulate matter from nutritive water that enters into
choanocyte lined spongocoel through minute pores. These pores act as simple but
effective sorting device that permit the entry of only the smallest particles. Each of the
-.
 Nutrition, Excretion and
choanocytes or collar cells possesses a single flagellum surrounded at its base by a Osmoregulation
protoplasmic collar. The flagella beat outwards from their base and move water
forward. The smaller food particles are ingested from the water by choanocytes and the
slightly larger ones by wandering amoebocytes and even by dermis cells. The digestion is
intracellular, within the food vacuoles (Fig.9.1). The partly digested food vacuoles from
choanocytes may also be passed on +oneighboring amoebocytes. Amoebocytes serve
also to store food materials. In sycon~dand leuconid sponges, with complex body form
choanocytes line small flagellated cham5 thereby increasing the area of ingestive
epithelium. Water is expelled from the spongocoel through osculum and thus the entry
of new nutrient rich water is facilitated.

Water current

Mesohyl amoeboid cell

Fig.9.1 : lntracellular digestion i n fresh water sponge.

Coelenterates
Coelenterates have tissue grade of organisation and therefore their feeding and digestive
mechanisms are also more complex. Coelenterates, in general, are macrophagous
carnivores. They use tneir tentacles to capture the prey. Their prey include all types of
small animals especially crustaceans. Some scyphozoans and anthozoans may even feed
on fish (Fig.9.2). However, certain anthozoans and scyphozoans are microphagous
suspension feeders. They either trap the prey in mucus secreted by the tentacles or filter
plankton from water passing through tentacular fringe. For instance the anemone
Metridium and the jelly fish Aurelia collect small organisms by the tentacles in a mucous
secretion and move the food to the mouth by ciliary action.

Macrophagous carnivorous coelenterates use cnidoblasts(Fig.9.3) for trapping the food

by tentacles. Cnidocil projecting from cnidoblast, as is found in Hydra, is believed to be
the sensory element. Each cnidoblast discharges one nematocyst. Each nematocyst has a- .
Cytoplasm
pear shaped vesicle and a thread coiled within it. When discharged the nematocyst
pierces into the prey and injects poison into it.

I Fig.9.3: Structure of cnidoblast.

Comparative Forms and The swallowing of food follows its capture and paralysis by nematocysts. The
Functions swallowing is done by enlarging the mouth and expanding the gastrovascular cavity.
These two actions facilitate the passage of food into coelenteron or gastrovascular cavity
(GVC) (Fig.9.4) - the site of digestion of food.

Fig.9.4: L. S. of Hydra showing the gastrovascular cavity and the endoderm.

Digestion in coelenterates is both extracellular and intracellular. During extracellular

digestion, the large prey is broken into smaller particles in the GVC. Some of the
endoderm cells are secretary in nature and they release the enzymes into GVC.

-Dlnctive mesenteries Slphonoglyph

Scsondary mcscntcries

Primary mesenteries

Fig. 9.5: (a) L.,S of a sea anemone (b) Cross section at the level of pharynx filament.
In Hydra: a prey like Daphnia is broken down into small particles within four hours of Nutrition, Excretion and
Osmoregulation
its ingestion by secreting the enzymes into GVC. Besides the secretory cells, there are
also absorptive cells which are concerned with intracellular digestion. An hour after
ingestion, during which time the prey is actively broken down, the absorptive cells
begin to ingest the food particles into small vacuoles and digestion is completed with in
these cells. Absorptive cells also store the reserve food materials and even accumulate
the undigested residues. The accumulated intracellular waste is finally released by the
fragmentation of the absorptive cells into GVC. From GVC they are ejected out through
the mouth. Thus a combination of intracellular and extracellular digestion takes care of
the nutritional requirements.

In anthozoan coelenterates also such a combination is found. You may refer to Unit 5 of
this course for a detailed account of an anthozoan organisation. You may recall that
GVC of anthozoans is subdivided by the mesenteries. The thickened edges of
mesenteries known as mesenteric filaments carry the secretory and absorptive cells
(Fig.9.5). In macrophagous anthozoans the prey is firmly held by the filaments, with tk e
help of nematocysts located there and the digestive enzymes are directly applied to the
food. This.process increases the efficiency, of the action of digestive enzymes.
Coelenterates, being carnivores, secrete mostly lipolytic and proteolytic enzymes. Very
weak amylase activity is also recorded. Other carbohydrases are not reported.
Flatworms
Free living flatworms are known to digest the food ( I ) intracellularly, (2) both
intracellularly and extracellularly and ( 3 ) exclusively extracellularly.

i) Exclbsive intracellular digestion is seen in acoelan Convoluta. The endoderm

appears as a solid syncytiurn that can protrude through the ventral mouth. The
syncytium functions as if it were a pseudopodium and engulfs the food to form food
vacuoles. Digestion occurs intracellularly.

ii) In the triclad worm Polycelis the food such as crustaceans is trapped in the mucous
secretion. The long protrusible pharynx (Fig. 9.6) is inserted into the prey and the
soft contents are withdrawn. The food is broken down to small particles
extracellularly while passing to the alimentary canal. On reaching the gut, the
particulate food is engulfed by digestive cells and intracellular digestion takes
place. Although Polycelis feeds on large organisms, the food ingested is actually
made up of very.smal1 particles and hence the organism is to be regarded as
microphagous.

Epidermis

Protruded pharynx -.
Fig.9.6: Diagrammatic L.S. of Polycelis to show the protrusion of pharynx. ,

iii) The polyclad worm Cycloporus feeds exclusively on ascidian colonies Botryllus and
Botrylloides, sucking the individual zooids with the help of protrusible pharynx.
The wall of the pharynx is ruffled and plicate. The food arrives almost intact in the
gut. In contrast to Polycel~swhere digestion is exclusively intracellular, in
Comparative Forms and Cycloporus it is exclusively extracellular. The food is homogenised and digested
@netions with the alimentary tract and there is no intracellular digestion.
In trematodes which are parasitic, the digestive system is highly branched. The mouth
leads into pharynx which is followed by a short oesophagus and a branched intestine L.
bearing a number of diverticula (Fig. 9.7). The highly ramified diverticula fill the most
of the interior of the body. A trematode such as liver fluke-feedson the biliary matter as
well as the blood of the host. The muscular pharynx aids in sucking of the food.
Digestive glands have lost their utility in these animals and therefore they are absent.
The food is already in a state ready for absorption. The branched alimentary canal helps
in the reaching of the food to all part of the body.

SAQ 2
State whether the following statements are true or false. ,

i) . The ingestion of food particles in sponges is done by choanocytes and wandering

amoebocytes.
ii) Coelenterates are in general microphagous, herbivorous feeders
iii) Cnidoblasts are food capturing cells in coelenterates.
iv) Coeleriterates digest the food extraceliularly.
v) All flatworms can digest their food intracellularly.
' 9.2.2 FEEDING AND DIGESTION IN ANNELIDS
Oligochaetes
Among annelids, the oligochaetes which comprise mostly earthworms, feed on dead,
decomposing organic vegetation. They also digest organic matter contained in the soil
which they swallow during burrowing. Aquatic forms feed on detritus, algae and
microorganisms. Freshwater forms like Aeolosoma collect detritus with its prostomium.
The food is collected by creating a partial vacuum when the ciliated ventral surface of
prostomium is placed against the substratum and the center is elevated by muscular
contraction. The partial vacuum thus created releases the food particles which are swept
into mouth by cilia. Some oligochaetes like Chaetogaster catch amoebae, ciliates,
. rotifers and trematode larvae by the sucking action of pha~ynx.

rsal bbod vessel

Intestinal caecum

Fig. 9.7: Digestive system of an oligochaete.

Oligochaetes possess a simple and straight alimentary tract (Fig.9.7). The mouth
situated at the base of prostomium leads into a buccal cavity which opens into a spacious
 Nutrition, Excretion and
and muscular pharynx. Iri aquatic forms, the pharynx is an eversible organ and is mucus Osmoregulation
covered to which food panicles adhere. In earthworms, it functions as a pump.

Pharyngeal glands secrete mucus as well as digestive enzymes. Pharynx is followed by a

narrow tubular oesophagus. The oesophagus is modified at different levels into a gizzard
and a crop. The muscdar and cuticle- lined gizzard functions as triturating organ and
grinds the food particles. If crop is present it functions as a storage organ. Rest of the
digestive system, starting anywhere from 1 81h and 27" segment is intestine. Anterior
half of the intestine is secretory in function and the posterior half is absorptive. Besides
carbohydrales, proteases and lipases, earthworms also secrete cellulase and ch~tinase.
The absorptive area of the intestine is increased by a fold of tissue called typhlosole
present in middorsal wall. A layer of yellowish peritoneal cells called chloragogen cells
around the intestine is,the site of intermediary metabolism in oligochaetes. These cells
have the same function as the liver in vertebrates and fatbody in insects, and is the site of
synthesis as well as storage of glycogen and fat.
.,
Polychaetes
Polychaetes include both free moving (errant) and sedentary species. The free moving 1
species are generally macrophagous and the sedentary forms are microphagous. Nereis is
an example of a free moving macrophagous polychaete. But Nereis can also burrow and
make use of its proboscis (buccal cavity and pharynx) for this purpose (Fig. 9. 8). The
eversible proboscis is a feeding organ. The pharynx is lined with hooked jaws and is
everted for seizing the prey. The genus Nereis includes species which are carnivorous.
Some species of Nereis are omnivorpus, feeding on diverse material such as algae, .
invertebrates or detritus on the substratum. Some are scavengers. The tentacles and
palps present around the mouth assist in the manipulation of food, besides serving
sensory function.

Mouth

, Oesophagus

Pharynx

Buccal cavity

Osophageal gland

Intestine

Fig. 9.8: The digestive system ofhrrrris.

Fig.9.9: Terebella in a feeding posture.

~ o m ~ a r a t i vForms
e and The detritus feeding lugworms (Arenicola and related species) swallow the mud or sand
unctions in which the food is contained. The swallowing is done by the sucking action of the
proboscis which is the anterior end of the alimentary cana!. Being burrowers, they lack
tentacles. Other sedentary polychaetes have evolved very fine mechanisms of ciliary-
mucous feeding. For example, terebellid worms (e.g. Amphitrite, Terebella) live in
, permanent mud tubes and are detritus feeders (Fig.9.9). From the tubes they extend their
long ciliated prostomial tentacle clusters over the surface of the substratum. The food
particles consisting of detritus trapped in the mucus. The cilia are actively involved in
the movement of food particles. The mucus-trapped food particles pass through the
ciliated grooves of the tentacles and enter into the mouth.

Sobella, a large polychaete, builds tubes that project from the surface of mud in the
littoral zone of the sea. The coordinated action of cilia present on the branchial crown of
tentacles (Fig.9.10) creates water currents and the food is extracted from the currents.
The branchial crown of tentacles form a wide funnel at the base of which the mouth of
the an~mallies. The tentacles or the filaments of the branchial crown bear rows of
outgrowths called pinnules. Toward the lower part of branchial funnel, the pinnules
rnterlock and form a filtering system in which the food particles are trapped on the
pinnules. There are three types of cilia on the pinnules. These cilia produce a current of
water. Food particles that enter the funnel along with the stream of water are directed
into a groove that runs along the inner edge of each pinnule. The cilia located at the
base of the pinnule drive the food particles to the base of pinnules from where they enter
into a ciliated longitudinal groove that runs the entire length of filament and then into the
mouth.

Gill pinnule
Gill filament

basal fold
basal web
Dorsal tip

Vental sac
Parallelfolds .
Dorsal collar fold '
Ventral collar fold

Ventral groove shield

Thoracic parapodium
Ventral uncini
Dorsal bristle bundle

Abdominal parapodium
Ventral bristle bundle
,Dorsal uncini
Vennal collar

Fig9.10: Sabeua with its branchial tentacles.

 Nutrition, Excretion and
In Chaetopterus, which lives in 'U' shaped tubes made of sand or mud (Fig.9.1 I), a Osmoregulation
branchial crown is absent. Water is drawn by the beating of three pairs of fan shaped
structures that are derived from parapodia as well as by another pair of wing like
outgrowths located anteriorly. Mucus secreted by these structures traps the food particles
and is drawn into a ventral groove. Here it is shaped into a conical bag and the food
particles are filtered through the bag. The filtered food is rolled intc a ball by a
secretion of the wing like structures. The cilia in the ventral groove fransport the ball of
food to the mouth.

Annelids generally exhibit extracellular digestion, although phagocytosis by the

alimentary epithelium and completion of digestion of the ingested food by wandering
amoebocytes are reported in Arenicola marina. Digestion is mostly extracellular in filter-
feeding terebellids. The digestive system in polychaetes is differentiated into an
oesophagus, a fore-stomach, a muscular hind-stomach and an intestine. Enzyme secretion
is therefore confined to fore-stomach and fore-intestine. In general digestion occurs in
fore-stomach and fore-intestine. The hind-stomach is actually a mixing region.
Absorption takes place in the hind-intestine.

Mouth
-7%

Fig.9.11: (a) Cltnefopfenis inside the tube (b) Enlarged anterior r f ~ i o nto show the food gathering
apparatus. >

Hirudinea includes free living and ectoparasitic leeches. LeTches are blood suckers.
The digestive system consists of a preoral chamber which accommodates a sucker at
the base of which lies the triradiate opening ,the mouth. The walls of the mouth are
embedded with jaws, three in number, of which one is mid-dorsal and the other two
ventro-lateral. Each jaw bears a row of minute teeth. The alimentary canal consists of a
thick muscular pharynx followed by a short oesophagus. Unicellular salivary glands are
found on either side of the pharynx. The oesophagus is followed by a crop, an extensive
sac consisting of 10 to 11 chambers, the chambers communicate with each other by
openings surrounded by sphincters (Fig. 9.12 ). Each camber is provided with a pair of
backwardly directed lateral caeca. There is a progressive increase in the size of caeca,
the anterior pairs being smaller in size, and the caeca of last chamber being the largest.
Following the crop, there is a small round stomach with a much folded inner wall, and
the stomach in turn leads into a narrow intqstine the inner wall of which irprovided with
horizontal and transverse folds tb increase the absorptive surface. The intestine is
followed by rectum that opens outside by anus.
 -
Comparative Forms and
Functions

Fig. 9.12: Digestive system o f a leech.

Leeches feed on blood of man and cattle. They attach themselves to the victim b, their
suckers and make an incision in the skin with their jaws. The muscular pharynx sucks
the blood. The salivary secretions contain an anti-coagulant, the hirudin which prevents
the clotting of blood and ensures a free flow of blood into the crop. The crop is thus a
reservoir for the storage of blood. Once a leech has a'full feed, it may not again feed for
several months. The food from the crop passes into stomach from time to time and the
digestion takes place in the stomach. The'digested food is absorbed in the intestine.

SAQ 3
Match the items givermnder A with the ones given under B.

1) Oligochaeta , a) macrophagous feeders

2) Errant (free moving) polychaetes b) ciliary mucous. feeders
3) Sedentary polychaetes c) chloragogen cells
4) Tube dwelling polychaetes d) phagocytosis
5) Arenicola e) hematophagous feeders
6) Leech. f) microphagous feeders

9.2.3 FEEDING AND DIGESTION IN MOLLUSCS

Lamellibranchs
We begin the study of feeding and digestion in molluscs with bivalves or lamellibranchs.
Lamellibranchs resort to ciliary-mucus type of filter feeding as witnessed in some
polychaetes. They are semi-sessile animals confined to their protective shells and inhabit
nluddy or sandy substrata. These are microphagous feeders and make use of their ciliated
gills or lamellae as food-gathering devices. The lamellae are structured to filter
suspended and deposited material from a current of \;later that enters into the animal
 Nutrition, Excretion and
through inhalant siphon and exits through exhalant siphon.. During this, waier current is Osmoregulation
maintained by the action of cilia present on the gills.

Each gill or ctenidium is formed of two delnibranchs attached to a central axis and each
demibranch has a parallel row of filaments. The structure of the filament varies in
different groups of bivalves. In primitive protobranch, the tilaments lack folds but in
filibranchs and eulamellibranchs, the filaments are folded to from ascending and
descending limbs (Fg.9.13). Figure 9.13 illustrates evolution of lamellibranch gills. In
Mytilus, a filibranch, adjacent filaments are joined by ciliary junctions. In Anodonta, a
eulamellibranch, the fila'ments are joined to each other by vacuolar interfilamental
junctions. In lamellibranchs cilia are arranged in frontal and lateral series on gill
filaments (Fig.9.14).

Fig.9.13: ( I ) Vertical section of lamellibranchs to show the differences in the structure of ctenidia. (A)
protobranch, (6) filibranch and eulamellibranch, (C) septibranch.

Water is drawn into the mantle cavity by lateral cilia. As water passes through the
filaments, the food is gathered by fronto-lateral cilia and thrown towards frontal cilia
(Fig. 9.14). The food particles now entangled in mucus are sweptvover the surface of gill
lamellae and find their way to food groove. This may be ventral marginal groove or
dorsal groove along the axis of the gill, depending on the species.

Mucus in . .
% t
food grhve
A B
Fig. 9.14: Structure of the gill in lamellibranchs (A) vertical section (B) horizontal section ofgill in
Mytilus. .

Food material is then carried to the two pairs of labial palps lying on each side of the
mouth and from there to the mouth. 'The ciliary tracts on gills as well as labial palps do
the sorting of food. Fine cilia on certain gill tracts convey the fine particulate food to the
mouth and the coarse cilia on the other tracts reject the large particles (Fig.9.15). The
sorting mechanism here depends on the weight of particles. The heavier particles that
settle down on $e palps are swept away by powerful ciliary currents. Lighter ones avoid
this current, sink to the bottom and are swept towards the mouth. The rejected particles
get entangled in mucus and are ejected out through exhalant siphon. ,
-
 --
combarative Forms ahd
, Functions

Upper margin
Outer pale face. -\\
- point where meterial is rejected
from palps

Proximal
otel groove

Lateral oral
groove

-. -
Base of demibranch of palps

Fig.9.15: Junction of palps and gills in 0sirea to show the route of water and food particles. X-the point
at which the material is rejected from palps.

Lamellibranchs are characterised by a structure called crystalline style, a very compact

and long rod formed by the consolidation of mucus. The style is secreted by the style
sac. The style sac is an extension of the stomach, either opening directly into stomach or
cut off from the stomach. The style sac is lined with cilia which rotate the crystalline
style and drive it forward into the stomach. The free end of the style is worn away by the
alkaline stomach contents as well as its friction against the gastric shield which is a
thickening of the cuticular lining of the stomach. During this process the style substance
releases amylase, a carbohydrate digesting enzyme. In some species cellulase is also
released. Thus extracellular digestion of carbohydrates is initiated in the stomach. The
rotation of the mucus-cum-food strand by the crystalline style as well as the pH of the
stomach contents(pH5-6) result in the continuous separation of the food and mucus into
fine particles and their detachment from the main strand. The detached particles are
sorted out by the stomach wall based on their size. The larger and heavier particles are
transported by cilia into the intestine where they are converted into faecal pellets and
removed, The finer particles are carried towards the openings of digestive diverticula or
gland which is a characteristic feature of molluscan digestive system. Each one of the
two diverticula is highly branched system of blind tubules. The ciliated epithelium lining
the tubules is formed of vacuolated cells which are phagocytic. The cells ingest the fine
particles into food vacuoles and digest them. Thus digestion in lamellibranchs is largely
intracellular, although secretions from style Sac initiate carbohydrate digestion
extracel lularly.
Gastropods
In most gastropods the digestion is extracellular. However, some herbivore gastropods
like Crepidula are ciliary feeders and has a digestive system similar to that of
lamellibranchs. In.them amylase is the only extracellular enzyme; the digestive
diverticula do not secrete any enzyme but are only absorptive in function. Crepidula
has also a crystalline style similar to those of lamellibranchs. Some herbivorous
gastropods belonging to Taenioglossa, Rhapidoglossa and Pteropoda also possess
crystalline style. It is confined to those herbivorous gastropods that are microphagous
feeders, including rasping feeders that use radula (Fig. 9.16). Patella, Haliotis, Aplysia
and Helix are some otthe herbivorous gastropods that do not possess a crystalline style.
Essentially in those forms where extracellular digestion has replaced intracellular
digestion, a crystalline style is absent.
 Nutrition, Excretion end
Radula sac Odontophore Osmoregult~tion
Salivary gland
...

Radula membrane
Radula teeth
Radula retractor

Odontophore retractor /'aiula

u- Mouth
wOdontophore protractor
protractor (a)

Fig.9.16: (a) Radula rpparatus of a gastropod; (b) and (c) protraction and retraction of radula.

Cephalopods
Cephalopods are carnivorous. Tentacles or arms are food capturing organs. The number
of tentacles vary in different cephalopods. Sepia has ten arms, Octopus has eight and
Nautilus has around 90 tentacles. Sepia, like other squids and cuttlefishes, has four pairs
of short and heavy tentacles called arms and a pair of long structures called tentacles.
The irner surface of the arms are provided with suckers which are stalked, cup-shaped
and adhesive in nature. The suckers are provided with horny rings and hooks. In the
mobile tentacles the suckers are present only at the flattened ends. The arms aid in
holding the prey tightly after it is captured. Suckers are present in the arms of Octopus
as well but they are stalkless and devoid of horny rings and hooks. Cephalopods, besides
radula possess a powerful pair of beak shaped jaws in the buccal cavity. The jaws are
used for tearing and biting the prey before the tongue like action of radula pulls the
food down and aids in swallbwing. Octopods inject the prey with poison or without a
bite with jaws and the prey is flooded with enzymes. The partially digested food ihen
passes into gut. While feeding on shelled gastropods, octopods drill a hole in the prey
with radula and then inject poison into the animal through the hole. Whereas cuttlefish
feed on surface inhabiting organisms such as shrimps and crabs, octopods feed on a
variety of prey including clams, snails and crustaceans. Nautilus is a scavenger-predator
feeding specially on decapod crustaceans, particularly hermit crabs.
I
'The buccal cavity leads into an oesophagus which conducts the food into stomach by
peristalticaction. In Nautilus and Octopus there is a crop, which is an extension of
oesophagus. The stomach of cephalopods is highly muscular and contains a caecum
attached to its anterior end. The walls of the caecum absorb the digested food and the
absorptive function is carried out to a certain extent by the intestine as well. . The
intestine opens outside by an anus (Fig.9.17).

In cephalopods, the digestion is exclusively extracellular. The digestive glands of

cephalopods include salivary glands -two pairs of them located on either side of buccal
cavity. The posterior pair of salivary glands secrete poison which are glycoproteins. In
Octopus they also secrete proteolytic enzymes. Cephalopods also possess pancreas and
liver. In squidi the two structures are separated from each other and the pancreas empty
their secretions into the duct of the liver. Enzymes of both the glands are emptied into
stomach, at the junction between stomach and caecum. In fact the 'liver' in Ocfdpus
Comparative Forms and performs three functions, the secretion, absorption and excretion, very similar to the
Functions hepatopancreas of other molluscs.

Stomach Liver
Oesophagus
Caecum
\ \ I / Lip

I I J

Fig 9.17: Digestive system of Loligo.

9.2.4 FEEDING AND DIGESTION IN ECHINODERMS

Echinoderms exhibit a variety of feeding habits. Most of them are carnivorous. But some
of them are suspension feeders, some are deposit feeders, others could be scavengers and
still others are grazers. We shall briefly study the digestive system and the nutrition of
. different group of echinoderms. .

In asterojds (e. g. starfish) digestive system is radial, and is modified with the flattening
of the body. The system is compressed onoral-aboral axis (Fig 9.18). The mouth lies in
the middle of a muscular peristomial membrane and leads into a short oesophagus,
followed by the stomach. The stomach shows two regions, a large oral chamber - cardiac
stomach and a small aboral chamber - the pyloric stomach. The pyloric receives ducts
from pyloric caeca which are the digestive glands. There are two pyloric caeca in each
arm. The caeca are glandular mass of cells suspended in the coelom of the arm by a
mesentery. The pyloric stomach leads in to a short intestine that opens on the aboral
disc by an anus. The intestine bears a pair of caeca, the rectal caeca.

Asteroids are scayengers and carnivores, and feed on invertebrates such as snails,
bivalves. Crustaceans, polychaetes and other echinoderms. When a starfish comes across
a prey such as a mussel, it bends its flexible arms over the body of the prey and attaches
the tube-feet to the two valves with the suckers. The two valves are pulled apart and the
soft body of the mussel is exposed. Then the cardiac stomach is everted through the
mouth and wrapped over the soft body of the mussel; the stomach engulfs the prey and
the digestive juices are on it from the pyloric caeca. At the completion of the
digestion, the stomach is withdrawn into the body, leaving the shell outside. The
digestion is thus extracellular and the products of digestion are stored in the cells of

Anus
Intestinal caecum ( R9ctum
M / Py loric stomach
Cardiac stomach
Stone canal ,
Py loric caecum

coelom

I
.
Tentacle
Oesophagus EW
9.18: Digestive system of a starfish.

pyloric caeca or passed through the caeca into the coelom for distribution. The
undigested waste is ejected through the anus by pumping action of rectal caeca.
 Nutrition, Excretion and
Ophiuroids, of which brittle stars are examples, are also carnivorous and they could be Ckmoregulation
scavengers, deposit feeders or filter feeders. The food consists of polychaetes, molluscs
and small crustaceans. Food is usually captured and brought to the mouth by arm
looping. The alimentary canal consists of mouth, a short oesophagus, and a saccular
I
I
stomach that ends blindly. The stomach is eversible. There is no intestine or anus and the

'1
system does not extend into arms. Extracellular and intracellular digestion as well as
I absorption occur mostly within the stomach. During filter feeding, the arms are waved in
water, and the planlcton and detritus adhere to mucous strands strung between the spines
of adjacent arms. The trapped food particles may be swept downward by a reduced spine
called tentacular scale by the ciliary action. Or the food particles may be collected by the
two pairs of tube feet located close to the mouth of each arm. The tube feet are then
scraped across the tentacular scales. The collected particles are deposited in front of the
scale, compacted into a bolus and passed along the mid-dorsal line of the arm towards the
mouth by the ciliary action.

Echinoids consisting of sea urchins, heart urchins and sand dollars are grazers and use
their teeth for straping the substratum on which they live. They feed on a wide variety of
plant and animal material. and the algae are most important food. The mouth leads into a
buccal cavity and then into a pharynx , both of which are surrounded by a masticatory
I apparatus, the Aristotle's lantern. 0esoihagus follows the pharynx and runs vertically
upwards close to the aboral end continues into stomach. The saccular stomach at first has
a downward course, reaches almost the oral end and then curves, runs anti-clockwise
closely apposed to the inner side of the test. The stomach then continues into intestine
w N h runs parallel to stomach clockwise (Fig.9.19). The intestine is followed by rectum.
The oesophagus, stomach and intestine are all suspended by mesenteries. Attached with
the stomach, at both ends, is a narrow tube, the accessory intestine or intestinal siphon.
One end of the siphon opens into stomach at its junction with the oesophagus and the
other end passes into the intestine. Digestion is extracellular, begins in the stomach and
1 is completed in the intestine. The function of the siphon is to remove excess water from
the food before the digestion commences in the stomach.

Anus,

Fig. 9.19: Digestive system o f a sea urchin.

Upper end of the tooth

Lower end of th; tooth

Fig.9.20: Aristotle's lantcrn o f r sea urchin.

Comparative Forms and Aristotle's lantern, the masticatory apparatus, is formed of five plates or jaws fitted
Functions together into an inverted pyramid (Fig. 9.20) surrounding the pharynx. Each plate is a
triangular framework, within which a long pointed tooth is present. Since there are fivr
jaws, there are five teeth as well. The tooth can slide up and down in the frame by
muscular action. In addition to the jaws and teeth, Aristotle's lantern also consists of a
number of smaller, rod like pieces at the aboral end. The lantern can be protruded from
the mouth by the action of the muscles. In addition to scraping the lantern is also useful
in pulling and tearing of food. Aristotle's lantern is absent in heart urchins.

Sea cucumbers belonging to class Holothuroidea are suspension or deposit feeders. The
alimentary canal consists of mouth, the buccal chamber, a wide oesophagus, an
indistinguishable stomach, a long intestine and the cloaca that opens to the exterior (Fig
9.2 I). The mouth is located in the middle of a buccal membrane at the base of the
tentaclular crown. When the animal is disturbed the mouth and the tentacles can be pulled
completely into the anterior end of the body. The branched tentacles are the food
capturing organs. They are either swept over the bottom or held out in'sea water. In both
instances the food particles are trapped on to the adhesive papillae located on the
tentacular surfaces. One by one the tentacles are stuffed into the pharynx and the food
particles are wiped off the adhesive papillae even as the tentacles are pulled out of the
mouth.

Radial ambulacral ues

Longitudinal muscles

I)orsal intestinal vessel

Ventral intestinal vessel

Circular muscles

Radial ambulacral vessel

b l o a c a l aperture
Fig.9.21: Digestive system of holothurian.

The pelmatozoan echinoderms, the crinoids, are suspension feeders. The alimentary
. canal consists of the mouth, a wide oesophagus, a spacious stomach with diverticula, a
coiled ascending intestine opening outside by anus. During feeding, the arms and
pinnules are outstretched and the tube feet are erect. The tube feet appear as small
tentacles and bear mucus secreting papillae along their length. The food particles trapped
in the podia are passed into the mouth by the ciliary currents.
9.2.5 FEEDING AND DIGESTION IN ARTHROPODS
Arthropods are known to lack cilia. Feeding in arthropods is carried out by the various
appendages. Virtually every pair of appendages may be modified in one or other group
of arthropods to aid in the feeding process. Whereas terrestrial arthropods in general are
macrophagous and predaceous (there are many herbivores among insects), aquatic
arthropods, mainly crustaceans have adapted themselves for filter feeding.
 Nutrition, Excretion and
Crustacea Osmoregulation

Branchiopods (Anacostraca) provide a good example for filter feeding mechanism. The
appendages are of foliaceous type(leaf-like) called phyllopodia and they beat
metachronalty. The inner edge of these appendages bears a series of endites, the most
basarendite being larger than the others. The backwardly directed endites bear large setae
which are also backwardly directed. The outer edge of the limb has several lobes. The
most distal of these lobes is the exopodite while the basal ones are epipodite and
protopodite.

The limbs perform different functions. They are locomotory, feeding and because of
their delicate structure can also function as respiratory organs. The beating of limbs
produces currents in surrounding water and brings in the food particles. In between the
limbs present on one side, there is an interlimb space (Fig.9.22 and 9.23). The food
materials are trapped on to the setae of endites which act as the filtering elements of the
food capturing mechanism. The filtered food materials are taken to the mouth.

Fig.9.22: Median view of three consecutive limbs of a branchiopod to show interlimb space, setae and
setules. I n limb I the setae are removed. The setules are shown by stippling the area. I n limb
r 2 the endites are removed to show the interlimb space and the chsnnel into which it opens.
Limb 3,shows endites with complete setae.

Copepods like Calanus resort to maxillary filter feeding. In these organisms feeding
current is created .by a swimming vortex.. The maxillary setae act as a filter and collect
the food when the water passes through them. The setae on maxillular endites and.
maxillipedes remove the food from the maxillary setae and pass it onto the mouth.

Crustaceans like other.arthropods have a ventral mouth that leads into almost a straight
alimentary canal. The foregut is essentially enlarged stomach that harbours a triturating
apparatus. Usually the apparatus is made up of chitinuous ridges, denticles and calcareous
ossicles all located on the wall of the stomach. The length of the midgut varies in
different graups of crustaceans and contain one to several pairs of caeca which increase
the absorpti;e surface of the midgut. ~e~atopancreas are spongy digestive glands
composed of secretory tubules and ducts and are the sourceof proteinases and lipases.
Both foregut and midgut are the sites of digestion. Absorption takes place across the
midgut walls and the tubules Hepatopancreas also serves as a storage organ. The midgut
Comparative Forms and leads into rectum that opens to the exterior by anus. Let us b;iefly look into the digestive
Functions system and nutrition of different groups of crustaceans.

Fig.9.23: A to E: the first to fifth pair of trunk limbs of Dtrplmicr: the g9nthobase of seeond pair of limbs
bears an elongate s e b that assists in the removal of food particles from the filtering setLe of
3rd and 4Jh pair of limbs.
In malacostracans the food is caught by chelipeds (the thoracic appendages) and is
passed to the third maxillipedes which in turn push the food between the mouth parts.
The mandibles and maxillae help in tearing the food into pieces before they are directed
into the mouth. The foregut consists of an oesophagus, a cardiac stomach and a pyloric
stomach all of which have chitinuous lining. Among the various decapod crustaceans
the stomach is variously thickened to from ossicles or teeth which together form the
gastric mill. The gastric mill together with the movement of stomach walls triturates the
food. The pyloric stomach is divided into two portions, the dorsal one that leads directly
into intestine and the ventral one, a bilobed gland filter which leads into hepatopancreas
by two large ducts. The hepatopancreas is a large bilobed structure. Besides secreting
the digestive enzymes, the hepatopancreas can also carry out intracellular digestion of
particulate food, absorption of digested food materials, storage of nutrients, and vesicular
packaging of indigestible wastes and their removal by exocytosis.

Decapods combine predatory feeding with scavenging. Large invertebrates like

echinoderms and bivalves are predated upon by crabs and king crabs. Many fresh water
and marine shrimps on the other hand are scavenging - detritus feeders. Some of the
crabs feed by scraping the algae from rocks and some others feed on detritus from the
surface of the sand. Fiddler crabs resort to filter feeding. This is also true of many
burrowing shrimps, pea crabs, porcelain crabs and mole crabs. Wood boring marine
isopod malacostracans feed on wood and their hepatopancreas secrete cellulase to diges.
the cellulose of wood.

Branchiopodcrustaceans are suspension feeders and the food particles are collected by
the setae of the trunk appendages. The collected food particles are transmitted. .to a
 Nutrition, Excretion and
mouth. In branchiopods, the oesophagus forms the foregut and it is the midgut that is ~hnore~~~lrtion
gnlarged to form the stomach. The intestine is coiled one to several times.

Ostracod crustaceans show a variety of feeding habits. They are either carnivores or
herbivores; some of them are scavengers and still others are filter feeders. Plant food
mostly consists of algae and the animal food includes other crustaceans, small snails and
,annelids.
I
Copepod crustaceans also have a range of feeding habits. Planktonic copepods are
suspension feeders feeding on phytoplankton. Some of the planktonic forms are
omnivores and predaceous as well. Bottom dwelling copepods feed on microorganisms
and detritus attached to sand grains, algae or sea grasses.

Cirriped crustaceans are suspension feeders or predators. In suspension feeders, the

beating of cirri create water currents and the food particles are trapped in the setae. The
first pair of cirri scrape the food particles attached to the setae on the other cirri and
transfer them to the mouth parts. Both mandibles and maxillae help to macerate the food.
Stalked barnacles such as Lepas capture copepods, amphipods and other relatively large
1 organisms and hence predaceous. Rhirocephalan cirripedes are parasitic and usually
parasitise crabs.
Insects
Among the terrestrial arthropods insects have developed a variety of adaptations in their
mouth parts to suit their varied feeding habits. A detailed account of the mouth parts of
insects is provided in unit 5 of this course. Briefly the mouth parts are the labrum,
mandibles, the first and the second maxillae and these are variously modified to suit the
I type of feeding that a particular group of insects has specialised. Mouth parts of insects
fall under two categories - the mandibulate type and suctorial type. Orthopterans,
dictyopterans, coleopterans are some of the groups that have mandibulate type of mouth
parts. Butterflies and mosquitoes are examples of insects with suctorial mouth parts. In
1 mandibulate type (Fig 9.24), the mandibles are well developed and the maxillae remain

Submentum

Mentum

Paraglossa

Labial palp

1st maxilla (Labium) sub-menturn .Labellurn

Fig:9.24 :Mandibulate type of mouth parts. Fig.9.25: Sbctorirl type of mouth parts.

in their typical unmodified form. The mouth parts are adapted for cutting, chewing and
crushing the food. In suctorial type (Fig. 9.25) the mandibles are more or less reduced
and the maxillae are modified into stylets used as piercing organs. Labium also known as
second maxillae may also be modified in different ways for piercing and sucking. The
two broad types can be further subdivided into five categories: (a) biting type with
toothed mandibles as is seen in cockroaches; -(b) sucking type consisting of a proboscis .
which are modified maxillae as found in butterflies suited for obtaining nectar from
flowers; (c) piercing and sucking type suited for piercing the tissues of animals and
plants, such as sucking blood formanimals and fluids from plants, as is found in
mosquitoes and bugs; (d) biting and sucking type as is-found in hoaey bees and (e)
sponging and suckirrg type-as is found in houseflies. .

-
In general, alimentary canal of insects is divided into three regions the stomdaeum or ,
foregut, the mesenteron or midgut and proctodaeum or hindgut (Fig 9.26). The foregut
-

55 1
Compar?tive Forms and
Funstions
arises as an anterior ectodermal invagination and the proctodaeum as a posterior 1
invagination. The two r'egions are lined with chitin. The midgut is a endoderm
derivative. The foregut consists of a short oesophagus, a sac like crop and a
proventriculus or gizzard that may harbour a masticatory apparatus, specially in solid I
food feeders. Gizzard is well developed in cockroaches and in grasshoppers and is known 1
as honey-stomach in honey bees. Fluid feeders may be provided with a reservoir that is
usually attached with oesophagus as seen in Diptera, Lepidoptera and Hymenoptera. I

The midgut is often tubular and is the major site'of digestion and absorption. The I
1
epithelium lining the midgut is formed of three types of cells. One type is secretory that
secretes digestive enzymes ; the second type ih generative that replaces the secretory
cells that are destroyed during secretion; and a third type of cells known as goblet cells of
uncertain function. In many insects the midgut lined by a membrane formed by loose
chitin fibres known as peritrophic membrane. This membrane protects the midgut
epithelium from abrasion and injury by contact with fuod particles. The Malpighian
tubules located at the junction ofthe mid and hindgut, mark the beginning of hindgut.
The hindgut shows three regions', a narrow ileum, a coiled colon and a broad rectum.
Rectal glands are found connected with rectum in many insects. Both rectum and rectal
glands play a role in resorption of water from faeces before it is ejected through anus:
i

Mesenteron

\
Ileum
Rectum colon
9.26: Digestive system of an insect.

Many groups of insects have salivary glands and the glands assume varied form and
structure. In Orthoptera the salivary glands are large structures and consist of four lobes
and reservoir. In lepidopterans the salivary glands are tubular filaments. In hemipterous
insects the glands are formed of four lobes and a reservoir. In hematophagous (blood
sucking) insects such as mosquitoes, the saliva contains an anti-coagulant that facilitates
free flow of blood in o the crop. In honey bees, the salivary secretions serve several
/t
pruposes - inverting sugars, forming preservatives like formic acid and aiding in pollen
digestion.
Myriapods and Arachnids
Very briefly, you will study the digestive system of other terrestrial arthropods.
Centipedes and millepedes, belonging to class Myriapoda have a typical arthropodan
digestive system. In centipedes the alimentary canal consists of a short foregut, a very
long midgut extending almost the entire length of the body and a short hindgut (Fig.
9.27). A pair of salivary glands open into the foregut. Most centipedes are predators and
feed on a variety of small arthropods. Some of the chilopods are known to feed on small
frogs, toads, snakes, birds and mite. Burrowing centipedes feed on earthworms, snails
and nematodes. Mandibles, first maxillae, second maxillae and labium are the mouth
parts (Fig 9.28).
 (b) First maxilla

(a) Mandible

Fig.9.27: Digestive system of a centipede. Fig. 9.28: Mouth pats of a centipede.

In millipedes the alimentary canal is almost very similar to those of centipedes. But
unlike centipedes, the millipedes are herbivorous and feed on decaying vegetation. The
mouth parts consist of besides a short seven segmented antennae, a mandible and a
structure called gnathochilarium (Fig. 9.29). 'The mandibles are composed of a proximal
cardo, a middle stipes and a distal mala mandibularis. The distal piece bears a large
movable tooth and a masticatory plate provided with toothed ridges. The gnathochilarium
resembles the labium of insects and is actually formed by the fusion of maxillae of two
sides. Mala gnathochilarii interior
Lingulal lobe i gnarliochilarii exterior

StlpeS gllathochilarii

L.ingua

Cardo gnatliocliilar~i

Prementum

A: Mandible

B: Gnathochilarium
Fig.9.29: Cnathochilarium of a millepede.
Comparative Forms and Arachnids are generally carnivorous and predaceous. They feed on small arthropods.
Functions The prey is captured and killed by the cephalic appendages the chelicera and pedipalpi.
Part of the digestion takes place outside the body of the animal. While the prey is held by
chelicerae, digestive enzymes from the midgut are poured on the prey. Partially digested
semisolid food is passed onto the oesophagus of foregut through mouth and pharynx.
The food then reaches mesenteron which has a central tube and a number of lateral
diverticula. All the diverticula are filled with partially digested food. Further digestion ,
occurs in the midgut diverticula. The digested food is absorbed by the absorptive cells of
the diverticula. The mesenteron extends throughout the length of the abdomen where in
'
the last part of it is sclerotised and forms the hind gut. The hindgut opens out through
anus.

SAQ 4
Choose the correct answer from the alternatives provided.
% , I) Lamellibranchs are ciliary- mucusfrasping feeders.
2) In eulamellibranchs the gill filaments lack foldsfare folded.
3) As the water passes through the gill filaments food is gathered by latero-
frontallfrontal cilia.
4) The crystalline style is formed of mucoprotein/glycoprotein.
5 ) Finefcoarse cilia on the gill tracts deflect the large food particles.
6 ) Crystalline style releases carbohydratefprotein digesting enzymes.
7) In most gastropods, digestion is extralintra cellular.
8) Herbivorous m~crophagousgastropods do haveldo not have a crystalline style.
9) Actively swimming echinoderms are predaceous macrophagous/ciliary-mucous
feeders.
10') Crustaceans are ciliary-mucousffilter feeders.

9.3 EXCRETION IN NON-CHORDATES

Gonad Nephridiopoie Protonphridium

1 I / SoIenocyte I

Intestine
Mouth

--7 Tr=/-=p
I Nephridiostome
genital pore
!.
Metanephridium Coelomoduct

Fig.9.30: Longitudinal section of a primitive annelid showing protonephridin on the left and
-
metanephridia on the right.
 Nutrition, Excretion and
Excretion as defined under Introduction of this unit refers to removal of the waste Osmoregulation. ,
products of metabolism - carbon dioxide and water released by the oxidation of energy
rich compounds and the nitrogenous wastes released by the metabolism of proteins and
nucleic acids. However, we will here limit ourselves to elimination of nitrogenous waste.
In small ones, especially aquatic ones , simple diffusion from body surface plays an
important role in elimination of nitrogen.0~~ wastes. Even in larger aquatic animals - Jme
diffusion takes place from body surface, but they have evolved specialised organs for
excretion, which play the major role in eliminationof nitrogenous waste material.
However, it has to be remembered, that many of these organs serve primarily
osmoregulatory function rather than excretory. Well defined excretory organs are seen
from the pseudocoelomate animals onwards. Many groups of metazoans have nephridia
as excretory organs. A nephridium develops from the ectoderm centripetally. 'The lumen
of the nephridium is formed by the hollow~ngout of nephridial cells. Thus nephridia are
intracellular. In primitive animals this lumen is closed internally but subsequently
acquires an opening into the coelom. The opening is called nephridial funnel or
nephrostome. Its opening to the outside is by nephridiopore (Fig.9.30).
9.3.1
1 ,
PROTONEPHRIDIA AND METANEPHRIDIA
Nephridia occur in two major forms - the protonephridium and metanephridium. Very often there are another entirely
Protonephridia are found in flat worms. The protonephridia1 canals end bIindly in diwerent set o f tubules, the coelomic
ducts, connecting the coeloni to the
structures called flame cells or solenocytes internally. Flame cells have central cavities outside. These develop centrifugally.
that are continuous with the cavities of tubules and contain a bunch of cilia - the flame and are lnesodermal derivatives.
(Fig.9.3 1). Actually, the flame cell interdigitates with the first cell of the tubule, by They are found as outgrowths o f the
means of finger shaped processes. In solenocytes, the cell lumen is prolonged into a wall o f coeloniic cavity. The tube is
lined by a layer o f cells and is hence
delicate tube and the flame is reduced to a single flagellum. Flat worms generally have not intracellular, but is i~itercellular.
a pair of protonephridia. The protonephridial canals are much branched bearing flame It opens into the coelom by a ciliated
ceIls at the end of branches. The flame cells are found scattered through out the funnel. This funnel is called
parenchyma. The flame cells do the filtration, the filtered fluid is propelled by flagella. coelomostome. The primary
function o f the coelomoduct is t~
The nephridiaI.epithelium carries out the firnctions of reabsorption and secretion. These carry the germ cells to the outside.
physiological mechanisms are comparable with those carried out by a vertebrate kidney. However, secondarily their structure
and function may get obliterated.

Binucleate
%tune ceB

(p~fferent
duct
Fig.9.31: A single protonephridium of r flat worm. .
I
In annelids, nedhridia frequently bear ciliated funnels at the inner end and are then
j termed as metanephridia. Metanephridia are open at both ends. The nephrostome located
1 in the coelom waft the waste materials by their cilia and pass them iito nephridial tubes

I
Comparative Forms and , where they find their way to outside through the nephridiopores. However in
Functions trochophore larva of certain polyachetes possess a pair of simple typical protonephridia
each with a flame cell bearing a single flagellum. But adults possess segmental
metanephridia with open ciliated channels. In certain adult polychaetes such as Nephtys
and Phyllodoce there are only protonephridia with solenocytes. Except in the
polychaete family Capitellidae where there are separate nephridia and coelomoducts, in
other families they become associated and form a compound organ called the
nephromixium. The nephridium in a nephromixium may be a protonephridium ot. a
metanephridium. In protronephromyxium (Fig.9.32a) the developing coelomoduct
grows backwards along side the protonephridial canal and an open communication ariys
between the two at sexual maturity. In rnefanephromixium (Fig.9.32b), the nephridial
component is a metanephridium (9.32~). In Arenicola the recombined structure forms
a mixonephridiurn serving both excretory function as well as transportation of germ
cells.
Protonephridium
, ,.
Ovarv
,- . - BI Point where coelomoduct
onens into ne~hridialc-1

Solenocytes /
Bo 4y wall
Nephrid'al canal / \
Germ eel 1 Nephridiopore
Intersegmental , , PAnlnmn
septum

Coelomo

Nephrjdio
Intersegmentd
septum. Nephridiowre

- 1 Point where r >elomoduct

Coelornostorne fJI, Coelomostome , opens into nephridial canal

Rudiment of
/ %canal Nephridial
9 .
nephridium
Coelomic -
epitheium
0 *\
Nephridiopore
Fig.932: Nephridia in Polychaetes. (A) Protonephromixium (B) metanephromixium (C)
Mixonephridium.

In oligochaetes the nephridia are all metanephridial type. The coelomoducts are also
present, but are restricted to genital segments, one duct being associated with each g ~ n a d .
A similar arrangement is seen in leeches as well.
9.3.2 COXAL GLANDS
In Onychophora (Peripatus) there is a pair of coxal glands, in almost all the segments
(Fig.9.33a). Developmentally; in each segment a hollow follicle or somite arises and in
the trunk region, they become divided into dorsal and ventrolateral portions. The
ventrolateral portion graws into the appendages and its cavity forms the end sac of the
coxal gland. The end sac opens'by a ciliated canal, the coelomostome, into acoiled
 Nutrition, Excretion and
excretory canal. The terminal portion of the canal is enlarged to form a vesicle or
, Osmoregulation
bladder. Thus the coxal glands are derived from coelomoducts. In arthropods there is a
similarity in the formation of coxal glands from coelomoducts and the formation of
gonads and their ducts.

a.0; of coxal organ c.0; of excretory organ e.0; of lingual gland 1.0; of maxillary organ
m.0; of salivary otgan s.0; Dorsal coelom d.c; genital pore g.p; (both sexes indicated in Decapoda and lnsecta)
Fig 933: The exeretory and genital coelomoducts in different arthropods. (a)Peripatus (b) scorpion (c)
Crustacea (d) Symphyla (e) lnsecta.

Arachnids and crustaceans also posses similar coxal glands except that the numbers are
greatly reduced (Fig.9.33b kc).

Bladder
Excretory pore

Fig.934: Antenna1gland of a crustacean.

Comparative Forms and
Functions
- In crustaceans they are found in third and sixth segments and t ~ k yare appropriately.
-

named depending on the site of their opening to outside. The one located at the base of
the third segment opens at the base of the antenna, hence called antennal gland (Fig
9.34). The one located in the sixth segment opens at the base of the second maxilla, and
is called maxillary gland. Antenna1 gland is present in the larval forms of
Branchiopoda, Ostracoda, Copepoda, Branchiura, Ci i edia and lower Malacostraca but
TP
the adults gf these groups have maxillary glands. The adult amphipod and decapod
crustaceans have antennal glands. Mysidaceae, a primitive group of crustaceans have
both antennal and maxillary glands functional.

Most arachnids have a pair of coxal glands opening in the sixth segment, opening at the
base of third pair of walking legs. (Fig.9.35b). In Limulus, there are four pairs of coxal
glands.
9.3.3 MALPIGHIAN TUBULES
Other arthropods such as insects and myriapods and arachnids have Malpighian tubules,
the outbrowths of alimentary canal as excretory organs. Malpighian tubules are totally
new structures, having no resemblance either to nephridia or coelomoducts. Generally,
the Malpighian tubules are composed of single layered epithelium and are bathed in the
blood of haemocoel. By the process of active secretion water passes into the lumen of
Malpighian tubule together with nitrogenous wastes and dissolved salts. The secretion
essentially occurs in the distal parts of the tubules. The proximal part of the tubule as
well as the rectum are the sites of absorption. Most of the terrestrial arthropods excrete
uric acid. The reabsorption of water results in the precipitation of urate crystals as
nitrogenous waste, hence the animals are known as uricotelic. For an account of
excretory products of invertebrates refer to Unit 4 of Block 1 of LSE-05 'Animal
Physiology' course.

Peripatus which is also terrestrial lacks Malpighian tubules. Nevertheless it is uricotelic.

The excretion of uric acid is achieved from the intestinal epithelium into the intestinal
lumen. It is removed from the intestine within a peritrophic membrane which is
sloughed off every 24 hours. There are also segmentally arranged coelomic excretory
organs. The coelomoduct of Peripatus can be regarded as vestigial organs of its aquatic
ancestry, with a much reduced activity.
9.3.4 COELOMODUCTS OF MOLLUSCS
In Molluscs, as in crustacea nephridia are absent. But certain of the larval pulmonates do
possess protonephridia suggesting that they have been secondarily lost in molluscs. The
excretory system is more or less similar in all tAe groups of molluscs. In molluscs it is
assumed that the two coelomic cavities meet dorsally to enclose the heart and their walls
proliferate into germ cells. Their cavities by further differentiation give rise to gonad
anteriorly, pericardial canal centrally and gonoduct posteriorly. The last segment, in
addition had an excretory hnction (Fig.9.35a). In Aplacophora the coelomoducts of
adults consist of a pair of tubular structures, leading from the coelomic cavity to the
outside and primitively constitute the genital ducts(Fig.9. 35b). But in other molluscs
modifications have arisen in the following lines:
i) There is the development of certain degree of asymmetry.
ii) There is a separation of genital and excretory organs.

In Polyplacophora the coelomoducts split in the region of coelomostome and the gonadal
cavities become closed off from pericardial coelom. The excretory coelom remains
connected with pericardial coelom (Fig.9.35~). In gastropods there is a marked
asymmetry in the coelomic complex. The left gonad disappears and the right gonad
opens into coelomoduct that has lost its renal function and its connection with the
pericardial coelom. The excretory organ is present only on one side(1eft kidney) and
that is large and thick walled. In prosobranchs, the excretory opening is in the posterior
part of mantle cavity and in pulmonates it opens outside the mantle cavity.

In lamellibranchs, though the complications of asymmetry in gastropods do not exist,

the genital and renal ducts get separated. In the primitive group protobranchs, the entire
coelomoduct has an excretory function. The gametes are discharged into renal organs
 Nutrition. Excretion and
(Fig.9.35d). .In filibranchs the coelomoduct is bent into a "U" shaped structure, the Osmoregulation
lower limb being glandular and the more distal upper limb forms a bladder (Fig.9.36e).
The connection has shifted to the posterior end of the kidney. In eulamellibranchs, the
two organs have developed separate openings (Fig.9.350.

In cephalopods also the separation of genital and excretroy components of the coelomic
complex has been achieved. The genital duct comes to run separately from the
renopericardial canal and kidney (Fig.9.35g).

Aplacophora cephalopoda

\ Filibranchia

Gonadoduct
I. Pericardial Eulamellibranchia

Excretory pore
Fig. 9.35 : Coelom and coelornoducts in Mollusca: (a) primitive mollusc, (b) Aplacophora, (c)
I Polyplacophora, (d) protobranch, (e) filibranch, (1)eulamellibranch, (g) Cephalopod.

SAQ 5
Match the following.
r
a) Flatworms 1) Malpighian tubules
b) Annelids 2) metanephridia.
c) Crustaceans 3) coelomoducts(~idneys) +
d) Insects 4) coxal glands
e) Molluscs 5) protonephridia

9.4 OSMOREGULATION IN NON-CHO'WATE

METAZOANS.
In unit 2 you have studied that in protozoans, particularly fresh water protozoans,
contractile vacuoles play a significant role in the regulation of water content of the body.
Fresh water organisms often face the problem of their body being flooded with water
and have to constantly regulate the water content of their body. Marine animals,
although, isotonic with the medium in which they live have to maintain an ionic
composition of their body fluids that may be unique to them. Terrestrial organisms -
confront the problem of water conservation and have developed adaptations that prevent
the water loss from the body. Essentially osmotic relationships-ofthe orgairism have to
be considered in relation to&e ion& composition of body fluids. Ionic regulation could
be defined as' maintenance in bodyfluid, of concentrations of ions differing from
those that would result from a passive equilibrium from the medium. In this section we
Comparative Forms and shall briefly discuss the mechanisms of osmotic and ionic regulation of metazoan
Funrtions organisms adapted to living in different habitats.
9.4.1 OSMOREGULATION IN FRESHWATER METAZOANS
Freshwater and brackish water animals live in hypoosmotic(of lower osmotic pressure)
environment and maintain a hyperosmotic (of higher osmotic pressure) condition in their
body fluids. They may be able to tolerate only a narrow range of salinity of the medium
in which they live. These are called stenohaline animals. If they tolerate a wide range of
salinity they are known as euryhaline animals. Essentially their living environments
are osmotically less concentrated than their body fluids. They face the problem of the
water continuously entering into the body and leaching of salts from the body. You have
already studied the role of contractile vacuole as water pumps in osmoregulation of
freshwater protozoans in unit 2 of block I of this course. Such vacuoles are present in
fresh water sponges as well. Protonephridia of freshwater flatworms, metanephridia of
annelids and coxal glands of crustaceans are other such water pumps that are capable of
removing large amounts of fluids from the body. In fact such organs have the primary
fu?ction of water balance rather than excretion of nitrogenous wastes.

In some animals there are no special organs for the removal of water. Hydra is one such
example. The regulation of both water and salt in Hydra is carried out by active transport
of sodium. In the absence of calcium or sodium in the environment the osmoregulatory
process breaks down in Hydra. The pumping in of sodium into the gut is followed by
the passive flow of water along the osmotic gradient. The mesogloea functions like an
extracellular fluid space. It is believed that two pumps may be operational in Hydra, one
transporting Na into mesogloea and the second that transports it into gut. Water taken
osmotically is expelled through the mouth. Active transport of sodium takes care of
both osmotic and volume regulation. Thus there is an influx of water into the body
through the external surface, and the excess water is removed through the gastrovascular
cavity, through the mouth. Fluid in the gastrovascular cavity is hypoosmotic to tissue
fluid. The gastrovascular cavity is thus supposed to act like'big contractile vacuole

Ability to produce dilute urine has been demonstrated in animals belonging to more
advanced phyla ( arthropods, earthworms and fresh water molluscs). Using the
techniques of micropuncture and clearance of tubular fluid in the metanephric tubules,
both filtration and active transport have been demonstrated. For instance, in the antenna1
gland of fresh water crayfish, the end sac functions as the site of filtration. Chloride is
reabsorbed as the filtered urine passes through the long tubule resulting in conservation
of salts and reabsorption of water (Fig. 9.36).

\ \ 1 I
End-sac Labyrinth Nephridial Bladder
canal

Fig.9.36: Role of the different segments of coxal gland i n the formation of hypoosmotic urine in the
crayfish Astacus.
f
 Nutrition, Excrcti6p and
Filtration in arthropods and molluscs is essentially carried out by the hydrostatic pressure Osmoregulatiou
of the blood. In arthropods, the wall of the coelomic sac is highly vascularised. In
molluscs the heart passes through the filtration cavity or pericardial sac. There is filtration
through the wall of the heart, into the pericardial cavity. From the pericardial cavity,
filtrate passes through the nep%rostome into the kidney. Generally the coelomic sac is
located near the heart or near the region of high blood pressure.
r

The observed dilution of urine in the distal tubule and ureter could be due to the addition
of water or to the reabsorption of salts. But the use of metabolic poisons that arrests the
active uptake has clearly demonstrated that absorption of solutes is responsible for the
excretion of hypoosmotic urine. It could be said that the capacity of excretory organs to
form hypoosmotic urine and to trap ions from ambient fluid played a significant role in
the colonisation of the fresh water environment.
9.4.2 OSMOREGULATION IN MARINE NON-CHORDATE
METAZOANS
Studies on the osmotic pressure of body fluids of marine organisms have shown that their
internal osmotic pressure is more or less similar to the sea water in which they live.
Marine invertebrates are isotonic with the seawater in which they live, but the ionic
composition of their body fluids may be markedly different from that of the normal sea
water. For instant.: the mesogloea of the coelentrates has a high potassium and a low
sulphate concentration as compared to the seawater in which they live. This is true of
polychaets and echinoderms as well.

Marine and brackish water animals have isoosmotic or slightly hyperosmotic body fluid.
Production of an isoosmotic or a slightly hyperosmotic urine may cause the loss of
valuable electrolytes. Therefore, there is a continuous regulation of electrolytes of body
fluids. Such regulation is achieved by several ways. Surface areas permeable to water
and ions are generally reduced to a minimunl Water pumps in the form of contractile
vacuoles and nephridial tubules are present. But the most important machinery that
remains a part of every cell is the active transport. In certain organs such as gills of
crustaceans, highly specialised tissues exploit the capacity of active transport of large
amounts of salts.

In decapod crustaceans and cephalopods, ionic regulation may extend to every ion. For
instance, in these organisms calcium and potassium are more concentrated in body fluids
than in the external medium, whereas magnesium, sblphate and chloride are less
concentrated. Reduction in anion concentration such as sulphate is compensated by an
' increase in sodium concentration. Thus in marine invertebrates including coelentrates,
the internal medium has a specialised ionic composition quite distinct from that of
external medium. Excretory organs play a role in ionic regulation.
9.4.3 WATER RELATIONS IN TERRESTRIAL ENVIRONMENT
Insects are the largest group of metazoans that have most successfully invaded
the terrestrial environpent. Besides, most arachnids, myriapods and isopod crustaceans
do not depend on the aquatic environment for their survival. Terrestrial a)thropods owe
their success to life on land to the presence of an impermeable cuticle that prevents
evaporation of water from the body. Their cuticle, a chitin-protein complex with a
hydrophobic wax layer on the surface is the water-proofing structure.

The cuticle is an important structure that had made possible to a large extent the
successful colonisation of land. Water loss through the spiracular openings is
minimised by keeping the spiracles closed whenever the inspiration does not take place.
The proximal region of Malpighian tubules and more importantly, the rectum play a
significant role in water resorption expellin$only dry faecal pellets with insoluble uric
acid as nitrogenous waste. Pulmonate molluscs which have taken to terrestrial habitat
have a calcareous shell that prevents desiccation of the soft inner parts of the body.
Physiological adaptations such as aestivation help them to overcome adverse climatic
conditions.
Comparative Forms and SAQ 6
Functions
State whether the following statements are true or false.
1) Osmoconformers can have an ionic composition of their body fluids very different
from that of the medium in which they live.
2) Euryhaline animals cafi tolerate only narrow ranges of salinities.
3) Marine animals excrete a hypo-osmotic urine.
4) Water resorption is more pronounced in fresh water metazoans than in marine forms.
5) Insects excrete dry pellets as urine.

915 SUMMARY
In this unit you have learnt:
The metazoans have evoIved a variety of feeding habits depending on the
environment in which they live and availability of the type of food. They may be
microphagous feeders or macrophagous feeders. If microphagous, they may be filter
feeders or feeding on deposited food materials. Filter feeders may use ciliary-
mucus feeding mechanism or make use of setae for gathering food.
Sponges make use of the canal system for filter feeding of particulate food materials.
Digestion is intracellular. Among coelenterates, the macrophagous and carnivorous
feeders use the tentacles and the cnidoblasts present in them for capturing prey.
Digestion is both intracellular and extracellular. Flatworms may digest food
exclusively intracellularly as in Polycelis or exclusively extracellulary as in
Cycloporus.
Oligochaete annelids feed on dead organic matter. The intestine of these organisms
secretes enzymes such as cellulase and chitinase . Among polychaetes free living
forms are macrophagous and the sedentary forms are microphagous. Mac]ophagous
feeders possess eversible proboscis for food capturing. Sedentary polychaetes
resort to ciliary mucus -feeding. Sedentary forms possess a branchial crown of
tentacles lined with pinnules which interlock and form a filtering device. Cilia are
borne on the pinnules and direct the food into mouth. Tube dwelling polychaetes are
efficient ciliary-mucous feeders. Annelids generally exhibit extracellular digestion.
Among molluscs, lamellibranchs are ciliary-mucus feeders. Ciliated gills or lamella
along with labial palps are food gathering devices. Coarse cilia reject the large food
particles and fine cilia direct the movement of food towards the mouth. Digestion is
mostly intracellular in IamelIibranchs. A unique structure called crystalline style
releases carbohydrate digesting enzymes. Some herbivorous particulate feeding
gastropods also possess cpst;.lline style. Both in gastropods and cephalopods
digestion is extracellular.
Arhropods lack cilia. Appei,,tages modified for feeding aid in food gathering.
Appendages are provided with setae that are used as filtering devices to collect
suspended particulate food. The metachronal movement of thoracic appendages, the
fringe of setae present in the endites, the inter-limb spaces, the mikiventralchannel
formed by the interlimb spaces and the maxillipedes and the maxillae of cephalic
appendages are the structures involved in the filter feeding process of aquatic
crustaceans. Insects, on the other hand, have developed specialised mouth parts for
feeding as discussed in Unit 5 of Block 2 of this course.
Metazoans have evolved a variety of 'excretory' strGctures. Many of these are
however primarily osmoregulatory. Nephridia which are ectodermal derivatives are
intracellular, formed by the hollowing out of the nephridial cells. Protonephridial
canals that end blindly, carry solenocytes or flame cells secrete fluid into lumen and
waft the materials towards nephridjopore. Metanephridia which are characteristic
of annelids are closely associated with coelomoducts. Coelomoducts known as
coxal glands are.the excretory structures in many arthropods. In primitive
'
arthropods, the coxal glands are present segmentally, but in higher arthropods they
are present in one or two segments. Depending on their position they are variously
named as antennary glands or maxillary glands. Insects have Malpighian tubules as
excretory structures. The epithelial cell lined tubules secrete fluid into their lumen in
the distal region and in the proximal segment there is reabsorption. lMuch of&
water from the excretory material is resorbed both by the Malpighian tubule and
 Nutrition, Excretion nod
rectum and dry insoluble uric acid is excreted. Molluscs have coelomoducts as Osmoregulatioo
kidneys and they are closely associated with genital ducts. Marked asymmetry in
the coelomic complex and gradual separation of excretory and genital structures are
important features of molluscan coelomoducts.
Regulation of water and ionic components of the body is the adaptation required for
survival in aquatic environment. Fresh water organisms live in a hypo-osmotic
environment and have to maintain hypertonic fluid in their body. They excrete a
hypo-osmotic urine and their excretory system resorbs the vital ions required by the
. body. Marine organisms are more or less iso-osmotic with the medium in which

they live; nevertheless the ionic composition of their.body fluids is at variance with
that of the medium. They resort to active uptake of ions to maintain the ionic
composition of the body fluid. Terrestrial metazoans face the problem of water loss
from their body and the problem has been solved by eyolving a number of water
conserving measures such as an impermeable integument, excretion of dry ~A
nitrogenous waste materials and keeping the respiratory apertures closed to minimise
water loss through them.

9.6 TERMINAL QUESTIONS

1. Describe the structure of a cnidoblast and its functioning.
......................................................................................................
......................................................................................................
,<

......................................................................................................
%
.
......................................................................................................
.......................................................................................................
......................................................................................................
2. Describe the mode of digestion in the flatworms Polycelis and Cycloporus.
......................................................................................................
......................................................................................................
......................................................................................................
......................................................................................................
.........................................................................................
(.............
........................................................................................................
3. Describe the mechanism of ciliary -mucous feeding in sedentary polychaetes.
......................................................................................................
......................................................................................................
......................................................................................................
......................................................................................................
.......................................................................................................
4. Describe the structure of lamella in a lamellibranch and its function as a food
gathering device.
......................................................................................................
......................................................................................................
......................................................................................................
......................................................................................................
.......................................................................................................
5. Describe the coelomoducts of different groups of molluscs. , .

.......................................................................................................
.
.....................................................................................................
-

......................................................................................................
i
Comparative Forms and
Functions

6. With suitable examples, describe how do the fresh water organisms maintain the
water and ionic content of the body. -

9.7 ANSWERS
Self-Assessment Questions
1) i) Macrophagy, ii) cilia, iii) suspension and filter, iv) sorting and rejecting,
v) radula 1
2) i) T, ii) F, iii), T iv) F, V) F

3) 1-c, 2 - a, 3-f, 4 - b, 5-d, 6-e

ciliary-mucous,
are folded,
Laterofrontal,
Mucoprotein,
Coarse,
carbohydrate,
Extracellular,
have,
Macrophagous predators,
filter feeders

Terminal questions

1. Refer to section 9.2.1

2. Refer to section 9.2.1
3. Refer to section 9.2.2
4. Refer to section 9.2.3
5. Refer to section 9.3.3
 UNIT 10 RESPIRATORY AND CIRCULATORY
SYSTEMS
/
i
Structure
10.1 Introduction
Objectives
10.2 Respiratory System
Respiratory Organs
Process o f Respiration
Respiratory Pigments
10.3 Circulatory System
Open type and closed type of circulatory systems
10.4 Summary
10.5 Terminal Questions
i 10.6 Answers

1 10.1 INTRODUCTION
- -- - - - -

In the previous unit you have made a general and comparative assessment of different
modes of nutrition, osmoregulation and excretion in various groups of non-chordates. In
the present unit you will study what a respiratory system is and also examine the many
types of organs that help in respiration, found in different groups of non-chordataes. In
small animals passive diffusion will serve the purpose. On the other hand, when the
animal becomes larger respiratory gases have to be transported between the surrounding
medium and the area of metabolic activity, for which thecirculatory system comes in
handy. Thus you will see that from a simple type, a complex and highly developed
system of respiration and circulation have evolved. The study is divided into two parts:
the first part deals with respiration, and organs involved in the process of respiration and
the respiratory pigments. The remaining part is devoted to the study of the types of
circulatory systems found in the different phyla of non-chordates.
Objectives
After studying this unit you will be able to:
point out what respiration is,
describe the role of respiratory organs in the process of respiration in different groups
of non- chordates,
discuss the importance of respiratory pigments in respiration,
describe the structural organisation of the respiratory organs in various groups of
non- chordates,
distinguish between open and closed types circulatory system among non-chordates.

10.2 RESPIRATORY SYSTEM

Respiration is an essential physiological process in all living organisms by which they
obtain energy for carrying out all the metabolic activities of the body. The term
respiration has several meanings. In Latin, from which it has been derived, it means 'to
\ breathe' or 'to exhale' and in this sense, respiration was initially applied to the exchange
of gases between an organism and its environment. It meant the activities of breathing or
their equivalent. As years rolled by, it became apparent that the fundamental exchanges
occurred at the cellular level and consequently the term internal respiration was applied to
this phase of gaseous exchange. The exchange of gases between the body surface and the
environment then came to be called external respiration. This division is quite useful even
today. Now a days however, respiration is frequently referred to the cellular processes
involved in energy production that is the chemical processes occurring within the cell to
liberate energy. We are here concerned with external respiration and transport of gases
between tissues and the respiratory site.

From our point of view we may say that the characteristic feature of respiration is the
intake of oxygen and the output of the carbon dioxide. The oxygen taken in is used in the .
oxidation of digested food in the cell to liberate energy. Carbon dioxide is produced as a
consequence of oxidation of food materials. Its presence in the body isharmful, and
hence it is removed from the body during this process.
Comptrrtive Forms and In lower organisms such as protazoans, sponges, and platyhelminthes, oxygen is taken
Functions directly from the surrounding aquatic medium and carbon dioxide is given out directly
into the surrounding medium. In larger animals, most of the cells are deprived of direct
contact with the external environment because of their complex structure and larger size.
Hence they require the help of respiratory and circulatory systems so as to facilitate
exchange of gases and distribution of oxygen in all parts of the body. The process of
respiration in these animals comprises the following phases.
i) External respiration is generally described as breathing. It involves organs which
bring oxygen from the environment to the respiratory surface of the body. The
system also serves the purpose of removing carbon dioxide from the body to the
environment. The respiratory surface may often be within the body of the animal.
The respiratory surface may be integument, gills, tracheae or lungs.

ii) The second phase of respiration comprises the transportation of oxygen from
respiratory surface to the body tissues and carbon dioiide from the tissues to the
respiratob surface. In higher animals the transportation of respiratory gases is often
carried out through blood.

iii) During this phase oxygen is consumed by the cells and carbon dioxide is produced
by them as a result of oxidative processes which liberate energy 'for physiological
activities of the body. It is the sum of enzymatic reactions, both oxidative and non-
oxidative processes, by which energy is made available to maintain the vital
activities. This phase is internal respiration or tissue respiration.
10.2.1 RESPIRATORY ORGANS
These are the organs concerned with the gaseous exchange, i.e., intake .of oxygen and
output of carbon dioxide. They have usually greater rate of gas exchange per unitarea
than the general body surface. Thus respiratory organ may be a part or special region of ,

the body or may be an organ particularly meant for this purpose such as lung.

In non-chordates various types of respiratory organs are found. These organs may differ
in their structure but all are provided with a large surface of contact with the surrounding
environment and are richly supplied with blood vessels and capillaries that ensure rapid
gaseous exchange between the external environment and the blood. The external
environment is usually water. Even when it is air, there is a thin film of water covering
the respiratory surface and exchange ofigases takes place through this aqueous film. The
respiratory organs which work in water are gills; those which work in air, are lungs. They
may project outwards (gills) of they may be invaginations (lungs). The kind of respiratory
organs found in different phyla of non-chordates are as follows.
i) General Body Surface
In protozoans respiration takes place by diffusion. Oxygen diffises into the cytoplasm
through general body surface and carbon dioxide diffuses outwards. Sponges do not have
special o$ans of respiration; they have a canal system which brings water to the interior
of the body. Gases diffuse through the surface of the cells. Sponges prefer places where
water is rich with oxygen. If kept in foul water or water with less oxygen content the
sponges undergo reduction in size and ultimately die. Similar results are obtained if the
dermal pores are clogged, which prevent water current. The oxygen consumption is

Mesoglea
I Gastrovascular cavity

Mouth Ciaitrodemis
(A)

Fig. 10.1 :Diagrammatic representations of the body form of a sponge and a Coelenterate. (A) radial
section of an asconoid sponge and (B) radial section of a coelenterate medusa. Arrows in (A)
and (B) indicate respiratory water currents through body cavity.
 Respiratory and Circulate
Syste
Capillary blood vessels Median dorsal vessel

~ e d i a nventral vessel
Fig. 10.2: Diagrammatic section o f Nereis to show the course of the main seg~rlentalblood vessels.

..-..,
A, Ventral view A, Dorsal view

& "Felt' setae

Fig. 10.3: The sea mouse, ~ ~ h r o r i iaculeata.

ta A l . Dorsal and A2. ventral view. The dorsal and lateral
I surfaces are covered by felt setae. B. Anterior end, including the first pair o f dorsal scales.
They live offshore where they burrow i n soh bottom muds, le&ing only their posterior ends

!
I
exposed at the sulrace. The ventral surface is free o f felt and forms a flat, muscular, creeping
sole. They ventilate their burrow by elevating the ventral surface, which pumps surface water
into the burrow and moves i t anteriorly along the sole. Thezole is then depressed, forcing the
water posteriorly i n a channel formed by thedorsum and its covering scale+ Movement o f the
scaled also helps create the exhaust flow.
Comparative Forms and dependent upon the rate of water current (Fig. 10.1). Thus if the oscula are partially
Functions closed it is reduced, but this is compensated when the oscula are filly open. Also in
coelenterates no definite organs of respiration are present. However, there is a water
current through the gastro;ascular cavity of these animals. So their epidermis and
gastrodermis are necessarily in contact with water. Gaseous interchange necessary for
respiration takes place by diffusion (10.1).

Free living flatworms are usually small and they respire by simple diffision through body
surface. Respiration of the helminth parasites is quite different. They usually live in an
environment whose oxygen content is very low or nil as in the case of gut parasites. So
they have no access to air or oxygen. They are anaerobic. Similarly most nematode
parasites also resort to anaerobic respiration. Lactic acid is the end product in such
anaerobes. Generally speaking, the amount of oxygen the endoparasites use depends upon
its availability.

In annelids respiration takes place through the entire body surface which is enormously
increased by thin flattened parapodia of many polychaetes (e.g. Nereis). Within the
parapodia there is an extensive capillary network. The dorsal and ventral tody walls are
provided with numerous such networks of capillaries, which lie very close to the surface
(Fig. 10.2). While passing through them, blood receives oxygen from the surrounding
water and gives up carbon dioxide collected from the tissues. The oxygen carrying
capacity of the blood of many annelids is increased by the presence of haemoglobin or
other similar blood pigments. Usually these blood pigments are found in the plasma
(fluid) instead of being contained within the cell. Gills are common among the
polychaetes, but they vary greatly in both structure and location indicating that they have
arisen independently within the class. Gills are never enclosed within th* protective
chambers. Many species which possess gills are already protected because they live in
tubes and burrows. Gills are lacking in those polychaetes which are very small or which
possess long, thread like bodies such as many burrowing Lumbrineridae, Arabellidae and
Capite!lidae. In the scale worms gas exchange is largely restricted to the dorsal body
surface which is roofed over by the elytra. Cilia present on the dorsal surface create a
current of water flowing posteriorly beneath the elytra. In felt covered sea mouse
(Aphrodita) cilia are lacking (Fig. 10,3), but a dorsal water current is produced by the
animal that tilts the elytra upward and then rapidly brings them down in sequence. Most
commonly the gills are associated with parapodia and in many cases these gills are
modified parts of the parapodium. The notopodium sometimes possesses a flattened
branchial lobe, which acts as a gill as in case of nereids (Fig. 10.4). Generally, the dorsal

Cilia r--:.4--:-

Dorsal cirrus ,

Longitudinal muscle
Gut blood sinus

Ventral blood vessel

~ h l o r o g o ~ ecells
n Segmental ganglion
Fig. 10.4 :Cross section of Polychacte trunk.
citrus of the parapodium is modified to serve as a gill (10.5) or the gills may arise from
the base of the dorsal cirrus. Cirratulids have long, contractile, thread like gills (Fig.
10.6), each attached to the base of the notopodium.

The gills are not always assoiciated with parapodia. Many sedentary species have gills at
the anterior ends near the opening of the tubes or burrows. For examp!e, the gills of some
terebellids such as Amphitrite (Fig. 10.7) are arborescent and are situated on the dorsal
surface of the anterior segments. The bipinnate radioles composing the fans serve as sites Respiratory and C4rtulabr
SyMrm
of gas exchange in the fan worms sabellids (Sabella, Fig. 10.8) and serpulids. Ventilation
may be provided by gill cilia or by gill contractions (Fig. 10.5). But many burrowing and

Anterior
ciliated band

Fig. 10.5 :Surface ciliation in two polychaetes Arrows indicate direction o f water currents. Flows on
trunk.surface are directed posteriorly A. Scoletepis squamata B. Phyllodoce Iaminosa.

Fig. 10.6 : Cirrafulus cirraius, a polychaete with long, threadlike, dorsal cirri (gills).

Fig. 10.7 :Amplritrife at the aperture o f its U-shaped burrow with tentacles outstretched over the
substratum.
P
Comparative Forms and
Functions

Fig. 10.8 :Anterior end o f the fan worm, Sobello, showing the filter-feeding currents and tube building.

Prostomium
Peristomium
\ I

Fig. 10.9 :The lugworm, Arenicolo, i n its burrow. Arrows indicate the direction o f water flow produced
by the worm. The worm ingests the column o f sand on the left, through which water is filtered.
The pile o f sand at the burrow opening is defecated castings.
 ~ e s ~ i r i t o and
r y Circulptor
System

Fig. 10.10 :Chaefopferusduring feeding. A, Anterior part of the body (dorsal view). B, Worm in tube
(lateral view). Arrows indicate direction of water current through the tube.

Fig.*] I : Posterior end of Dero showing circlet of gills around the anus.

: Fish leech (Piscicolidae). Ozobranchu~showing la~teralgills.

Comparative Forms and tube dwelling polychaetes drive water through their burrows or tubes by undulating or
Functions
peristaltic contractions of the body e.g. Arenicola (Fig. 10.9), Chaetopterus (Fig. 10.10).
Worms which ventilate by muscular activity typically exhibit a spontaneous ventilating
rhythm in which a period of ventilation alternates with a period of rest. It is demonstrated
that ventilation activity increases the worm's oxygen requirement as much as 15 fold, but
theie is approximately a 20-fold increase in oxygen uptake. Earthworms and leedhes are
also devoid of special respiratory organs. However, some aquatic oligochaetes such as
Dero, (Fig. 10.1 1) and Aulophorus have gills at the posterior knd of the body. The fish
leech Ozobranchus (Fig. 10.12) has lateral gills.The skin which has rich blood supply
acts as respiratory surface for the exchange of gases.

In echinodermata, there are finger like evaginations of the coelomic cavity called dermal
papulae (dermal branchiae or branchial papulae) that serve as respiratory organs (Fig.
10.13). A major part of respiration takes place in echinoids, through ten branched gills

Lateral canal connecting radi

Perivisceral coelom

ORAL SIDE

Fig. 10.13: %midiagrammatic representation of major structurer tnvolved i n gas exchange i n a starfish.
(A) The general plan ofthe water vascular system. A circular canal, the water ring, sends out
a radial canal along the length o f each arm. The water ring and radial canals are situated on
the oral side of the body cavity (B)Diagram o f part of an arm, with the aboral-lateral
integument cut away on one side. The ampullae lie i n the perivisceral coelom and connect with
tube feet that project through the integument on the oral sidc. Each ampulla connects with the
radial canal of the arm through a valved lateral canrl. Two dieestive (pylor~c)caeca and two
gonadal branches r u n along each arm i n the perivisceral coelom; only one o f each o f these is
shown. The branchial papulae are thin-walled evaginations of the perivisceral coelomic wall
and appear externally as minute, fingerlike projections. Solid arrows show movements o f
ambient water, water vascular fluid and coelomic fluid. Dashed arrows indicate diffusion o f
gases between the ampullar fluid and perivisceral coelomic fluid.

situated in the area surrounding the mouth (Peristomial gills), one pair in each angle
between the ambulacral plates (Fig. 10.14). The tube feet in echinoderms are also
respiratory in function.
 Rcspiratory and Circulator
System

podium

Fig. 10.14 :The regular urchin (Arbociopunclulolo) Oral view showing the presence of gills.

ii) Lungs
In arachnid arthropods like scorpion and spider. respiration takes place by means of book
lungs (Fig. 10.15). There are four pairs of these structures in the scorpion, opening at the
four pairs of slit like stigmata on the ventral sternum. Each book lung takes the form of a
small hollow sac (pulmonary sac) filled with clusters of lamellae, 130 to 150 in number.

ium

Air flowing
in through
(b) spiracle opening
F i s 10.15: The book lung of a spider is a type of diffusion lung. (a) Position of ohe member of the pair of
book lungs within the abdomen of a spider. (b) Diagrammatieseetionthrough a book lung
showing diffuswn of gases or air flow between lamellae.
Comparative Forms These lamellae are disposed like the leaves of a book. The leaves are borne on an axis.
Functions Each leaf is hollow and the blood to be oxygenated flows within the narrow slit like
cavity, separated from the air by membranous walls. The spiracle opens into an atrial
chamber. Each lung is enclosed in a pulmonary cavity. The lamellae contains respiratory
air derived from the atrium and blood circulates in the spaces between the lamellae. It is
here that excange of gases takes place through the thin walls of the lamellae. Air enters
the lamellae by diffusion, but in some species the atrium is reported to produce a
ventilating action by means of muscles. .
iii) Tracheae
Tracheal respiration is characteristic of insects. Onychophorans, arachnids, diplopods and
chilopods also use tracheal system for respiration. In this type of respiration air is carried
directly to the tissues without the intervention of blood. Tracheal system typically

Fig. 10.16 :A, A spiracle with atrium, liltering apparatus, and valve, B, A tracheal system of an insect, C,
Diagram showing relationship of spiracle and tracheoies to tracheae, D, An air sac.
I
 consists of a large number of interconnected small tubes, the tracheae (Figs. 10.16). These Respiratory and Circulator
open outside through minute pores called spiracles, which are located on either side of the System
body. Air is pumped into and out of tracheae through these spiracles by the ventilating
movements of the body-and gaseous exchange takes place directly in the individual cell.
Single spiracle may serve for both inspiration and expiration; but usually there are
numerous spiracles, some of them for inflow and others for outflow of the air.

The pattern of the internal tracheal system is variable but there exists a pair of
longitudinal trunks with cross connections (Fig. 10.16). The tracheae are supported by
thickened spiral rings of the cuticle, the taenidia. The rings resist compression (i.e.,
prevent collapse) but permit stretching of the tube. The tracheae divide and redivide to
formminute branches called tracheoles. These ramify through the different tissues of the
body. In some insects the tracheal tubes are dilated to form air-sacs (Fig. 10.16 D). A
number of tracheoles may be formed by a single tracheole cell. In flight muscles of some
insects, the tracheoles even push into the fibrils. The tracheole cuticle is not shed during
molting as is the case of tracheae and after molting new tracheae are joined to old
tracheoles.

Exchange of gases through the tracheae is known to occur primarily by diffusion.

However, spiracles remain closed most of the time and exchange probably takes place as
a result of both diffusion and ventilation. Studies have demonstrated that the spiracles
open very briefly and not all at once in response to a localized reduction in hemocoelic
pressure. The spiracle is literally sucked open and a "gulp" of air is taken in. The pressure
drop results from intersegmental muscle contraction and is under the control of nervous
system which in turn may be regulated by the oxygenlcarbon dioxide tension of the
blood. More spiracles are therefore open during flight compared with the insect at rest.
Ventilating pressure gradients result from body movements, largely abdominal
movements which bring about compression of the air sac and the longitudinal extension
and contraction of trachea. Ventilation is facilitated by the sequence in which certain
spiracles are opened and closed.

Gases are exchanged by diffusion down a concentration gradient. Tracheoles are

permeable to liquids, and in most insects their tips are filled with fluid. This fluid seems
to be involved in the final transport of oxygen. Some of the small insects such as
collembolans and proturans which live in moist surroundings lack tracheae and gas
exchange occurs over the general body surface. Some immature aquatic insects also lack
tracheae, particularly during early stages of development. Tracheae are also usually
present in adult insects which live in water. The adults merely utilize air from air bubbles
or films held against the body surface by special "unwettable (hydrofuge) hairs. But the
nymphs and larvae of certain groups may possess special adaptations for gas exchange in
water.

Fig. 10.17 :A damselfly, Isclmeuru cervulu (Coenagrionidae). (A) Adult male and (B,C) Nymph dorsal
. --
..- .-_ and lateral views. '
Compamtive Forms and Damselfly nymphs and mayfly nymphs possess abdominal gills (Figs 10.17 and 10.18).
The gills are provided with closed tracheae and gas exchange occurs across the gill
surface between the water and tracheae. Dragonfly nymphs pump water in and out of
rectum, which contains rectal gills (Fig. 10.19) supplied with tracheae. Generally in
immature insects gas exchange occurs across the general integument between the tracheae
and water. Some larvae, such as those of mosquitoes, have a few functional spiracles
associated with one or more breathing tubes. The larva rises to the surface periodically
and obtains air through the tube.

Fig. 10.18 :MayflyH~xcrgeniaIitnbata (Ephemeridae), (A) Adult and (B)Nymph.

Gill

I" leg
A

Body wall -\
Ih~Wl1

Sternum
D

Fig. 10.19 :A-C, Paths o f water circulation through the gill chamber o f three decapods, showing
progressive restriction o f openings into the chamber. A. Shrimp, Water enters along entire
ventral and posterior margins o f the carapace. B. Crayfish. Water enters at the bases o f the
legs and at the posterior carapace margin, C. Crab. Water enters only at the base ofthe
cheliped. D. Cross section through the gill chamber o f a crab.

In Onychophorans, the organ of gas exchange is tracheae. The spiracles are minute
openings and are present in large numbers all over the surface of the body between bands
o f tubercles. Each spiracle opens into a very short atrium, at the end of which arises a tuft
of minute tracheae. Each trachea is a simple, straight tube and extends directly to the
tissue that is supplying.
 iv) C ~ I I ~ ' Respiratory and Circulator
System
Gills are the specialised respiratory organs of many aquatic animals. They are found in
mollusis and also in many crustaceans. Typically gills are.filamentous structures richly
supplied with blood. The gills found inside the body are referred to as internal gills
whereas those found outside the body are referred to as external gills. Both are meant for
respiration.
I
In crustaceans, gills are usually associated with appendages. They may however vary in
form, origin and location.

In crayfish, lobster, crab etc, the gills are filampntou., outgroivths from certain thoracic
I appendages, namely of epipodites and are enclosed iq a chamber, covered by carapace
,(Fig. 10.19). These gills are usually ventilated by the paddle like movements of special
appendages such as the scaphognathites. Many aquatic insects have gills. These are
usually abdominal or caudal and are called tracheal gills. These occur in nymphs of
Odonata, Trichoptera and in larvae of some beetles. These gillsare supplied with fine
tracheae instead of blood capillaries. The tracheal gills of the nymphs of the dragonfly
Aeschna lie in the rectum (Fig. 10.20).

~ e c t agills
l
Fig. 10.20 : Rectal gills of an insect.

Suprabranchial cavity .
Posterior retractor muscle
. Mouth (beneath palp
Posterior adductor muscle
Anterior retractor muscle

Anterior adductor muscle

Fig. 10.21: Gill structure and ventilation in eulamellibranch clams as exemplified by the freshwater
mussel Anodonta. A specimen ofAnodonla with the Ien shell removed (semidiagrammatic).
One of the four gill lamellae suspended in the mantle cavlty is shown. It has been sectioned
longitudinally near its dorsal attachment with the body to reveal the water channels within.
I Molluscs possess a variety of gills (ctenidia). The lamellibranchs possess two pairs of
'
ctenidia which show various modifications (Fig: 10.21). In chiton there are 6 to 80 gills in
each pallial groove. In gastropods the gills are relatively simple pairep, plume-shaped and .
are located in the mantle cavity (Fig. 10.22), but in cephalopods the gills are large and
richly vascularised (Fig. 10.23).
ComprrHve Forms and
Functions

,Hinge ofshells
Shell Mantle cavity 1
A

Mantle cavi
Shell

Mantle cavity
hell
Gill
Enlargement of u#I( Shell
gill leaflet
A Head
Foot Foot
(1) Longitudinal section (2) Head on vlew
view (D)
lUl

Fig.10.22: Schematic tepraentaHonaof the arrangement of the mantle, mantle cavlty and gills In several
group of molluacr (A) Tranaverrie pcction of a chlton. (B)Transverse section of a lamelllbranch
clam. The mantle cavlty is relatively capacious, and the gills a n suspended in the eavity. (C)
Diagrams of the condition i n many prosobranch gastropods. There is only one gill, the IeR and il
has become modified and fused to the mantle assuming the form of many triangular gill leailcts
that hang Into the mantle cavity something like the pages of. book. (D) Longitudinal aection of a
terrestrial pulmonate gastropod. Gills are lacking and the walls of the mantle cavity have b~eomc
richly vascularized, transforming the mantle eavity into an air-breathing lung. The mantle cavi9
opens to the outside only through a small porelike oriflce. When the mantle cavity is ventilated,
air passes both i n m d out through this pok.

Afferent branchial vessel

. ,

Fig. 10.23: Anatomy of Octopus showing gills.

Gaseous exchange is carried on by both mantle and the gills. The gills of most bivalves Respiratory and Circulator
are highly modified for filter feeding. They'are derived from the primitive ctenidia by
lengthening of the filaments on each side of filamentous axis (Fig. 10.24). As the ends of
long filaments became folded back towards the central axis; the ctenidial filaments took
the shape of a long slender 'W'. The filaments lying beside each other became joined by
ciliary junctions or tissue fusions, forming plate like lamellae withmany iertical water
tubes inside. Thus water enters the incurrent siphon, propelled by ciliary action, then
enters the water tubes through pores between the filaments in the lamellae proceeds
dorsally into a common suprabranchial chamber (Fig. 10.25) and then out of the excurrent
aperture.

Connection of

filament halves Food groove

Efferent branchial vessel adjacent filaments

Fig. 10.24: Evolution of lamellibranch gills. (a) Primitive protobranch gill (position relative to foot- .
visceral mass and mantle i~~dicated in cross section). (b) Development of food groove to
produce the lamellbiranch c ~ n d i t i o ~(c)
t . Folding of filaments at @od groove to produce the
lamellibranch condition. (d) Small section of lamellibranch gill; arrows show direction o f
water flow. (e) Tissue connections that provide support for the folded lamellibranch filaments.
 Suprabranchial chamber
Ventricle
I A

Afferent
Alae
tubes

Efferent

- Interlamellar junct~an

Efferent branchial vessel

Fig. 10.25 : A, Section through heart cgion of a freshwater clam to show relation of circulatory and
a

respiratory systems. Re?,)'.-utorywater current% water is drawn in by'cilia, enters gilt pores
and then passes up wat:.. t ~ b r to s suprabranchial chambers and out excurrent aperture.
g r ? ~ dioxide for oxygen. Blood circulation :ventricle pumps blood
Blood in gills e r c l ~ i l ~ ~carbon
forward to sinuses cif. foot &.id : iscera, and posterioily .to mantle sinuis. Blood returns from
mantle to auricles; it returns ttom viscera to'the kidney, and then got? to the giUs, and finally
to the auricles. B, Filibranch gill. a, Five adjacent filameoh ( d c c 3-I) view). b, Frontal

Frontal section.
.,
section, c-d, Eulamcllibranch gill. c, Five fused, adjacent f i l a n ~ q b(surface 3-Dview). d,
,:
56.
L -4
9
10.2.2 PROCESS OF RESPIRATION Respiratory and Circulator
System
Gaseous exchange, i.e. intake of oxygen and output of caEbon dioxide, takes place at the
respiratory surface because the surface is richly vascularised. The respiratory surface
could be the general integument, a lung or a gill. As soon as the source of oxygen
(atmospheric air or water containing dissolved oxygen) comes in contact with the
respiratory surface as a result of various ventilating mechanisms, exchange of gases, takes
place, e.g., oxygen is absorbed into the blood and carbon dioxide from the blood is
released into the environment.

The exchange of gases takes place by simple diffusion, which is caused due to the partial The partial pressure of any
pressure of the respiratory gases. Gases move from the medium with high partial pressure gas in gaseous mixture is
to the medium having low partial pressure. In the environment (air or water) the partial proportional to its concentration
in the mixture. For example,
pressure of oxygen is comparatively higher than in the blood. Therefore, oxygen diffises oxygen constituting approximately
from the environment to the blood through the respiratpry surface. Similarly the partial 2 1% of the air at a barbmetric
pressure of the carbon dioxide in the blood is higher than the immediate environment. So pressure of 760 mm Hg exerts a
partial pressure of about
it diffuses outside from blood through the respiratory surface. 160 mrn Hg (0.21 x 760).
Transport of Respiratory Gases This is called theS'oxygen tension"
o f air.
In the more primitive animals, which are usually smaller, respiratory gases (Oxygen and
Carbon dioxide) directly d i f i s e between the body and the environment. But in higher
animals which are usually larger, respiratory gases are transported with the help of a
;irculatory system in which usually blood is the circulatory fluid.

Nater normally dissolves only about 0.5 volumes per cent (i.c., 0.5 cc per 100 cc)
)xygen. Carbon dioxide is soluble in water to a greater extent, but not more than 5
lolumes per cent (i.e., 5 CC per 100 CC). Blood (plasma) in itself is normally a rather
boor carrier of respiratory gases. It is capable of dissolving in it only as much as water or
aline can. This will suffice for the purpose of small animals or even some larger but
edentary animals with very low metabolic activity. Larger ?on-chordates (and of course
lost chordates) with high metabolic activity usually have respiratory pigments. These
igments, also known as chromoproteins transport and store oxygen. In addition, these
igments (and some other substances) serve as buffers increasing the capacity of blood to
arry carbon dioxide. Carbon dioxide can also combine with some chromoproteins to
)rm carbamino-compounds, which may alSb play a small but significant role in
ansportation of carbon dioxide.

hus you have seen blood itself is only a rather poor carrier of the respiratory gases, but
:ry often in nonchordates it contains one of the respiratory pigments which acts as
.rrier of oxygen. These pigments take up oxygen when it is abundant and transporting it
rough blood readily parts with it to tissues where it is poor. Thus, respiratory proteins or
~romoproteinsplay very important role in transport of respiratory gases, whereever they
;cur. However, respiratory pigments are by no means present in all invertebrates. In
ost insects and in many other arthropod groups for example, these are absent. When
.esent, the nature of respiratory pigment varies in different animals. They may be found
ther in solution in blood plasma or in corpuscles. You will study now the various
spiratory pigments found in different invertebrates, briefly.
1.2.3 RESPIRATORY PIGMENTS
gen dissolves in blood and other body fluids in accordance with the physical laws.
amount of oxygen carried in solution per unit volume increases in proportion to the
:vailing oxygen tension. However, at physiological tensihs, the dissolved oxygen
lcentration of body fluid is never great. In animals which depend on the circulation of
jy fluids for oxygen transport, the capacity of the fluids to cany oxygen can be
reased to well above that permitted by simple solution by the presence of specialized
igen transport compounds. These compounds may be either dissolved in blood plasma
nay be in the floating corpuscles in the circulating body fluid. These transport
npounds undergo reversible, M s e chemical combination with oxygen. Haemoglobin
ich is present universally among vertebrates is an example. Chemical combination of
.gen molecules with haemoglobin accounts for over 98% of the oxygen carried by the
3d whmmkavingthe lungs; less than 2% is in simple solution.

en we say that a compound undergoes reversible combination with oxygen, we mean

it tends to take up oxygen when exposed to high oxygen tensions and to release .
Comparative Forms and oxygen when oxygen tensions are low. The compounds in the body fluids of animals that
Functions function in this way fall into ;Q groups: haemoglobins, haemocyanins, haemerythrins
and chlorocruorins. All these compounds are collectively called the respiratory pigments
or respiratory proteins; these are also sometiples called chromoproteins, because these are
often coloured.
't

The process of combining with oxygen is termed oxygenation, whereas that of releasing
oxygen is called deoxygenation. The prefixes oxy- and deoxy-, when appended to the
name of a respiratory pigment, refer respectively to pigment that is combined or not
combined with oxygen. For example, haemoglobin combined with oxygen is called
oxyhaemoglobin (red), whereas haemoglobin devoid of bound oxygen is termed
deoxyhaemoglobin (purple).
Haemoglobins
Haemoglibin is the best known of all respiratory pigments. The basic molecular unit of
haemoglobin consists of a haem group (Fig. 10.26) bound to a protein or globin moiety.
Haeme is a metalloporphyrin, to be more specific, ferrous protoporphyrin. This serves as
the prosthetic (serving as a prefix) group, with which is linked the protein. Haemoglobins
of different species differ in their chemical and physical properties. These differences are
attributable to differences in their globin moieties. The molecular weight of each unit
molecule is typically about 16000 - 17000. The four-unit haemoglobin of vertebrates thus
have molecular weights of approximately 64090 - 68000. Relatively huge haemoglobin
molecules are found in some non-chordates. For example, in numerous annelids including
earthworms (eg. Lumbricus) and in the polychaete Arenicola the blood haemoglobin has
a molecular weight of nearly 3 million. Such large molecules often contain around 100 or
more oxygen-binding sites.

Alpha chains
(a-elobins)

CH, CH=CH,
I I

CH3- C-CH, '

COOH -(CHh,. C -CHZCH,

Fig. 10.26 : \%'hen r protein consists of two of more polypeptide strands, as haemoglobin does, it is said to
have a quaternary structure. Each of the four polypeptides in hremoglobin is joined to an
iron-col~tainirrgmolecule, the hneme, but on!! two ofthe haemes can be seen in this view of
the molrrulr.

I-iaemoglotin~~ are quite cornmon,in ~nnelida.In these, haemoglobins are usually

c~Issolvedin the blood.

They are, however, sometimes fol~ndeven in tissues. Among arthropods, haemoglobins

are found in a number of species of small branchiopod crustaceans (e.g., Daphnia,
Artemia) and raiely in insects (larva of the midge, Chironomus). Here also it is usually
in the dissolved form, in the blood. Among molluscs, haemoglobin is found dissolved in Respiratory and Circulator
the blood of some planorbid snails and in the blood corpuscles of some bivalves. System

Chlorocruorins
These pigments are found in four polychaete families: Sabellidae, Serpulidae,
Ampharetidae and Chlorhaemidae. The pigment is found in plasma solution and bears
close similarities to the extra-cellular haemoglobins so commonly found dissolved in the
blood of annelids. Chlorocruorins are large molecules, with molecular weights nearly 3
million. In dilute solution chlorocruorins are greenish, but in concentrated solution, they
are red.
Haemerythrins
The haemerythins are rather rare. They occur in some animals belonging to the minor
phyla such as the sipunculid worms, some brachiopods, priapulids and in the polychaete
Magelona. Haemerythrins are located intracellularly, usually in Cells of the coelomic
fluid, or in blood cells as in Magelona. Despite their name, haemerythrins do not contain
haee groups. They however, contain ferrous iron, bound directly to the protein. They have
a molecular weights of about 75,000. These pigments are colourless when deoxygenated
but turn reddish violet when oxygenated.
Haemocyanins
This pigment is found in some arthropods and molluscs. Haemocyanins do not contain
haee groups. The metal they contain is copper, which is bound directly to the protein.
Haemocyanins are found dissolved in the blood plasma and are always of large molecular
size. Although colourless when deoxygenated, the oxygenated form is blue.

Among molluscs, haemocyanins are found in cephalopods, chitons and in many

gastropods. Among arthropods they occur in malacostracan crustaceans (crabs, lobsters,
shrimps, crayfish), horseshoe crabs, spiders and scorpions. The molluscan pigments have
a molecular weights of 4-9 million.
Functions of respiratory pigments
As you have studied in the preceding sub-sections, ttie pigments are the carrier of oxygen.
In the absence of respiratory pigments the blood can carry oxygen only in solution as in
water or saline and the amount carried will be very low. All the pigments show great
affinity towards oxygen with which they can combine at high partial pressure and
dissociate at low partial pressure. Thus these pigments also show reversibility in their
action.

(Under High Partial Pressure of oxygen)

Respiratory
organs
Haemoglobin + oxygen \ Oxyhaemoglobin

Tissues
(Under low Partial Pressure of Oxygen)

Respiratory pigments found in tissues like muscles may also serve the function of storage
of oxygen. In addition, they serve as buffers, increasing the capacity of blood to carry
carbon dioxide.
SAQ 1
(a) Tick mark (J)the correct statement and put cross against the wrong ones.
i) Respiration is a chemical activity taking place mainly in the nucleus of the cell,
liberating energy.
ii) In insects air is delivered directly to the tissues through the tracheal system.
iii) Sponges prefer places where water is rich in oxygen.
iv) Earthworms and most leeches are devoid of special respiratory organs.
v) Gills are found in arthropods like scorpion and spider.
vi) In echinoderms, tube-feet are not respiratory in function.

(b) Fill in the blanks with appropriate words from the text.
(i) Gases diffise from high ........................ to low partial pressure.
-
Comparative Forms and (ii) The quantity of oxygen carried by haemoglobin is affected by the partial
Functions pressure o f . ....................... in the blood plasma.
(iii) Respiratory pigments found dissolved in the blood plasma have usually
........................ molecular weight from those found within the cells.
(iv) Haemoglobin contains ......................... as the prosthetic group.
(v). Chlorocruorin is a pigment of ........................colour found in
.............................. annelids.

- -

In sponges, coelenterates and platyhelminths nutrients, respiratory gases and waste

materials can easily diffuse through the intercellular spaces. i ow ever, in higheranimals
the circulatory system provides a medium through which oxygen and hutrients from the
environment reach all the tissues of the body and carbon dioxide as well as nitrogenous
waste materialsare eliminated from the body into the external environment. The
circulatory system consists of usually blood, the blood spaces and a series of tubes called
vessels.

Circulatory systems can be divided into a number of components with similar functions.
In addition to the ciruculating fluid, these consist of:
i) A main propulsive organ, usually a heart which forces blood around in the body,
ii) An arterial system which serves to distribute' blood and as a pressure reservoir,
iii) Capillaries, or in some animals blood spaces, in which transfer of materials take
place between blood and tissues, and
iv) The venous system that acts as a blood reservoi; and as a system for returning blood
to the heart.

One or more of the above component may be absent in various'animals.

10.3.1 OPEN AND CLOSED TYPE OF CIRCULATORY SYSTEMS
There are two types of circulatory system found in higher metazoans. In one type the
original blastocoel continues to be the main perivisceral space. Either there is no true
coelom at all as'in pseudocoelomates, or the coelom may be highly obliterated or reduced
and restricted to gonads and excretory organs as in arthropods and most molluscs. In
these animals there is no closed network of capillaries connecting the arteries and veins.
The arterial blood flows into large spaces or sinuses (or into smaller spaces called
lacunae), in which various tissues or organs are bathed by blood. From these spaces blood
passes into large open veins or through ostia into the heart directly. This is called open
type of circulatory system.

Dorsal vessel
\ Circumoesophageal vessels

Ventral nerve cord

Ventral blood vessel
'(b)
Fig. 10.27 :The circulatbry system of annelid worms. (a) k lateral view of the anterior part of the system
in an earthworm. (b) Vascular system within a segroenf of Nereis. Arrows indicate the .
direction of blood flow.
88
In the other type, a system of closed capillaries connect the smaller arteries with veins, Respiratory rod Clrculrtor
fluid containing nutrients filter through the capillary walls into tissue spaces, from which System
cells in turn derive their nutrients. This is called closed type of circulatory system. This
type of cuculatory system is found in annelids, cephalopod molluscs and chordates (Fig.
10.27). ,

a) Open Type
Many nonchordates have thus an open type of circulation - a system in which blood
pumped by the heart empties via an artery into an open sinus filled with blood which lies
between the ectodern and endodenn. The blood filled space is referred to as haemocoel,
especialy in arthropods. The fluid contained within the haemocoel, referred to as
haemolymph or blood, is not circulated through capillaries but bathes the tissue directly.
Figures 10.28 show the organisation of the main vessels from two groups of non-
chordates having open circulation, a cray fish and a bivalve mollusc.

Arterial valve Ostium with valves

branchial vessel

Ster;lal sinus branchial vessel

A

Ne~hridialvessels Ventricle

Ctenidial vessel

Fig. 10.28: Invertebrate circulations (A) simplified diagram of the circulation in the crayfish, (B)
Simplified diagram of the circulation in bivalve mollusc.
In many animals the haemocoel is large and constitutes 20 - 40% ofbody volume. In
contrast, chordates with a closed circulation have a blood volume of 5 - 10% of the total
body volume. Open circulatory systems have low pressures, seldom exceeding 5 - 10 mm
Hg..
Animals with an open circulation generally have only limited ability to alter the velocity
and distribution of blood flow. As a result, in bivalve molluscs [Fig. 10.28 B) and other
animals which have an open circulation and use blood for gas transport, change in oxygen
uptake is slow and rate of oxygen transfer is low per unit weight. In other words, their
metabolisy is very low, Inseots have, however, avoided this problem by evolving a
tracheal system in which gas transport to tissues occurs direct throu~hair filled tubes or
tracheae (Fig. 10.29) that bypass the blood. Consequently, although insects have an open
circulation, they have greater metabolic rate.
b) Closed Type
Some non-chordates, such as cephalopods (octopuses, squids), earthworms, polychactes
and all vertebrates have a closed circulation, with blood flowing in a closed circuit of
tubes from arteries to veins through capillaries. There is better separation of functions in
closed circulatory system than in open system. In a closed circulation as shown in figures
 --
'
Comparative Forms and 10.30'the heart is the main propulsive organ, pumping blood into arterial system and
Functions
maintaining a high blood pressure in arteries. The arterial system, in turn, acts as a

fl
A - HEMOCOEL

Thoracic muscles

Alary
muscl
~erivisceral - = &d' .''A
blood flow
cavity
,

Blood space
I Chamber of heart

Fig. 10.29: A, Ciculatory system a) lateral view of hypothetical insect indicating directional blood flow; b)
dorsal view of heart. B, Schematic transverse section to show open blood vascular system of
an insect. C. Blood is kept moving in the blood space by the heart. The heart sucks it out
through the ostia in the sides and pumps it through the aortal opening front.

pressure reservoir forcing blood through the capillaries. The capillary walls are thin, thus
allowing high rates of transfer of material between blood and tissues.

As shown in Fig. 10.30 A blood returning from the systemic circulation is directed to the
gills. Much of the returning blood flows through the two lateral vena cava via the kidneys
.before arriving at the bases of the gills. Near the base of each gill is a bulbous accessory
pulsatile organ, the branchial heart, which receives the systemic venous blood. This heart
pumps the blood into an afferent branchial vessel, from which it passes through
capillaries in the gill to the efferent branchial vessel and return to the systemic heart.

Physiologically the circulatory systems of cephalopods (squid and Octopus) resemble

those of chordates far more than to those of the other molluscan groups. As for example,
both cephalopods and chordates have closed circulatory systems; the blood volume of
octopus is relatively small, about 6% of the body weight, well within the range of that of
, vertebrates but radically lower than in non-cephalopod molluscs.
 Respiratory and Circulator
System

C C

Systemic
Circulation

Afferent branchial
vessel
Gill
Renopericardial canal
Abdominal vein
Branchial heart appendage Branchial heart

Lateral vena cava

Efferent branchial vessel

Ventricle 'cephalic aorta

Fig. 10.30: A, Schematic representation of the circulatory plan in squids and octopuses. Stippled parts
carry relatively deoxygenated blood. B. Circulatory systems ofOctopus.

SAQ 2
Match the animals given in column A with the type of circulatory system found in them
in column B.

A B
i) Helix Open type
ii) Octopus
iii) Squids
iv) Snails Closed type
v) Insects
vi) . - Crab
vii) Earthworm

10.4 SUMMARY
In the end you will try to recapitulate what you have studied.in this unit.
Respiration is an essential physiological process of living organisms by which they
obtain energy for carrying out all the metabolic activities of the body. The term was
initially applied to the exchange of gases (taking up oxygen and giving away carbon
dioxide) between an organism and its environment. Sometimes, this process is now
called external respiration as opposed to the chemical processes occurring within the
cell to. liberate energy, which is called internal respiration.
In lower non-chordates such as protozoans, sponges and platyhelminths, oxygen is
taken up directly from the air or from the aquatic medium surrounding them and
carbon dioxide is given out directly to the surrounding medium.
Comparative Forms and In arachnids like scorpions and spiders book lungs serve as respiratory organs
Functions
whereas in insects, onychophorans, diplopods, and chilopods, tracheae are the
respiratory organs.
Gills are the respiratory organs of a number of aquatic animals like many
polychaetes, molluscs and many crustaceans. These gills are typically filamentous
structures richly supplied with blood. The gills found inside the body are referred to
as internal gills whereas the ones found outside the body are referred to as external
gills.
To facilitate gaseous exchange i.e., intake of oxygen and output of carbon dioxide,
the respiratory surface is richly supplied with blood vessels.
Blood itself is a rather poor carrier of respiratory gases. But often it contains
respiratory pigments such as haemoglobin, haemerythrin, haemocyanin and
chlorocruorin. These respiratory pigments or chromoproteins act as the carrier of
oxygen. They also serve as buffers increasing the capacity of blood to carry carbon
dioxide.
The circulatory system in higher animals ensures the exchange of substances
between various tissues of the body on the one hand and the external environment
on the other. It also transports various substances from one part of the body to the
other.
Many non-chordates like some crabs, snails, insects etc. are said to have an open
type of circulation a system in which blood pumped by the heart empties via an
artery into open sinus filled with blood. The various tissues and organs are bathed
by blood in this sinus. There are no closed net work of capillaries connecting the
arteries and veins in these animals.
Some other non-chordates such as cephalopods (octopus and squid), annelids and all
vertebrates have a closed circulation, with blood flowing in a continuous circuit of
tubes from smaller arteries to veins through closed capillaries.

10.5 T E ~ I N A L
QUESTIONS
1. List the various organs found in non-chordates for respiration. Discuss the mode of
respiration in scorpion.

2. What is the purpose of respiration in animals? Explain briefly the mechanism of

respiration in molluscs.
3. Discuss briefly the mechanism involved in the transport of respiratory gases in Respiratory and Circulator
multi-cellular animals. System

.......................................................................................................

4.. What do you mean by open and closed types of blood circulation?
......................................................................................................

10.6 ANSWERS
Self-Assessment Questions
1. a) i) false ii) true
iii) true iv) true
v) false vi) false

b) i) partial pressure ii) oxygen

iii) Higher iv) haeme
v) green, polychaete

2. Open type (i), (iv), (v) and (vi)

Closed type (ii), (iii) and (vii)
Terminal Questions
1. In non-chordates respiration takes place through various organ such as (i) general
body surface, (ii) lungs, (iii) tracheae and, (iv) gills. In scorpions respiration takes
place in book lungs. Each book-lung is a small hollow sac (pulmonary sac) filled
with clusters of lamellae. The lamellae contain respiratory air derived from the
atrium and blood circulates in the spaces between the lamellae. It is here that
exchange of gases takes place through the thin walls of the lamellae. Air enters the
lamellae through diffusion.
2. Respiration is an essential physiological activity of all living organisms for obtaining
energy to carry out various metabolic activities of the body. In molluscs respiration
takes place through gills.
3. Multicellular animals with high metabolic activity usually have respiratory
pigments. These pigments are also called chromoproteins which transport and store
oxygen. Besides, these pigments serve as buffers increasing the capacity of blood to
carry carbon dioxide. Oxygen dissolves in blood and other body fluids in accordance
with physical laws. The respiratory compounds in the body fluids of animals are of
four types (i) haemoglobin, (ii) haemerythrin, (iii) haemocyanin and (iv)
Comparative Forms and chlorocmorins. The process of combining with oxygen is termed oxygenation
Functions whereas that of releasing oxygen is called deoxygenation.
4. There are two types of circulatory system found in higher metazoans. In open type
the original blastocoel continues to be the main perivisceral space. There is neither
true coelom at all as in pseudocoelomates or the coelom may be highly obliterated
or reduced. In these animals there are no closed network of capillaries connecting
the arteries and vein. Whereas in the other type, a system of closed capillaries
connect the smaller arteries with veins, fluid containing nutrients filter through the
capillary walls jnto tissue spaces from which cells in turn derive their nutrients. This
is called closed type of circulatory system.
 UNIT 11 NERVOUS SYSTEM AND SENSE
ORGANS
Structure
11.1 Introduction
Objectives
' 11.2 Organization of Nervous System
Nerve Cell: the Basic Unit
Neuroglia
Ganglia .
Nerves
I 13!. Primitive Nervous System: Nerve Net
11.4 Advanced Nervous Systems
Platyhelminthes
Annelids and Arthropods
Molluscs
1 1.5 Giant Nerve Fibre
11.6 lnformation Processing
1 1.7 Receptors
Properties of Receptors
Mechanoreceptors
Chemorecepton
Photoreceptors
11.8 Summary
11.9 Terminal Questions
11.10 Answers

11.1 INTRODUCTION
You have by now studied that non-chordates also carry out a variety of activities such as
feeding, digestion, locomotion etc. For this purpose, they have corresponding organs and
organ systems working in a coordinated way. Moreover, the environment of the animal,
both internal and external, is never stable and the animal has to change its activities in
relation to the changing environmental conditions. This includes escape from adverse
climatic conditions or from a pursuing predator, catch food to overcome hunger, digest
the food for energy requirements, excrete waste material, regulate the respiratory rate and
so on. Towards this purpose, the concerned organs have to be coordinated in an efficient
and purposeful manner. This is brought about mainly by the nervous system.

Necessarily, the animal has to perceive any change in the environment, compute the
changes and ultimately translate these computations into requisite actions in a manner
most profitable and adaptive to the animal. Towards this goal the nervous system has
receptor components consisting of sense organs, coordinating centres in the central
nervous system and the motor components controlling the motor elements. The system is
connected throughout so to say, by cables of nerves through which messages or impulses
v flow as if on a telegraphic system. The receptororgans receive messages from the
environment, which are transmitted to the central nervous system. The central nervous
system is the decision centre, which sends requisite orders to the effector organs through
motor neurons. Before you proceed to study this unit, you will have to go through the unit
on physiology of the nervous system, to have a clear understanding of the working of the
nervous system. In the present unit you will study essentially how the basic component of
the nervous system, the neuron, has become specialised into sense organs, and how some
of the neurons have got concentrated into central nervous system for effective control and
1 coordination, during the process of evolution of nonkhordates.
Objectives
After reading this unit you should be able to:
name the basic unit of the nervous system,
differentiate the three structural types of neurons,
describe the chief variants of mechanoreceptors of non-chordates and point out
how they function,
distinguish between olfactory receptors and contact chemoreceptors of insects,
illustrate the parallelism in the structure of the cephalopod eye and vertebrate eye,
elucidate what is nerve net and why it is supposed to be the primitive condition,
Comparative Forms and describe how the central nervous system has originated and trace its evolution
Functions
among non-chordate metazoans,
explain what giant fibre is, and
briefly point out how information processing takes place in the central nervous
system.

11.2 ORGANISATION OF NERVOUS SYSTEM

Nervous systems are composed of nerve cells or neurons and glial cells. In the latter half
of the 19" century it was strongly believed that nervous systems are composed of
complex, continuous meshworks of cells and processes in protoplasmic continuity with
each other. This theory was known as the reticular theory and it was strongly supported
by Golgi's School. However, during early 2oLhcentury this theory was gradually
superseded by the neuron doctrine. The champion of the neuron doctrine was Cajal, who
demonstrated convincingly that neurons do not physically join or continue with one
another but are closely apposed at synapses. The debate on continuity versus contiguity
persisted until 1950s, when electron microscopy permitted resolution of cell membranes
at neuronal endings, thereby demonstrating discontinuity of neurons. The neuron doctrine
is the cornerstone of our understanding of nervous system. According to this doctrine,
neurons are discrete cells forming the basic units of the nervous systems.
11.2.1 NERVE CELL : THE BASIC UNIT
The basic unit of the nervous system is the nerve cell or neuron. A neuron is to the
nervous system what a brick is to the building. It has a cell body known as perikaryon and
one or more processes known as neurites. You will see that, functionally, there are three
types of neurons. Certain neurons are specialised for perceiving changes in the external or
internal environment and transforming or transducing these stimuli into nerve impulses.
We call them sensory neurons. Some neurones are specialised for receiving nerve
impulses from other neurones and transmitting these impulses to effector organs like
muscles or glands. These neurones are called motor neurones. Usually, intervening
between sensory neurons and motor neurons are one or more association neurons.
Association neurones receive impulses from sensory neurones and other association
neurons. They also transmit impulses to other association neurons and motor neurons.
Thus one or more association neurons may intervene between sensory neuron and motor
neuron. However, sensory neuron may also transmit impulses direct to motor neuron.

Y i
= 9
Dendrites

A UNIPOLAR
Terminal
arborization

B BIPOLAR

Fig. 11.1: Neurons found in the insect nervous system. Arrows indicate direction o f impulse conduction.
You have studied above that neuron has usually one or more neurites. Of these, dendrites Nervous System and Sense
Organ
usually conduct impulses towards perikaryon. From the perikaryon, impulses are
generally transmitted away, through the axon. So you will see that it is the direction of
impulses travelling along a neurite which determines whether it is a dendrite or an axon.

structurally, it is customary to categorise neurons into three: unipolar, bipolar and

multipolar (Fig. 1 1 .I). Unipolar neuron has only one neurite arising from the perikaryon
and that is the axon. Dendrites join it as collaterals. By far most of the neurons of
invertebrates belong to this category. They may be motor or association neurons. Bipolar
neurons have two neurites, one being the dendrite and the other, the axon. Sensory
neurons are bipolar.

Multipolar neurons have more than two processes connected with the perikaryon, one of
which is the axon and the others are dendrites. These are association neurons. Most
metazoan axons are surrounded by a thin sheath cell (Schwann cell) membrance. They,
however, lack the so-called thick whorls of myelin sheath characteristic of many
vertebrate axons (Fig. 11.2 A and B).

Epithelial cell 1

Sensory
Sensory neuron
neuron

/I Muscle cells

One-neuron system Two-neuron system

body

Bipolar neuron

Invertebrate
sensory neuron

Fig. 11.2: A. Drawings of neuronal organizations found in midarians and insects.

11.2.2 NEUROGLIA
We may consider neuroglia as the connective tissue of the nervous system. It includes all
elements of the nervous system other than neurons. You will see that neuroglia fills the
space among the neurons and extends processes into the central region of the ganglia
(neuropile). It also includes the covering of the ganglia (perineurium) with the-outer, non-
.Comparative Forms and Crayfish neuron Vertebrate neuron
Functions

, Fig. 11.2 B. Non-chordate neuron and a vertebrate, neuron.

Sensory
neuron

Central nervous
system

Fig.lI.3: A drawing of the neuronal organization in a primitive worm with a central nervous system.
celTular, fibrous neurilemma or neural lamella; These coverings, though thin, extend Nervous System and.Sense
along the nerves too. .The Schwann cells too form part of the neuroglia. Organ

11.2.3 GANGLIA
Among higher non-chordates with a central nervous system, you will see that the
association neurons and motor neurons are concentrated in masses called ganglia, and the
bipolar, sensory neurons are distributed peripherally in association with sense organs
(Fig. 11.3). Invertebrate ganglia have a cellular rind or cortex and a central mass called
neuropile (Fig. 1 1.4). The neuropile is made up of nerve fibre mass, criss crossingjn all
directions Axons of sensory neurons terminate in the neuropile and those of motor
neurons arise from it.

Nuclei of perineuri

Fig.ll.4: Cross sections through (A) abdominal ganglion and (B) interganglionic connective to show
general structure.

11.2.4 NERVES
Because of the pattern of distribution of neurons as described above, there are bundles of
nerve fibres, called nerves, connecting the central ganglia on the one hand, and sensory
neurons or the effector organs on the other (Fig. 11.3). The nerves may contain sensory,
motor or both types of fibres, and depending upon this the nerves may be sensory, motor
or mixed respectively.

Similarly, there are connectives running between ganglia of the same side, and
commissures between ganglia of the opposite side. They mostly contain association
fibres.

SAQ 1
Draw a diagram involving sensory, motor and association neurons to show possible
pathways of transmission of an environmental stimulus resulting in motor action.

SAQ 2
Given below are structural and distributional characteristics of sensory, motor and
associationneurons. Choese most appropriate structural and distributional characters.
Type of neurons I. Structural 11. Distributional
Characters Characters
i) Sensory neurons a) unipolar A. Peripheral
Comparative Forms and
Functions ii) Motor neurons b) bipolar B. Central (within ganglia)

iii) Association neurons c) multipolar

11.3 PRIMITIVE NERVOUS SYSTEM: THE NERVE

NET
A nervous system makes its appearance for the first time in the phylogeny among
Cnidaria. In this group the nerve cells form an irregular nerve net orplexus (Fig. 1 1.5). In
hydras we have a simple condition. The nerve net is made up of bipolar or tripolar and
rarely multipolar neurons. The sensory nerve cells are distributed among the epithelial
cells; they send their nerve fibres to the subepithelial nerve plexus. The plexus is situated
beneath the epidermis. In hydroid polyps the plexus is concentrated around the mouth,
suggesting the beginning of a centralised nervous system. Many cnidarians have more
complicated nerve nets; for example they may have also a gastrodermal nerve net within
the gastrodermis in addition to the epidermal nerve net as in sea anemones. A double
nerve net in the same body layer is also common in some coelenterates. There is quite a
lot of autonomy, in that, isolated parts of the animal behave as if parts of the body. There
is no central controlling mechanism apparently because there is no distinct central
nervous system. It is also poor in specific responses to particular stimuli. Transmission is
unpolarised, in that impulse can be transmitted in either directions along the synapse
unlike in higher animals. This hap?ens because both terminals secrete transmitter
substance. Thus the nerve plexus in this group shows diffuse, unpolarised transmission,
autonomy of parts and scarcity of reflexes.

Neuron Epidermis Sensorv neuron

Sensory neuron' '~astroderrnis' ~eso~lea

Fig.11.5: A. The nerve net of Hydra and other cnidarians is the simplest organized nervous system. N o
central control organ and no definite neural pathways are present; B. Surface view of a
cnidarian nerve net (details of synapses not shown); C. Body wall of Hydra showing nerve nets
in both epidermis and gastrodermis. Sensory neurones are restricted to the gastrodermis in the
column (left), and to the epidermis in the tentacles (right).
Before studying the nervous system of echinoderms, you have to bear in mind the
peculiar organization of these animals. Here also, the nervous system is primitive,
retaining the nature of the network pattern beneath the surface epithelium. The epithelium
itself is strewn with sensory cells. The network is however, concentrated into
ganglionated nerve cords. These retain the general radiate pattern of the group. You will
see three systems here:
 (i) The oral or the ectoneural system beneath the oral epidermis, made up of a ring Nervous System and Sense
around the oesophagus and a ganglionated strand extending from this ring along Organ
- each arm (Fig. 11.6). Each cord is made up of tracts of fibres. This is the main
system in most echinoderms.

Fig. 11.6: Sea star A. The nervous system consists of a central ring with major nerves radiating into each
arm. B. Cross-section of the starfish arm. Nerve fibres from the ectoneural system terminate on
tJe surface of the hyponeural system but there is little or no direct contact between the two
systems.

(ii) The deeper hyponeural system and

(iii) The aboral or apical system are weaker but organized on the same pattern as the ,
ectoneural system. Thus it is simple in structure and with close relationship with
the epidermis, lacking ganglia or brain. However, it has undergone certain amount
of concentration, with circular and longitudinal cords. This marks the beginning of
a central nervous system.

SAQ 3
Given below are some characters of echinoderm nervous system. Point out advances (A)
or similarities (S) of the system compared to that of coelenterates, by denoting respective
alphabets against each character.

The system a) has close relation to epidermis; b) lack ganglia or brain; c) has undergone
some amount of concentration with circular and longitudinal cords.

11.4 ADVANCED NERVOUS SYSTEMS:GENERAL

TREND IN EVOLUTION
In higher groups of animals you will notice clear trends towards centralisation of the
nervous system, to maximise efficiency. The diffuse peripheral network becomes less
conspicuous or unrecognizable giving place to nerves made up of nerve fibres. The motor
neurons and association neurons become mostly concentrated and centralised into masses
known as ganglia. The increasing dominance of the assoc:ation neurons which are the
links in reflex pathways, makes possible more complex central pathways which form the
structural basis of advances in integration. The sensory neurons become relegated to the
periphery. This results in differentiation of functional tracts of neurites and nerves
through which stimuli are brought to and carried out of the central nervous system. Thus
better association or integration of afferent sensory stimuli with meaningful efferent
motor outputs become possible, resulting in more intricate and efficient behavioural
, patterns. Perikarya become lin~itedto the rind or cortex of the ganglia, the interior being
made up of nerve fibre mass called neuropile in which conspicuous pathways or tracts
already evident in the lower groups, become developed. The development of giant fibre
system associated with the central nervous system further enhances the efficiency
resulting in enhanced speed of propagation of impulse and hence efficiency of response to
stimuli. With one end of the animal always directed forward during locomotion, an
anterior end and bilateral symmetry become established. The anterior end becomes more
important with most sensory organs becoming associated with this end. Naturally, the
ganglia become better developed and more highly organised in this region, giving rise to
the structurally and physiologically more complicated brain dominating over the whole
. organism. This is the direct result of cephalisation.
 Comparative Forms and Now, we will discuss about the tre..d of development in the nervous system in different
' Functions phyla of the invertebrates.
11.4.1 PLATYHELMINTHES
Among Platyhelminthes, you will see that turbellarians have a brain and three to five
pairs of longitudinal nerve cords extending from the brain to the posterior end (Fig. 11.8).
They are positioned along dorsal, lateral and ventral aspects. The cords are connected
together by means of commissures. The commissures are in turn connected to a
submuscular nerve plexus. The system is thus not only sunk deeper into the submuscular
position, but it does not correspond to the real primitive plexus at all. The motor and
association neurons are mostly confined to the brain and the cords all along their
periphery surrounding the central neuropile. One pair of these longitudinal cords become
conspicuous, the others tending to become weaker ultimately disappearing thus giving
rise to the typical ladder like nervous system.
Primitive brain

Fig. 11.7: Planarian flatworms have a ladder-type nervous system. Cerebral ganglia in the head region
serve as a simple brain and, to some extent, control the rest oithe nervous system.

With the development of the central nervous system, movements become more co-
ordinated. Parts of the animal can maintain spontaneous movements and show co-
ordination of responses in most species of this group, only if it contains portion of the
nerve cord. The brain appears to have started exerting a dominating influence in this
group.
11.4.2 ANNELIDS AND ARTHROPODS
From the ladder-like nervous system of Platyhelminthes is derived the meta,nerically
arranged nervous system of annelids and arthropods (Fig. 11.8). Neurons become
concentrated metamerically into ganglia, instead #ofbeing distributed throughout the
periphery of the longitudinal cords. Anteriorly the cords continue as circumoesophageal
connectives which end dorsally into a brain made up of cerebral ganglia. The two
longitudinal ladder-like cords present in certain annelids and in some primitive
crustaceans show a tendency to cor..e closer together. Ultimately the metarneric ganglia
of the two cords fuse into a single ganglionic mass in each metawere. The two cords may
even completely fuse together into a single cord having lost all indications of their double
nature externally.

From each ganglion, varying number of paired nerves are given off. There may be further
concentration by fusion of the ganglia in different groups of animals.

In Polychaetes, the brain is iq the prostomium and may extend back through the anterior
few body segments. The ventrkknerve cord ertends through the length of the body. The
brain is betterdeveloped, with sensory centres, because of the presence of the well-
developed sense organs like palps, antennae, eyes and'nuchal organs. However, in leeches
and eartKworms lacking well developed sense organs on the head, the brain is less
developed. In the leeches, there is the nerve ring and there are also two ventral nerve
cords but each pair of ventral ganglia is fused. In oligOckaetes also the brain is shifted a
few segments posteriorly; the ventral nerve-cords are however fused into a single nerve Nervous System and Sense
cord. Organ

nerves

Cerebral ganglion

Circumoesophageal
connectives

Thoracic ganglion

B
Fig.ll.8: A. The segmented ventral nerve cord in leeches is made up of well-differentiated ganglia.
containing the cell bodies, and connectives composed of nerve fibres. The sub-oesophageal
ganglion is derived from a numbersf segmentrl ganglia. B. Highly condensed central nervous
system in a decapod crustacean. C. Nervous system of gasshopper.

In arthropods there is a higher degree of cephalisation. The brain becomes

comparatively larger with better developed sense organs on the head, like eyes and
antennae. The brain has mostly three regions: the anterior protocerebrum, the middle
deutocerebrum and the posterior tritocerebrum, with well developed sensory, motor and
association centres.
Motor Control
In annelids and arthropods in general, individual metameric ganglia of the ventral nerve
cord are capable of initiating and maintaining locomotor movement in the concerned
segment. The suboesophageal ganglion exerts a control over these; in fact it keeps the
animal in hyperexcited condition. The brain on the other hand exerts a moderating
influence on the suboesophageal ganglion and suppresses the concerned centres in the
suboesophageal ganglion. Thus many of these animals show hyper excitability and an
increased tendency for locomotion when their brain is removed. On the other hand, when
their suboesophageal ganglion is removed, locomotion is considerably inhibited or even
stopped.

SAQ 4
What are the main trends in the evolution of the nervous system? Give four points.
r7omparative Forms and 11.4.3 MOLLUSCS
Functions
You will see that the central nervous system of different molluscs has reached different
degrees of development depending upon the adaptive radiation of this group. The
molluscan nervous system shows conspicuous difference from that of annelids and
arthropdds in that it is not metamerically segmented. The primitive system is itlustrated
by that of Chiton (Fig. 1 1.98). In this group, as in Platyhelminthes. though there is
centralisation,there is very little concentration of nerve cells into ganglia. Even cerebral
ganglia are lacking. But there is a nerve ring with longitudinal nerve cords consisting of
pedal cords along foot and pallial cords innervating the mantle and the visceral mass, with 1
transverse commissures forming ladderlike nervous system. The pair of buccal ganglia
concerned with movement of radula are the only ganglia in these animals. Otherwise the
nerve cells are scattered along the ring and cords.

Fig.li.9: A. Nervous system of Chiton, B. The mollusc, Patella, showing the principal ganglia and torsion
I
I of the visceral connectives.

-
Buccal ganglion

ererag a

Pedal nerve cord

Visceral nerve cord

n g i o Pleural
nPedal ganglioni:
From this condition, the central nervous system has not increased very much in
organization either in gastropods or bivalves, which are mostly rather inactive animals
resorting to retreat in their shells. In gastropods (Fig. 1 1.1 1 B) a pair of pleural ganglia

Buccal ganglion
ganglion

Parietal ganglion
erebral ganglion

Visceral ganglion
lnfraparietal ganglion
Supraparietal ganglion

Visceral ganglion

,
'
Pedal ganglion 'pleural aanglion

Environment

Fig.ll.10: A, Hypothetical pretorsiqn nervous system. B, Posttorsion nervous system. C, Lateral view of
gastropod showing positic. af nervous system.
appear at the anterior end of the pallial cords. The pallial cords form a visceral loop with Nervous System and Sense
a pair of parietal ganglia and visceral ganglia. Further concentration of nerve cells led to a Orgrn
pair of cerebral ganglia in the nerve ring and a pair of pedal ganglia, with pedal cords
disappearing. The torsion of gastropods results in twisting of the visceral loop, the
original left and right parietal ganglia becoming the subintestinal and supraintestinal
ganglia (Fig. 1 1.10). Higher gastropods show an increasing trend in further concentration
of the ganglia. Thus the subintestinal ganglion and the supraintestinal ganglian are drawn
into the main nerve ring, resulting in shortening of the visceral loop. In
Opisthobranchia, reversal of torsion (detorsion) leads to untwisting of the nerve loop.

The predatory and active swimming modes of life and locomotor dexterity of
cephalopods have led to a high degree of cephalization among this group. The sharp sense
of Sight and touch has resulted in development of corresponding centres in the brain. The
organization of the brain in cephalopod has attained perhaps an unparalleled level among
invertebrates (Fig. 1 1.1 I). All of the ganglia characteristicof the molluscan central
nervous system are concentrated and more or less fused into a brain encircling the
oesophagus. As a result in Oclopus for example about thirty lobes of the brain have been
recognised, each with a layer of nerve cells and a central mass of neuropile, concerned
with definite functions.

1st order giant fibre /Yd order giant fibre 3'* order giant fibre

Cell bodies of

Palli

stellar nerves

Fig. 11.11: (a) The giant fibre system of the squid Loligopeuleii and (b) the synapse in the stellate
ganglion.

11.5 GIANT NERVE FIBRES

We now know that giant nerve fibres (Fig. 1 1.1 1) occur in the central nervous system of
many polychaetes, oligochaetes, insects cephalopod etc. They are broader and longer than
the other, ordinary nerve fibres which are about 2 pm or so only in diameter. In the
squid, they may attain a diameter of 700 pm or even more. Their size makes them ideal
material for electrophysiological studies, as recording electrodes can be inserted inside
the axons and action potentials measured. They are of immense advantage to the animal
because, with the increase in diameter of the axon, resistance for propagation of action
potential is decreased, whereas velocity of propagation is increased. They conduct for
longer distances with fewer or no synapses which also further facilitates conduction. On
the whole they are pre-eminently suited for rapid conduction. These Fhres are involved in
escape reactions of the animal. The squid for example is rapidly propelled in water by
expelling the water within the mantle cavity through the fun,- I, by rapid and
simultaneous contraction of the muscles of the mantle innervated by the giant fibres using
Comparative Forms and the principle ofjet propulsion. The animakcan escape from its enemies by this method
Functions within a fraction of a second.
- - - - -- -

11.6 INFORMATION PROCESSING

pp

Yod have already seen that the sensory neuron transmits the information it receives, as an
action potential along its axon. We call them an impulse. The intensity of the stimulus is
trahsmitted aldng the nerve fibre by changing frequency. A stimulus of greater intensity
will result in greater frequency of firing along the axon, one of lower intensity will have
lower frequency. Let us see what happens when one of the first pair of legs of a
cockroach is touched. Stimulation of the mechanorecepors of the first leg of a cockroach
by touch will be carried by the axon to neurons in the first thoracic ganglion where it
makes connection, in simplest case, through synapse, with a motor neuron directly. In
more complicated cases, the motor neurone may be informed through one or more
association neurons. These neurons may be receiving at the same time a number of other
stimuli from a large number of presynaptic fibres. Some of them may be excitatory and
some of them may be inhibitory, thus carrying opposing instructions. This type of
connection also enables higher centres of the central nervous system to exert control over
the action, so that, if inappropriate, it can withhold from exercising the reflex, like
running away. Thus the postsynaptic cell receiving often contradictory messages, decides
whether to fire or not, on the basis of all the instructions it receives. In case the neuron
'decides' to fire, the stimulus can result in the release of neurotransmitter at its terminals
making the concerned muscle to contract, causing movement of the leg and the animal
run away.

This kind of receptor-effector loop is the lowest level of information processing by

central nervous system and is called a reflex. In lower animals this constitutes their entire
behaviour. However, in higher animals especially in those like cephalopods and insects,
with a well developed central nervous system, varying degrees of higher levels of control
are exerted on the reflexes, with increasing hierarchy of the central nervous system
exerting its influence. These animals have larger and complex type of brains by
invertebrate standards; their large eyes and other sense organs feed a variety of stimuli
into the brain. There are various motor centres in the central nervous system ultimately
controlling activities through various motor centres. For example,~horacicganglia of
insects are under the control of suboesophageal ganglion; the suboesophageal ganglion in
turn is under the control of the brain.

, We must not forget that most animals have also the capacity of learning. Learning
provides the animal with a record of information on which it can draw and act. The
memory provides this information and the central nervous system can select the motor
activity best suited for the animal's life.

1 1 7 RECEPTORS
In the previous sections you have studied how the nervous system is organised. Now you
will study in detail, more about the receptors.
11.7.1 PROPERTIES OF RECEPTORS
The receptors collect information about the change in the environment such as
temperature, illumination etc. In lower animals, sensory perception is mediated through
undifferentiated nerve endings. These are their sense organs. However, the receptors or
sense organs become more specialized and elaborate. Such elaborate sense organs are
present even in Cnidaria, and they become more complicated in higher animals. You have
already seen that sensory receptors convert one form of energy into another, electrical. So
all receptors are transducers. Even one molecule of odour substance is often capable of
evoking a small electrical disturbance in a chemoreceptor. This local current is usually
the result of amplification too, and is propagated along the axon of the sensory receptor.
The receptor is thus extremely sensitive to change in the environment. You have to also
note that each sensory receptor responds to only one kind of stimulus. For example,
chemoreceptors will respond to only chemical stimuli, but not to mechanical or light
stimuli. Another important fact you have to bear in mind is that the frequency of nerve
impulse increases with strength of the stimulus. In other words, information regarding
intensity of the stimulus is transmitted by frequency code. Thus the code of information
travels as impulse in axaxonsof all receptors, be it mechanoreceptor, photoreceptor or
chemoreceptor. But because each type of receptor is sensitive to one type of stimulus only Nervous System and Sense
and because these axons cany impulses to particular regions of the central nervous Organ
system, brain correctly interprets the information reaching it. Depending upon the type of
stimulus, receptors may be divided into mechanoreceptors, chemoreceptors and
photoreceptors.
SAQ 5
1. A moth : (Write "Yes" or "no" against each capacity)
a) sees a flower;
b) hears the sound produced by a bat;
c) smells its mate.

2. Do you think:
i) the three pieces of information will reach the brain of the moth as electrical
impulse? (YesMo)
ii) the moth will recognize the three objects above, even if the nerves concerned
are severed from the brain? (YesMo)
iii) the moth will recognize the three objects above correctly, even if the nerves
concerned are connected to the brain improperly (for example, optic nerves
to antennal centre; antennal nerves to optic centre, and auditory nerve to
optic centre) ? (YesMo)

Mechanareceptors
Mechanoreceptors include those receptors involved in perception of touch, pressure,
tension, hearing, vibration, gravity, muscle tension etc. These would appear to us at first
sight as rather heterogeneous assemblage of sensations, but the basic mechanism involved
in the perception of all these is the same: they are all sensitive to corrtact with objects and
are mechanically deformed temporarily by all the above stimuli. In the simplest case, they
involve free nerve endings on the body surface. But often mechanoreceptors are more
complex.

A mechanoreceptor of wide occurrence in the nonchordate metazoans is the statocyst

(fig. 11.12). It is primarily an organ concerned with perception of gravitational force and
is a balancing organ found in almost all phyla, including coelenterates. It consists
essentially of a particle of calcium carbonate called statolith. The statolith is carried in a
vesicle lined by a layer of sense cells which have hair-like processes. The statolith rests
on the sense cells. The vesicle is of course filled with a fluid. The animal can appreciate
the direction of gravity or tilt, from the sense cells on which the statolith comes to rest.

Attachment stalk

Nonmotile cilia

Fig.ll.12: Gravitation and equilibrium receptors. A. A statocyh of a mollusc.

In insects, as in other arthropods, since the body surface is covered by a rigid cuticle,
mechanical sensation should be transmitted through cuticle to the sense cell situated
within the body. In the trichoid sensillurn, for example, a bristle or seta is articulated in a
socket. The tip of a sensory cell is attached to the base of the seta (Fig. 11.13). The
movement of the seta generates an impulse in the sensory cell.
Comparative Forms and
Functions

Base of hair

Articular membrane

s;lopoid body

<;LuY:~mrna ~~
cell
~.--

Fig.ll.13: Simple trichoid sensillum.

On the other hand, in the case of campaniform sensillum of insects (Fig. 11.14) there is
no projecting hair. It consists of a small elliptical, thin, dome like, arched, cuticular area
with a thickening or rib along its long axis. To the middle of this is attached the sense
cell. The campaniform sensilla are distributed widely in insects, especially at the base of
wings and on legs, in groups. Mechanical deformities of the cuticle cause slight bulging
of the roof of the sensillum outward pulling the hair and stimulating it.
&
Dome-shaped plaque
7

Accessory cell

Sense cell

/ : k x ~ e u r i l e m m acell

A B
Fig.ll.14: (A) Campaniform sensillum and (B) section through tip of campanlform sensillurn to show
stiffening rod of cuticle running along cuticular plate present in some species.

Stretch receptors (Fig. 11.15) capable of perception of muscle tension or stretch are
distributed in the connective tissue associated with muscles of many groups of animals.
This consists of a multipolar neuron with free nerve endings embedded in the fibrous
connective tissue. Stretch of the muscle is capable of stimulating this receptor.

Many insects have a fme membrance or tympanum stretched across a cavity with air on
both sides. In the locust (Fig. 1 I . 16), for example, a pair of such tympanal organs are
present in the fvst abdominal segment, the inner side of the membrance held over the air
sac of the tracheal system. Against the inner side of the tympanum are held a group of
mechanoreceptors. Sound waves between 500-1 1,000 c/s causing vibration of tympanum
can generate impulses in the tympanal membrane. You will be surprised to know that
'
many insects like moths are even sensitive to ultrasound which we cannot perceive.
 Nervous system' and Sense-

)L Nerve
.Organ

Connective tissue
$f( 1 Intersegmental
membrane\,

recepror nucleus

Fig.11.15: Stretch receptor of male mosquito.

~ F l i e hmuscles
t

Fig.11.16: A. Diagrammatic horizontal section through the base of the abdomen of Locusta, showing the
positions of the tympanic membranes, the air-sacs and the a8sociated spiraclu. B. Diagram to
show the method of attachment of the auditory ganglion to the inner surface of the tympanum
of Locusta. The f ~ l d e dbody, styliform body and elevated process are cuticular structures. The
orientations of t kscolopidia are indicated by the arrows.
 Comparative Forms and
1 Functions
I The following are apparently different types of sensations:
I
I
1) Sound
2) Touch
3) Gravity
4) Muscle tension

Examine Wether it js correct to club the coresponding receptors together as

mechanoreceptors? Why?

'Chernoreceptors
These are receptors involved in perception of chemical stimuli. You will see that there are
three types of chemoreceptors among metazoans:
i) those concerned with general chemical sense perceived by undifferentiated nerve
endings or unspecified receptors. These are distributed all over the surface of the
body and are common among lower animals such as cnidarians, worms and
molluscs. This the most primitive type of chemoreceptor and results in escape
reaction of the animal.

ii) Chemotactile sense perceived by contact chemoreceptors involving contact with

molecules in dilute solution; they are usually present in large numbers in special
regions of the body. In mammals these are situated on the tongue and we call the
sensation taste or gustation. But in Octopus they are concentrated around the rim
of thexckers, and in insects they are present on mouthparts and in flies, they are
found in abundance also on tarsal segmentof foot. So one may say that the fly
"tastes with their f e d (Fig. 1 1.17).

LThinwalled cavity
Thick-walled cavity
I
-Dendrites of
chemoreceptors

\ ,Socket of hair

Lw-,Hypodermis
enclosing dendrites

Fig.ll.17: Chemosensory hair (taste receptor) from the mouthparts of an insect.

iii) Olfaction involves distant chemical sense and is the most sensitive type of Nervous System and Sense
chemical sense. Olfactory receptors are capable of perceiving molecules of Organ
extremely low concentration in air and we usually call it smell. They are
distributed in high density on the antennae of insects. The combined electrical
recordings of the antenna1 nerve produced by the odours on the olfactory receptors
of the antennae are termed electroantennograms. Certain insects can recognize
their mates even at a distance of a couple of kilometers, from the odour
(pheromone) molecules they emit.

You can easily distinguish chemoreceptors structurally with comparative ease among
arthropods, especially insects, as they are associated with well differentiated cuticular
outgrowths, usually setae, bristles or peg like structures. In them Olfactory receptors
(Fig. 11.18) often have many sense cells; their dendrites highly branch and end close
beneath the surface of the peg in association with large number of tiny pores, in a liquid.
The odour molecules from air are dissolved in the liquid, and on capture by the dendrites,
set in an impulse in them. Contact chemoreceptors in insects have however a smaller
number of sense cells associated with the bristle and their dendrites do not branch but
terminate at the tip of the hair in association with a pore. Chemoreceptors play an
important role in the life of animals, such as finding food, recognition of mate, selection
of oviposition site etc.

T z n i n g s in cuticle
t ~ a s a spot
l

'
1-
Lw
Scolopale

Dendrites
k z ~ r i c h o ~ cell
e n

ensory neuron
Epidermis

Basal lamina
Fig.11.18: Diagram of a peg-shaped insect olfactory sensillurn in longitudinal section.

SAQ 7
How would you.distinguish between the contact chemoreceptors and olfactory receptors
of an insect, structurally?
Comparative Forms and Photoreceptors
Functtonr
Photoreceptors are involved in absorption of light by photosensitive pigments. The
chemical change involved generates the ihpu~seconcerned in the nerve cells. The
pigments involved are caroteiioid derivatives consisting of retinine (vitamin A aldehyde)
combined with a protein (opsin). Rhodopsin is one of the common pigments.

Simple eyes are very common; highly differentiated compound eyes have evolved in
many arthropods, and well differentiated camera type eyes, comparable to vertebrate eyes
occur in cephalopods. You will see among many non-chordates a variety of eyes ranging
betweerl these extremes.

Simplest eyes in Platyhelminthes and annelids are in the form of pigment spots consisting
of sensory cells associated with pigment cells (Fig. 1 1.19). They serve differentiation of
light from darkness. In many polychaetes they reach higher level of organization. The cup
may often secrete a lens. The cephalopod eye (Fig. 11.20) is the result of highest
elaboration of vesicular eye. The eye in Octopus in fact reaches the structural complexity
of that of vertebrate eye, resembling it in considerable details. However, the cephalopod
retina is direct type with sensory ends of the receptor cells directed towards the source of
light whereas the retina of vertebrate eye is inverted with the sensory ends turned away
from the source of light.

Epidermis

cell

erve

Fig.ll.19: Simple invertebrate eyes. A. ~cellus'ofplanarian worm. B. Eye of a polychaete worm. C. Eye-
cup with sensory neurons in the leech.
 Nervous System and Sense
Organ

Fig.11.20: The octopus has a pair o f large image-forming eyes. Each eye is a camera-like eye i n which the
receptive ends of the retinal cells are directed toward the light source.

We find among arthropods simple eyes like the median eye of nauplius larva, dorsal
ocelli of insects, stemmata of many insect larvae and ultimately highly advanced
compound eyes of many insects and crustaceans. The compound eye (Fig. 1 1.2 1) is
made up of large number of units called ommatidia. Often you may be able to recognize
hexagonal patterns on the surface of the compound eye; each of these represents an
ommatidium. Each ommatidium consists of a dioptric (optic) unit and a sensory unit. The
dioptric unit consists of: (i) a biconvex, colourless cuticular lens secreted by a pair of
corneagen cells. When secretion of the lens is completed, the comeagen cells are pushed
apart and are converted into primary pigment cells. (ii) A crystalline cone secreted by
four Semper cells. These two lenses determine the focal length of the ommatidium. The
sensory components consist of eight receptor (retinula) cells arranged around an axis.

I +corneal lens

Lamina ganglionaris

Fig.ll.2 1: Diagrams illustrating the structure of the compound eye. A. Section through part of an eye
showing the arrangement o f ommatidia. B. Surface view of part o f the eye o f an aphid, which
consists o f a small number of ommatidia, showing the facets well separated b y v d
cuticle. C. Surface view of part of the eye o f a syrphid, which consists of a large number of
ommatidia with the facets crowded together. D. Detail of a single ommatidium.
Comparative Forms and Each retinula cell forms an inner fibrillar rhabdomere. The rhabdomeres of the eight
Functions retinula c'ells together form the rhabdome. From the base of the retinula cells arise a nerve
fibre. The visual pigment is situated in the rhabdomere and hence the rhabdomeres are
comparable to the rods and cones of vertebrate retina. Photostimulation of rhabdomere.
results in nerve impulse which is transmitted along the nerve fibre. There are also a few
secondary pigment cells which isolate each ommatidium from its neighbours. You have
to bear in mind that, unlike photopigment distributed in rhabdomes, the pigment granules
in.the primary and secondary pigment cells are not photopigment but serve to cut off light
entering one ommatidium into the adjacent ommatidium. The nerve fibres of the rtinula
cells enter the optic lobe. The electrical recordings of the response of the eye is called
electroretinogriim.

There are two types of compound eyes: (i) the apposition eyes and (ii) superposition
eyes. The apposition eye is characteristic of diurnal species. The receptor cells are highly
elongated, extending distally as far as the crystalline cone. The image formed by the eye
as a whole is a mosaic of areas (Fig. 1 1.22 A). Nocturnal insects have superposition eyes
(Fig. 1 1.22 B). In them the retinal cells are widely separated from the crystalline cone;
each receptor cell is stimulated by light entering through a number of gnlmatidial lenses.
This type of eye is suited for vision in dim light. The over-lapping images, however, give
poor resolution. The pigment of the primary and secondary pigment cells move in strong
light to screen retinal cells so that they can be stimulated only by light entering along the
ommatidial axis. The eye is then called light adapted, protecting it from light of high
intensity. In dim light, pigment moves to expose the retinal cells, which can now be
reached by light from a wider source. Now the eye is called dark adapted.

~ o r n e a lens
i Distal pigment
/Corneal lens,

J%rystal~ine cone
Nocturnal species
Dark adapted Light adapted
L /
v
Diurnal species

Fig.ll.22: A. Insect ommatidia. showing a diurnal type (left) and r nocturnal type (right). In the diurnal
type, the pigment is shown in two positions: adapted for very dark conditions on the lefi side,
and for relatively bright conditions on the right. B. Nocturnal type of eye adapted for dark
conditions, showing how light can be concentrated upon one rhabdome from several lenses. I f
the pigment moved downward. light from peripheral lenses would be screened out.

SAQ 8
Which of the following cells of the compound eye contains photosensitive pigment?
a) Primary pigment cell
b) Secondary pigment cell
c) Retinula cell
d) Rhabdomere
 -
-

Nervous System and Sense

Organ

11.8 SUMMARY
After reading this unit you have learnt that
The nervous system is made up essentially of nerve cells called neurons. They are
sensory, motor or association neurons, and may be unipolar, bipolar or multipolar.
A nervous system is present from Cnidaria onwards. In Cnidaria, it is in the form of a
subepithelial nerve net, with very little centralisation.
During evolution of the metazoa, clear trends in the development and evolution of a
central nervous system are visible. These consist of: 1) Disappearance of
subepithelial nerve net, giving place to a central nervous system where moior and
association neurons are grouped together. 2) Initially, the central nervous system is
in the form of thickened nerve cords with uniformly distributed new? cells along
the periphery of the cords. 3) Subsequently the cells become clustered together into
ganglia with commissures and connectives. 4) Finally, with cephalisation,the brain
has evolved which has assunied gradually greater importance and better
organization.
Giant fibres have evolved in many groups of non- chordate metazoans to facilitate
rapidity of conduction of impulses. They are involved in escape reactions of
animals.
Receptors are sensory neurons. They perceive various kinds of stimuli and transform
them into electrical impulses. Different types of receptors are specialised for
perception of different kinds of stimuli. Accordingly, they are mechanoreceptors, I

chemoreceptors or photoreceptors.
Receptors may be scattered throughout as individual cells; they usually tend to get
concentrated as for examples in eyes, statocysts etc. which frequently occur among
animals.

11.9 TERMINAL QUESTIONS

I . How would you identify a neuron, structurally?
......................................................................................................

2. What is neuroglia?
......................................................................................................
1 Comparative Forms and
Functions

3. Mention the three major types of receptors; what are their functions?
......................................................................................................

'\ a'
4. What i r the basic difference between t h ~ ~ o p eye
g and
d vertebrate eye? Cite
one important difference.
......................................................................................................

5. Why do yatl consider epithelial nerve net as the primitive type of nervous system?

6 . Why do you consider the cephalopod brain as highly evolved? Give one good
reason.
......................................................................................................
 Nervous System and Sens.
11.10 ANSWERS Orgal

. Self-Assessment Questions

i) b A
ii) a B
iii) c B

The peripheral, subepidermal nerve net disappears; the nervous system becoming
mainly submuscular, with nerves.
While sensory neurons are retained at the periphery, the association neurons and
motor neurons become concentrated into central nervous system;
iii) The neurons become further concentrated into ganglia, connected together by
connectives and commissures;
iv) With cephalisation, the brain develops, which becomes more and more important
with evolution.

1) a) Yes
b) No
c) Yes

2) i) Yes
ii) No
iii) No

6.
Yes. Because all the four types of sensation produce their effect by bringing about
temporary deformation of the receptors concerned.

Contact chemoreceptors Olfactoty receptors

1. Have smaller number of sense have larger number of sense cells.

cells associated with them

2. Dendrites do not branch dendrites highly branch

3. Single pore at the tip of the sensillum numerous pores all over the sensillum
Comparative Forms and 8.
Functions C) retinular cell. (note: though rhabdomere contains pigment, it is not a cell. It
is only part of the retinular cell).

Terminal Questions
1. Neuron has a cell body called perikaryon and one or more processes called neurites.
2. Neuroglia is the connective tissue of the nervous system, filling the space among the
neurons. It also includes Schawann cells, covering the neurites, it includes the
perineurium and the neural lamella covering the ganglia as also the nerve sheath.
3. Mechanoreceptors: sensations of touch, gravity, sound, tension.
Chemoreceptors: taste, odour.
Photoreceptors: vision, sensation of light.
4. In the cephalopod eye, the receptor cells of retina are directed towards the source of
light, in the vertebrate eye they are directed away from the source of light.
5. The epithelial nerve net is found among the most primitive metazoans, where this is
the only type of nervous system. It is not found among higher animals.
6. Various centres (association, motor and sensory) of the brain are highly developed
and localised in large and distinct lobes of the brain.
 UNIT 12 ENDOCRINE SYSTEM
\I ructure
Introduction
Objectives
Endocrine versus Neural Integration
Endocrine Organs
Ne~~rosecretoryCells and the Concept of Ne~~rosecretion
Neurosecretory Systenis
Other Endocrine Organs
Hormones in Growth and Reproduction
Annelida
Mollusca
Artliropoda
Hormones controlling other functions
Summary
Terminal Questions
Answers

12.1 INTRODUCTION
- - - - - - - - -

In the previous unit you have learnt that the nervous system brings about integration and
co-ordination of various activities of the animal. The afferent stimuli from various sense
organs are brought to the brain, which is the coordinating centre. From the brain
appropriate efferent stimuli are transmitted to various motor or effector organs resulting
in coordinated motor activities of the animal. You have seen that these stimuli travel
through a closed system of circuits comparable to telegraphic cables, called nerve fibres,
I
in the form of electrical disturbances. There are indeed many minute gaps in between the
connections where neurotransmitters make and break the circuits between gaps. In the
present unit, you will learn that there is another equally important system, which also
serves transmission of messages form one part of the body to another, bringing about
integration and coordination of various activities of the animal. This is the endocrine
system. The endocrine system brings about communication through chemical substances
called hormones produced by endocrine glands. The hormones are transported to their
I
target tissues or organs by the blood stream. The nervous system and the endocrine
I system form an integral part of the coordinating system of the animal. In fact the two
systems are closely associated with each other and the branch of study of the two closely-
knit interacting systems is known as neuroendocrinology. Before you begin the study of
I endocriye system, you may quickly review what you have studied under nervous system.
Objectives
After studying this unit you should be able to:
describe the differences between the neural and hormonal integration,
explain the concept of neurosecretion,
distinguish between an ordinary neuron and a neurosecretory cell,
name the major endocrine structures in annelids, molluscs, insects and crustaceans,
describe the process of hormonal control of moulting and metamorphosis in insects,
elucidate the mechanism of hormohal control of colour change in crustaceans, and
discuss the importance of hormones in the life of non-chordates.

12.2 ENDOCRINE VERSUS NEURAL INTEGRATION

A question that might come across your mind is, " what is the need for two types of
integrative mechanisms, the neural and the endocrine"? Each of these systems has
specific properties and functions and therefore confers specific adaptations to the animal.
In the previous unit you have learnt that the nervous system responds rapidly to the
various stimuli. So, when a rapid action is called for, nervous system will be more
effective. On the other hand, in functions controlled by endocrine system hormone
concentration is built up gradually, and, only when the titres reach a threshold level, the
function of the cell or tissue or organ is carried out.

Essentially hormone concentration is built up gradually, the specific function is carried

out and then the hormone is metabolised 2nd broken down slowly. As a result, the effect
of the hormone is sustained and continued for a long duration. Thus when rapid
responses are required, such as the escaping from a predator, the animal resorts to neural
Comparative Forms and
mechanism; but when a siow but sustained effect is called for, for instance, during the
Functions
growth of the animal or the development of the oocyte, the hormonal mechanism takes
over.

SAQ 1
Some activities exhibited by non-chordates are given below. Try to assess whether the
type of regulation involved is neural (N) or endocrine (E).
i) A prawn changes colour pattern of its body to suit its surroundings.
ii) Yolk deposition taking place in the eggs of cockroach.
iii) A resting moth flies away to escape from a lizard.
iv) A honeybee visiting flowers.
v) A snail retracting into its shell, in response to fouling of the water in which it lives.
vi) Transformation of a caterpillar into a butterfly.

ENDOCRINE ORGANS
Endocrine organs, as described earlier are those organs or tissues, which release chemical
substances directly into blood stream. Unlike exocrine glands they do not have ducts to
carry their secretion, and are hence considered ductless glands. Table 12.1 lists the
endocrine glands of the higher invertebrate groups, the secretions they produce and the
functions of the secretions. Among non-chordates neurosecretory cells (NSCs) form an
important constituent of the endocrine system. These are nerve cells specialised for
hormone production. So they combine neural and endocrine properties. Apart from this,
endocrine glands may arise from the nervous system, by complete modification. Corpus
cardiacum of insects is one such gland. Many non-chordates have also epithelial.
endocrine glands derived from embryonic ectoderm, mesoderm or endoderm layers. The
optic glands of cephalopod molluscs, androgenic glands and Y-organs of crustaceans, and
the corpus allatum and prothoracic glands of insects are some examples. You will be
studying these structures and their functions in detail. But before we proceed further, let
us look into the concept of neurosecretion a little more elaborately, since neurosecretion
plays an important role in regulating metabolic and reproductive functions in non-
chordates.
12.3.1 NEUROSECRETORY CELLS AND NEUROSECRETION
We have earlier said that the neurosecretory cells are an important component of the non-
chordate endocrine system. Of course, they are also present in chordates. But unlike
chordates, among non-chordates there are fewer epithelial endocrine glands, and so they
have to depend more heavily on neurosecretory cells for chemical coordination.

Berta Scharrer and Ernst Scharrer are considered progenitors of the concept of
neurosecretion. Put in a simple way, it is the concept of neurones taking to secretary
activity and producing hormones. We can easily locate and identify these NSCs in
histological sections of the brain or ganglia, because they contain plenty of stainable
material or colloids, unlike ordinary neurones, which do not contain any stainable
colloids. We believe that these colloids are carrier substances for hormones. Very often
we can prove the presence of such hormonal principles in such cells by experimental
means. Unlike the neurosecretory neurones, the ordinary neurones do not contain or
secrete any hormones. Though they release neurotransmitters at synapse and
neuromuscular junctions, these are not released into blood and are short lived, unlike the
hormones secreted by NSCs. In the live brain of certain insects, in dissections, we can
easily see the NSCs as tiny bluish white specks through the brain sheath, using a
binocular dissection microscope.

'The neurosecretory material containing the hormone is produced mainly in the cell body,
and is transported through the axons to the finger-like axon terminals (Fig 12.1). If we
follow these axons, we see that these axonal endings often form s,wellings in association
with blood spaces at a distance from the neuronal cell bodies. The secretory material is
stored at the swellings and hormones are released from them in@ blood stream. Such
organs seen in associatiqn with blood spaces and containing the swoHen nerve endings of
NSCs are called neurohaemal organs. You will see that there are neurohaemal organs in
some non-chordates. Sinus gland in crustaceans and corpus cardiacum in insects are
examples of neurohaemal organs. Vertebrates also have their own neurohaemal organs -
the neural lobe of pituitary is one such neurohaemal organ.
 Endiirim Syitem
able 12.1 :
Animal Group Endocrine gland Some of the hormones Some of the functions
isolated
Annelida Brain NSC Stimulates growth and regeneration

lnfracerebral gland
I Inhibits sexual maturation and
epitoky
Stimulates male and female gonads
i) Cephalopoda
ii) Gastropoda NSC of brain Growth hormone stimulates body
growth.

Growth retarding hormone inhibits

body growth and stimulates egg
production.

Dorsal bodies Stimulate development of male.and

female phases of reproductive
cvcle.
Arthropoda
i) Crustacea X-organ - sinus gland Red pigment Colour change; moult inhibition;
system; Other NSC;Post- concentrating hormone inhibition of ovarian maturation
commissural organs

Pericardial organs Regulates heart beat.

Y -organ Ecdydone Stimulates moult

Androgenic gland Stimulates spermatogenesis and

development of male secondary
characters

ii) lnsecta
Pars intercerebralis NSCs Prothoracotropic hormone Stimulates prothoracic glands to
produce ecdysone.

Prothoracic glands Ecdysone Stimulates moulting.

,.Corpus allatum Juvenile hormone Regulates metamorphosis by

retaining juvenile characters.
Stimulates oocyte growth and
vitellogenesis.

1 I Co~uscardiacum I Adipokinetic hormone ( Stimulates fat metabolism. I

Release

Fig.12.1 :The concept of neurosecrction.

I Comparative Forms and
Functions
Scientists have isolated hormones from neurosecretory systems. These are mostly
peptides, both in non-chordates and chordates. They contain a few mino no acids and
hence we call them oligopeptides. , I

An example is the prothoracicotropic hormone produced by the brain NSCs of insects.

Red pigment concentrating hormone of crustaceans is another example. However, as the
material available from non-chordates is very little, isolation and identification of
hormones from non-chordates has been very difficult. But presence of hormonal mater~al
in their neurosecretion has been demonstrated by experimental means in many non-
chordates and their functions are also known.

Before we proceed to the next section on neurosecretory systems of non-chordates, you

may answer the following SAQs.
SAQ 2
Which of the following statements are true?
i) All neurones are neurosecretory because they all secrete transmitter substances.
ii) All neurones containing stainable colloids are neurosecretory.
iii) ~euronesf which secrete hormones, are neurosecretory.
iv) Neurosecretory cells are those cells associated with neruohaemal organs.
12.3.2 NEUROSECRETORY SYSTEMS
Almost all groups of multicellular animals have been investigated to find out if
neurosecretory cells are present in them. Such investigations have shown that these cells
are located either in brain or ganglia. However, discrete neurosecretory systems made of
neurosecretory cells with their axonal tracts and axon termini forming neurohaemal
organs have been f o u ~ damong non-chordates only in certain groups. These systems are
best known among arthropods, especially in insects and crustaceans.

Neurosecretory cells
of the pars ~nterccrebralis

Corpora cardiaca
Corpora aliata ~aacou~phic
hormone
hormone ii
I
1
Ecdysone
Ecdysone
I

LARVA

PUPA
ADULT

Fig.12.2 : Brain neurosecretorycell -corpus cardiacum -corpus allatum system of a generaliscd Insect.

Among insects (Fig. 12.2) we can see conspicuous groups of NSCs in the median region
of the protocefebrum of the brain, known as pars intercerebralis, on either side of the
median line. If we follow their axonal tracts we can see that they cross each other in the
brain and emerge out of the brain as corpus cardiacum nerves (nervi corporis cardiaci).
These nerves enter the corpus cardiacum situated just behind the brain. Thereafter their
neurosecretory fibre terminations branch and rebranch, ending close to aortal wall. The
neurosecretory material elaborated in the NSCs of the brain are transported and stored at
these axonal endings. At the time of necessity the hormone is released from these
structures into the aortal blood. So we can call the corpora cardiaca as neurohaemal
organs. Frequently we can also see other well-defined neurosecretory systems in
association with ventral ganglia, forming the perisyrnpathetic system in many insects (Fig
12.3).

Among crustaceans (Fig. 12.4) we can see a tightly clustered group of NSCs situated on
the margin of optic ganglia. These constitute the X-organs. Some authors describe more
'
than one X-organ on each side, in crustaceans. The neurosecretory axons from x-qrgan
 Endocrine System
traverse the eye stalk ganglia and converge to neurohaemal organ, which is called the
sinus gland. It is a white iridescent structure on the surface of the optic ganglion. As the

Medial cells

m . Corpus allaturn

Fig.lt.3 :Brain neurosecretory cell -corpus cardiacum -perisympathetic system of a generalised insect.
Neurosecretory cells of the ventral ganglia are also shown.

name indicates, the sinus gland is situated close to a blood sinus. This is the most'
common picture in the crustaceans that have been studied. Usually there are no NSCs in
the sinus glands.

X-organ
Sinus gland

X-organ

Gecarcinus lateralis

Fig. 12.4 : X-organ - sinus gland system of a crustacean.

In crustaceans we can commonly see oth& neurosecretory systems also (Fig.12.5). Fur
example, the post-commissural organs are found attached to the circumoesophageal
connectives. These are also the neurohaemal organs. Their neruosecretory cell bodies
appear to be situated in the brain. So also, the pericardial organs floating in the
pericardial sinus of crustaceans are neurohaemal organs. Most of their neurosecretory
cells are situated in the thoracic ganglia.
Comparative Forms and
Functions

Cerebral ganglion (brain)

Circumoesophageal connectives

Postcommissural organs

Sub-oesophageal ganglion

Last thoracic ganglion

\ Q )\ First abdominal ganglion

Fig.125: The endacrine organs of a generalised crustacean. Except the androgenic gland and the male
reproductivcsystem,other glands are common for female also.

In millipedes and centipedes also we can see cerebral NSCs leading to a neurohaemal
organ, the cerebral glands on each side (Fig. 12.6). In some %illipedes, in addition, there
is another neurosecretory system with NSCs usually in the tritocerebrum and connective.
body as the neurohaemal organ. The connective body is situated close to the post oral
connectives whtre it joins the circumoral commissure.

Neurosecretory cells

ner
cerebral

Fig.12.6 :Neurosecretorysystem of a millepede.

12.3.3 OTHER ENDOCRINE ORGANS

Corpus cardiacum: We earlier mentioned that corpus cardiacum (pl. corpora cardiaca)
is a neurohaemal organ in insects. It is gland of neural origin. They are considered as
transformed ganglia. Apart from these, we will discuss in the ensuing section a number
of other endocrine glands. Most of them are, however, of epithelial origin. The corpora
cardiaca are situated close behind the brain dorsally, on either side of the dorsal aorta or
remain in close association with it. They appear as a white bluish organ in dissections
and so we can easily locate them under dissection microscope. Though they are
neurohaemal organs in insects as they contain fibre terminations of NSCs situated in the
pars intercerebralis of the brain, they have their own cells too (Fig. 12.2). That is, they
contain intrinsic cells, which synthesise hormones different from.the ones produced by
the NSCs of the pars intercerebralis. So they have a dual function. They are the
 Endocrine System
neruohaemal organs as well as endocrine glands in their own right. They receive usually
two or sometimes three nerves from the brain in front and are connected to the corpora
allata (sl. corpus allatum) behind, by allatal nerves. In some insects there may be also
ordinary nonsecretory neurones in the corpora cardiaca.

Corpus allatum : The corpus allatum is located in the neck region of insects(Fig 12.2).
It is usually a rounded, solid and translucent structure. Nerves to corpus cardiacum in
front connect it. The allatum is densely packed with small nuclei with little cytoplasm
around them. It produces juvenile hormone. The juvenile hormone is a straight chain
terpenoid (a type of lipid) compound of 16 to 19 carbon atoms.

Prothoracic gland: The prothoracic glands (Fig.12.7) are usually made up of loose
strings of cells situated in the prothorax of immature insects. They produce the steroid
hormone ecdysone. These glands undergo degeneration in the adult.

Prothoracic gland

*~an~lion (thoracic)

-Ventral nerve cord

Fig.12.7: Prothoracic glands of an insect.

The Y-organs: The Y organs Fig.12.5) are a pair of endocrine glands in crustaceans. We
see them either in the antennary or in the second maxillary segment. They produce the
- 'moulting hormone ecdysone in crustaceans.

The androgenic glands: The'androgenic glands (Fig 12.5) are the endocrine glands
found only in male crustaceans: 'They are attached to the hinder end of the vasa
deferentia and their hormone is thought to be a peptide or protein.

The optic glands: The optic glands of cephalopod molluscs (Fig.12.8) are the endocrine
glands found in the optic stalk, internal to the eye and are well innervated. However, the
gland is not part of the neurosecretory system.

Fig.12.8 :Optic glands of the cephalopod mollusc Octopus.

Among the gastropod molluscs (Fig.12.9) as for example in the fresh water snail Lymnea
stagnalis, there are structures called tbe dorsal bodies and the lateral lobe containing
follicle glands; both of them are associated with,the brain. They are important structures
in these molluscs.
Comparative Forms and
In polychaete annelids, there is an infracerebral gland close to the brain. This endocrine
Functiohs
gland has a complex structure.

Mid dorsal body

Latero-
body
lobe

Fig.12.9 :Endocrine system ofa gastropod, Lymneastagnalts'showingMedian and Lateral Dorsal bodies
as well as Lateral lobes. Also shown are the brain neurosecretory cells.

SAQ 3
Indicate which of the following statements are true or false.
i) Neurosecretory cells usually secrete steroid hormones.
ii) Neurosecrefory cells usually secrete peptide or protejn hormone.
iii) Prothoracic glands secrete steroid hormone.
iv) Juvenile hormone is a straight chain lipid.
v) Androgenic glands secrete a lipid hormone.

12.4 H~RMONESIN GROWTH AND REPRODUCTION

The growth and reproduction of higher invertebrates are known to be regulated by the
endocrine secretions. The regulatory mechanisms have been well demonstrated in
annelids, arthropods (specially in crustaceans and insects) and molluscs. We shall briefly
study the hormonal regulation of growth and reproduction in chosen non-chordate
groups.

&Axon tracts

infracerebral gland

v B l i o d vessel

Fig.12.10 :lnfracerebral glands of a polychaete annelid.

 Endocrine System
Annelids
Studies on polychaetes have shown that the endocrine glands play a key role in growth
and reproduction. In addition to the brain neurosecretory cells, we see that in annelids
there is an infracerebral gland (Fig.12.10). This is situated in close association with the
brain 'immediately beneath or behind it. The infracerebral gland has its own secretory
cells. In addition, the neurosecretory cells of the biain have their fibre terminations in the
brain, close to the infracerebral gland, or even passing into the gland. Nereis and
Nephtbs are examples.
The polychaetes are capable of rapid growth when young, with proliferation of new
segments. They are also capable of extensive regeneration. However growth rate and
regenerative ability decline with age. An interesting phenomenon in polychaetes is
epitoky. This takes place during sexual maturation in some nereids. Sexual maturation in
them is accompanied by changes in certain regions of the body. The changes include
enlargement of parapodia and development of modified chaeta. The animal is
transformed into heteronereid phase (Fig. 12.1l), adapted for rapid swimming and sexual
swarming.

Anterior portion of mature nereid

I

Immature nereid

(c) Parapodium of mature nereid

Fig.12.11 :Changes undergone by the immature nereid during transformation into heteroneried worm.

In polychaetes growth, regeneration as well as sexual maturation accompanied by epitoky

are all controlled by a hormone of the brain-infracerebral gland complex. This hormone
is required for growth and regeneration, and is secreted in large quantities at the early
stage of the animal's life, which is the growing stage. However, the same hormone
appears to inhibit sexual maturation and epitoky. So sexual maturation does not take
place at this stage. When growth period terminates the hormone secretion drops and the
animals become sexually mature.

127
Moonlight synchronises swarming in these worms. In moonlit nights we can see these
mature worms swarming in the sea in large number and they shed their germinal products
(sperm and ova) more or less simultaneously. This brings about successfil fertilisation.
Molluscus
We know fairly well about the endocrine mechanism of growth and reproduction in the
freshwater snail Lymnea stagnalis. This is a gastropod, and like other gastropods,
Lymnea is protandric hermaphrodite. That means, in these groups of animals, male
gonads mature first and this is followed by the maturation of the female gonad. The
dorsal bodies produce hormones, which promote development of both male and female
phases of reproductive organs. In the female phase these glands produce pronounced
effects. They stimulate maturation, yolk deposition in the eggs, ovulation and oviposition.
They also stimulate growth and differentiation of the female accessory sex organs. In this
animal, certain groups of neruosecretory cells of the cerebral ganglia produce a hormone
(growth hormone) which stimulates growth and differentiation of the female accessory
sex organs. In this animal, certain groups of neurosecretory cells of the cerebral ganglia
produce a hormone (growth hormone) which stimulates growth of the body. On the other
hand the lateral lobes attached to the brain produces a growth-retarding hormone. This
hormone not only retards growth of the body but also causes an increase in egg
production.

The optic glands of the cephalopod mollusc Octopus control reproduction in the male and
female. The glands secrete a gonadotropic hormone at the onset of sexual maturity. This
stimulates development of the genital ducts and maturation of germ cells.
Crustaceans
Arthropods have a tough and often hard exoskeleton made up of cuticle composed of
I
chitin and protein. The cuticle imposes restrictions on the growth of the animal. Unless
the cuticle is shed, growth is not possible to any extent. The periodic shedding of cuticle
is known as moulting. So, moulting is closely related to growth in arthropods. In
crustaceans, two hormones control the moulting process. The Y-organ produces the
moulting hormone, the ecdysone, a steroid. Under the influence of ecdysone the animal
moults. So it is a moult-inducing hormone having a positive influence on moulting.
However, moulting in crustaceans is also under the inhibitory control of another hormone
produced by the X-organ and released at the sinus gland. This hormone has a negative
influence on moulting. If the inhibitory hormone concentration in the blood dominates,
the animal moults. It is the balance between the two hormones that determines moulting
in crustaceans. However, it is not certain whether the inhibitory hormone acts on the Y
organ decreasing the release of moulting hormone ecdysone or it acts directly at the level
of the epidermal cells where ecdysone also acts. Anyway, the effect is same. You will
see that in many animals, two or more hormones of antagonistic effects may control very
often one function, the balance determining the result.

Reproduction in crstaceans is also under hormonal control. The X-organ-sinus gland

complex is involved here also. It is easy to remove the X-organ-sinus gland system of
crustaceans, which are situated in the eyestalk. Removal of the eyestalk, known as
!
eyestalk ablation, would result in the removal of the endocrine structures. Removal of
the eyestalk from the adult female crustacean results in the acceleration of yolk deposition
in the oocytes; size and width of the ovaries will therefore increase. Sometimes it results
in precocious egg-laying too. Injection of an extract of the eyestalks of the adult females
into & eyestalk-ablated animal will reverse the trend. Such types of experiments led to
the conclusion that in normal animals ovarian growth is inhibited by a neurosecretory
hormone from the eye stalk, namely, X-organ-sinus gland complex. When the hormone is
absent, the ovaries start growing, depositing yolk in the oocytes. Even though a number
of other hormones controlling ovarian and testis activity have been postulated, suff~cient
evidence to prove their existence is tacking. An exception is the androgenic gland I

hormone of some crustaceans.

The androgenic glands are normally present only in the male crustaceans and are
necessary for production of sperms by testes. These glands stimulate spermatogenesis in
the male. Even.though it is normally absent in the female, you can imp1 t the
8"
androgenic gland into a female and watch the effects. Implantation o f t e androgenic
gland in the female can convert the ovaries into testes. The ovaries ultimately produce
sperms. The androgenic gland hormone is required for the differentiation of male
 Endocrine System
secondary sex characters also. You have studied the phenonmenon of testicular
feminisation in crabs occurs due to parasitisation by Sacculina. The parasite causes the
degeneration of the androgenic gland. In the absence of the androgenic gland hormone
responsible for maintenance of maleness, the sex reversal takes place.
Insects
In insects hormones regulate moulting and metamorphosis. The larvae or nymphs that
hatch out of the eggs undergo regular moulting which is followed by growth, as in
crustaceans and ultimately they become adults. The change in form from larva to adult is.
known as metamorphosis. In insects like cockroach, grasshopper etc., (hemimetabolous
insects), the change in form is gradual. However in some other insects like moths,
butterflies, houseflies etc., (holometabolous insects), the change is more conspicuous
during the later period of the life history and the adult which emerges from the pupa is
quite different. As in crustaceans, the hormone which brings about moulting in these
animals is also ecdysone but in insects it is secreted by prothoracic glands.

How is metamorphosis in insects brought about? In insects the hormone that is

responsible for preventing the animal from metamorphosing is juvenile hormone secreted
by the corpus allatum. In fact, as the name implies, juvenile hormone keeps the insect
juvenile. So in effect it inhibits metamorphosis. We have already seen that the
prothoracic glands in immature insects secrete ecdysone. This hormone causes the insect
to moult. As long as the larva moults in the presence of high titres of juvenile hormone, it
moults into another larva (Fig.12.12). If the moult takes place when the titres of JH are
low in the blood, it results in a pupa. Finally when there is no JH circulating in the blood,
the pupa moults into adult. Thus the concentration of JH in blood determines the type of
resulting individual. It is also known that JH causes the repression of those genes, which
are responsible for adult differentiation.

Karel Slama, an insect endocrinologist from Czechoslovakia while working in the

laboratory of Carroll Williams in United States found that the bugs he brought from
his country were not undergoing normal metamorphosis. Many of them did not reach
adulthood and died as nymphs. Since it was a laboratory working with hormones,
they suspected a possible contamination of juvenile hormone (that prevents
metamorphosis) and care was taken to avoid any such possible contamination.
Everything, right from a glass-dropper was kept clean; but the problem persisted and
the insects did not complete the metamorphosis. Slama then undertook the laborious
task of screening all the materials in the laboratory for the possible cause of
disruption of metamorphosis in the bugs. Much to his surprise he found that the filfer
papers that were used to line the petri dishes in which the bugs were reared were the
cause for the problem. An analysis of the filter paper showed that they contained
juvenile hormone like substances (JH analogues) in it. Slama and Williams called the
substance the 'paper factor'. But how did the paper come to possess juvenile hormone
activity? In fact the JH activity was traced to the trees from the pulp of which the
paper was made. Subsequently, a number of trees were screened for JH activity and
the activity was found in several of them. Here is an instance of plants having
evolved a certain defense mechanism from the possible attack by the insects. The JH
analogues would not allow the insects which feed on the trees.which possess such
analogues to moult into adults and the insects die as juveniles, thus saving the trees
from the subsequent attack of insects.

Yet another type of an interesting defense mechanism which plants have evolved
relates to the presence of anti-juvenile hormone substances in them. The anti-JH
substances have the property of suppressing the juvenile hormone secretion. If an
insect nymph or larva feeds on a plant, which possesses anti-JH substance in it, the
substance causes the inhibition of the corpus allatum of the insect, thereby causing
the lowering of the juvenile hormone titres ir. the blood. Under such circumstances,
the insect, instead of undergoing a normal metamorphosis, undergoes an abnormal or
precocious metamorphosis. Such a metamorphosis results in abnormally developed
insects called adultoids, which die without reproducing. And the substances in plants,
which cause a precocious metamorphosis of insects, have been termed as precocenes.

Let us now examine the hormonal control of reproduction in insects. While studying the
mechanism of metamorphosis in insects, we have seen that adult differentiation takes
Comparative Forms and
place when there is no JH in blood stream. Essentially at the time of pupal adult moult the
Functions
corpus allatum is inactive. Once the adult emerges, the corpus allatum becomes once
again active and produces JH. In female insects the JH has been shown to function as a
gonadotropic hormone. Under the influence of the JH, the fat body of certain insects '
synthesise yolk protein precursors', the vitellogenins, which are subsequently releasedjnto
the haemolymph. Also JH is shown to promote the uptake of vitellogenins from the
haemolymph by the developing oocytes. Inside the oocytes the vitellogenins are
transformed into vitellins, the yolk proteins. Besides regulating the oocyte development
and vitellogenin synthesis and uptake, JH has been shown to promote oviposition as well.
Besides JH, the neurosecretory substances produced from the neurosecretory cells of the
pars intercerebralis region of the brain are shown to regulate the oocyte development and
oviposition in locusts and grasshoppers.

Brain
Corpus cardiacum

a-
Suboesophageal
ganglion

Thoracic ganglion 1

0-Thoracic ganglion 2

Thoracic ganglion 3.
Abdominal ganglion 1

/f&.:. .$ Abdominal ganglion 2

) .!::L Abdominal ganglier. 4

q-:gy-

Terminal abdominal
ganglion

Fig.12.12 : Hormonal mechanism of moulting and metamorphosis in insects.

In mosquitoes such as Aedes, ecdysone is shown to be the gonadotropic hormone. Ovary

has been shown to be the source of ecdysone that promotes viltellogeneni'n synthesis. In
hymenopterans such as honey bee, an interaction between JH and ecdysone promotes
vitellogenin synthesis. While the JH primes the fatbody for vitellogenin synthesis, the
synthesis per se depends on ecdysone. Thus insects exhibit a variety of mechanisms of
hormonal regulation of yolk protein synthesis and a generalised mechanism does not
prevail.

Spermatogenesis,the formatlun of mature sperm from the primordial spermatogonial

cells has not been shown to be under hormonal control. But in many insects JH has been
shown to regulate the synthesis of the male accessory gland substances. These
substances, mainly proteins, are transferred to females as a part of the seminal fluid at the
time of mating. They are known to perform a variety of functions, such as the formation Endocrine System
of spermatophores, inhibitionof receptivity of once mated females, enhancing their
fecundity and acting as oviposition stimulants. In certain grasshoppers, they were also
shown to contribute to the yolk formation in females.

12.5 HORMONES CONTROLLING OTHER

FUNCTIONS
Cru ;taceans exhibit pronounced capacity for physiological colour changes. It is known
th-t the colour changing mechanisms in Crustacea are regulated by hormones. The
anl:nals have a variety of chromatophores containing different pigments: the
melanophores usually contain black pigments.

Many crustaceans are brilliantly coloured animals and they have the capacity to change
their body colour. Body colour is often due to coloured pigments present within certain
cells. The cells concerned are called chrornatophores. Colour change may be (1) either
due to synthesis of new pigments or destruction of existing pigments. In other words it is
due to a quanititative change in the pigments of the body. This type of colour change
called morphological colour change is very slow, taking considerably long time to
accomplish. (2) However, animals can also change their colour without recourse to
synthesis or destruction of pigments. That is, there is no change in the quantity of
pigments present. This involves movement of already existing pigment granules within
the chromatophjores. When the pigment granules within the chromatophores are
concentrated at a point, the cells appear pale or blanch; when the pigment granules are
dispersed throughout the cell, it appears brightly coloured. This type of colour change is
called physialogical colour change, and is comparatively more rapid than morphological
colour change.

Colour change is of immense advantage to the animal. By adapting its colour to the
surroundings, it can escape from its enemies; it serves to attract opposite sexes when the
colouration is sexually dimorphic; sometimes the colour change may be usefid to the
animal to adjust to the environmental temperature.

Similarly red, orange, yellow and white chromatophores are also present in these animals.
These chromatophores are controlled by a number of hormones. The source of these
hormones may be the neurosecretory cells of the X-organ -sinus gland complex or the
isolated scattered neurosecretory cells of the brain or of the thoracic ganglia or the post-
commissural organs. Some of these hormones have been isolated, purified and their
structure elucidated and they have even been synthesised. The red pigment concentrating
hormone of Pandalus borealis has thus been found to be an octapeptide, a peptide
conatining eight amino acids. Under its influence red pigment in the chromatophores
concentrates and the cells appear pale. So the animal which earlier appeared red will now
appear pale.

How can the opposite effect be produced? The red pigment, which is now in a
concentrated condition, will disperse, usually under the influence of a red pigment
dispersing hormone. The chromatophores will now assume red colour and the animal
becomes red. Sometimes even the very disappearance of the red pigment concentrating
hormone from the blood stream may result in dispersal of the pigments. In that case also
the chromatophores will appear red and the animal too will appear red.
We have now seen that non-chordates too have endocrine glands. In annelids, molluscs,
crustaceans and insects, growth and reproduction are under hormonal control. In insects
they also control metamorphosis. In crustaceans the hormones bring about colour
change.. Besides these functions, a variety of other functions are also under hormonal
control in them. Crustaceans and more so, insects, have been studied extensively fiom
this point of view. The metabolism of carbohydrates and fats are known to be regulated
by hormones in insects. Adipokinetic hormone in insects is known to regulate the fat
metabolism in insects. Similarly hyperglycemic and hypoglycemic factors from brain
and corpus cardiacum have been shown to regulate carbohydrate metabolism.
Prothoracic glands, which secrete ecdysone, are themselves under the control of a
hormone fiom the brain neurosecretory cells. A number of other hormones controlling a
variety of functions in non-chordates are already known. Also there is accumulated
evidence to show that there is the possibility of the existence of yet unidentified
Comparative Forms and hormones in non-chordates. In short, we may sayathathormones play a very important
Functions
part in integration of a multitude of physiological activities in non-chordates.
. .
SAQ 4
Correlate the following endocrine ~ r ~ a n s ' ~ i under
v e n A with their direct positive
functions given under B.

1. Prothoracic glands a) moulting

2. Y-organ b) development of gonads
3. Optic gland c) colour change
4. Androgenic gland d) regeneration
5. Post-commissural organs e) prothoracic gland stimulation
.6. Pars intercerebralisNSCs f ) male sex characters
7. Brain-infracerebral complex

12.6 SUMMARY
In this unit you have studied the various endocrine glands of non-chordates, the
'
hormones they release and the functions they discharge. The glands are either
neurosecretory, neural or epithelial in nature.
Neurosecretory cells are important constituents of the multicellular non-chordate
endocrine system as they have fewer epithelial endocrine glands. The neurosecretory
cell is a neurone specialised for hormone production.
Corpus cardiacum and sinus gland are the main neurohaemal organs of insects and
crustaceans respectively. Corpus cardiacum is also a neural endocrine gland. Copus
allatum, prothoracic glands, Y-organs, androgenic glands and optic glands are some
of the epithelial endocrine glands.
The endocrine glands play important role in annelids, molluscs, crustaceans and
insects, in coordinating and con@lling growth, reproduction, colour change and
metabolism.

12.7 TERMINAL .QUESTIONS

1. Give two differences between neural and endocrine coordination.

2. Give two importa?' characters of neurosecretory cells.

......................................................................................................
 What is a neurohaemal organ? Name two neuro-haemal organs in non-chordates. Endocrine System
3. . .
......................................................................................................

......................................................................................................
.........................................................................................................
4 . Explain how moulting and metamorphosis are controlled by hormones in insects.

5. Compile a table showing the main endocrine glands of non-chordates you have
studied, the hormones they produce and the functions they discharge.

12.8 ANSWERS
Self-Assessment Questions
1) i) - E, ii) - E, iii) - N, iv) -N, v) -N, vi)- E

2) i) - F, ii) - F, iii) - T, iv) - F

4) i) -a, ii) - a, iii) - by iv) - f, v j - c, vi) - e, vii) - d

Terminal Questions
1) Neural coordination is (a) rapid and (b) momentary whereas hormonal coordination
is (a) slow and (b) sustained.
2) Neurosecretory cells are (a) neurones (b)which produce hormones. a

3) Neurohaemal organs are organs containing fmgerlike, highly branched, fibre

terminations of neurosecretory cells, situated close to blood sinuses, and store
neurosecretory material and release hormones into blood on requirement. (a) Pars
intercerebralis NSC -corpus cardiacum system (b)X-organ-sinus gland system.
4) Prothroracic glands secrete ecdysone. When the concentration of ecdysone reaches a
certain level, moulting results. Corpus allatum secretesjuvenile hormone. When
Comparative Forms and moulting takes place in the presence of high concentrations of juvenile hormone
Functions
another larval-larval transformation takes place; when moulting occurs in the
presence of relatively low concentrations of JH, the larval-pupal transformation
occurs. Finally When the corpus allatum is totally inactivated and no JH secretion
occurs, then the pupal-adult transformation occurs. Thus the metamorphosis is
controlled by the titres of JH in blood.
UNIT 13 REPRODUCTION IN NON-CHORDATES
Structure
13.1 Introduction
Objectives
13.2 Asexual Reproduction
Binary Fission
hultiple Fission
Fragmentation
Budding
Strobilation
Formation of Special Reproductive Units -The Gemmules
13.3 Regeneration Leading to Asexual Reproduction
Fragmentation and Regeneration
Autotomy and Regeneration
Epitoky
Polarity and Regeneration
13.4 Asexual Reproduction - its Prevalence and Significance
13.5 Sexual Reproduction
The gametes and significance of sexual reproduction
The two sexes and sexual dimorphism
Patterns of sexual reproductio~i
The reproductive organs
Accessory sex glands
Mating and fertilization
Ovipary, vivipary and ovovivipary
13.6 Hermaphroditism ,
13.7 Parthenogenesis
I 3.8 Alternation of generations
13.9 Reproduction, Life Cycles and Larval Forms
13.10 Summary
13.1 1 Terminal Questions
13.12 Answers

13.1 INTRODUCTION
-- -- - -- - -- - - - - -

In previous units you studied the body systems and life processes of non-chordates which
are essential for keeping an individual organism alive and enabling it to hnction
normally. There you noted that although each of these systems performed the same
overall common function, yet it differed considerably in the different phyla of the non-
chordates and even in the s u b - g r ~ ~ of
p sthe same phylum. These differences are evidently
correlated with mainly two things - the structural organisation of the organisms and, the
environment.which they occupied. The same is true about the differences in the
reproductive patterns. Since reproduction also is a significant phenomenon of life
enabling a species to continue generation after generation, it is logical to find that the
non-nhordates have evolved no less diverse methods of reproduction. Such methods range
from simple splitting into two, to very complicated methods of sexual reproduction,
parthenogenesis, etc. You will learn that asexual reproduction, combined with sexual
reproduction has evolved complicated life cycles and several lar\,al forms that are entirely
different from the adult forms. This unit is designed to make you familiar with such
/
varied types of reproduction among the non-chordates.
Objectives
After studying this unit you will be able to:
illustrate the various types of asexual reproduction in non-chordates giving examples,
define regeneration and describe how it is contributory to asexual reproduction,
describe sexual reproduction and its importance,
cite examples of hermaphroditism and relate the phenomenon with the conditions of
life of the hermaphrodites,
describe the phenomenon of parthenogepesis, its significance and its various types in
the non-chordates,
explain the importance of alternation of generations in some phyla of non-chordates
differentiate between direct and indirect life cycles and
describe some of the important larval forms found in invertebrates.
Comparative Forms and
Functions 13.2 ASEXUAL REPRODUCTION
Reproduction may be considered as production of true copies. Most of the animals which
are quite familiar to us produce male and female gametes (sperms and eggs). These
gametes unite together to form a zygote that subsequently develops into an adult, which is
a copy of the parents. This kind of reproduction is called sexual reproduction. There are
also a large number of animals, particularly among non-chordates, which reproduce from
body parts other than gametes. This does not involve sex or sex-cells or meiotic division.
It basically involves both growth and regeneration of missing parts. Such reproduction is
categorised as asexual reproduction and it is very common among coelenterates,
plananians, polychaetes and 01igoch.aetes. Asexual reproduction is also called vegetative
reproduction; this is because it involves some body part or any special unit produced by
it, not related to the gamete-producing organs (ovaries and testes). In fact, asexual
reproduction is the most primitive one -the first organisms that evolved on the earth were
protistans and they probably just divided into two - the simplest form of asexual
reproduction. Even today, this is the most common type of reproduction among
protistans.

Main forms of asexual reproduction

The following are the main types of asexual reproduction.
Binary fission
Multiple fission
Fragmentation
Budding
Strobi lation
Formation of special units (or bodies)
13.2.1 BINARY FISSION
Binary fission is a process in which an organism divides mitotically into two equal
individuals which are normally identical to each other. This process is widespread in
protozoans. Amoeba, Euglena and Paramecium are the most familiar examples. All the
three are freshwater forms. Amoeba (Fig. 13.1) is an irregularly shaped unicelluar
organism. %en full grown and when the conditions are favourable it slows down its
movements and produces short fine radial pseudopodia. The contractile vacuole stops
functioning and may disappear. The nucleus elongates, becomes dumb-bell shaped and
undergoes mitotic division. The cell constricts in the middle to finally divide into two
smaller daughter amoebae each possessing one daughter nucleus (Fig. 13.1).
Subsequently, the pseudopodia become normal and a new contractile vacuole is formed in
each. The entire process of fission in Amoeba may take about 30 minutes at moderately
warm temperature. Amoeba thus divides regularly, almost every 24 hours.

Fig. 13.1 :Binary fission in freshwater amoeba. -* '

Euglena, a freshwater flagellate, divides longitudinally (longitudinal binary fission). The

kind of division in it is said to be symmetrogenic, that is, producing gymmetrical or
mirror-image daughter cells (Fig. 13.2). The centrioles first divide into two and the
nucleus then begins to divide. Each centriole produces a new basal body and a new
flagellum. The contractile vacuole also divides into two. As mitosis (nuclear division)
reaches completion the cytopharynx (gullet) also begins to divide and finally when the
organelles have duplicated the anterior end starts dividing as a fork, which deepens
posteriorly to ultimately produce two daughter Euglena.
 ~eproduetionin
Non-Chordates

Flagellu~n Fission Daughter individuals

Fig. 13.2 :Binary fission in Euglena.

Paramecium (Fig. 13.3) divides transversely into an anterior half and a posterior half. The
fission here is called homothetogenic, meaning looking like the letter "8" i.e., "theta" of
greek alphabet. The macronucleus divides amitotically and the micronucleus divides
mitotically. A second cytophaynx is produced by the posterior half. Two new contractile
vacuoles are formed one going to each half. In the meanwhile a constriction is formed at
the middle which deepens to finally separate the two independent daughter paramecia.
The entire process of binary fission in Paramecium may take about two hours. According
to one estimate as many as 600 generations can be produced in a single year. It may be
seen that all theiprogeny thus produced from a single original parent as a result of binary
fission either in Amoeba, Euglena or in Paramecium have the same genetic composition.
They are identical in all respects. The term clone is used to refer to such a population.
-
Plasmotomy is a variant form of fission found in some multinucleate protozoans like
Opalina (Fig. 13.4), a ciliate that lives as a commensal in frog's rectum. In this kind of
asexual reproduction the multinucleate protozoan divides into two or more parts without Fig. 13.3 :Transverse binary
any nuclear division and the existing nuclei are distributed among the new individuals. fission in
The new, still multinucleate, daughter opalinas produce more nuclei as they grow. Paramecium.

I 2 1 4
i Fig. 13.4 :Multinucleate Opalina which reproduces asexually by plasmotomy. Opalines usually divide
longitudinally, between the rows of cilia.
I
13.2.2 MULTIPLE FISSION
Multiple fission is a variation of fission in which the parent divides mitotically into a
number of smaller units, which are the daughter individuals simultaneously. The nucleus
in this case divides very fast, repeatedly. Subsequently the cytoplasm surrounding each
I *. daughter nucleus begins to form separate units leading to formation of small uninucleate,
masses. These units become independent. This method of reproduction is also known as
schizogony and it occurs typically in Sporozoa and some Sarcodina. Schizogony is also
known as sporogony if it is associated with production of sporozoites after formation of
zygote. A very good example of multiple fission is seen in Plasmodium where
sporozoites are introduced by the mosquito in the human host and where they enter the
liver to undergo several schizogonous cycles (pre-erythrocytic schizogony). The
merozoites liberated from liverenter erythrocytes to undergo another cyc!e of schizogony
in which the schizont inside the human RBC undergoes multiple fission to produce
merozoites (refer to Unit 2 of this Course). Can you visualise here that the parasite is
exploiting the food resource to very rapidly multiply its numbers? That is one very
significant advantage of asexual reproduction.
Comprrtive Forms and
In addition to the different types of fissions described above, a modified type of fission in
Functions
the form of budding occurs in suctorians. The adults bud off one or more smaller
daughter cells. When buds are given off from the outer surface, it is known as external
budding. When budding takes place inside a chamber, buds arising from the wall of the
chamber it is called internal budding (Fig. 13.5). The buds, when released, possess
ciliated bands and they swim around. But they soon attach to the substratum and become
adults.

Fig. 13.5 :Suctorian budding in Tokophlya lemnariim. The bud escapes from an internal pouch through a
narrow pore (a) and swims away. (b) I t attaches to the substratum to produce a stalk. (c) The
cilia are lost and tentacles appear. (d-e)

13.2.3 FRAGMENTATION
Fragmentation is a phenomenon in which a parent animal spontaneously (on its own
accord) splits into two or more fragments. Each one of such fragments then regenerates
the missing parts to form a new individual. This method is reported in certain sea-
anemones and some worms. Sea anemones sometimes break off part or whole of their
pedal disc while the main body moves away (pedal laceration). The pedal disc may put
out lobes which pinch off and regenerate into small sea anemones. Sometimes a sea
anemone leaves behind its pedal disc alongwith a few mesenteries at the site of
attachment, in such cases the part left behind regenerates into a new sea anemone and the
other main part regenerates the missing pedal disc. Fragmentation may be considered as a
type of fission. Moreover longitudinal and transverse fission occurs as a normal method
of asexual reproduction in many sea anemones.

Spontaneous separation is beautifully illustrated in certain freshwater planarians

(Platyhelminthes, Class Turbellaria). A common example is Dugesia. The fission occurs
when the animal ha5 grown to the maximum size. The fission plane usuaily fcrms behind
the pharynx.

Fig. 13.6 :A plannrian reproduciog asexually by fragmentation.

At that time the hind part of the animal is firmly attached to the substratum and the front Reproduction in
Non-Chordates
part continues to move forward until the worm breaks at the constriction into an anterior
and a posterior segment (Fig. 13.6). Each segment next regenerates the missing parts to
form a new small worm. In some planarians such as Microstomum,chains of individuals
"
may be formed as a result of repeated fission. When these individuals develop and attain
a certain degree of differentiation, they separate from the chain. (Fig. 13.7).
13.2.4 BUDDING
In reproduction by budding, small groups of cells form buds in some part or parts of the
parent animal. The bud grows, differentiates and ultimately forms a new adult individual.
Thiq adult may either remain attached to the parent as in colonial forms (sponges and
many coelenterates) or p a y separate from it to form an independent individual, as in
Hydra. Fig. 13.8 shows budding in the common freshwater hydra. It fmt starts as a
simple bulging of the ectoderm. Then both ecto- and endoderms project.outward in the
form of a bud. Thereafter, the coelenteron also extends into this little protuberance. The
bud begins to grow in size, the tentacles start forming and a mouth appears. When the bud
practically assumes the shape of a miniature hydra a constriction appears at its base which
gradually narrows to a small point and finally the little bud hydra separates off to settle
Fig. 13.7 :Microslomumwith a
somewhere else as an independenthydra. chain of zooids.

I
Fig. 13.8 :Budding in Hydra.
Asexual reproduction by budding is quite common in many other cnidarians also. It is
prevalent both in the polypoid stages (as in hydra) as well as in medusoid forms
(jellyfishes). Budding at times takes the form of stolons that grow along the ground from
which are budded upright stems; this is quite common in colonial cnidarians.
How budding and fission are similar and how these are different.
Both budding and fission are similar in at least one way in that the young ones produced
by these processes are the result of direct splitting off from the body of the parent. But the
two are also clearly different in many respects. These differences are listed in the Table
13.1 given below.
'Table 13.1 :Differences between budding and fission.

The parent individual persists after the daughter The parent individual loses its identity after splitting into two
individual has budded off. or more daughter individuals.

size before it pinches off but it is still smaller than the in structure but all are smaller in size than the parent. Later
they grow to attain their normal dimensions.

Budding is rather slow and gradual. Fission is rapid and instantaneous.

~ o n i b r a t i v eForms and
Functions 13.2.5 STROBILATION
Strobilation is a kind of asexual reproduction in which successive segments are separated
off from the body one after another. It is well illustrated in some jellyfishes (Scyphozoa)
and in tapeworms. In the life-history ofAurelia, a scyphozoan, the planula larva develops
into a hydra-like stage called hydratuba or scyphistoma. From the stolon, the
scyphistomae'may produce new scyphistomae. After a certain period of life this
scyphistoma undergoes transverse division producing buds of medusae. This gives it a
kind of pattern of a pile of saucers arranged one above the other. In this pile, the youngest
(smallest) saucer lies towards the base and ihe oldest and the largest one at the free end.
This "pile of saucers" stage is called the strobila. The transverse discs or young medusae
- buds separate one after another from the strobila, each resulting in a kind of larva (called
ephyra) which gradually metamorphoses to become the adult Aurelia (Fig. 13.9, also refer
to Fig. 14.7 of Block 2 of this course).

Ephyra

Fig. 13.9 :Strobilation of the scyphistoma ofdurelh.

A situation somewhat similar to the above also occurs in the tapeworms. New segments
(proglottides) are co~ltinuouslydemarcated off from the neck by a sort of transverse
fission; Here also, it is the oldest proglottid at the free end which breaks off and the
process keeps on repeating (Fig. 13.1Ci).
Scolex

, .
-
Fig. 13.10 :Strobilation in tapeworm.

:O
 13.2.5 FORMATI~N
OF SPECIAL REPRODUCTIVE UNITS - Reproductlon in
Non-Chordates
THE GEMMULES
All freshwater sponges (Spongillidae) and some marine species, under dry conditions or
low temperature produce asexual bodies called gemmules. These are formed before the
onset of the adverse conditions. The parent sponge disintegrates, leaving behind, the
gemmhles, which can withstand drought and winter. In spring, when temperature begins
to get wanner the gemmules undergo development and each of them produces a new
Layer of amphidisc
sponge. Micropy:I lspicules
Formation of gemmules
During the formation of gemmules, masses of food - laden amoebocytes called
archeocytes feed on other cells and lay down yolk within them. They get surrounded by
spongiocytes (another kind of amoebocytes). The latter secrete a two layered hard
covering. A tiny pore, the micropyle, is left as a small opening in this protective shell.
Thereafter, some scleroblasts (spicule-producing cells) come to lie between the two layers
of the shell and produce spicules (Fig. 13.1 1). The gemmules are now completed. The
parent sponge may now disintegrate leaving behind the gemmulus. On revival of
favourable conditions, the gemmules start development. The centrally located archeocytes
emerge out through the micropyle as a mass which undergoes cell division and
differentiation to produce a new sponge.
,
Inner membrane
- - Archeocytes
SAQ 1
I. Match the modes of asexual reproduction with the appropriate examples. Fig. 13.11: Section of Gtmmule
of Spongilla. Throughout
Mode of reproduction Example ' dormancy which may last for
several years, the cells of the
1. Binary fission (a) Hydra inner mass retain the
2. Fragmentation , (b) Aurelia ultrastructure of active cells.
Within 24 hours of favourable
3. Plasmotomy (c) Plasmodium conditions arising, the membrane
4. Budding (d) Planaria below the pore is digested by
5. Multiple fissim (e) Opalina enzymes and cells start to escape
6. Gemmule formation (f) Paramecium out.
7. Strobilation (g) Spongilla

11. Tick (J ) mark the correct statements.

i) The most primitive mode of reproduction in animals is by binary fission.
ii) Multiple fission and cell division are one and the same.
iii) Asexual reproduction by fragmentation depends on power to regenerate.
iv) Budding is a slow and gradual process.
v) ~sexua~re~roduction by gemmules is a regular method found in all sponges.

13.3 REGENERATION LEADING TO ASEXUAL

REPRODUCTION
Regeneration is defined as the replacement of the lost parts of the body of an organism.
This capacity is present in almost~allorganisms but in many cases, especially in most
higher vertebrates and some non-chordates, this capacity is limited only to replacement of
cells, but not to organs or major parts of organs. But regenerative capacity in many non-
chordates, especially the lower forms, is tremendous a d can lead to the formation of a
completely new individual from any broken bit or the fraction of the parent's body, and in
this manner it almost becomes yet another method of asexual reproduction.

Regeneration as a process leading to asexual reproduction can occur in two situations: '

1. Animals natural$fragmenting (spontaneous separation) followed by regeneration of

the missing parts in each fragment.
2. Animals accidentally cut or broken into pieces by injury followed by regeneration in
each bit to produce as m.any complete individuals.
' 13.3.1 FRAGMENTATION AND REGENERATION
Both the situations mentioned above i.e., whether occurring naturally or accidentally, can
in general, be ~ategorisedas fragmentation. Of these, the first one, of natural or innate
fragmentation followed by regeneration producing complete individuals has alreqdy been
 Comparative Forms and
covered in the previous sections. We shall here describe the second manner in which J
Functions
accidentally produced body fragments regenerate all the missing parts to produce I
complete individuals.
1 I
Regenerative capacity is fairly well-developed and extensive in'echinoderms, but to a
different degree in different groups and species. Though echinoids or sea urchins are
poorly endowed with regenerative power, star fishes (asteriods), brittle stars (ophiuroids)
and sea lilies (crinoids) have extensive capacity to regenerate. Most of these, especially
many star fishes and brittle stars, not only regenerate lost arms or part of the central disc,
but even an arm can regenerate a whole animal, including the other arms and the central
disc. Some starfishes eg., Linckia are able to cast off their arm, which will regenerate into
a starfish (Fig. 13.13). A number of starfishes and some crinoids (eg. Ophiactis) even
normaly reproduce asexually. This involves division of central disc so that the animal
divides into two. This is known as fissiparity. The two halves will regenerate the missing
1
half. Certain holothurians (sea cycumbers) are unique in showing what is called
evisceration. On encountering any immediate danger of some intruder or enemy these
can throw out the large masses of "tubules of cuvier" attached to respiratory trees, or
some times even almost all of their viscera through their cloaca and regenerate them later.
In some species evisceration is a normal seasonal phenomenon.

Spontaneous fragmentation of the body followed by regeneration of the body parts to

foun a new individual is a common form of asexual reproduction in polychaetes. In
many syllids the point in the septa where the segment will fragment is predetermined and
is different from other septa. In such species fragmentation and regeneration are highly
organised and each fragment can develop into a complete individual. In some cases the
Fig. 13.12 :'Regeneration in original somite remains large and a head and tail regenerate at each end. These break OX
starfish Linckia leading to
and the tail portion grows a new head, while the head grows a new tail.
asexual reproduction (a).
A single arm regenerates the
central dise as well as the
13.3.2 AUTOTOMY AND REGENERATION
remaining four arms (b).
Shedding of body parts in self-defense to divert the attention of the predator-enemy or in
any other emergency is a kind of autotomy (auto: self, tomy: cutting). The most familiar
example of autotomy is a breaking off of the tail in the common house lizard among the
chordates, but the phenomenon is far more common among the non-chordates. You have
already seen that this occurs is many echinoderms. The star fishes and brittle stars can cast
off their arms which can regenerate into whole animals. A very interesting case of autotomy
followed by regeneration is found in a polychaete annelid worm called Chaetopterus.
Antenna
Ciliated groove .N

Fan-like parapodia outh

Fig. 13.13 :A tube-dwelling annelid Chaetopterus (a). tt can regenerate the lost front end of the body, as
well as the entire worm from a single isolated body segment in the figure from a single fan
segment (b).
 It lives permanently inside a U-shaped parchment tube in the muddy bottom in shallow Reproductionin
Non-Chordates
sea water (Fig. 13.13 a). If the anterior end of this worm is pulled by a predator a
constriction between 12th and 13th segments breaks the body into two pieces. The worm
loses its head region to enemy while the remaining hind portion is left behind in the U-
tube. This hind portion contains the gonads and regenerates the front part required for
normal feeding activity. Chaetopterus has been found to have a remarkable power of
regenerating a complete worm from a single isolated body segment i.e., segment 14 (Fig.
- 13.13 b).
13.3.3 EPITOKY
This is a highly specialised form of asexual reproduction where an asexual worm known
as "atoke" buds off a sexual form. Certain polychaete annelids occur in two distinct
phases - a non-reproductive (asexual) and a reproductive (sexual) phases. Nereis irrorata
is a simple example. This worm is a bottom dweller in shallow sea. In the non-breeding
phase it lives mostly hiding among rocks and crevices or inside tubular formations to
creep out only at night for feeding. During breeding season, it undergoes marked
structural changes in both males and females. The body gets differentiated into an
anterior half and a posterior half - heteronereis (Fig. 13.14). The anterior half called
atoke remains almost unchanged while the posterior half called epitoke undergoes a few
radical changes. The changes in the epitoke primarily include (i) the parapodia, which get
enlarged to become oar-like for efficient swimming and (ii) the gonads, confined only to
this region, grow profusely and the gametes fill the body cavity. At certain fixed nights,
determined by environmental factors such as the moonlight in a particular month of the
year, the posterior sexual parts (epitokes) of the worms get detached and swim around.
Large number of such male and female epitokes produce a dense swarm on surface of the
sea during these night.

These epitokes then burst and disintegrate releasing sex cells in the sea where they Fig. 13.14: Heteronereis, male
undergo\fertilization. The non-swarming body parts (atokes) regenerate the lost segments showing the atoke and
epitoke regions.
to repeat the process once again the next year. The term swarming is used here to
emphasize the fact that the rising of the epitokes to the surface of the sea is collective
constituting sometimes even millions of such individuals, both males and females,
together,. When they all burst to release the sex cells, the chances of fertilization are very
high.

Fig. 13.15: a) Aufolytus budding offn chain of epitoke, budding occurs in both males and,females.
Producing(b) male or (c) female epitoke.
 Comparative Forms and There are numerous variations in qitoky. For example in Syllis vittata the epitoke
Functions,
develops the head and a pair of eyes even before fragmentation from the atoke, and
similarly, the atoke begins to grow the hind part before the epitoke has separated.

In the porychsete worm Autolytus the asexual worm developed from the ovum gives rise
by a process of constrictions and posterior proliferations to new (one or more) zooids .
which may remain connected in a string before getting separated. The sexual zooids later'
develop into matuie male and females (Fig. 13.15).

Syllis ramosa is another peculiar polychaete annelid that lives inside the cavities of
certain deep sea sponges. The annelid produces lateral buds repeatedly from various
segments. Thus it presents the appearance of a branching colony (Fig. 13.16). The
individuals produced by such budding get separated. They are sexual zooids.

Amongst poriferans any piece cut away from the main body of a sponge can grow into a
complete sponge, but the process is fairly slow and months or even years may elapse
before full size is reached. Some sponges reproduce asexually by constriction of branches
and their falling off, which regenerate into new sponges. Sponges can even regenerate
from masses of isolated cells. When a sponge is squeezed through fine silk cloth it gets
Fig. 13.16: Syllis showing
branching individuals,
broken up into isolated cells or cell-clumps. Through random amoebid movements of the
amoebocytes the isolated cells form aggregates. These are the reunition masses. These -
budding off from rearrange and ultimately grow into new sponges.
parapodia.
Hydra and many other CnidarianJ like Tubularia, Obelia, and even sea-anemones, if cut
into pieces, tan regenerate producing new individuals. For example, if Hydra is cut into
three equal pieces, the piece with tentacles regenerates the pedal disc; the basal piece
regenerates the oral disc along with the tentacles; the middle piece regenerates the oral
disc from its apical end and the pedal disc from its proximal end. (Fig. 13.17).

Fig. 13.17 : Regeneration in Hydra cut into pieces.

Planarians ( Phylum Platyhelminthes) exhibit remarkable property of regeyeration. When

cut into half, the missing half will regrow in each piece to produce a complete planarian.
Many interesting expe~imentshave been performed on kegeneration in plarlarians. A two
headed planarian can be produced by splitting only the head and two-tailed specimen can
be produced by splitting the tail only (Fig. 13.18).

Fig. 13.18: Regeneration in a planarian.

 Reproduction in
13.3.4 POLARITY IN REGENERATION Non-Chordates
A distinct polarity or gradient exists in planarians (and in other animals). It means that in
any piece or segment of the body; its anterior end (head end) represents one pole and the
posterior end (tail end) represents the other pole. Regeneration is normally correlated with
this polarity. In any excised piece the cut surface towards the head end produces a new
head and the cut surface towards the hind end produces a tail.

In the remaining non-chordate phyla (i.e. Aschelminthes, Arthropoda and Molluscs)

regeneration of missing parts is minimal, and asexual reproduction involving regeneration
is almost non-existant.

13.4 ASEXUAL REPRODUCTION - ITS PREVALENCE

AND SIGNIFICANCE
Having studied the various aspects of asexual reproduction in the non-chordates we can
now make a few generalisations.

A. Prevalence of asexual reproduction

1. Asexual reproduction is far more prevalent in the lower animal phyla than in the
higher ones.
2. Asexual reproduction is largely associated with the regenerative capacity.
3. Lower animals with relatively less differentiated cells and tissues possess higher
regenerative power than the higher animals and thus contribute to asexual
reproduction.
4. In higher animals regenerative capacity in largely restricted to organ replacement (In
chordates it is even less and is limited to tissue regeneration).

8. Significance of asexual reproduction

Asexual reproduction is advantageous to the species in many ways:
1. Asexual mode of reproduction is a surer method of reproduction, it normally does not
involve chance or risk factors which are usually inherent in the sexual method.
2. Asexual reproduction, specially by fission and budding, is a method of rapid increase
in numbers and thus leads to a faster growth of population.
3. Animals reproducing asexually are able to take maximum advantage of favourable
environment by rapid multiplication.
4. Asexual reproduction by formation of gemmules in certain sponges is a means to tide
over unfavourable conditions.
SAQ 2
I. Fill in the blanks in the following sentences:
' I. In starfishes even a single separated arm can regrow a complete
as well as the remaining four .......................
............................
2. Evisceration followed by regeneration of the lost viscera is frequently found in
....................
3. Sponges can regenerate from masses of isolated cells often termed as
.......................... masses.
4. A two headed planarian can be produced by ......................... the head
longitudinally.
5. Production of ..................... in sponges is a method to tide over condition's of
drought or cold.

11. Give one main reason why asexual reproduction is more prevalent in lower non-
chordates than in the higher ones.
. Comparative Forms and I

'Functions 13.5 SEXUAL REPRODUCTION

All metazoans, whether or not capable of ropagating asexually, produce special cells
called gametes. The gametes maybe eith male gametes (sperms) or female gametes
(ova). Syngamy follows during which a sperm fertilizes an ovum producing a zygote. The
zygote develops and ultimately produces an adult. During syngamy, there is doubling of
chromosomes; the zygote and its derivative cells are diploid. To avoid repeated doubling
of chromosomes, there is meiosis or reduction division during the production of gametes.
In this process chromosome number is reduced to half (haploid number). The cycle is
repeated, and the chromosome number is maintained, (refer to sexual reproduction in the
courses LSE-05 and LSE-06)
13.5.1 THE GAMETES AND SIGNIFICANCE OF SEXUAL
REPRODUCTION
If chromosome number is thus maintained constant in a species as a result of sexual
reproduction, then what is the use of this process involving syngamy and subsequent
meiosis? The answer is, that there is mixing of maternal and paternal chromosomes in the
zygote. This, together with recombination between homologous chromosomes taking
place during meiosis, gives ample opportunity for genetic variation of the individuals
produced. On this genetic variation, selection can work and evolution can proceed.

It must be clearly noted here that sexual reproduction is a fundamental Goperty of

protozoans as well, though there are important differences between the two group as we
shall see when we proceed further in this section.
Following are some general characteristics of gametes:
I. The gametes of opposite sexes are usbally unequal in size - the ones which are
generally larger are the eggs or the female gametes, and the others that are smaller
are called the spermatozoa (or sperms) or just the male gametes.
2. The female gametes are generally laden with food material, the yolk, while the male
gametes are without food reserve.
3. The female gametes are usually nonmotile or immobile (cannot move about actively)
while the male ones are motile.
4. Both the kinds of gametes possess half the quantity of the genetic material (DNA),
i.e. half the number of chromosomes of the parent. This is expressed by saying that
the gametes are haploid (n) possessing only one out of each pair of chromosomes of
the species, whereas the parents which produced them are diploid (2n).
5. The gametes usually cannot develop singly. Instead, they must undergo syngamy
involving fertilization in which the male and the female gametes fuse together to
, form the zygote.
6. The zygote is still a single cell which has to undergo a process of cell division and
cell differentiation to gradually acquire the adult form and structure.
7. The sex cells or gametes which fertilize to produce a new individual usually come
from different parents (males and females). This provides for the combination of
genes from the two parents, and the offspring, due to such chance combinations of .
genes, may be better equipped (adapted) for life.
13.5.2 THE TWO SEXES AND SEXUAL DIMORPHISM
-
The two sexes male and female : In most animals, each species has two types of
,- individuals the male, producing male gametes or sperms and the female producing ova or
eggs. However, in many animals, especially many non-chordates, both types of gametes
are produced in the same individual. Such individuals are called hermaphrodites (recall
from LSE-05 Unit 14). You will read more about them in section 13.6. Apart fiom
producing different types of gametes, usually the males and the females c k be
distinguished by other characters. This is called sexual dimorphism. Sometimes sexual
dimorphism may be very conspicous, sometime the differences may not be very
pronounced. But practically all animals above the level of Cnidaria show varying degrees
of sexual dimorphism. Some examples are:
1. The blood fluke of the genus Schistosoma, for example, Schistosoma haematobium
(Phylum Platyhelminthes, Class Trematoda) is a parasite in human blood vessels.
The male is large and thick: it bears a long canal along its ventral surface, which is
formed by the body folds. The female is also long but thin and is firmly held inside
the male's ventral canal - gynaecophoric canal (Fig. 13.19).
 Reproduction in
Non-Chordates

Acetabuluin
/

Gynecophoric canal

Female
Place of fusion

Yolk g l a n d . 4

Intestine
I

Fig. 13.19 :Schistosotna haemzfobium, the male holding the female inside the gynaecophoric canal.
2. Ascaris lumbricoides (Phylum Nematoda or Nemathelminthes) is a fairly common
intestinal parasite of humans; males are smaller than the females and can be readily
distinguished by their curved posterior end looking like a hook (Fig. 13.20a).

i
' Fig. 13.20 :Ascaris lumbricoides. (a) - Male, (b) - Female.
Comparative Forms and
In female, the posterior end is straight (Fig. 13.20b). In males a pair of needle like
Functions
chitinous structures called penial setae project out from the transverse anus which
also serves as the reproductive aperture.

3. Bonellia is a member of a group of marine worms allied to annelids. It shows very

marked sex dimorphism. Its female has an ovoid body about 5 cm or more long,
usually lodged in crevices of rocks. A highly extensible long proboscis, which may
be about a meter long, projects from the main body. The male is a dwarf about
0.5mm long. It is ciliated all over and has no proboscis. While still young and
immature it enters the body of the female, and after attaining sexual maturity it lives
permanently inside the uterus or coelom of its female partner. There are also
co.nsiderable differences in the structure of the male.

Male Female

Fig. 13.21 :Bonellia. Male, tiny microscopic (0.5mm long), with most internal organs other than those
related to reproduction, degenerate. Female is large (5 cm long) which harbours its male
partner inside the uterus or coelom.

4. The crayfish Palaemon shows the following sex differences.

i) The male is smaller than the female.
ii) The second pair of chelate legs in males are much longer and are more densely
covered by spines and setae.
-
iii) The second abdominal appendage in males has an extra process'called appendix
masculina.

5. Insects : Most insects show sex differences, especially with regard to external .
genitalia. The common cockroach is easy to collect and examihe. The male .
cockroach Periplaneta americana bears a pair of anal styles projecting behind the
ninth sternum. These are missing in the female. The female has a much enlarged
(boat-shaped) VII sternum, largely concealing the sterna of thecVIII& 1X segments
forming a genital pouch (Fig. 13.22 a).

In mosquitoes the antennae of the males are more hairy and densely plumose (Fig.
13.22 b). In Bombyx male, it is pectinate (comb-like) (Fig. 13.22 c) and in some
other male moths (eg. Saturnia) it is bipectinate.

Many dipterans have much larger eyes (holoptic) almost touching each other in
males, while they are smaller and clearly separated in the females.

6. Among molluscs, a cephalopod Argonauta, popularly called the paper nautilus,

shows conspicuous sexual dimorphism. The male is small just about 2 cm and has
no shell, whereas the female is about 8 times larger and possesses a thin transparent
shell (Fig. 13.23).
. L
 E&production in
' Non-Chordates
I.
d eye

Coxa (firstjoint of leg)

'\BIwingm

Male Female

Male Female

Fig. 13.22 :~eriplanetaamrlcana (a)- Ventral view ofthe male, distinguished by a pair of anal styles
projecting behind 1X sternum. Ventral view ofthe tip ofthe female showing enlarged V11
sternbm and no anal styles. (b)- Antennae of male and female mosquito. (c) - Antennae of
Bombyx.

Male

Fig. 13.23 :Argonaura. female afld male,

Comparative Forms and
Functions 13.5.3 PATTERNS OF SEXUAL REPRODUCTION
Highly diverse non-chordates varying from Protozoa to Echinodermata, show numerous
patterns of sexual reproduction, the major types are:

I. Syngamy - Sperm fuses with the egg. This results in both the union of the paternal
nucleus with the maternal one (karyogamy), as well as the fusion of the cytoplasms
of the two gametes (plasmogamy). Syngamy leads to fertilisation producing a zygote
which develops into a new individuals,

Depending upon the size and shape of the gametes involved, syngamy can be
subdivided into three types (Fig. 13.24).

i) Isogamy :The gametes are morphologically similar although they may differ in
their physiological and biochemical properties. For example, the gametes
produced from the male and female gametocytes of Monocystis.
ii) Anisogamy : The gametes differ in size and structure and are collectively
known as anisogametes. Of these, the smaller ones are usually more numerous
and motile. They are called the male gametes (or the micro-gametesa s in
protozoans and the sperms as in metazoans). The fusion of micro - and
macrogametes is known as anisogamy. It is frequently found in protozoans as in
Plasmodium and Vorticella. In higher phyla the term fertilisation is used instead
~f anisogamy.
iii) Oogamy : In oogamy one gametes type is always motile and usually small (the
sperm) and the other is always nonmotile and large (the egg). All metazoans
exhibit oogamy. The eggs of most fully terrestrial non-chordates such as insects
have shelled eggs. The shell bears a minute pore (micropyle) for allowing the
entry of sperms for fertilization.

Fig. 13.24 :The three patterns of fertilsation. (a) Gametes may be of the same size and motility (isogamy)
(b) of dimerent sizes but with motility (anisogamy) or (c) of dimerent sizes and one being
motile (oogamy).

11. Conjugation -Conjugation ip a temporary union between two individuals of the

same species, during which the original macronucleus disintegrates, a new
macronucleus is built from micronucleus, and the conjugants exchange nuclear
material. The male nucleus of one moves over to the opposite individual and fuses
with its stationary female nucleus. The process is reciprocal and the conjugants
separate. This is followed by fission of the conjugants. Paramecium is a good
example. You can refer to Fig. 2.12 subsection 2.3.7 of Blgck-l for a detailed
description of conjugation. Conjugation in such cases provides a kind of
rejuvenation. Repeated asexual reproduction by fission througIpome 350 or more
generations leads to onset of senility and weakens the race that may otherwise die
out.

111. Autogamy - It is known to occur in many ciliate protozoans including Paramecium.

Autogamy involves same nuclear behaviour as in conjugation, but it takes place in a
 --

single individuals. Therefore, there is no exchange of micronuclear material between Reproduction in

Non-Chordates
two individuals (Fig. 13.25), as only a single individual is involved.

i Fig.13.25: Autogamy in Paramecium. The macronucleusdegenerates and micronucleus undergoes meiosis

I to form.8 or more nuclei (1). Two of these nuclei fuse together to form a zygate nucleus (2). The
others degenerate. The synkaryon divides to form a new micronucleus and macronucleus (3).

13.5.4 THE REPRODUCTIVE ORGANS ,

1
I
I
In most metazoans, gonads are well-defined. Testes and ovaries take up different shapes
and anatomical dispositions. In cases of certain hermaphrodites (see section 13.6) the
same reproductive tissue from one part may produce sperms and from another part the
1
I ova.
I
1 In sponges, gonads are poorly defined. Sperms and eggs arise from choanocytes and
I archeocytes which are transformed into spermitogonia. Sperms leave the sponge through
4 water currents. They enter choanocytes of other sponges which transport the sperms to
the egg. Thus fertilisation occurs in situ by the intervention of choanocytes.
1
In cnidarians, testes and ovaries are distinct. In some species of Hydra both tests and
ovaries may be formed in the same individual (monoecious, bisexual) or in different
irrdividuals but in most species they are found in different individuals (dioecious
unisexual).

I Female
Male

Fig.13.26: Sexual reproduction in Hydra. Sperms from male Hydra testes fertilise the single ovum of
female Hydra.

i
. Comparative Forms and
Functions
A single ovum produced in the ovary (Fig. 13.26) is fertilised in situ by the sperms .
usually coming from a different hydra. In colonial cnidarians the gonads are formed on
the medusae from interstitial cells beneath the radial canals. They are formed in the
epidermis of sub-umbrella. Fertilization is usually external, in water (refer unit 4, Block -
2, ~ i g4.14).
.

In almost all higher non-chordates there are three kitids of structures associated with
reproductive syste s -the gonads (testes or ovaries), the gonoducts (vasa deferentia or
7
oviducts) and the c pulatory apparatus. A few examples of male and female reproductive
organs in some nonCchordates.are shown in Fig. 13.27 to 13.32.

Median -eao
. for A
cardiopyloric strand

Coiled mass of

Flapcovering
walking leg' male genital pore
Male Female

, Fig. 13.27: Poleomon (prawn), male and female reproductive systems.

Fig. 13.28 :Scorpion, male and female reproductive organs.

13.5.5 Accessory sex glands

ilise thkir eggs internally usually have accgssory sex glands that are
some key aspects of their reproductive physiology. For example,
coating of eggs with protective and nutritive substances; transport of sperm in a liquid
suspension or in encapsulated form, and for storage of sperm in the females in special
chamber, and for supply of nutrition to sperms.
-
 Reproduction in
Non-Chordates

Testis 7
Fig. 13.29 :Cockroach. male and female organs.

[)igcstive gland.

part of
'l'\l(>~~lar
vas detkrens
.:,.,.;.. ..

.
i
.:,
2.
.._:
..
...
+.:

,
.

;,-T$

,'?:
..:;:

,t'.;..
..
. .. ..,
.. .
.....
....~

.:.*..
:.,,:
:.::;
:.:....,.
,F.
gonapophysis gonapophysis

Glandular part genital pore

Anus of vns deferens

Fig. 13.30': Pulmonete snail reproductive organ organs.

.ory
:ntal gland

Male
Female, '
Comparative Forms and
Functions

Fig. 13.32: (a) Earthworm,reproductive organs (Hermaphrodite). (b) Tapeworm, a mature proglotid
showing both male and female reproductive organs. (Hermaphrodite).

These accessory glands are distributed generally along or in the genital ducts and some in
'
the gonads. Many of them may be outside the genital system but pour their secretion into
the genital duct.

As a rule, accessory sex glands are either absent or reduced in groups that shed their
gametes into the surrounding water (Cnidaria and Echinodermata). On the other hand
they are well developed in internally fertilizing groups. (eg. Platyhelminthes, Molluscs,
Annelida, Arthropoda).
13.5.6 Mating and fehilisation
In all animals sperms are motile and must move and seek the eggs to fertilise them. For
this an aqueous environment is necessary. This requirement of a liquid medium has led to
two basic mating patterns.

1. External fertilisation - mating partners come in clohd proximity in water and

simultaneously shed their eggs and sperms in water.
2, Internal fertilisation - mating partners come into physical contact and copulate,
wherein the male transfers the sperms directly into the reproductive ducts of the
female. The ova coming down the ducts get fertilised. Internal fertilisation is
pharacteristic of terrestrial animals, but also occurs in several aquatic forms. Internal
fertilisation usually leads to development of an intromittent organ or penis in the
male partner. A variety of such copulatory organs are found in non-chordates, such
as the c ~ m ins certain worms, complex phallus (penis) of cockroachetc.
In several non-chordates there are spermathecae for storing sperms received during Reproduction in
Non-Chordates
mating. As the eggs pass down the ducts sperms are released from the spermathaca to
feitilise them. Honey bee queen mates just once when it receives enough sperms to
fertilise the many thousands of eggs she is going to lay in 4-5 years of reproductive life.
Spermatophores
Many non-chordates do not release free sperms during copulation. They have a
mechanism to bundle and enclose a number of sperms in a sheath usually made of
gelatinous material. Such bodies are called spermatophores. Most insects, the centipedes
and certain molluscs produce spermatophores. Male centipede emits a spermatophore and
places it on a web already made. The female picks up the spermatophore and takes it into
her genital opening. In many male cephalopods such as Loligo and Argonauta one of the
arms gets specialised for the transfer of spermatophore into the females. Such'an arm is
called hectocotylised arm. The spermatophore, after having been deposited in the female,
finally releases the sperms which move into the female ducts or the spermatheca.
13.5.7 Ovipary, Vivipary and Ovovivipa&
In all cases of external and in some cases of internal fertilisation, development of the
fertilised egg (zygote) takes place outside the mother's body. The egg is laid either in
water or in soil, on plants, or inside the body of their hosts in case of certain parasites. Ail
such animals or called oviparous (laying eggs).

Some exceptional cases of non-chordates showing, vivipary i.e., giving birth to young
ones nourished by thegenital tracts of the female, are found in some insects such as tsetse
fly (Glossina).

A third condition is that of ovovivipary. In this case, after fertilisation the zygote is
retained in the female reproductive system without any direct connection with the
maternal tissues for providing nourishment. Here it undergoes development. Ultimately
the eggs hatch still inside the female and the young ones are "born". This happens for
example in certain flies.

SAQ 3
1. List any three signifiiant differences between eggs and sperms.
......................................................................................................

.2. Match the features in column I with the animals in column 11.
Column I Column I1
Features
(1) Anal styles (a) Ascaris (male)
(2) Curved tail end (b) Schistosoma (male)
(3) Shell (c) Mosquito (male)
(4) Gynaecc?horic canal (d) Cockroach (male)
(5) Plumose antennae (e) Argonauta (female)

3. Name one example from non-chordate animals for each of the following:-
i) Epitoky
ii) Isoeamy
iii, iiectocotylized arm
iv) Viviparous condition.
 C
Comparative Forms and
Functions 13.6 HERMAPHRODITISM
You would have fi~llyappreciated now that sexual reproduction is definitely the more
advanced type of reproduction. It provides for combination of genes from different
genetic stocks and imparts the potentiality of greater adaptation to the environment and
evolution. The involved sex cells (gametes) i.e., the eggs and sperms come froni different
individuals, namely the female and the male. Species having separate sexes are called
dioecious (di =two, oikos = house) such species are also called unisexual - composed of
individuals of any one sex. But a different condition pervails in many non-chordates in
which both types' of organs (testes and ovaries) are found in the same individual. This
condition is called bisexualism or more popularly hermaphroditism, and such
individuals are termed monoecious (mono: single, oikos; house). The term hermaphrodite
comes from Greek mythology in which Hermaphroditos the son of Hermes and Aphrodite
possessed both male and female sex characters.

Hermaphroditism is common in non-chordate groups. In the most incipient form we can

see it even id the ciliate protozoans such as Paramecium. From the account of
conjugation of Paramacium, it is clear that it has both a male and a female pronucleus.
Thus each Paramecium serves as a male and a female.

In Paramecium during autogamy, the same micronuclear changes as seen during

conjugation occur in the single individual. But here as there is no second individual
involved, it is comparable to self-fertilisation.

Among metazoa, hermaphroditism is the rule in flatworms, (Platyhelminthes), pulmonat

among gastropods, earthworms and leec!es (Annelida) etc. In addition, hermaphroditisim
occurs among other animals also, as in some bivalve molluscs.
Kinds of hermaphroditism
Depending on the timing of maturity of the gonads, hermaphroditism can be of two
types:-
I. Simultaneous hermaphroditism. Both male and female gonads mature at the same
time. Some common examples are liver flukes (Trematoda), tapeworms (Cestoda)
earthworms, leeches, pulmonate gastropods etc.
2. Protandrous hermaphroditism. (gkprotos: first, andr: manlmale). Male gonads
mature first. This condition is common in some molluscs like limpets and slipper
shells (Crepidula).
Copulation and fertilisation in hermaphrodites
Copulation among hermaphrodites is common. Tellatworms possess a cirrus and penis
that is inserted into the uterine vaginal opening of the other worm. After copulation, the
sperms are stored in a seminal receptacle. In earthworms (Refer to Fig. 5.11 in Block 2 of
this course), the genital papillae help iqmating, the sperms released from the male genital
pore of one earthworm are received and stored in s e ~ n areceptacles
l (spermathecae) of
the other earthworm. Sperm transfer is thus mutual.

Similarly cross fertilisation is probably the rule in tapeworms when there are adjacent
individuals inside the hosts gut, but self fertilisation between two different proglottids of
the same tapeworm is also known to occur.

Majority of hermaphrodites tend to be protandrous i.e. male gonads maturing first, and
this is a condition to ensure cross fertilisation. In protandrous hermaphrodites the gonads
(testes and ovaries) may be separate organs and they mature at different times. In some
rare cases as in certain gastropod molluscs there is a single gonad called ovotestis which
produces both eggs and sperms. But the gonad first produces sperms and later the egg.
SAQ 4
Mention whether the following statements ire True or'False.
i) Earthworm may be termed bisexual, hermaphrodite or monoecious.
ii) Autogamy in Paramecium is a kind of cross fertilisation.
iii) In.most hermaphrodites the female gonads mature first.
 --
~e~rbduetionin
13.7 PARTHENOGENESIS Non-Chordates

Parthenogenesis means development from a female gamete (egg) alone, without

fertilisation. Among non-chordates this condition is found in several groups such as,
some crustaceans and a number of insects including aphids (plant lice), thrips, ants,
honeybees etd.
There are different types of parthenogenesis.
1. Obligatory parthenogenesis. It is found in certain insects and some branchiopods
(Crustacea). Here males are unknown and parthenogenesis is the only way of
reproducing.
2. Facultative parthenogenesis. In this case ova can develop into adults whether
fertilised or not. The fresh water flea Daphnia, for example, may produce eggs
developing parthenogenetically into females for several generations. Some times
males may appear. Then fertilised eggs are produced. Some insects like the aphids
(plant lice) have a complicated pattern of reproduction. In them parthenogenesis
occurs alternately with sexual phase.
The queen of the honey-bee (Apis sp.) may lay unfertilised and fertilised eggs. The
fertilised eggs give rise to females (workers and queens) and the unfertilised ones to
males (drones).
, 3 . Arrhenotoky. In this case the haploid egg does ndt undergo fertilization and gives
- rise only to male.
'4. Thelytoky. The unfertilised eggs give rise to only females.
Artificial parthenogenesis
I1
, .Inseveral cases, the eggs that normally develop only after fertilisation, can be
'
' experimentally induced to develop parthogenetically by certain treatments. This is
artifi~ig~artheno~enesis. For example, sea urchin eggs develop parthenogenetically if
they are subjected to certain temperatures, electric shocks, ultraviolet light, ether, alcohol,
lactic acid, mechanical stimuli like pricks etc.

I SAQ 5
1. Name the following:-
i) The kind of parthenogenesis in certain species that may or may not occur,
depending on environmental conditions.
i The kind of parthenogenesis that leads to the production of males only.

t 2. Name any three non-chordates in which parthenogeneis is very,common.

13.8 ALTERNATION OF GENERATIONS

b
The term alternation of generations (metagenesis) in animals refers to a phenomenon in
their life-histories, in which asexual and sexual generations alternate with each other.
-. Some well-known examples are given below:
Protozoa
I. A foraminiferan Elphidium (=PolysromeIla) found in shallow sea water occurs,in two
forms - macrospheric and microspheric. Both forms have a many chambered shell
(Fig. 13.33). The macrospheric form has a larger proloculum or initial chamber and a
large nucleus. The microspberic from has a smaller proloculum and numerous small
nuclei. The macrospheric individual represents the sexual phase and produces
flagellated anisogametes (haploid). Pairs of these anisogametes fuse to form a ,
zygote, which begins life with a small initial chamber but slowly grows a
multichambered shell and its single nucleus divides several times to produce
numerous small nuclei. This is an agamont. The full grown microspheric form then.
undergoes cytoplasmic divisions (asexual reproduction) and the resulting sirtgle-
Comparative Forms and
Functions
nucleated "amoebulae" or spores are liberated. Each of these establishes the initial
macrospheric chamber and subsequently grows into the macrospheric form. Thus
here, there is an alternation of macrospheric (sexual) phase with microspheric
(asexual) phase.

Fig.13.33 :Elphidium (a) macrospheric and (b) microspheric forms.

Macrospheric

Amoebulae (2n) SEXUAL Gametes(n)

Fig.1334: EIplridium, schematic life-history indicating alteqnation of sexual with asexual generation.
2. Malarial parasite Plasmodiium (Refer to Unit-2 Block-1) carries out its sexual cycle
inside the female mosquito starting with the micro- and macrogametes received from
human host. These then fuse together to produce the zygote. T h U t t e r grows and
undergoes sporogony (an asexual cycle) to produce spgrozoites. The sporozoftes are
released into human host through mosquito bite. These sporozoites first undergo
schizogony in liver cells to produce merozoites. Next these may undergo andther
schizogony inside the red blood cells. Id this way, fhe sexual cycle in the mosquito is
followed by two kinds of asexual cycles in human~host.
Cnidaria
The mature medusoid jellyfishes such as Aurelia are the sexual stages. Sperms and eggs
produced in the different (male and female) individuals ale released ir. Hater, which then
fuse to form a zygote. The latter develops into a fixed polypoid scyphistoma which
undergoes several transverse fission to produce ephyras by the process called strobilation
(see page lo), a kind of asexual reproduction. Thus the sexual generation of the adult
jellyfish is alternated with an asexual generation i.e., scyphistoma. You would recall from
unit 7 of this course that alternation of generation is quite common among hydrozoans, in
which the highly branching sedentary colonies like those of Obelia are asexual
generations. They give off medusaewhich are sexual generations. The medusae give rise
to fhe colonies and vice versa, Atemating with one another.

SAQ 6
Fill in the blanks:-
i) In Elphidium the macrpspheric individuals r~presentthe .................. phase
whereas the ~nicros~hkric'ones represent ............... phase.
ii) In Plasmodium sporogony is followed by,.... ........ .:. ........ '
iii) Strobilating .................. !.. represents the asexualphase in the life-history of
 Reproduction in
13.9 REPRODUCTION, LIFE CYCLES AND LARVAL Non-Cmrdates

FORMS
You are well aware by now that meiosis in metazoans takes place only during the
formation of haploid gametes. The diploid condition is restored with the formation of
zygote after fertilisation. However, from earlier sections you can see that a new diploid
individual is not always formed through sexual reproduction alone. Offspring of
invertebrates can also be formed without recombination of genetic material. Asexual
reproduction combined with the capacity for sexual reproduction leads to complex
patterns of life cycles.

What pattern of reproduction, is followed by a species (i.e. whether asexual, sexual or

through parthenogenesis), depends on the environment it lives in.

Planula -p Echinoidea
Pluteus

PLATYHELMINTHES

Muller

Brachiolaria

Trochophore W'
Nectochaeta

Holothuroidea Doliolaria

MOLLUSCA A
CRUSTACEA
I
- . -
Nauplius

Cirripedia ' Nauplius Cypris

Crabs and related Zoea Megalopa

groups

Fig 13.35: Characteristic larval stages of major marine invertebrate groups along with their adult forms.
Note that the adult forms are drawn to a much smaller seale than the larval forms.
Comparative Forms and "
For example,ma~ineinvertebrates discharge their gametes into the surrounding medium
Functions
where fertilisation may take place but freshwater and terrestrial invertebrates do not do
so. Externally fertilised gametes of invertebrates usually have little yolk or stored sources
of nutrition and develop into motile independent post-embryonic stage called the larva,
which is typically tiny and very different from the adult. The larvae may feed on an
external source of nutrition and are independent from their parents. They are usually the
means of wide dispersal for the species. Larvae never reproduce sexually, and often it is
impossible to establish the sex into which the larvae will grow as is the case with most
caterpillars of insects. After a period of time the larva settles down and undergoes
metamorphosis to attain adult form and life style. Such a life cycle with larval forms is
said to be indirect and is found in a majority of marine invertebrate species and
platyhelminths, annelids and arthropods. The major disadvantage of such a life cycle is
the high mortality of the planktonic eggs, embryos and larvae. Many marine invertebrates
have overcome this by depositing their eggs in protective envelopes and the hatching
occurs at larval stage (see Fig. 13.35). The larvae may be long lived, feeding on diatoms
and other minute organisms may be planktonic permiting adequate dispersal. The
majority of invertebrate phyla have external fertilisation and pelagic larvae. They exhibit
a pelagic benthic life cycle. This condition of having pelagic larvae is considered to be a
primitive trait.

Other species have dispensed with larvae altogether. The entire development takes place
inside the protective egg envelope and the young hatch as miniature adults. Fewer eggs
are deposited by such species and they contain large amounts of yolk. Such a
developmental pattern is called direct development and is characteristic of many marine
and most freshwater and terrestrial invertebrates. However, amongst terrestrial
invertebrates, most insects have a larval stage during development. Some important larval
forms have been described below.
Larval forms
Porifera
In the majority of sponges the larval stage develops within the body of the parent. The
larva is usually at the blastula stage of development. Figure 13.36 shows two kinds of
poriferan'lawal stages, amphiblastula and stereoblastula in which monociliated cells
cover the outer surface except the posterior pole. The interior of the larva commonly
contains most of the cell types found in the adult except choanocytes. The larva breaks
out of tkparents' body wall and has a brief free swimming existance.

Fig. 13.36: (a) amphiblastula of @con showing flagellated miromeres and non-flagellated macromeres.
-
(b) Demospongian larva stereoblastula.

Cnidaria
The common larval stage found in cnidarians is the planula which forms following
gastrulation. The planula is elongated and radially symmetrical with anterior and
posterior ends. The surface is covered by ciliated cells. A mouth may be present ,
sometimes at the posterior end of the larva and the larva atta~hesby its anterior end. The
planula may settle down to form a polypoid scyphistoma that ultimately farms the
medusa. In most hydrozoans the planula forms the actinula larva that transforms into the
medusa.
Platyhelminthes
Development in parasitic platyhelminthes is generally complicated with several free
swimming (Fig. 13.3 1) to parasitic larval fonns. The progression is from miracidium to
 sporocyst to redia and cercaria. You can refer to Unit 4 of Block 2 of this course for iife Reproduction in*
Non-Chordates
histories and larval forms of parasitic platyhelminths.
Polychaete and Moliusca
t?
The polychake have a trochophore larva. A typical trochorphore larva is top shaped
bearing a tuft of cilia at the apical end and a band or girdle of cilia called the prototroch
which rings the body about one third to one half the distance from the apical tuft. It is the
swimming organ and collects suspended food partitles. The gut is a complete tube and
mouth opens posterior to prototroch. Trochophore larva is characteristic of polychaetes
and molluscs (Fig. 13.35).
Echinodermata
Almost all larvae of echinoderms are planktonic (Fig. 13.35). Unlike the adults the larvae
do not show pentamerous symmetry and do not have tube feet. They move by bands of
cilia over the body and its projections. After a time of planktonic life a rudimentary
skeleton begins to form and the larvae sink to the bottom to develop the adult shape and
fom.
Arthropoda
1 Some of the planktonic larvae of crustaceans are shown in Fig. 13.35. The larvae of
crustaceans occupy different niches from their parents and there is no competition for
t food among the larvae and adult forms.

1 Moult

Moult

Spiracle

Moult

Moult

I
Moult and pupation

Moult and
metamorphosis

1 Metamorphosis

Aduk grasshopper
(a)

Fig. 13.37 : Life cycle of insec:ts. a) exopterygota, b) endopterygota.

Comparative Forms and
Functions
In the life cycles of insects we see that larval and adult stages have different patterns of
activities. Insects grow through a series of moults or instars. The simplest life history is
one in which there is a graduzl transition from embryonic to adult condition and both
larva and adult exploit the same food resource. These are the exopterygote insects and
their larvae have externally visible wing pads although, only the adults have hnctional
wings and the period of inactivity between the final moult and adult is not much longer
than that between the larval moults. This simple life cycle is seen'in grasshopper (Fig.
13.37 a).

More than 80% of living species of insects including Coleoptera (beetles), Lepidoptera
(butterflies and moths), Hymenoptera (wasps, bees and ants), Diptera (flies) are
endopterygotes. Their larvae include caterpillars, grubs and maggots. They never have
externally visible wing pads and are very different from the adult (Fig. 13.37 b) in form
and diet. There is a resting stage called pupa which may I@t from a few hours to many
months before the adult emerges from the final moult.

Thus we see that the different types of life cycles demonstrate the diverse sets of
reproductive strategies found among invertebrates. Sexual reproduction is the dominant
way of reproduction and often alternates with asexual reproduction in complex life cycles
seen in many cnidarians, flatworms, annelids, small crustaceans and some insects.
-

13.10 SUMMARY
In this Unit you have learnt that:
Reproduction ensures continuation of a species, generation after generation. Non-
chordates with their diverse body organisation and different ways of life present
equally diverse modes of reproduction. The most primitive and simplest of these is
binary fission, i.e.; just a kind of cell division. Some protozoans divide transversely
and some longitutjinally.Multiple fission in which the parent cell divides
simultaneously to produce numerous daughter individuals is also a kind of asexual
reproduction. Some animals undergo spontaneous fragmentation, each fragment
regenerating the missing part to make a h l l animal as in certain sea anemones and
some worms. Asexual reproduction commonly occurs by budding as in hydra and
some other cnidarians. Some non-chordates such as certain sponges reproduce
asexually by formation of gemmules under unfavorable conditions.

Regeneration is a very common phenomenon that in many non-chordates leads to the

formation of new individuals. It is thus closely associated with asexual reproduction.
Regeneration of missing parts following natural fragmentation or accidental breaking '
leading to forming of new individuals is very common. Most organisms exhibit
polarity in regeneration. Some non-chordates resort to autotomy-primarily in self
defence, and regeneration may lead to reproduction.

Sexual reproduction involves production of gametes that are usually distinguished as

the smaller, motile spermatozoa and the larger non-motile eggs. Gametes we
produced through meiosis, and fertilization restores the normal chromosome number.
The greatest advantage of sexual reproduction is the consequent mixing of genes that
lead to better adaptation and to evolution. Very often the two sexes are distinctly
different exhibiting sexual dimorphism. Sexual reproduction takes up various
patterns such as isogamy, anisogamy, conjugation, autogamy etc. The reproductive
organs include the gonads, gonoducts, accessory glands and certain structures that
facilitate copulation. Patterns of gonadal shape and disposition varies considerably.
Some non-chordates shed gametes in water (external fertilization), in some cases
sperms are released in the form of a spermatophore. The fertilised eggs may be
released (ovipary) or sometimes retained inside the female (vivipary, or ovovivipary)
till development is partial or hll, before being released.

Many non-chordates, specially sluggish and parasitic forms are hermaphrodites.

Hermaphroditism can either be simultaneous or protandrous. Many non-chordates
reproduce parthenogenetically.
 Asexual reproduction in many cases may lead to a generation of sexual individuals. Reproduction in
Non-Chordates
Often there may be an alternation of sexual and asexual generation in the life history
of animals. This is called alternation of generations or metagenesis.

Asexual reproduction combined with sexual reproduction leads to complex patterns

of life sycles. What pattern of reproduction is followed by an organism depends on
its environment. Marine invertebrates often have external fertilisation and one or
more larval stages very different from the adult. Such life cycles are called indirect.
Other species have dispensed with larval stages and have direct development. ~ i f e
cycles in insects have different patterns of development involving a series of molts or
instars.

13.11 TERMINAL QUESTIONS

1. List any six methods of asexual reproduction in non-chordates and writenot more
than two lines about each.
...................................................................................................

2. What are the advantages of asexual reproduction to the animals that practice it?
...................................................................................................

3.: Give any three examples of conspicuous sexual dimorphism in non-chordates.

.....................................................................................................

4. What is the advantage of sexual reproduction?

....................................................................................................

, 5. Describe the adaptive value of hermaphroditism.

6. Differentiate between:-
i) Obligatory and facultative parthenogenesis
...................................................................................................
Comparative Forms and
ii) Arrlienotoky and thelytoky.
Functions
...................................................................................................

7. Write briefly about the alternation of generations in malarial parasite.

...................................................................................................

8. Give,an important difference between the role of insect larvae and marine
invertebrate larvae in the life cycle of the species.
...................................................................................................

: *
13.12 ANSWERS

Self Assessment Questions

1. 1. I-f, 2-d, 3-e, 4-a, 5-c, 6-g, 7-b
11. Correct ones - (i), (iii) and (iv)

i) Central disc, arms

ii) Holothurians (sea cucumbers)
iii) Reunition
iv) Splittinglincising
v) Gemmules
Asexual reproduction is largely hssociated with regenerative capacity which is
stronger in lower animals in which the cells and tissues are less differentiated'
and specialised.

3. 1. Eggs are (i) nonmotile, (ii) larger, and (iii) usually laden with food material
2. 1-(d), 2-(a), 3-(e), 4-(b), 5-(c)
3. i) HeteronereislNereis
ii) Monocystis
iii) LoligolArgonauta
iv) Glossina (tsetse fly)

4.. 1. i) True, ii) False, iii) False, iv) True.

5. 1. i) Facultative
ii) Arrchenotoky
2. Aphids, honey bee, wasp, thrips, daphnia (fresh water fka)

6. i) sexual; asexual
ii) schizogomy
iii) scyphistoma
Ter*minalQuestions 'i Reproduction in
Non-Chordates
Six methods of asexual reproduction - Binary fission, multiple fission,
fragmentation, budding, strobiliation, formation of special bodies like gemmules.
Binary fission usually takes place in protozoans. These divides into two
longitudinally or transversely; in multiple fission the animal divides into more than
two. In fragmentation, a metazoan splits into two or more parts each of which
regenerates the missink part. Budding is a kind of outgrowth that break off from the
parent to grow into an adult. Strobilation is a serial buddinglcutting off of segments
from the parent. Special units like Gemmules are formed to tide over unfavourbale
4
environmental conditions.
Advantages of asexual reproduction - rapid multiplication accompanied by surer and
quicKer exploitation of 'favourable environment.
~ l o o d f i k e s- Male large and thick, with gynaecophoric canal to hold the long thin
female.
Ascaris - Male smaller with a curved tail and carrying a pair of penial setae.
Bonellia - Microscopic male living parasitically inside the nephridium of several
times larger female.
Advantage of sexual reproduction. Gametes from opposite sexes combine resulting
in mixing of genetic characteristics of male and female parents. This enables the
progeny to be better adaptated to changes in the environment. Sexual reproduction
tends to revitalise the population.
Hermaphroditism usually expresses itself in sessile, sluggish and parasitic species
that may otherwise have poor chances of meeting other individuals (opposite sexes).
With hermaphroditism, any two individuals would serve the purpose of reproduction.
i) Obligatory parthenogenesis is a condition where parthenogenesis, is the sole '

method of reproduction;facultative parthenogenesis is one in which the eggs,

may develop parthenogenetically or by fertilisation.
Malarial parasite shows two generations - (1) schizogony either in human RBCs or
liver cells or in both; (2) gamogony in mosquito. The sporozoites released into the
human body enter the liver cells first (pre-erythrocytic);the cryptomerozoites
released from liver cells enter RBCs ro carry out erythrocytic schizogony. These
merozoites may repeat the erythrocytic cycle, several times before they are taken in
by the mosquito. Inside the mosquito gamogony (sexual cycle) takes place. The
zygote subsequently passes through gamogony or sexual phase inside mosquito.
In case of insects, the adult is mobile and can disperse to greater distances than their
larvae; while in case of marine invertebrates the larvae are the main dispersal stages
as they are planktonic. The adult mollusc or echinoderm is relatively slow moving or
sedentary.
Comparative Forms and
Functions GLOSSARY
Aerobic respiration : Respiration involving the use of oxygen.
Afferent blood vessels : Blood vessels which carry deoxygenated blood to the respiratory
surface i.e., the gills.
Afferent nerve: A nerve which carries impulses from the receptor to the central nervous
system.
. Aorta: The main artery springing from the heart and leading blood away from the heart.
Arteriole: It is a fine artery formed after repeated branching of main artery.
Autotrophy : Capability of organisms to synthesise organic nutrients from simple
inorganic substances. All green plants are autotrophs.
Axon: It is one of the protoplasmic processes of a neuron which conducts impulses away
from the cell body.
Capillary: A tube of very small diameter. In the circulatory system, capillaries connect
the ends of the smallest arteries (arterioles) to those of smallest veins (venules).
Combplate One of the plates of fused cilia that are arranged in rows for ctenophore
locomotion.
Dendrite: The process of a neuron which conducts impulses towards the cell body.
Diffusion: It is a process in which molecules of gas or substance are transported from
high concentration tq low concentration.
Epipod, epipodite A lateral process on the protopod of a crustacean appendage, often ,
modified as a gill.
Exopod,exopodite Lateral branch of a biramous crustacean appendage.
Exopterygote Insect in which the wing buds develop externally during nymphal instar,
has hemimetabolous metamorphosis.
Extracellular digestion : Occurrence of digestion in the cavity of digestive system, the
cavity being lined by cells. The digestive enzymes are secreted into the cavity by the
I
secretory cells.
Flagellum Whiplike organ of locomotion.
Ganglion: A collection of nerve cell bodies.
Haematophagous feeders :Animals which feed on the blood vertebrates, especially
mammals.
Haemoglobin: An iron containing respiratory pigment concerned with the t~arnsports~of
oxygen by the blood.
Halter In diptera, small club shaped structure on each side of the metathbrax
representing the hindwings, believed to be sense organs for balancing, also called
balancer.
Heterotrophy :Refers to the dependency of organisms on other organisms for their
nutritional requirements. Heterotrophs can not syntehsise food on their own.
Hypertonic : Refers to a situation where the body fluid of an organism has a higher
osmotic pressure than the medium in which it lives. ,
Hypotonic :Refers to a situation where the body fluid of an organism has a lower ,
osmotic pressure than the medium in which an it lives.
Intracellular digestion : Occurrence of digestion within specified cells of the digestive
system; theses cells secrete the necessary enqmes for digestion.
Macrophagous feeders :Organisms which feed on large food particles.
Maxilla :One of the head appendages in arthropods.
Maxilliped :One of the pairs of head appendages located just posterior to the maxilla in
crustaceans, a thorasic appendage which has become'incorporated into the feeding
mouth parts.
Microphagous feeders : Organisms which feed on microscopic food particles.
Nematocysts : Stinging cells found jg-cnidarians and used for defense, capturing prey
and for anchorage.
Nerve: A bundle of nerve fibres.'
Nerve fibre: It is an elongated protoplasmic process of nerve cell.
Neurilemma (Neural lamella): It is the outermost codring of the nerve fidre.
Neurohaemal organ :A vascular structure which temporarily stores\ho@onal
substances secreted by neurosecretory cells before they are released intopfood.
Notopodium :Lobe of parapodium nearer the dorsal side in polychaete hnnelids.
Parapodium :One of the paired lateral pmcekses on each side of most segments in
polycjlaete annelids, variously modified for locomotion, respiration or feeding.
Pedipalps : Second pair of appendages of arachnids.
Plasma: It is a fluid part of the blood which is distinct from the blood cells corpuscles.~.
*
1 Pleopod :One of the swimming appendages on the abdomen of a crustacean. , Reproduction in
Non-Chordates
Respiration: A chemical activity taking place within the protoplasm of the cell an4
results in liberation of energy.
Saprozoic feeders :Animals which feed on dead and decaying organic materials.
Uricotelic :Refers to organisms that excrete uric acid as main excretory product of
metabolism.
Uropod :Posteriormost apppendage of many crustaceans.
Vein: The blood vessel which transports blood towards the heart from the capillaries.

Further Readings
General and Comparative Physiology, William S. Hoare, 1.983
Biology of Animals: A Text Book for Degree Students, B.B. Ganguli, A.K. Sinha
and S. Adhikari, New Central Book Agency, 1977.
The Invertebrates: Functions and Form, Irwine W. Sherman and Vilia G. Sherman,
Macmillan Publishing Co., Inc., 1976.
A Life of Invertebrates, W.D. Russell-Hunter, Macmillan Publishing Co., Inc., 1979.
Text-Book of Zoology: Invertebrates, Ed. by A.J. Marshall and W.O. Williams (The
edition of A Text Book of Z,ooIogy Vol. 1 by J. Jeffery Parker and William A.
Haswell), The Macmillan Press Ltd., 1972.
Invertebrate structure and Function. E.J.W. Barrington, ELBS, 1979.
-
Barnes, R.D. (1980) Invertebrate Zoology. Holt Saunders, Tokyo.
Barrington, E.J.W. (1979) Invertebrate structure and function (Chapters 14,15 and
16). ELBS & Van Nostrand Reinhold (UK), 2nd Ed.
Ramsay, J.A. (1968) Physiological approach to the lower animals, Cambridge
University Press.
NOTES
UNIT 14 ADAPTIVE RADIATIONS
Structure
Introduction
Objectives
Solitary and Colonial Forms
Colonial Forms Among Protozoans
Colonial Forms in Metazoans
Adaptive Radiations
Adaptive Radiation in Annelida
Adaptive Radiation in Arthropoda
Adaptive Radiation in Mollusca
Flight in Insects
Migration in Insects
Summary
Terminal Questions
Answers

14.1 INTRODUCTION
--- - - - --

You have read in our course on Taxonomy and Evolution (LSE-07) that living organisms
are products of evolution. In simple words evolution may be defined as 'descent with
change'. Here change is the basic factor. Evolution may be in the nature of the
environment or in the form and fhnction of the organism. The nature of the environment
has a strong bearing on the form and function of the organism. You will see in Section
1.3 of this course (Unit 1, Block]) that the organisms always tend to establish harmony
with their environment. This process, called adaptation, enables the organism to face the
vagaries of the everchanging environment. Adaptation actually sums up the whole result
of evolution.

You might have noticed that when organisms belonging to different groups come to
occupy similar environment, they develop striking similarities in structure and.behaviour,
giving a false impression of closer relationship. For example, fishes and whales which
belong actually to different classes (Pisces and Mammalia, respectively) are found in
similar habitat i.e., they are both aquatic. They look so alike that for a layman both are
'fishes'. This condition where organisms belonging to different groups adapt themselves
to the same environment and look and act alike, is called adaptive convergence.
C

On the other hand, there are situations where organisms belonging to the same or closely
related groups may occupy different environments due to which they develop varied
adaptations. It gives rise to diverse evolutionary lines. You know that house- lizard,
snake, tortoise and crocodile belong to the same class, Reptilih But they look so different
from each other that one may be tempted to place them in separate classes. This situation
in which animals belonging to same or closely relqted groups occupy different habitats
and acquire different functional adaptations, is cafled adaptive divergence or adaptive
radiation. In this unit you will study how adaptive radiations have evolved in different
non-chordate groups.

Objectives
After studying this unit you should bd able to:
distinguish between solitary and colonial forms of animals and explain the needs for
the evolution of true colonies in lower non-chordate groups,
differentiate behveen adaptive convergence and adaptive divergence or adaptive
*-
radiation and identify the different ways in which adaptive radiations have occurred
in Annelida, Arthropoda and Mollusca,
.
describe the structure of insect wing and explain the mechanism of fligMm insects,
describe the meaning, process and significance of migration in insects.

114.2 SOLITARY AND COLONIAL FORMS

4nimals may lead their lives either as individuals or in groups. When they exist as
t
iiidlviduals, they are called solitary, but if they live in organised colonies, we name then
Adaptation and Bchaviyral colonial. Colonies are a form of intraspecific association in which the interests of an
mtern individual are subordinate to those of the whole group. In true colonies the individuals are
organically connected together by living matter or through material secreted by them. The
degree and extent of closeness among individuals in a colony may vary considerably.
True cqlonies are found only in primitive groups with simple organisation, such as
protozoans and coelenterates. In sponges it is difficult to ascertain whether the branched
aninial is an individual or a colony. Colonial forms mostly reproduce asexually. Actually
the colony results due to the failure of the individuals to separate. Each individual in a
colony is called a zooid.

14.2.1 Colonial Forms among Protozoans

Many well known colonial form9 occur among protozoans. Simplest colony formation is
seen in Choanoflagellates, which have a collar around the base of the flagellum (hence
the name). There are a Few zooids in a colony as for example, in Codosiga (Fig. 14.1 a).
Each zooid leads an independent life though remaining attached to a common stalk. The
volvocales form more complex coloni8s. Fbr example, Gonium form!+plate like colonies
of 4-16 individuals; Pandorina forms spherical colony of 16 individuals; Eudorina is a
spherical colony of 32 zooids arranged on the surface, Pleodorina has 128 zooids. An
advanced form of colony is seen in Volvox,which is a plant-like mastigophore, or
phytoflagellate. In this colony thousands of individuals remain embedded (Fig. 14.1 b) on
the surface of a spherical jelly-like substance secreted by the zooids. Here also each zooid
leads an independent existence except for a co-ordinated flagellar movement which helps
in swimming. There is also connection among zooids by protoplasmic threads (Fig. 14.1).
It is of interest to know that these colonies always swim with a particular side forward
i.e., they possess polarity, an attribute of fundamental importance for colonial existence.
In Volvox and Pbodorina some sort of division of labour is also noticed. The anterior
zooids do not reproduce while those elsewhere are reproductive some of which become

Somatic zooids

Reproductive

(a) (b)

Fig. 14.1: a) Choanoflagellates :the simplest protozoan colony codosiga b) Volvox: an advanced colony.

Many ciliates like Epistylis and Zoothamnium also form colonies. Each of these colonies
has bell-shaped zooids united by their stalks (Fig. 14.2 a and b) to a common stem. In
Epistylis all zooids are alike while Zoothamnium shows polymorphism in zooid structure.
There are four types of zooids in the colony (Fig. 14.2 b) a single terminal macrozooid
which transforms into macro- conjugant; median axillary microzooids that can become
migratory ciliospores; terminal branch microzooids which can form microconjugants and
vegetative microzooids that can transform either into ciliospores or microconjugants.

~ b let
w us examine the advantages of the colonial life in protozoans. Most of the
protozoan colonies are autotrophic i.e. they obtain their food in a plant-like manner by
photosynthesis. Therefore no apparent nutritive advantage is provided by the colonial
way of existence. Then what are the benefits? We can observe following advantages:
 Adaptive Radiation
1. Association of individuals in a jelly-like ground substance, especially when
iI interconnected by protoplasmic strands, facilitates transmission of nutrients.
2. Combined flagellar activity of many zooids gives locomotory advantage.
3. Most of the colonies being spherical, minimum surface area for a given volume of
1 sphere is exposed to the surrounding water, due to which the resistance offered by
water is least.
4. In colonies of ciliates the group association provides protection and more efficient
exploitation of food-supplies.
Before we pass on to colonial life in metazoans, let us see what you have learnt so far.

e,~.:. e r m i n a macrozooid
l

Terminal branch
microzooids

Myoneme,m ;$ ~ e d i a naunillary
microzooids

Fig. 14.2: a) EpisWL :Colony having similir zooids, b) Zoothamniurn :a polymorphic protozoan colony.

SAQ 1
i) Fill in the blanks using the correct word given in parenthesis below:
(environment, different, alike, adaptive convergence, adaptive divergence)
a) The condition where organisms belonging to widely different phylogenetic
groups adapt themselves to the same ................................
and look and act
alike is called .........................................
b) When animals belonging to phylogenetically closely related groups occupy
different habitats and acquire different adaptive modification, it is called
....................................................
I ii) Indicate whether the following.statementsare true (T) or false (F):
I a) In true colonies rooids a?e not ?rganically connected with each other.
b) True colonies are found only in+organismswith comparatively simple
organisation. .
C) Colonies in Carchesium show polymorphism.
d) Colonial life provides nutritive advantage in all colonial forms.

1 143.2 colonial POTS in Metazoans

Among metazoans, true colonial fornis are met with in coelenterates, though the term
'colony' is often used in relation to sponges and insects, also. In sponges the 'colony' is
arbitrarily defined on the basis of the number-ofoscula. OJle osculum-one-individual is
Adaptation and Bchavloural the rule. However, this definition is more a matter of convenience. In insects the word
Pattern
'colony' is employed to denote the complex societies that are fo ed by them, as in the
case of honey bees, ants, termites etc. But these social groups strictly fall within
the definition of the term colony.

Growing point
I
Gonangium
(gonozooid)

Fig. 143 ;a) Obetia, the hydroid colony (monopodial colony). (b) Plumularia colony showing monopodial
growth. (c) Sympodial growth ofHdecium colony. (d) Hydraciinia colony where polyps grow .
directly from the mat of stolons.
I Coelenterate Colonies
'Adaptive Radiation

Members of the class Hydrozoa and Anthozoa form true colonies. Zooids with well-
defined individuality are present in a colony, the shape and pattern of which varies from
species to species. Floating colonies are formed by siphonophores. In the case of
sedentary hydrozoan colonies, the colonies are attached to the substratum by living
thread-like horizontal stolon or hydrorhiza (Fig. 14.3 a). From the stolon arise vertical
branching stems called hydrocauli. Each hydrocaulus gives off lateral branches, which in
their own turn bear branches of the third order. Zooids are borne on these branches (Fig.
14.3 a).

The branches and .zooids in a colony are connected by a living coenosarc, over which a
non-living horny perisarc is present. The coelen'teroncontinues throughout. A zooid is
cylindrical or umbrella-shaped, diploblastic structure with a body cavity or coelenteron
opening to the exterior by a mouth situated on hypostome. A circlet of tentacles is usually
present around the hypostome. In some cases the perisarc forms a cup like or capsule-like
covering, variously named hydrotheca or gonotheca, around the zooids. The zooids
covered with theca are called thecate; those without a thecal covering are naked ones or
athecate.

The growth of the hydrocaulus is of two types viz. monopodial and sympodial'(Fig. 14.3
b and 14.3 c). In the former the main axis maintains a single line of growth i.e., each
branch ends in a permanent terminal zooid which is oldest of that branch. Below the base
of this tetminal zooid is the zone of growth. This zone lerigthens the branch. The growth
zone may also give rise to lateral buds, which may elongate in their turn. In the sympodial
growth the primary polyp does not continue to elongate. But itproduces lateral polyp.
This also stops growth, but produces lateral buds again. Here the terminal polyp is the
youngest; The main axis is formed by the combined hydrocauli of many polyps (eg.
Halecium). In some forms like Hydractinia (Fig. 14.3 d) zooids arise directly from the
stolon in an irregular fashion.

Polymorphism is another characteristic feature of coelenterate colony. The zooids exist in

.many forms and show division of labour. ~ i u a lsome
l ~ zooids are concerned with
feeding; these are called gastrozooids. The blastozooids or gonozooids undertake
reproduction, budding off medusae which are another type of zooids, while
dactylozooids are meant

..
i medusa
(b)

I
(a) .
Fig. 14.4 :a) A typical submergent siphonophore Muggiaea; b) Part of another siphonophore Physalia
colony showing the zooid polymorphism.
Adaptation and Behavioural The saucer-shaped zooids called medusae produce gametes and reproduce sexually. YOU
Pattern may recall here the siphonophores and other hydrozoan colonies which you have already
studied in previous unit. Siphonophora shows extreme case of polymorphism, for
exqmple, Muggiaea and Physafia (Fig 14.4). The group includes pelagic zooids. The
colony comprises gastrozooids, dactylozooids and gonozooids. The medusae remain
attached to the parents and modify to form gonophores (reproductive zooids),
nectophores (locomotory zooids) and gas- filled pneumatophores (floating zooids).

In one of the previous units dealing with coelenterates you have studied in detail, corals
and coral reefs. You may recall that section here. The Hydrocorallina (the hydrocorals;
eg. Stylaster, Miflepora) (Fig. 14.5) are colonial, polypoid, hydroid corals which may
attain considerable size contributing to coral formation. Their skeleton is calcareous,
internal and epidermal. Though not accompanied by polymorphism, many anthozoans are
also Colonial, eg. Palythoa.

/ Mouth

~astrivascularcavity
(c)

Fig. 14.5 :Hydrocoral MiUepm; a) The colony, b) part of the colony magnified showing the pores, c)
polyp ofMUqwra emerging froin the porcs in the ekeleton.
 Adaptive Radlrtkn
), I

Fig. 14.6 : Part of the coral colony showing contracted and expanded polyps.

Many true corals, though not all, are also colonial. Conspicuous among the colonial
anthozoans are in fact many coral formers. The colony-forming scleractinian corals or
stony corals, also known as hexacorals and madreporerial corals (Fig. 14.6), include
Astrangia, Montastrea and the brain coral. Here the polyps are interconnected by
horizontal connections. The skeleton is epidermal and external (Fig. 14.7). There are
solitary corals also among them, like Fungia (Fig. 14.8). The colony-forming octocorals
(alcyonarians) among anthozoans include th: common sea pens, sea rods, sea pansies, sea
fans, whip coral, pipe corals etc. (Fig. 14.9). The arneobocytes secrete their skeleton
which supports the colony. The skeleton of these octocorals is therefore internal and is
part of the tissue.

From this account you will see that corals have developed from different groups of
coelenterates. This is convergence in their adaptation.

ieptum

plate

Fig. 14.7: Vertical section of the coral polyp in its theca.

Adaptation and Behavioural
Pattern :

Fig. 14.8: Fungia.

(c) ((1) .

Fig. 14.9: a) Sea rods; b) Sea fan; c) Organ-pipe coral; d) Sea pansy;
 -

Adaptive Radiation
Thds the benefit of colonial life is also clear: competition for survival among individuals
is replaced by co-operation. This provides distinct nutritive, protective and reproductive
advantages to them.

SAQ 2
i) Match the words in list A with those in list B.
List A List B
1) Dactylozooid a. Feeding
2) Gastrozooid b. Floating
3) Pneumatophore c.. Protection
4) Medusa d. Sexual Reproduction

I . ii) Fill in the blanks in the following statements on the basis of what you have studied
irl- subsection 14.2.2.

k
The growth of the hydrocaulus is of two types viz.., ....................................
or
.....................................In the former each branch ends in an oldest terminal
I zooid, below the base of which is the zone of growth that lengthens the branch and
I gives rise to buds.

14.3 ADAPTIVE RADIATIONS

In the beginning of this unit, you have studied that when the animals belonging to the
same or closely related groups adapt for different modes of life, they are said to show
I adaptive divergence or adaptive radiation. This concept was known to early zoologists
I like Lamarck and Darwin. Larnarck called it embranchment and Darwin called it
divergence. The idea was, however, concretized in the form of a law by Osborn, the law
of adaptive radiation. Though Osborn postulated the law of adaptive radiation on the
basis of his studies on mammals, it applies equally well to other animal groups, whether
non-chordates or chordates. There is always a brief period in the history of all animal
groups when the rate of evolution is very rapid. It results in the emergence of many new
major lines of evolution or adaptation. Further evolution'of each of these lines is
comparatively slow. This process of breaking up of the parent stock into diverse lines
which continue their own evolution is adative radiation.

Two fundamental needs of the organisms are mainly responsible for adaptive radiation.
These are need for food and need for safety. A major factor in the animal evolution has
been their habit of food gathering. Two basic ways of this habit may be recognised. There
were animals which would just wait for the food to come their way. Secondly, some
animals would actively seek the food and go after it. The former acquired sedentary
habits and radial symmetry. Radial symmetry would'atlow the fixed animals access to
food in all directions. These animals evolved a special feeding device called filter-
feeding. On the contrary, a typical food- seeker has to resort to movement for food
gathering. They would go after the food. This gave rise to antero-posterior polarity, -
cephalisation, and bilateral symmetry. The anterior end may have developed into a
distinct head, the seat of mouth-parts, sense organs and brain. (see earlier unit).

The second major factor in animal evolution has been the need for safety from the hostile
abiotic and biotic components of the environment. All this has aIso led to exploitation of
new habitats and acquisition of appropriate adaptations to suit the new surroundings.
These modifications may have been in the habits, morphology and physiology of the
animals. The evolutionary history of various lines or animal groups show that functional
adaptations in different groups are different. Therefore you will find different lines of
adaptive radiations in different groups of animals.

14.3.1 Adaptive Radiation in.Annelida

Phylum Annelida includes three classes; Polychaeta, Oligochaeta and Hirudinea. Of
these, the Polychaeta do not h?ve clitellum; the Oligochaeta and Hirudinaria are
clitellate. Now you should recollect the classification and characters of Annelida which
you have studied in Unit 4, Block 1 of this course. The early annelids are supposed to
I have been marine worms burrowing in the'bottom, in sand and mud on the shore.
I Adaptation and &havioural Polychaetes comprise the marine species which continued their life in the sea diversifying
Pattern
into various niches there; Oligochaeta include the line which led to the fresh water forms
and to the earth worms; Hirudinea includes the leeches which arose from some fresh
water oligochaetes.

Adaptive Radiation in, Polychaeta

Having evolved from some small, annelid worm-like creatures adapted for burrowing and
crawling life in oceans, the group diverged into two main branches on the basis of their
food habits; a group of active food seekers and another line of sedentary animals. The
former actively sought after the food either scavenging or preying upon it whereas the
laver gave rise mainly to burrowing or tubicolous forms. In this section we will discuss
the polychaete groups adapted to various modes of locomotion, habitation and nutrition.

Errant Polychaetes: Worms such as nereids or hesionids are rapid crawling worms that
crawl beneath stones and shells in rock and coral crevices and among algae and sessile
animals. In these worms the head is well dt <elopedhaving one to four pairs of eyes, upto
five antennae and a pair of palps (Fig. 14.10).

Antenna

Prosto~niuin

Pel.istomial cirri

Pcristomium

Dorsal vessel

Fig. 14.10: The herd region of the polychaeteNereis.

These worms have large parapodia that help in crawling. Many of these polychaetes are
crhivores and feed on small vertebrates including other polychaetes. Food habits like
scavenging, algae eating and detritus feeding have also evolved in some of these
polychaetes. The pharynx of the errant polychaetes is muscular and eversible and possess
teeth or jaws which vary in number in different families (Fig. 14.1 1).

Fig. 14.11: The head of the polychaete Nereh with everted pharynx (lateral view) the jaw is open when
pharynx is everted and closed when pharynx is retracted.

Pelagic Polychaetes: Certain families of polychaetes are adapted to life in oceans*and

thus are pelagic or planktonic. In these worms head is well developed and parapodla are
large that are used as paddles to aid in swimming. Like other planktonic animals these
polychaetes are pale or transparent. These worms are generally carnivores, e.g.
Rhynchonerella angelina (Fig. 14.12), Tomopteris renata.

Gallery Dwellers: These polychaetes are adapted to live in burrows made of sand or
mud. The gallery dwellers make extensive burrow system or galleries that open to the
surface at many points (Fig. 14.13a). These burrows are lined by the mucus secreted by
the worms that prevent the collapsing of burrows. The prostomium of these worms can be
a simple lobe or of conical shape and lack the eyes and other sensory organs (Fig. 14.13
b).
 Adaptive Radiation

Fig. 14.12: Pelagic polychaete Rhynchonerella angelina (a) and its parapodium (b).

Fig. 14.13: a j Burrow system of a gallery dweller worm GIycem olbo showing the worm lying in wait for
the prey. b) Anterior end of the gallery dweller Drilonereis showing the conical prostomium
that lacks eyes and sensory appendages.

The worms usually move in the burrows by peristaltic movements, and the parapodia are
reduced and help to anchor the segments by gripping the walls of the burrow. The septa
and circular muscles are well developed. However, some of the gallery dwellers may
crawl with the help of parapodia. They may be chrnivorous or non- selective d$posit
feeders and consume the substratum through which the burrows are made.

Glycera, is the best studied gallery dwelling polychaete and is also used as fishing bait.
These worms lie in wait in the gallery system and when the prey moves across the surface
creating pressure waves, the worm moves to a nearby opening. Blood worms have a long
proboscis that is shot out with explosive force to seize the prey (Fig. 14.14).

Sedentary Burrowers: Certain polythaetes make simple burrows that have only one or
two openings to the outside (Fig. 14.15 b). These worms move about very little when the
move, they also go by peristaltic contractions only. Parapodium is reduced to hook like
setae that help in gripping the burrow wall. The prostomium is devoid of most of the
sensory structures. However, special feeding appendages may be present. Some of the
sedentary burrowers are nonselective.deposit feeders while others are selective deposit
feeders. L-ug worms (Arenicola) (Fig. 14.15 a) nonselective deposit feeders, live in L-'
shaped burrows and ingest the sand at the bottom with an eversble pharynx. At fixed
intervals the worm comes out of its burrow and defecates at the surface as a casting. After
this the worm resumes feeding and ventilating. Ventilation occurs because peristaltic
contractions bring in the water current.
=-,
Adaptation rod Behavieural
Pattern

Fig. 14.14: Anterior end of Glycem showing the everted pharynx. .

Peristomiurn, Prostomium

Not

Gill

Fig. 14.15: The Lug worm (ArenlcoIa) with its pharynx everted (a). Worm in the burrow (b)
In selective deposit feeders there is no eversible pharynx. Instead they have special head
structures that pick bp the organic matter from the surrounding sand grains. For example
Amphitrite has a great mass of long tentacles that spread over the surface from the
 Adaptive Radihtion
opening of the burrow (Fig. 14.16). The detritus material adheres to the mucus'on the
tentacles and is then passed on to the mouth in a ciliated tentacular gutter with'the help of
tentacular contraction.
Tentacles

Fig. 14.16: Anphitrite in its burrow with outstretched tentacles on the surface. Also shown is.the food
particles trapped in the tentaclesthat are rolled up to form a ciliary gutter.

Tube-Dwelling Polychaetes: Tube dwelling is more widespread habit among the

polychaetes as compared to other animal groups. The worms can make the tubes in the
sand or in firm and exposed substrates such as algae, rock, coral or shell. The tube may be
completely made up of hardened material secreted by the worm or composed of foreign
material cemented together. Thus, a tube will remain intact ylhen dug out of the sand.

Mouth

Fig. 14.17: Bamboo worm that lives upside dawn in the tube.
Adaptation and Behavioural Tube dweller (tubicolous) polychaetes show structural diversity that i s correlated with
Pattern
their different modes o f feeding. Majority of the tube-dwellers are sedentary and move
about within the tube lazily with the help o f peristaltic contractions. They lBck sensory
structures, although feeding appendages may be present. Parapodia are reduced to ridges
with hooked setae for gripping. lnfact these adaptations are similar to those for sedentary
burrowers as these two habitations are similar to some extent. Some families contain both
burrow dwelling and tubi~colousspecies.

The nonselective deposit feeders like bamboo worms (Clymenella, Axiothella) live in
sand grain tube by keeping their head down and. ingest the substratum at the bottom o f the
tube with the help o f eversible pharynx (Fig. 14.17). Periodically the worm comes back to
the surface and defecates.

Filter feeding is another mode o f nutrition that has evolved in several families o f
sedentary and tube dwelling polychaetes. One kind o f filter feeding i s seen in fan worms
or feather duster worms, where the prostomium palps have developed to form a funnel -
shaped or spiral crown consisting o f pinnate processes called radioles, for example in
Sahella (Fig. 1'4.18). During feeding the particles are first trapped in the mucus o f the
radioles surface and then passed on to the mouth with the help o f cilia. When the worm
pulls back its anterior end into the free end o f the tube, the radioles are rolled and closed
up together.

Fig. 14.18: Filter feeding in fanworrn Sabella (a) Water current passing through rrdioles. (b) Water
current and ciliary tract over a part of mdiole.

Chaetopterus exhibits another mode of filter feeding. It feeds by filtering water through a
mucus bag. These worms live in U- shaped tube made by secreted parchment like
material (Fig. 14.19). In the middle o f the body of the worm there are three piston like or
'fan' like parapodia thatdrive water through the tube. A pair o f long winglike anterior
notopodia secrete a film of mucus that i s rolled up into bag like shape with the help o f
cilia. The water current driven in the tube passes through these mucus films. Periodically
 Adaptive Radiation
the mucus secretion is stopped and the mucus bag containing trapped food is rolled up
into a ball which is passed along the ciliated groove to the mouth.

Fig. 14.13: Chaetoprerus in its tube (lateral vlew).

'
Till now you have studied about the adaptations in annelida. Try the following SAQ
before we proceed further.

SAQ 3
i) Mark ( J )the correct alternatives in the following statements :
a) Law of adaptive radiation was postulated by LamarcklOsborn.
b) Adaptive divergence was called 'embranchment' by DarwinlLamarck.
c) Warms with well developed head having eyes, antennae and a pair of palp are
gallery dweller polychaeteslerrant polychaetes.
d) Large parapodia in polychaetes that are used as paddles are adaptations to life in '
oceanslsand and mud.

i) State whether the following statements are true (T) or false (F).
a) Sedentary burrower Polychactes are essentially carnivores.
b) Filter-feeding is mostly found in sedentary tube-dwelling polychaeta.
c) Polychaets are mainly fresh water animals.
d) Sedentary and tube-dwelling polychaetes mostly lack well-developed sensory
organs.

ii) Name the two fundamental needs responsible for adaptive radiation.

Adaptive Radiation in Clitellate Annklida

Polychaeta, being aquatic, lay their eggs in water and most of them develop through a
trochophore larval stage. Aquatic environment offers many advantages: animals require
less energy expenditure for supporting their body weight, water acts as a cushioned
envelope providing protection from jerks and strains, temperature gradient in water is
least and it provides an ideal medium for egg-laying and development. These advantages
are available to marine as well as fresh-water forms. However, fresh water habitats have
one big drawback. These are not permanent like marine habitats. They may dry up during
some part of the year. Therefore the fresh water forms and terrestrial ones face a two fold
problem to different extent. One, they are denied the benefits of permanent and .
continuous access to the aquatic medium and secondly, they are exposed to the risk of
Adaptation and Behwioural desiccation. This problem has been solved by clitellate annelids successfully. Let us
Pettern
examine how. .

The clitellum-bearing Annelida, the Oligochaeta and Hirudinea, mostly inhabit fresh
water and terrestrial habitats. They have no larval stage and the development is direct.
Glandular cells of some of their body segments become active during breeding season
and form a conspicuous belt-like clitellum (Fig. 14.20) which produces a cocoon. Eggs
are laid and develop within these cocoons. The clitellum may be permanent (as in
earthworms) or temporary (as in leeches). These annelids are hermaphrodites, though
reciprocal.copulation in earthworms is well known. The clitellates are devoid of antennae,
palpi or parapodia.

Prostomiurn Male gonopore . Genital chaetae

Fig. 14.20: Earthworm showing clitellum

Oligochaeta appear to have evolved directly from the marine annelids, independently of
polycheates. Some species of tubulificids and enchytraeids especially from littoral and
intertidal zones and estuaries have been reported from marine waters.

Fig. 14.21: T u b w that lives head down in long tubes. The posterior end waves about in water to facilitate
gas exchange.

-
Fig. 14.22: (a) Aedosoma the cilia around the mouth sweep in the food particles. (b) Slylaria -
prostornium drawn out into long snout.
 Adaptive Radiqtion
Tubifex (Fig. 14.21) and Limnodrilus are reported to thrive well in sewage-polluted
waters. Tubifex which lives in stagnant water in mud, builds tubes. It projects posterior
part of its body from the tube and waves it about in water facilitating gas exchange. Some
like Aeolosoma and ~tyiaria(Fig. 14.22) are fresh water forms. Aulophorus canstructs
tubes in mud and debris. Aquatic forms like Dero (Fig. 14.23) and Aulophorus have
finger like gills at the posterior end. Branchiura and Branchiodiilus have filamentous
gills on the body. Aquatic forms are generally small; but they have longer setae. Some
like enchytraeids are transitional between aquatic and terrestrial habitat, and live in
marshes. These include lumbricids, megascolecids and moniligastrids. Earthworms are
burrowing animals and are known to increase land fertility.

Fig. 14.23: Dero has ciliated analI gills.

.sucker

sucker

Flg, 14.24: a) Giossiphoniid I m h Glossbhoniaconplantua; b) piscicolid leech Cysobmnchus; c)

hirudinid leech Him& me&cinu& d) haemadipsid I m h Huamdlpsa.
.Hirudinea includes leeches. Many of the leeches are ectoparasites of vertebrates
[glossiphoniids, piscicolids, hirudinids, haemadipsids (Fig. 14.24)]. The parasitic
Adaptation and Behavioural
adaptations of the leeches are: presence of sucking pharynx and a post-anal sucker,
Pattern
provision for the secretion of mucus layer over the body by the skin glands to prevent
dehydration, secretion of an anticoagulant hirudin to facilitate feeding on blood, and
provision of food storage in the spacious crop. One full meal by a leech may last it for
about four months. Leeches and oligochaetes have a common ancestry. Leeches are
mostly fresh water animals. However, some have become adapted to terrestrial life
(Haemadipsa). Some have also become secondarily adapted to marine habitat.
. .
SAQ 5
i) Indicate whether the following statements are correct or not.
a) All members of the phylum Annelida develop through a larval stage called
trochophore.
b) Clitellum is a well developed permanant structure in eanhworms.
c) The skin of leeches contains gland cells which secrete a mucus layer over the
body which prevents desiccation.
d) Tubifx is very comfortable in ponds polluted with sewage.

I
14.3.2 Adaptive Radiation in Arthropoda
i
~ i o l o ~ i c aArthropoda
ll~ is the most successful group in terms of numerical strength,
adaptive diversity and extent of territorial distribution. They are supposed to be I
polyphyletic in origin, with a number of (three or four) independent lines of evolution. I
There is also a tendency among them for reduction of number of segments by fusion or
loss. Arthropoda, literally meaning joint- footed (Gr. arthros =joint; podos = foot), is
i
represented by horse-shoe crabs (Subclass Xiphosura), prawn, lobsters, crabs (Subphylum I
Crustacea), spiders and scorpions (class Arachnida), centipedes (class Chilopoda),
millipedes (class Diplopoda) and insects (Class Insecta). Formerly Peripatus
(Onychophora) also used to be included in this phylum. Arthropods have certainly
evolved from marine Annelida by acquiring an armour of chitin over the body and paired
appendages on almost all body segments. The armour not only provided suppoi and
pt'otection to the animal but also pr&ented entry of excess water and salts in the body in
aquatic forms and desiccation in terrestrial ones. But it interfered with smooth gas
exchange through the body surface and hampered growth. The adaptive radiation in
Arthropoda is mainly related to the evolution of suitable respiratory mechanism,
appropriate limb modifications and flight.

Respiration
The annelids respire by the general body surface or by gills. In Arthropoda while gills are
retained by aquatic forms, terrestrial members of the group have evolved book-lungs
(scorpions) and tracheae (centipedes, milipedes and insects). Xiphosura and Crustacea
are almost exclusively aquatic. They evolved in water and remained there. There are also
sqme water-dwelling arthropods, which had actually invaded land and acquired terrestrial
adaptations. 'They re-entered aquatic medium and made it a second home. This includes
many adult insects (water-bugs, water-beetles, etc), which respire by tracheae while
living in water. Thus, aquatic respiration in Arthropoda may be by gills (usually called
branchial respiration) or by tracheae (tracheal respiration).

Aquatic Respiration by Gills

In Xiphosura, Crustacea and many larval insects respiration takes place by gills. A gill is
a vascular outgrowth of the bodywall. It remains bathed in water and gaseous exchange
occurs on its surface. In Limulus (Xiphosura) five pairs of book-gills are present on the
ventral surface (Fig. 14.25). These are flat, lamellate abdominal appendages. Each gill
supports about 150 gill lamellae arranged in a manner which gives it an appearance of the
leaves of a book, hence the name.

In Crustacea the gills or branchiae are arranged as lateral extensions along the central
axis. Gills may be of three types: Phyllobran~hsare simple, leaf-like lobes set on either
side of a main axis (Fig. 14.26 a), Triochobranchs have filaments arranged around a
central axis (Fig: .14.26 b) and dendrobranchs are modified phyllobranchs with each
lateral lobe being subdivided (Fig. 14.26 c). Gills have a supply of haemocoelomic
 Adoptive Radiation
channels. A continu~ussupply of water is maintained in the gill chamber. This ensures
proper gas exchange.

Gill
Gill

Fig. 14.25: Ventral view of horseshoe crab ~ & f ~ s p ~ & ~ h ~ m u s .

Blood channels
Blood channe'ls

Blood channels

Fig. 14.26: Gill types i n crustace8ns.h) Phyllobranch, b) Triochobranch. c) Dendrobranch, (i) Lateral '
view (ii) Transverse view.
Adaptation and Behavioural Aquatic Respiration in Insects
Pattern
Two modes of aquatic respiration in insects are recognised. In one, the insects obtain .
oxygen dissolved in water. This may be affected either through the general body surface
or by means of different types of gills. Tracheal gills of the aquatic larvae of mayfly,
stonefly and caddisfly are the lateral outgrowths of the bodywall and contain tracheal
branches. Rectal gills are present in the rectum of Odonata larvae. The stonefly larvae
(Fig. 14.27) possess tracheal gills on various regions of the body and analgills on each
side of the anus. The blood-gills of some dipteran larvae are blood-filled out-growths of
the bodywall. The blood'of Chironomus larvae is red due to haemoglobin, giving the
name blood-worm to the larvae.

gills

Fig. 14.27: Stoncfly larva showing tracheal and a n h ~gills.

There are some insects which live in water but still breathe air. They have devised
modifications to obtain supply of fresh air at periodic intervals. We will discuss more
about these forms when we deal with respiration in terrestrial'insects.

Respiration in Terrestrial Arthropoda

In land-dwelling Arthropoda respiration is effected either by book-lungs (Arachnids) or
by tracheae (Myriapoda and Insecta). The book-lungs seem to have been modified from
the book-gills of the ancestral arachnids, the Merostomata. Scorpions possess four pairs
of book-lungs, one inside each mesosomatic segments 3-6 (Fig. 14.28 a). A book-lung
has a ventral atrial chamber and a dorsal pulmonary chamber (Fig. 14.28 b). The
former opens to the exterior by stigma (plural-stigmata) and the latter contains about 150
vertically placed lamellae giving the whole structure an appearance of the leaves of a
book that explains the nomenclature. In spiders the respiratory organs may be either
primary book-lungs or some modified structures such as tube- trachea and sieve-
trachea.

Respiration by Tracheae
In most animals, except small and simple integument-breathers, oxygen is supplied to the
different body parts through blood stream. Blood and air come in contact either in the
gills or in the lungs.' However, the land arthropods like Myriapoda and Insecta evolved an
entirely unique system for oxygen transport in the body. In their case a series of pores,
called spiracles are present on either side of the thorax and abdomen. These pores lead
into a network of branching tubules, the tracheae (Fig. 14.29), which ramify throughout
the body. Finer branches of tracheae called tracheoles reach out almost every cell. Air
enters the trachea via spiracles and directly reaches the tissues. Thus, blood does not carry
respiratory gases in them.
 Walking legs

Stigma' i . stemurn

Fig. 14.28: Ventral surface of scorpion showing spiracle of book-lungs (a). Vertical section of a book lung (b).

Fig. 14.29: Tracheal system in insects (Dorsal view).

 1

-- - -
Adaptation and Bchavioural While talking about respiration in aquatic insects it was mentioned that some insects live
Pattern
in water but breathe air. Let us now see how they do that. The tracheal system in such air-
breathers is little altered and the spiracles are open. But these insects acquire various
modifications to renew the supply of fresh air. The Odonata and mosquito larvae
periodically come to the water surface to take in fresh one. In many water-bugs and water
beetles there are tufts of water-proof (hydrof@ge)hairs on different parts of the body. The
air trapped among these hairs is used for breathing. In the water-beetle Dytiscus the air is
enclosed between body and the forewings (elytra). The adults of water-bug Nepa or the
larvae of Eristalis (Diptera) possess a long respiratory siphon (Fig. 14.30), which remains
in contact with the air.
Spiracles
\
, surface
--
- -
- P

Water

Fig. 14.30: Respiratory siphon in Eristulis larva.

SAQ 6
i) On the basis of what you have read in subsection 14.3.2 supply the missing words in
the following sentences.
a) In scorpions the respiratory organs are ...................................,of which there
are ............................................pairs.
b) Respiratory organs of Limulus'are ........................................
c) In Crustacea three types of gill$ are found. These are .....................................
-

........................................and .........................................
~

d) Rectal gills are present in the aquatic larvae of .............................................

e) The adults of the water-bug, ..............................has a long respiratory siphon.

hiodifications of Limbs 1

I
You may recall here what you have already studied under "Animal Diversity" on
arthropod appendages. The ippendages in Arthropoda have undergone three major
functional modifications: (a) aS sense organs, (b) as mouth parts, and (c) as locomotor
organs. Out of these, maximum adaptive divergence is shown by the locomotor organs
and mouth parts. Locomotor organs are modified in most of the arthropod groups. On the
whole, primitive groups have more appendages. The locomotor appendages tend to
become reduced in number as they become specialised and modified for various
functions. This increases their maneuverability as well as speed. Two distinct lines of
adaptations are recognised: aquatic and terrestrial. On the contrary, mouth parts show
highest adaptive divergence only in insects, the details of which you have already studied
in Unit 4, Block 1 of this course. Therefore, we will presently discuss the modifications
of locomotor appendages only.

Locomotion in Aquatic Arthropoda

The aquatic arthropods are mostly adapted for crawling or walking on the substratum an4
for swimming. The horse-shoe crab Limulus is a coastal dweller. Its locomotion is
effected by walking on sand or mud bottoms. For this it has five pairs of walking legs.
The fifth pair of walking legs is specially modified for removing the mud while
burrowing. The distal larnellae of this pair spread and can push against the floor to
prevent sinking in the loose mud.
 Adaptive Radiatson

Retractor mus
Brood chamber
Ovi
(b)
(a)

Petrolisthes
cabrilloi
(Porcelain crab)

Lernaeodiscus
porcellanae

Fig. 14.31: a) Balanus a sessile barnacle; b) Lepas, a stalked barnacle. Barnacles; the marine animals'
belong to order Thoracica of subclass Cirripedia, that are usually enclosed i n a shell of
calcareous plates. I n these arthropods head is reduced, abdomen is absent and the thoracic
legs are long and many jointed cirri with hair like setae c) Lernaeod&cusporcellunue,the
rhizocephalanbarnacle, parasitic on the porcelein crab.

Most Crustacea, except the sessile and parasitic cirripedes (Fig. 14.3I), are active
swimmers. Their thoracic and abdominal appendages are adapted for swimming. These
are oar-like and are usually provided with fringed setae which increase surface area of the
swimming organs. In Branchiopoda (small fresh-water crustaceans) all appendages are
adapted for swimming and respiration (Fig. 14.32).

Shell gland Thorax

style

., appendages

Fig. 14.32: Branchiopoda: appendages used for swimming and mpimtion

Adaptation and Behavioural The water fleastswimby strong second antennae. The free living tiny copepods also use
Pattern
mainly their second antennae for swimming (Fig. 14.33 a). Most crustaceans have
become crawlers, though they can also swim and burrow. Whereas many parasitic
copepods (Fig. 14.33 b) have become highly modified for parasitic mode of life, the
prawhs, lobsters, crabs and many others possess well-formed swimming and walking
appendages. The walking in prawns and lobsters is effected by five pairs of walking legs,
which are the posterior thoracic appendages (Fig. 14.34 a). For swimming, these animals
have six pairs of abdominal appendages, named pleopods and uropod (Fig. 14.34 b). The
crabs have abandoned swimming and are adapted for walking. ConsequertIy they have ,,,
their abdomen shortended, abdominal appendages absent and five pairs or thoracic legs
developed for walking.

Fig. 14.33: a) A copepod Mocmeyclopsdolbus; b) So*nl~colos d m ~ ~ ae paraslHc

a cepepod. The mature
female ir attached to the gill of Europcan rrlmon.

Crustaceans have exploited all types of aquatic niches thus exhibiting high degree of
adaptive radiations. They are the dominant arthropod group in marine environment and
also share dominance of fresh water habitat with insects. Invasion of terrestrial
environment is, however, much limited. The most diverse class is Malacostraca and the
most abundant group is Copepoda. Both these groups include planktonic suspension
feeders and numerous scavengers. Copepods also include parasites of both vertebrates
and invertebrates. Cirripedes includes sessile and parasitic crustaceans. Parasitic
copepods exhibit varying degree of modifications as compared to the free living ones. In
most parasitic copepods the adults are parasitic exhibiting free swimming larval stages.
Parasitic cirripedes also show modifications as compared to free living ones. Their body '
is saccular anzthe mantle is devoid of calcareous plates. There is also the absence of
appendages and segmentation;-
 Adaptive Radiation
In aquatic insects.swimming is effected by variously modified legs (Fig. 14.35 a). In
many larval fonns hairy bristles help in swimming e.g. rudder-bristles on the ninth
abdominal segment of mosquito larvae (Fig. 14.35 b).

Dactylus

Dactylus

Propodus *a

Carpus

Ischium
Basis
Coxa
Non chelate leg ' Chelate leg I Chelate leg I1 Chelate leg
.
11,
,

: (Male) (Female)
1-1

(a)

. .,

Typical Pleopod First ~ ~ e o ~ b dSecond Pleopod Uropod

or male
!a)
?
Fig. 14.34: Walking legs of pqwn (a) Swimming appendagesof prnwn (b).

Locomotion in Terrestrial Arthropoda

In most'adult terrestrial arthropods well developkd walking l e e are present, which are
adapted to the needs of their habitats. Their number and structure are variable. Scorpions
possess four pairs of walking legs (Fig. 14.28 a) meant forrunning fast. In spiders also
four pairs of walking legs are present'and all of them are used in walking. These
arthropods cari move very rapidly for short periods.

Centipedes and millipedes are adapted for living in soil and among litter--d in crevices
among stones, logs and bark. Centipedes have one pair of legs per segment while in
'.
. millipedes there are two pairs per diplosegment. Unlike other arthropods these legs are
short and stumpy. They adapted for crawling, swift waking a n d . m i n g . Millipedes
can also effectively push into soil.

'MY
Adaptation and Behavioural Adult insects, as a rule, possess three pairs of legs and are appropriately called hexapoda,
Pattern i e, six-footed. An insect leg is attached to the thoracic wall by means of a ring-shaped
coxa. Besides coxa it has five more segments viz. trochanter, femur, tibia, tarsus and
pretarsus, (Fig. 14.36). The tarsus may have three to five sub segments called
tarsomeres. The insect legs not only serve the function of locomotion but also undertake
other roles like jumping (hind-legs in grasshopper, Fig. 14.37 a), swimming (hind-legs in
Gyrinus, Fig. 14.35 a), digging (fore-legs in mole-cricket, Fig: 14.37 b), grasping (fore-
legs in praying- mantis, Fig. 14.37 c) and grooming (toilet-organ in the hind- legs of
honey-bees, Fig. 14.37 d).

Tarsus \ Pretarsus

91habd~minal'se~ment
Water surface
Respiratory_siphon Rudder bristles
Tracheal gills
Comb on 8Ih abdominal segment
~ i n t r a brush
l

1st abdoni inal segment

Fig. 14.35: An insect leg adapted for swimming (a); Rudder- bristles of mosquito larva (b).

Tarsus
Pretarsus

Tarsomeres

Fig. 14.36: Insect leg adapted to different functions.

 Adaptive Radiation

Distal end of tibia

Fig. 1437: Insect legs adapted for a) jumping; b) digging; c) gnsping; d) toilet o w n of honey bee for
grooming.
Insect Wings: The outstanding success of insects as terrestrial animal is, to a great
extent, on account of their ability to fly. For this purpose most of them usually bear two
pairs of wings on their thoracic segments. We will discuss the wing structure and the
, mechanism of flight, in insects in section 14.4.

I
I SAQ 7
I i) Indicate whether the following statements are correct or incorrect.
1 .
a) Mouth parts in Arthropoda are modified segmental appendages.
b) In prawns and lobsters walking is effected by abdominal appendages.
! In aquatic insects swimming is brought about by six pairs of abdominal appendages.
c)
d) The hind-legs in grasshoppers are adapted for jumping.
e) In centipedes two pairs of legs are present in each segment.
f) Rudder-bristles in mosquito larvae are used for respiration.

14.3.3 Adaptive Radiation in Mollusca

t
Presence of sh 1, mantle, radula and foot distinguishes mollusca from the other animal
phyla. Present- ay molluscans are represented byforrns like Neopiiina
(Monoplacophora), chitons (Polyplacophora), Dentaiium (Scaphopoda), s~failsand slugs
(Gastropoda), mussels and oysters (Pelecypoda), and squids and octopuses- - -
(Cephalopoda). Respiration in auatic form takes place bymean-s of gills and in
Ad~ptatlonand Behavioural terrestrial molluscs by lung. The adaptive modifications in Mollusca are chiefly reflected
Pattern in the shell, foot and respiratory apparatus. You have already studied extensively the
structure and types of molluscan shell in Unit 4 Block 1 and the various modifications of
the foot in Unit 7, Block I 1 of this course. You may recall those portions here. We will
,
now d(scass the structural modifications of respiratory mechanism in Mollusca in the
following paragraphs.

Respiration in Mollusca
Molluscs are mostly marine. Some gastropods and pelecypods are found in fresh-water
while the pulmonate gastropods occur on land. Aquatic molluscs employ gills or ctenidia
(singular- ctenidium) for respiration. The terrestrial forms, on the other hand, breathe by
nteans of the pulmonary chamber, usually referred to as the 'lung'. In some molluscs
exchange of respiratory gases takes place through the general body surface. Thus we have
branchial, pulmonary and cutaneous respiration in Mollusca.

Branchial o r ctenidial respiration occurs in aquatic molluscs. Formed as an outgrowth

of the bodywall the ctenidia are present in the mantle cavity. In all molluscan groups the
basic structural plan of the ctenidium is the same. A gill has a horizontal main axis, which
remains attached to the body. The axis possesses on one or both sides a row of delicate,
flexible respiratory lamellae (singular-lamella) with their surface covered with ciliated
epithelium (Fig. 14.38). When the lamellae are present on one side only, the gill is called
monopectinate and if the lamellae are present on both sides, bipectinate. The ciliary
movement drives a continous flow of water over the richly vascular gills, which receive
deoxygenated blood through inlet veins or afferent branchial veins. The gills return
oxygenated blood through outlet or efferent branchial veins. The direction of flow of
water current over the gills is always opposite to the direction of blood-flow within the
gills (Fig. 14.38 c). This countercurrent flow ensures maximum and efficient gas
exchange.

Rectum. Hypobranchial glands

Efferent blood vessel fferent blood vessel

Gill axis Abfrontal cilia

'Skeletal rod

Filaments

skeletal rod
F A :.2:

~i~~'14.3
Transverse
8: sections through the body of a mollusc at the level of mantle cavity a). ~ A n t a l
, section throvgh the gill showing gill lamellae b). Transverse section through current and
blood flow c). the gill lemellae showine dir*rtinnsnc--tpr current.
 -

Adaptive Radiation
The rtenidial arrangment differs in different classes of Molluscs. In Pelecyimda the
gills subserve not only respiration but help in feeding also. In Monoplacophora there are
five pairs of monopectinate gills with finger-like lamellae (Fig. 14.39 a). The position of
the gills in this class shows segmental nature of the Mollesca, which otherwise is not
apparent in other classes. In Polyplacophora the chitons have six to eighty bipectinate
gills arranged in a row within the two mantle cavities (Fig. 14.39 b). While in the
Aplacophora (Solenogastres) the gills are reduced or absent.

Cerebral ganglion Preoral tentacie

Dorsoventral duscle 8

-
Fig. 1439: Mollusun gill a) Monoplacophora (Nwpilina);b) Chiron; c) Sea slug (A.eo1is).
Gastropods has three subclasses :Prosobranchia, Opisthobranchia and Pulmonata. In
Prosobranchia the gills are shifted in front along with the mantle cavity due to torsion and
there may be one monopectinate gill (as in Pila) or two bipectinate gills (as in Haliotis).
In Opisthobranchia the mantle cavity and the organs it contains shift to the right side
due to detorsion (see earlier unit for torsion). Forms like Aplysia (sea-hare) possess one
ctenidium on the right side while Doris and Aeolis (Nudibranchia) have altogether lost
true gills. Instead, they have acquired secondary gills which are present either around the
anus or on the lateral edge of the mantle or in rows on dorsal body surface (Fig. 14.39 c).
In pulmonates, gills are absent. Manfle cavity is on the right side. This becomes a
vascularised "lung" for air breathing.
Adaptation and Behavioural Pulmonary Respiration: In terrestrial gastropods the mantle cavity is transfbrmed into a
Pattern pulmonary chamber or lung, the roof of which is richly supplied with blood vessels.
The evolution of the pulmonary sac is a land adaptation. Alternate muscular contraction
and relaxation of the mantle floor causes the air to enter in and pass out of the pulmonary
sac through a small aperture guarded by a valve. The exchange of gases occurs through
the mantle wall. In some forms the pulmonary sac may also help in aquatic respiration.

The gills in Pelecypoda have a complex structure. Besides breathing, they also help in
collecting food and serve as a brood-pouch. There is one pair of bipectinate gills in the
mantle cavity, one on either side of the body. These extend from the anterior to the
postrior end of the animal. On either side of the axis in each gill, long filaments extend
ventrally and then bend upward like a hairpin (there being two "hair pins" on each side)
(Fig. 14.40). There is an ascending and a descending limb in each "hairpin". These
filaments may hang freely or the adjacent ones may be joined by inter filamer~tar
junctions forming a gill-plate or demibranch. There are two gill plates on each side, an
outer and an inner one. Each gill-plate has two larnellae each made of an ascending and a
descending limb. The outer and the inner lamellae are joined together by interlamellar
junctions (Fig. 14.40). The gill-plates divide the mantle cavity inlo an upper
suprabranchial and a lower infrabranchial chamber. The former opens to the exterior
by excurrent (exhalant) or dorsal siphon which drains the water out. The infrabranchial
chamber has an incurrent (inhalant) or ventral siphon through which the water enters the
mantle cavity. The action of the cilia present on the gills maintains a continuous water
current over their surface in the mantle cavity, where exchange of gases take place.

Afferent blood vessel

c=rJ w
Efferent blood vessel Food groove

Connection of filament tipst. A

with mantle
:tion of filament tip
lnterlamellar junctions foot-visceral mass
between folded

Interfilamentous junctions kd -
between adjacent filaments
(d)
Fig. 14.40: Evolution of Lamellibranch gills in Pelecypoda. a) Primitive protobranch gill b) Food groove
development to create lamcllibraneh condition. e) At food groove the filaments fold to form
lamellibrrnchs conditions. d) Lamellibranch gills with tissue junctions providing support to
the folded filaments.

Pelecypods are sedentary feeders and wait for the food to come their way. The constant
inflow of water through the incurrent siphon into the mantle cavity brings in food
particles which include micro-organisms and organic debris. When water enters the
mantle cavity the heavier particles sink down and afe expelled. The lighter food particles
passover the outer surface of the gill lamellae where they get entangled in mucus
..
 -

Radiation
secreted by the gills. The mucus-mixed food particles pass into food grooves on the
ventral edges of the gills, which take these towards the mouth. Near the mouth the labial
palps further sort out the particles according to their nature. Smaller digestible particles
are taken to the mouth while larger indigestible ones are thrown out of the mantle cavity.

Cephalopods have simple bipectinate gills situated on either side of the anus. The leaf-
like lamellae are arranged in a linear row on the axis. There are no cilia on the gill surface
and the flow of water is regulated by the ~nuscularmantle, funnel and the inlet-valves.
There are two gills in the cuttlefish, squids and octopuses and four in nautiloids.

Cutaneous Respiration
In Scaphopoda, Aplacophora and parasitic or terrestrial Opisthobranchia respiration
occurs through the moist integument of the mantle cavity or through the general body
surface. It is called cutaneous respiration.

SAQ 8
i) '2;;: in the blanks in the following sentences using words given in the parenthesis
below:
(deoxygenated, efferent branchial, ctenidia, afferent branchial, oxygenated, flow,
drives, vascularised)
a) Formed as an outgrowth of the bodywall, the .................................... are
present in the mantle cavity.
b) The ciliary movement drives a continuous flow of water over the richly
...................................gills, which receive .........................................
blood through inlet veins or ................................... veins and return
.................................blood through outlet veins or .............................
veins.

ii) Indicate whether the following statements are true (T) or false (F).
a$ In Pelecypoda gills also serve food capture.
b) There is one monopectinate gill in Pila.
c) Aeolis and Dorrs do not possess true gills.
d) The gill surface in Cephalopoda is ciliated.
e) Pulmonary chamber is found only in aquatic Molluscs. '

f) In Opisthobranchia the gills are anteriorly placed.

14.4 FLIGHT IN INSECTS

Insects are unique among non-chordates to have evolved the ability to fly. For this
purpose most adult insects possess one or two pairs of wings on their thoracic segments.
The wings form an important basis of insect classification. There are chiefly two types of
insects : winged and wingless. Wingless insects may be primarily wingless or Secondarily
wingless. In the former (primarily wirigless insects) the wings have not evolved. The
primarily wingless insects include silverfish and springtails. Secondarily wingless insects
lost the wings during their evolution from winged insects. The ants, lice and fleas fall in
the category which has secondarily lost wings. The dragonflies, butterflies, houseflies,
mosquitoes, bugs, beetles etc. are winged insects. The wings of insects evolved as lateral
outgrowths of the body.

Structure of Wings
The wing (Fig. 14.41) arises as dorso-lateral outgrowths of the bodywall on mesothorax
and metathorax. It is a thin membrane and is supported by a system oftubular veins. The
membrane actually consists of two layers of closely apposed integument. The \teins are
the heavily sclerotised regions where the two layers remain separate. The veins have
branches of nerves and trachae. Blood circulates through the veins in the wing.
 Adaptation and Behaviourall Costal margin A ~ i c a anale
l
, Pattern.

Humeral angle
I A

Anal margin Anal angle

Fig. 14.41: Wing of'an insect.

Fig. 14.42: A gall gnat showing wings and halhres. Haltere are responsible for eqaillibrium during the
flight.

The forewings sometimes become hardened sewing to protect the hindwings, as in the
beetles. In the dipterans (eg. housefly and gnat) the hind wings have become modified
into a sense organ called haltere (Fig. 14.42).

Fig. 14.43: Stonefly showing the wings folded at rest.

I In most insects however thb two wings on each side are membranous and can be folded at
Adaptive Radiation

i1 rest (Fig. 14.43). They are coupled to function as a single wing functionally. This
increases their efticiency during flight.

Wing movement
Flight in insects is the result of co-ordinated wing beats. These movemints are highly
complex, involving various components like elevation (upward movement), depression
(downward movement), and forward as well as backward movements. These wing
movements are produced mainly by (I) muscles directly inserted into the wing base
- (direct flight muscles); (2) distortions of thorax brought about by muscles which are not
directly associated with wings (indirect flight muscles) (Fig. 14.44 a-d); (3) elasticity of
wing hinge. Once the muscles have moved the wings in a particular unstable position, the
elasticity movements result in bringing the wings automatically into the stable position,
with great force. This is known as "click" mechanism. Presence of a protein called
"resilin" with considerable elastic property, in the hinge joints of insects, enables insects
to bring about this "click movement.

I Insects thus fly with a wing beat frequency of 4-20lsecond as in butterflies or I9OIsecond
in honeybees and house flies. Some small dipterans may show a wing beat frequency of
1000/second.

IDirect flight muscles of locusts and dragon flies Indirect flight muscles of flks and midges

Ig. 14.44: (a) and (b) Flight muscles of Insects sucb as dragonflies and locust where upward stroke is by
indirect muscles and downward stroke is by direct musclu. (e) and (d) Flight muscles of
insects like bees where both upstroke and downstroke are by indirect muscles.

'The wing movements are very complicated. The whole mechanism of flight may be
described in simple words as follows : The various wing movements create a low air
pressure zone above and a high pressure zone below the wings due to which the body of
the insect lifts above in the air: Similarly the wing twisting creates an area of low air
pressure ahead and high air pressure behind the insect. This provides a forward thrust to
the body. Many insects can remain stitionary during flight. In their case wing movements
create only lift and no thrust. Steering during flight is effected by shifting the centre of
gravity or by altered wing- beats.
Adaptation and Behavioural
Pattern SAQ 9
i)' Mark the correct alternative in the following statements.

a) The insects in which wings have not evolved at all, are called primarily1
secondarily wingless insects.
b) Dragonflies are wingedfwingless insects.

ii) Match the words in list I with those in list 11.

List I List I1
1. Silverfish a. Secondarily wingless
2. Ants b. Primarily wingless
3. Haltere c. Modified hind wings
4. Resilin d. Help the click movement

MIGRATION IN INSECTS
In insects mainly two types of flight activity occurs. Trivial flight serves routine activities
such as feeding and mating while during migration flight activity dominates. Migration is
essentially dispersal. Whenever in a partciu lar habitat any environmental factor hinders
feeding or breeding activity, the insects fly out to explore new pastures for food and
reproduction. This is what we call migration. In the adult life of many insects there is a
particular phase when this activity predominates. It is called the migratory phase and
varies from a few hours as in many insects, to many days (as in some Coleoptera and
Lepidoptera). In migration usually the animals move out from the feeding ground at the
end of that activity in search of breeding ground and afier breeding, they come back to the
old habitat.

Since the main aim of migration is dicpersal, females invariably participate in it, while
males may or may not do so. In the locust, Schistocerca both sexes are included in
migratory flights, but in the bug, Eurygarter males as well as females migrate from
breeding to feeding grounds but only the females return to the breeding ground.'In
Rhyacionia, a lepidopteran, females are fertilized before they start migration to the
breeding ground. The males do not migrate.

Direction of Migration
Direction of migration is mainly influenced by the wind-speed and direction of the wind.
Wind speed increases as one moves higher in the air. The speed of insects in flight in
relation to air is called air-speed. Wind-speed is comparatively lower closer to the ground.
This forms what is called boundary layer. Air speed is greater than the wind speed at the
boundary layer. However at higher levels, wind speed exceeds air speed. The insect can
determine the direction and course of migration on its own in the boundary layer. For
example, the moth Ascia monuste in Florida (US) flies 1 to 4 m above ground level. It can
easily proceed against a wind current of 10 kmlhour velocity. The direction of migration
in this moth is determined by availability of flowers in the area. In other insects, factors
responsible for aetermining the direction of migration may include position of the sun,
landmarks such as roads, coastline etc. In the boundary layer the migration may be
initiated by a number of factors but ultimately the position of the sun. pattern ofthe
polarized light in the sky and the visual landmarks decide the migratory path.

Migration outside the boundary layer is seen in many insects. Sometimes those insects
which usually migrate in the boundary layer, are seen flying above the boundry layer.

Ascia monuste has been reported flying at a height of 1500 m and above in Argentina. At
higher altitude the insects fly in the direction of the wind current. Denser swarms of
locusts at higher levels, in higher wind-speeds, fly in the direction of :he wind. Aphids
with a low air speed (0.6 mlsecond) find it difficult to fly against wind currents or to
migrate within the boundary layer. They ascend up in the air due to positive phototactic
reaction to ultra violet rays, and are then carried to long distances by wind currents. Many
other insects such as dragonflies, beetles, butterflies and rnotlls also move down-wind (in
the direction of wind) at higher levels. Once the insects are carried to higher altitudes by
 Adaptive Radiation
higher wind currents (convection current) these carry the insects over long distance in a
shorter duration. It is observed that swarms of locust Schistocerca cover a distance of '
1200 km within 24 hours at 700 m above ground at a wind-speed of 45 kmlhour.

Return Migration
Some insects show to and fro migratory movements. The monarch butterfly, Danaus
I
plexippus in the United State migrates in autumn from the north where winter temperature
I
becomes too low and food scarce, to the south where temperature is moderate and food .
supply plenty. In February these insects start return migration northward. This sentence is
not fitting properly. Such two way migration by the same individuals is exhibited by
Agrotis infusa (Lepidoptera) in Australia and Hippodarnia convergens (Coleoptera) as
well as a number of other insects.

Locust Migration
Locusts exhibit mass migration or swarms. The swarms of the desert locust, Schistocerca
: gregaria may cover an area of 10 to 250 square km. You may be surprised to know that a
swarm spread over about 20 square km may contain about 100 crore individuals. The
swarms may cover a distance of up to 100 km a day.
' L O C U form
~ S two types of s w a n s vir. stratiform and eumuliform. In the former the
locusts fly flat in the form of the thin layer within few meters above the ground and there
may be 1 to 10 individuals per cubic meter. In cumuliform swarms locusts fly in a tower-
like column extending up to 1000 m above the ground, with a low density of only 0.001
to 0.1 individual per cubic metre. 'The stratiform swarms are formed in the absence of
convactive cm-rents to take them up while the cumuliform ones occur when there is
convection current.

An interesting aspect of locust swarms is that all individuals in a swarm$onot face

forward. Their heads face in different directions. This is called random orientation.
I However, the locusts at the edge of the swarm face towards the body of other locusts.
This helps to maintain the integrity of the swarm.

Beginning and End of Migration

Let us now examine the cause of migration. Migration is often initiated not by the actual
onset of adverse environmental conditions. For example, in the monarch butterfly, the
southward migration begins before the onset of cold conditions in the north, and locust
swarms leave their habitats while plenty of food is still available. This shows.that
migration is an evolved adaptation in these cases and does not result from adverse
environmental stimuli as such. Migration begins even before the onset of adverse
conditions. This may be called spontaneous migration. On the contrary, in some cases
migration may be stimulated by some physiological or behavioural phenomena which put
the insect into a state of readiness to migrate. This type may be called facultative
migration. Photoperiod, temperature and food supply are some such factors. Once the
insects are kept in a state of readiness to migrate, the actual take off may be stimulated by
another set of factors, like light of a particular intensity, wind speed, temperature etc.
Similarly, it is not the physical exhaustion which brings migration to an end, but different
wave-lengths of light being reflected by leaves (as in aphids), smell of salt-marshes (as in
Ascia) and odour from host trees (as in beetle Melolontha) etc. may be responsible for the
termination of migration.

Significance of Migration
Migration enables the species to cope with, the changes in the location of its habitats. It is
more common in those insects which occupy temporary habitats. For example, many
species of Odonata, which live in permanent streams do not migrate whereas more than
half of those living in temporary pools do so. The temporary nature of habitats may be
due to changes in temperature, humidity, rainfall, etc. Migration is a way of over coming
the adverse environmental conditions.
 1
Adaptation and Behavioural
Pattern a
SAQ 10
i) Indicate whether the following statements are true (T) or false (F).
Trivial flight serves feeding and mating.
In the locust Schistocerca, only males are included in migratory flights. ,
Boundary layer of the air is near the ground and within this layer the air-speed is
greater than the wind-speed. .
Monarch butterfly, Danaisplexippw in the US migrates in winter from south to
north.
At higher altitudes in the air the insects fly downwind i.e. in the direction of the
wind.
Spontaneous migration is initiated by one or the other environmental factor.
Physical exhaustion of the insect brings migration to an end.
Migration is common in those insects which live in temporary habitats.

14.6 SUMMARY .
In this unit you have learnt that:
Animals which lead their lives a$, individuals are called solitary and those living in
organised groups are known as colonial. True colonies in which individuals or zooids
are organically connected by living matter, are present in protozoans and
coelenterates. Polymorphism and division of labour are some of the important
features of colonial life.
If the animals of the same or closely related groups adapt for different modes of life,
they are said to show adaptive radiation or adaptive divergence.
The basic needs of animals viz. food and safety, lead to adaptive iadiation. Among
non-chordates Annelids, Arthropoda and Mollusca exhibit clear adaptive radiation.
Adaptive radiation is chiefly refle.cted in method of feeding and exploitation of
different habitats in Mnelida, fespiratory modifications and limb modifications in
Arthropoda as well as modifications of shell, foot and respiratory apparatus in
Mollusca.
Wings are unique acquisitions of insects. Formed as dorso- lateral outgrowths of the
body wall, these are moved by direct and indirect flight muscles as well as by
elasticity of the thorax, of flight muslces and of wing hinge. They impart capability
of flight to the insects.
' Two types of flight activity are shown by the insects. Trivial flight for routine
activities like feeding mating etc., and migration for dispersal. Migration is common
in many species of insects and may be either spontaneous or facultative.

14.7 TERMINAL QUESTIONS

1. Differentiate between adaptive convergence and adaptive divergence. Write the
answer in two or three lines in your own words.

2. Name three flagella-bearing Protozoa which form advanced colonies. Does any of
these show polarity and if yes, explain why you think so. )
 Adaptive Radiation
3. Define polymorphism. Why do you say that Siphonophora colony is polymorphic?
......................................................................................................

4. Mark the correct alternative in the following statements:

a) The mimal: which would wait for the food to come their way, acquired
radial/bi!,te,dl symmetry.
b) Filter-feeding has evolved in sedentarylactive food- seekers.
c) Eversion of proboscis affects feeding in predatorylparasitic Polychaeta.
d) Earthworms lacklpossess antennae and palpi.
e) One full meal by a leech may last for four hourslmonths.

5. Give t v o advantages and two disadvantages of the hard and tough body cover in
Arthropoda.

6 . Match the words in the List A with the most appropriate ones in the List B.
List A List B
1. Branchiae a. Dipteran larvae
2. Tracheae b. Stonefly larvae
3. Anal (trachealj gill:, c. Insects
4. Blood-gills d. Crustacea

7. Molluscans are believed to have evolved from annelidan ancestors, though they show
no trace of segmentation. Give two grounds on the basis of which their ancestry may
be linked with Annelida.
......................................................................................................

8, Indicate whether the following statem,ents are correct or incorrect.

a) Migration in insects is meant for feeding only.
b) In a stratiform swarm locusts fly in a column at about 1000 m above ground.
c) In random orientation all individuals in a locust swarm face forward.
d) The course and direction of the insect migration is determined by the position of
the sun alone.

14.8 ANSWERS
Self Assessment Questions
1. i) (a) environment, adaptive convergence.
(b) adaptive divergence.
ii) (a) F, (b) T, (c) FY ( 4 F

2. i) (l)c,(2)a,(3)b,(4)d.
ii) monopodial, sympodial, permanent.

3. i) Osborn, ii) Lamarck, iii) errant polychaetes, (iv) oceans

Adaptation and Jhhavioural ii) (a) food, (b) safety.
Pattern

5.' i) and ii) incorrect, iii) and iv) correct.

6. i) booklungs, four, ii) book-gills, iii) Phylio-branch, trichobranch and

dendrobranch, iv) Odonata, v) Nepa.

7. i) Correct, ii) and iii) incbrrect, iv) Correct, v) and vi) incorrect.

8. i) (a) ctenidia, (c) vascukised, deoxygenated, afferent branchial, Oxygenated,

efferent branchial.
io (a) T, (b) T,(c) T,(dl F,ie) F, ( 0 F.

9. i) (a) primarily, (b) winged,

ii) - -
l b;. 2 - a; 3 c; 4 - d

lo. i) a)T,b)F,c)T,d)F,e)T,f)F,g)F,h)T.

Terminal Questions
1. In adaptive convergence animals of unrelated groups adapt for the same habitat while
in adaptive divergence those belonging to same or closely related groups adapt for
different habitats.
2. Volvox, Pleodorina and Pandorina form advanced colonies among flagella-bearing
A protozoans. All of them show polarity because they swim always with a particular
side facing forward.
3. Polymorphism is the occurrence of zooids or individuals in a colony in many forms,
which exhibit division of labour. Siphonophora colony consists of gastrozooids for
-
feeding, dactylozooids for protection, and three types of other zooids gonozooids
concerned with reproduction, nectophores for locomotion and gas-filled
pneumatophores for floating.
4. Correct alternatives are : (a) radial, @) sedentary, (c) predatory, (d) lack, (e) months.
5. Advantages:
1) Provides support and protection,
2) Prevents desiccation,
Disadvantages:
1) Hampers growth
2) Hinders smooth gaseous exchange through general body surface:
6. (1) e, (2) d, (3) b, (4) a.
7. The two grounds are :
1) A trochophore larval stage occurs in Polychaeta among Annelida as well as in
Mollusca.
2) The arrangement of paired gills in Neopilina (Monoplacophora) points to the
segmental nature of Mollusca.
8. All statements are incorrect.
 UNIT 13 BEHAVIORAL PATTERNS
Structure
15.1 Introduction
Objectives '
15.2 Taxis and Kinesis
Taxis
Kinesis
15.3 Biological Rhythms
Control of Biorhythms
Biological Clock
15.4 Communication Behaviour
Visual Signals
Mechanical Signals
Chemical Signals
Communication among Honeybees, the Dance Language
15.5 Courtship Behaviour
Need for ~ourtshipBehaviour
Sex rlifferences in Courtship Behaviour
V:sual, Mechanical and Chemical Displays
Nuptial Gilts
Sperm Competition and Mate Guarding
Alternative Tactics of Mate Competition
Rejection and Deceit in Courtship
15.6 Social Organisation in Insects
Advantages and Disadvantages of Social Behaviour
Social Wasps
Ants
Honey Bees
Termites
15.7 Parasitism
Types of Parasites
Effects of Parasitism on Parasites
15.8 Summary
15.9 Terminal Questions
15.10 Answers

INTRODUCTION
Behavioural patterns of animals are the patterns of their gestures and movements in
response to stimuli in their environment. Behaviour patterns are purposeful and may be
for procurement of food, finding mate, for locating convenient and sheltered site, or
1
communication with animals belonging to the same species or to different species.

When an animal moves its body in response to a stimulus, the movements are termed
taxes and kineses. Behavioural activities occurring with clock work precision at regular
intervals are called biorhythms. Elaborate ritualistic behaviour patterns are associated
with courtship and mating, communication with members of one's own species as well
with other species. Certain groups of animals live in groups to form societies and exhibit
social behaviour. They also possess special means of communication with members of
their own species. Some communication signals are for self defence. In this Unit you
I
shall learn about the various taxes and kineses, rhythms, social organisations, courtship
I
and communication behaviour and the behaviour of parasites, with reference to non
) chordates.

I Objectives
After studying this Unit you shall be able to:
distinguish between taxis, kinesis,
explain endogenous and exogenous biological rhythms,
describe different types of communication methods in non-chordates,
I
illustrate how non-chordates attract their partners and mates,
I
describe the caste system, division of labour and advantages of living in social
groups,
o distinguish between different types of parasites, and
discuss parasitic adaptations.
Adaptation and Rchavia~~ral
Pattern 15.2 TAXIS AND KINESIS
Atiitnals are characterised by great mobility. Stimuli from the environment direct these
movements. These movements may be either taxis or kinesis.

15.2.1 Taxis
Taxis (plu Taxes) is a directional movement, either toward or away from a source of
stimulation. The animal is oriented along a line that runs through the source of
stimulation and the long axis of the animal's body. Taxis is termed positive if the
movement of the animal is towards the stimulus and negative, if away from it. Taxis is a
behavioural response that cannot be modified by learning. A moth flying towards light
(Fig. 15.1) is a classicql example of taxis. So is the migration of an earthworm to the
surface of soil after a heavy rain. [Taxes are easily demonstrable in animal - like protists
such as Amoeba and Paramecium (Fig. 15.2).]

Taxes are classified according to the nature of stimulus. Table 15.1 shows the various
-
types of taxes thermotaxis, phototaxis, thigmotaxis, rheotaxis, galvanotaxis and
geotaxis.

Table 15.1: Different kinds of Taxes.

Fig. 15.1: A moth flying towards a Nature of Name of Type of taxis. ' Examples
source of light is an Stimulus Taxis (positive + o r negative -)
example of taxis.
Teniperilture Thermotaxis +or - Animals thrive in different ranges of
temperature. Optimum range
20-2S°C, cold blooded animals avoid
temperatures above and below the
range of temperature that,they can
tolerate.

Light Phototaxis Hydra, Musea (housefly), Ranaha

(an aquatic insect) move towards
light.

Earthworms, mosquitoes,
cockroaches, woodlice move away
from light.

Mechanical Thigmotaxis Contact with food causes positive

taxis in most animals.

Contact with barriers brings

about avoidance reaction.

Chemicals Chemotaxis Odour from chemical ingredients

of food orient house flies towards it,
also Hydra;
Mosquitoes avoid mosquito
repellents Animals show
negative response to injurious
chemicals.

Water and Rheotaxis Moths and buttedies fly

wind currents into wind current.

Planaria, the free living flatworm

moves against water current.

Electric Galvanotaxis Hydro reacts to weak currents

current of electricity, it bends toward: the
anode.

Gravity Geotaxis + Cnidarian larva planula swims

towards sea bed.

Ephyra larvae ofjelly fish swim

away from sea bed. Drosophilids
( h i t fly) fly up against gravity to dry
parts of a iar.
 - .

Bchavioural Patterns

THIGMOTAXIS

THERMOTAXIS

PHOTOTAMS

I
Fig. 15.2: Reaction to various stimuli in Amoeba, arrows indicate the direction of movement.

15.2.2 hinesis
Cinesis is a non-directional movement. Here the animal's body is not oriented with
espect to the source of stimulation, but the rate 03 speed of movement changes with the
ntensity of the stimulus. Hydra moves its .tentacles at random in search of food but if
ood is kept close to tentacles, they are moved faster.

Fig. 15.3: Kinesis in Woodlice.

r
Adaptation and Behavioural
Pattern If the woodlice Porcellio scaber are given a choice between humid and dry areas, they
tend to collect in humid areas gradually. h i s is due to their non-directional movement. In
other words, they do not seek out humid areas, but movement is random. What happens
here, is that their movement increases on dry areas but their movement decreases when
they occupy humid areas. The random movement with speed is an attempt to fmd out
optimal conditions. Once they reach humid areas, their speed slows down and they settle
there (Fig. 15.3).

SAQ 1
1. What is taxis?

.......................................................................................................
2. What is kinesis?
......................................................................................................
.......................................................................................................
3. What are flte terms used for the following taxes
(i) movement towards light.
................................................................................................
(ii) movement away fiom electric current.
................................................................................................
(iii) movement in response to gravity.
.................................................................................................

15.3 BIOLOGICAL RHYTHMS

Many behavioral.activitiesare carried out by animals at regular intervals of time. Most
animals are active during the day and rest at night (diurnal animals), while some animals
such as cockroaches are active during the dark hours of the night and rest during the day
(nocturnal animals). Similar behavioral activities occurring with cyclical regularity
constitute biorhythms.

Some biological activities .which showf rhythmic oscillations are feeding, mating, egg
laying, emergence from pupa (in case of insects) and migratory.behaviour. The rhythmic
activities are co-ordinated with the cycles of nature such as day and night cycles, annual
seasons, lunar cycle of one moon-rise to the next. This kind of rhythmi5 activity is to
avoid adverse environmental factors and to hlly make use of favourable factors. Fot
example, it will be beneficial for bees and other diurnal insects like butterflies, to be
active during the day when the flowers they visit open so that they can collect nectar and
1 pollen. Thus organisms have evolved their own rhythms which co-ordinate their activities

with environmental rhythms. Daily rhythms such as feeding, drinking and sleep follow a
cycle of approximately twenty four hours and are termed circadian rhythms (Latin circa:
approximately; diem, day). Many littoral shore animals become active when the tide
- leaves them exposed. This is tidal rhythm eg. fiddler crab emerges fiom its burrow to
feed at low tide. Certain intertidal snails release eggs at very high tides which occur once
in two months. The palolo worm and some other polychaete annelids show lunar
rhythm. They rise to the surface of the sea t'o spawn (lay eggs) during certain phases of
'
the moon. Certain animals show courtship behaviour, mate and reproduce once a year
(circa annual rhythm). Many animals migrate to and from breeding grounds twice a
year. Many insects or their stages go illto a state of dormancy or diapause during winter
when the climate is not congenial. For example, eggs of the mosquito Aedes, larvae of
flesh fly (Sacrophaga) and-certaindragonfly nymphs show diapause in winter. Thus
biorhythms are behavioural activities performed at regular intervals.

153.6 Controll of Biorhythms

Certain activities require an external stimulus to maintain them at regular intervals. These
rhythmic activities are termed exogenous rhythms. A major external factor regulating
rhythmic activities is photoperiod or relative length of day (or light hours) and night (or
dark hours). Temperature and humidity are other such factors which may control rhythms
 Behavioural Patterns '
in animals. The palolo worm swarms and mates once a year, on the first day of the lasr
lunar quarter of the year. Lunar cycle is the exogenous factor triggering this activity.

However an internal biological clock exists in almost all eukaryotes which can detect the
passage of time even if the environmental cues are absent. Behavioral activities controlled
by the biological clock within an organism are termed endogenous rhythms. Behaviour
of many terre@al insects appears to be controlled by endogenous rhythms related'to
photoperiods. Drosophila always emerge from pupae at dawn. Insects generally have an
inbuilt biological clock. Circadian or diurnal rhythms are the most common of these
biorhythms and they are controlled by endogenous biological clocks.

How would you distinguish whether the rhythm exhibited by an animal is exogenous or
endogenous? For example, cockroaches are nocturnal, they begin their activity by the
onset of night, and stop their activity before daybreak. Is this rhythm exogenous (i.e.
controlled by outside darkness in this case) or by its own internal clock? An easy way to
find it out, is to transfer cockroaches to continuous darkness or to continuous.light. You
will then see that irrespective of whether they are under conditions of constant dark or
constant light conditions, they will exhibit aperiodicity or rhythm of approximately 24
hrs. This indicates an endogenous rhythm, independent of external lighttdark conditions.

15.3.2 Biological Clock

Biological clocks are internal mechanisms that provide a means of measuring time. These
internal clocks are set to cyclic events in nature'such as day and ni,ght, temperature
I
changes during the seasons, high and low tides (in case of marine organisms) etc.
Animals accordingly show feeding behaviour and other activities, sleep and rest or
migrate (in case of migratory animals), controlled by the internal clock. A biological ,
1 clock is therefore a necessity for most organisms.

iI Entrainment,?nd free running of biologic~lclock

I Biological clock is autonomous and does not vary in its time keeping property. However,
if the environmental cycle is changed as happens when animals travel long distances
during migration or are transported for experimentalpurposes to a different continent, the
internal clpck sets into phase with the external clock prevailing in the new place. The
biological clock is then said to be entrained. Entrainment is like setting a c!ock to correct
time so that it does not give wrong signals at wrong times. In other words, entrainment is
setting of the biological clock in phase with the environmental cycle. Once the biological
I clock is set, it continues to run on set time for a while even if environmental conditions
are suddenly changed. As time passes, biological clock resets or entrains andgets into
b phase with the new set of environmental conditions.
I

OBSERVED
RHYTHMS

ENTERTAINMENT CLOCK
PATHWAY MECHANISM
Locomotory
f Dattems

Sunlight or
Sensory Feeding
environmental cues receptors behaviour

Honnone
release pattern

I \ Other rhythms
Fig. 15.4: A master clock may, in some species, act as a pacemaker to regulate tRe many other clocks that
control the circadian rhythm of the organism.
If animals are isolated from environmental influences then the cycle does not stick to a 24
hour rhythm. For example cockroaches are nocturnal, but if they are kept,in_constant
darkness their rhythmic activities continue but instead of an exact 24 hour cycle, they
may exhibit a slightly different cycle of say 23.8 hour. This property of the cycle
becoming slightly shorter or longer than the exact 24 hour schedule is referred to as free
running of the biological clock Free running period is the internal clock's repetitive
Adaptation and Bel~avioural
Pattern cycle when an animal is isolated from environmental cycle and kept under constant
conditions.

Zeitgeber
The environmental stimulus which entrains a biological clock is called Zeitgeber (from
German: Zeit - time; geber - giver). Light, temperature and tides are important Zeitgebers.
Several environmental factors have been shown to act as Zeitgebers (Fig. 15.4).

Where is the biological clock located?

Researchers have tried to locate the biological clock in the nervous system. But wl.at
generates the rhythms is as yet unknown. From some experiments on cockroaches and the
fruit fly, it is believed that the rhythm originates in the optic lobes of the brain. Optic
lobes, therefore, fonn the pacemaker of biological clock (Fig. 15.5) in these organisms.

Pig. 115.5: Nervous system in cricket. Visual information is relayed to the optic lobes of the brain. If the
optic lobes are surgically disconnected from the rest of the brain, the crirket.losesits ability to
maintain its circadian rhythm.

Nature of biological clock

The nature of biological clock seems to be biochemical. This biochemical mechanism
neither slows down in cold weather nor speeds up during warm days, inspite of the fact
that in cold blooded or poikilothermic animals, biochemical activities double with each
rise in 10' C. Internal clocks are little affected with changing temperature conditions.
Thus biological clocks are said to be temperature compensated.

Characteristics of the Biological Clock

1. Biorhythm or biological clock has repeating units called cycles of activity and rest,
sleep and wakefulness etc.
2. Each cycle takes a particular time period.
3. It shows peaks and troughs a phase of peak activity followed by a phase of low
activity.
4. Rhythms are said to be temperature compensated. That is, rhythm (or biological
clock) keeps the sime time irrespective of a rise or fall in temperature outside.
5 . Metabolic inhibitors do not affect biological clocks or biorhythms.

SAQ 2
I. Define biorhythm.
.....................................................................................................
2. Match the terms in Column 1 with words or statements in Column !T. Behavioural Patterns

Column I Column I1
i) Circadian rhythm a. light hours
ii) Photoperiod b. rhythm under constant darkness
iii) Zeitgeber c. marine invertebrates.
iv) Lunar rhythm d. environmental stimuli that set the biological clock.
v) Free running e. 24 hour cycle of earth's rotation.
3. What is the difference between exogenous and endogenous rhythm?
....................................................................................................... - - -

4. What does entrainment of the biological clock mean?

5. Which organ system, seems to control the biological clock?

&
.

15.4 COMMUNICATION BEHAVIOUR

Animals need to interact with their conspecifics (members of the same species) as well as
animals of other species. Interaction requires effective means of communication. Various
means of communication among animals have thus evolved. Communication behaviour is
highly developed in animals that exhibit social behaviour. Though there are many
definitions for communication for our purposes the following definition of
communication between two organisms can be acceptable.

An action on part of one individual organism which alters the pattern of.behaviour
in another organismis known as biological communication. The action is in the form
of a signal from one animal to another and the sender of the signal usually benefits from
the response of the receiver.

Human beings usually communicate through language made of words. Words can be
rearranged-in infinite ways to construct numerous messages. Language of animals, other
than humans is in the form of signals. These mutually recognisable signals may be visual, .
auditory, tactile or chemical. Signals are exchanged between individuals and these
influence the behaviour of each other. One signal may convey one or more messages.

Types of Signals and Their Purposes

Signals for communication may be of four types.
-
1. Visual Which can be recognised by seeing.
2. Mechanical - Which are recognisable through the tactile sense.
3. Auditory - These are sound signals of various frequencies which convey different
messages.
4. Chemical - Signals are due to secretions. The secretions include pheromones.
Signals are used to communicate various messages. The messages may be regarding
i) availability of food.
ii) need for defence from predators
iii) availability of mating partners.

An animal may utilise a channel of communication for more than one purpose. For
example, many spiders respond to captured prey through web vibrations. Through web
vibration they also communicate with prospective mates. Jumping spiders use visual
signals to stalk and capture prey. They utilise these signals for courtship also. All signals
of display are behaviour patterns established through evolution to become effective for
communication. This is called ritualisation. Through ritualisation, simple movements or
traits become more intense, conspicuous and precise and their original undifferentiated
function acquires a signal value.

Communication behaviour often occurs in regular sequences. These sequences become

stereotyped and then they are termed fixed action patterns. Most stereotyped signals are
for courtship and territoriality.
Adaptation and Behavioural
Pattern
signal
SIGNALLER b-
visual, tactile
auditory, chemical

1. Sender ready to mate

Meaning of Signal 2. Sender anxious, predator in sight.
3. Prey or food in vicinity
4. Predator beware, sender is harmful
5. Sender%territory, tresspassers flee

15.4.1 Visual Signals

The posture of the body and the colour patterns of the animal are the two major visual
signals among invertebrates. In general molluscs and arthropods have well developed
eyes. Their efficient eyes and well developed nervous systems are able to discriminate
visual signals. Hence these animals communicate a lot through visual signals. Many of
these animals can distinguish shapes and movements.
In nocturnal animals and deep sea inhabitants, communication by visual signals is limited
to flashes of light or bioluminiscence.(Fig. 15.6 A,B).
Visual signals are mainly used for
i. frightening the predator
ii. luring the prey and
iii. for attracting a mate
The advantage of visual signals is that if the receiver can see the signal, the sender can be
located immediately. But visual signals cannot be used at night except for
bioluminiscence. They cannot pass many physical barriers unlike auditory and chemical
signals. Another great disadvantage is that the sender can easily be detected by its
predator.

Fig. 15.6: Bioluminiscent invertebrates: a) c),prldina (crustacean), b) Firefly is actually a beetle. Tbe
white abdominal segments with photocyts are supplied oxygen by the abundant trachioles .The
oxygen dxidises the luciferin +ATP in the presense of Mg ions and enzyme luiciferase.
There are various examples of communication through visual signals among the
invertebrates,.which have been listed for your knowledge you may observe and add more
examples to this,list.

-
1. To startle the predator, some moths the eye like spots on the hind wings (Fig. 15.7).
I

3.The large coloured eye spots on the caterpillar of the swallow tail butterfly gives it
Fig. 15.7: Moth with eyespots on
wings. .
., the appearance of assnakehead (Fig. 15.8) which scares the predator.
3. The ant Crematogaster adopts an alarm and defence posture by curving its abdomen
upwards which warns and sends alarm signals to the predator, of its toxic sting (Fig.
15.9).

<n
 Behavioural Patterns

Fig. 15.8: Snake head caterglar.

Fig. 15.10: Lama and adult monarch butterfly showing aposemalic colouration warning predators ofthe
presence of cardiac glucosides in their bodies.
4. The monarch butterfly Danaus and its caterpillar &e brightly coloured. The bright
colouration keeps predators away as they associate bright colours with the presence
oftoxic cardiac glycosides (Fig. 15.10).

5. The reef squid Sepioteuthis sepioidea manipulates its chromatophores to exhibit

colour patterns which, along with the body posture &d position of tentacles, a l m s
the predator.

6. The crustacean Cypridina (Fig. 15.6 a) comes to feed at night and releases
bioluminescent substances (luciferin) into water. The bioluminescence attracts the
prey.

7. Female of the polydhaete Odontosyllis emits green light continuously during their
mating period. Circles of light attract male worms. They in turn respond by emitting
intermittent flashes of light. Eggs and sperms are deposited in water, resulting in
fertilisation.

8. The male and female fireflies of @e genus Photinus (15.6b) have bioluminescent
organs. They emit flashes of light. These serve as visual signals for mating.

Deceit in visual communication -.

Colouration and mimicry are antipredatory behavioural patterns. Kallima the dead leaf
butterfly, which resembles (mimics) a dry leaf when it rests folding its wings (Fig. 15.1 I),
is an example of mimicry for it mimics a leaf in this case. .

The leaf insect, is green in colour and shaped tike a 1ea.f;the spotted leaf katydid
resembles the leaves on which it feeds; the stick insect resembles the twig on which it
rests (Fig. 15.12 a and b). There are many other examples of mimicry. These insects
 .daptation and Behavioura~
Pattern canvf y visil:vl!\i that they are inedibll; or inanimate. The deceiving visual signals protect
these inst1c:> itom the predator.

Fig. 15.11: a) Kallima looks like a dry leaf when it folds up its wings (1)Note the fine lines on its wings
suggesting the veins of a leaf, b) a preserved specimen of kdlima with its wings open.

Fig. 15.12: a) The spotted leaf katydid is exactly like the leafon which it rests, b) Stick insect can hardly
be noticed. Note the spines on the body of the insect that resemble the spines on the plant on
which it rests.

Some bioluminescent fireflies even deceive the prey (species of tireflies belonging to
another genus) through visual signals. For example, the predatory female of the firefly
Photuris answers the courtship flashes of light given by the males of the firefly Photinus
species. If the Photuris female succeeds in attracting male of the species, she grabs him,
kills and eats him.

15.4.2 Mechanical Signals

Mechanical signals of communication include the tactile signals in which communication
is through touch and auditory or acoustic signals in which sound is generated through
a movement of wings or special stridulatory organs.

A,good example of tactile signal is the food begging behaviour of the larvae of the
Formica ant. When touched by an adult worker ant's mouthparts or antennae, the larva
tries to make contact with the head and mouthparts of the worker by tapping its mouth
parts with the mandible of the worker as though begging for food. The worker,
 Relirrvioural Patterns '

immediately regurgitates a food droplet and offers it to the larva. When an adult Formica
ant taps another worker with its antennae, the signal is to stop moving about.

Auditory Signals from the sender reach the receiver through airborne sound waves.
Special sound producing organs and organs of hearing are common among Insects So.~!;d
signals are effkient means of advertising the presence of the signaller. They inay thus be
used as (i) courtship songs; warning devices, advertising temtorial claims. Sound can
pass round comers and a great deal of information can be sent by modifying frequencies
of sound produced. The disadvantage of sound signals is that they can easiiy be exploited
by predators. For example the male cricket sings to lure the female into hrs buwow. but
the predator receives the sound of his song, locates and preys upon the cricket.

Among invertebrates, insects as a class have very effectiv, sound comrnurrication system.
Everyone is familiar with the buzz-of the bee, chirps of crickets, loud noise of cicadas,
and buzzing of mosquitoes at night. In insects sound is usually generated by frictional
methods, rubbing specialised surfaces of two parts of thz body, like thorax, abdomen,
wings and legs. This is called stridulation. Sound reception is by hair like
mechanoreceptors distributed in connection with the tympana1organs.
, Scraper 1

Left .
. teimen , Right tegmen

Projection
Fig. 15.13: Stridulatory organs of crichet.

Courtship singing is a daily programme with male crickets and begins in the evening.
Wings are moved so that a ridgeon the edge of one elytrn (forewing) forming the
"scraper" moves across the toothed files on the underside of the other elytron (forewing)
like a bow across the strings of a violin creating various patterns of sound (Fig. 15.13).
Song patterns are specific for a species as also specific m their meaning. The commonest
song is the calling song to beckon and guide the sexually receptive females into their
burrows. Once the couple meet, a courtship song ensues. After mating, the male sings a
triumphal song.

The courtship movements and songs of Drosophila are likewise, characteristic. The male
touches the female and extends its wings. It then moves the body and makes a
characteristic sound which the female of the same species alone can r-cognise by its
antennae. If the female 'is not ready for mating, it generates a buzz with its wings and the
male turns away.

15.4.3 Chemical Signals

.
Certain chemical compounds are secreted by animals which act as signals for olfaction
(smell). These serve to communicate with other members of the same species or with
other species. Chemical signals serve a variety of purposes. dome of these, especially in
moths, act as sex attractants. In social insects, chemical signalling is very well developed
and is used for various purposes.

Chemical signals diffise.through the environment and the speed and direction by whicb
the scent of chemicals disperses depends on the wind. But the advantage lies in the fact
that a small moth may not be seen or heard at long distance. But the species specific scent
of the chemical molecule can be detected by the mate several kilometers away. Chemical
signals are persistent and continue to signal even when the sender moves away. They are
Adaptation and Behrvioural
Pattern
effective even in the dark, can pass round obstacles and disperse over great distances.
Such chemicals released as signals for commmunication are called pheromones.

pheromones are chemicals which are released by one organism and produce a response in
conspicifics (members of the same species). Pheromone communication has been
discovered in unicellular organisms as well as in nearly all animal groups. Many kinds of
chemicals act as pheromones and many types of signals can be conveyed by pheromones.
Pheromones generally act as releasers, (signalling pheromones) stimulating the recipient
to show a very specific but transitory type of behaviour. Sex phenomenes of insects
generally belong to this category. Some pheromones act as primers and evoke slower but
longer lasting physiological responses from the receiver. An example is the chemical in
the "queen substance" that inhibits ovary development in the worker honey bees.

Communication in social insects by pheromones.

Trail marking pheromones are used by some insects to find mates, communicate
information on the location and quantity of food and to ensure that migrating grsups
retain their integrity. Amongst social insects, for example in ants, trail marking
pheromones are deposited on the ground as foraging workers return to recruit other
workers to a food source (see Fig. 15.23). Most trails fade away soon unless reinforced
continuously. The chemical nature of relatively few trail marking pheromones is known.
In termites and some ants they appear to be long chain acids, alcohols, aldehydes or
hydrocarbons.

Pheromones also help in recognition of members of the social group in insects. Alarm
pheromones warn members of a species about foreign intruders. If termites from another
nest enter a colony of termites, soldier members release an alarm pheromone which
attracts other soldier termites who chase away foreigners.

The nature of alatm pheromones is varied but specific for each group. In honeybees, the
pheromone is released from thestingshaft that is buried inside the invader. This attracts
other bees to attack.

'ln a honey bee colony, the queen secretes the "queen substance", by the mandibular
glands. It has two components oxydecanoic acid and 9 - hydroxy decanoic acid, which act
synergistically on the gonad development within the workers and their construction of
special cells in which larvae would develop as queens. These primer pheromones inhibit
the workers from raising a new queen and also inhibit the development of ovaries in
workers.

Sex attractants or sex pheromones in moths

Virgin Female silk moths have special pheromone glands whose secretions act as
chemical sex attractants. Males "smell" these chemicals with their large bushy antennae
with numerous sensory hairs functioning as olfactory receptors. These receptors are
highly sensitive to Bombykol, the pheromone produced by the female silk moth. The
female rests at a place and releases her pheromone, a little bombykol, which travels
downwind and reaches the male antennae. Immediately the male gets stimulated and flies
upwind to find the female. He moves randomly till he is close to the female flying
towards her by the smell of bombykol, finds her and mates with her. p i s is clearly then a
releaser pheromone.

In the gypsy moth, a similar releaser pheromone, gyplure is involved in bringing the two
sexes together.

15.4.4 Communication Among Honey Bees - g he Dance Language

The Austrian ethologist Karl von Frisch received Nobel prize for his research on
communication in honey bees. ,

The honzy bee, Apis exhibits social behaviour. Worker bees collect enormous quantity of
\
pollen arld nectar to feed the the colony mates. This is accomplished through co-operative
effort whereby the worker bee who discovers a rich source of pollen and nectar, comes
back to tfie hive and colnmunicates the informition regarding the source though specific.
dancing patterns, to the other workers. For instance; when a forager bee finds a garden of
n -...--- c-:-~-.
-I--- &I.- L:..- --.. ...:+L:, ,A : - ~,
,,,,ccn ,,
t
,, :+ ,P.c~.....~ the -.,..
..A
 Behavioural Patterns
dance (Fig. 15.14 a) at the hive. In the round dance, the bee turns in alterndte circles to
the left and right. The bee penetrates the hive and runs in a circular fashion excitedly.
Other workers follow this worker bee. 'The dancing bee ultimately regurgitatesthe nectar.
From the taste and the scent picked up from the dancing bee, other worker bees locate the
food sourde.

If the food source is beyond 50 meters from the hive, the bee performs the waggle dance.
'This is a modified round dance, which incorporates in it a straight run. During the straight
run, the worker kaggles (moves) its abdomen from side to side. Greater the distance to

/I the foraging site, longer is the straight run component in the waggle dance (Fig. 15.14 b).
The waggle dance thus indicates the distance and direction of a nectar source.

Fig. 15.14: Honey bee workers dance in different patterns to communicate the distance of the food from
the hive to the workers.
Sickle Dance is also another modification of the round dance, and is performed by the
Italian bee. She moves to make a flattened figure of eight, when the source is at
intermediate distance from the hive. The opening of the sickle faces the source of food
(Fig. 15.14 c). A dancer is always followed by her hivemates.

SAQ 3
1. Define biological communication.
'

......................................................................................................

2. What is ritualisation? ,
......................................................................................................

3. What do 'fixed action patterns' mean?

......................................................................................................
4. Name aoy two purposes for which
a) visual signals are used.
b) auditory signals are used.
~oaptationand Behaviooral
Pattern

5. Which head zppendage of the insect generally has sex pheromone receptors.
......................................................................................................
..................... ...............................................................................
7. Wli.li !.,: tksc:: siy-.ii;l:a&e of (a) round dance (b) waggle dance in honey bees.
.............................................................
...............‘.............. i

- .- --- -- - - -

15.5 C . O-U R ~ ~ X I P - B E H A V I O ~
---
Specid1l~i"l: h"lia\~j~,ar;lpatterns of animals ~OF-attractingthe opposite sex for
reprolil.ct, ,e t~rlrp~,, 'rl is termed courtship behaviour. These behavioural patterns may
be in rr~eh r : r ~~f diiplay of bright colouration or accessory morphological structures, or
irk the forlo of offe-ing nuptial gifts, or making specific gestures in front of the opposite
?ex rcsurtsili,- behaviour culminates in mating unless the prospective mate refi~sesto be
cat 8s:;. n ;v7 '.; ti,: courtship behaviour.

1 1 - ' .:male which shows various kinds of elaborate courtship behaviour,

,.-,.? A - rrange, to attract the female. The female rarely shows courtship behaviour,
r .as ro his overtures. Choice of mate seems to be her privilege. Interactions
'I. +. 1 - exes cease with copulation.

a " ':, Need for Courtship Behaviour

" ..::.!,hip behaviour serves the following purposes.
: ':ynchronisatipn and orientation of sexual partners.
%
~:c:issfulfertilisation, sperms and ova have to be discharged by the two sexes at the
P -Ie. The male and female of the same species have to be properly oriented. The
- .tj behaviour of the male attracts the sexually mature female ultimately
,.:. .ir\ating in fertilisation or inseminati~n.

2, Persuasion
Tile malt exhibits courtship behaviour patterns and the female examines and decides
about the fitness of its mate based on these patterns. More often than not the behaviour of
the male persuades her to accept him as the mate.

3. Mating with conspecifics

Courtship patterns are species specific. The female of a particular species can recognise
the courtship behaviour of a male belonging to her own species. This ensures mating
between members of the same species.

4. Reproductive isolation
As a consequence of specificity of courtship behaviour, mating with members of another
species (interspecific mating) is prevented. Courtship is thus a means of reproductive
isolation. Courtship patterns that prevent interspecific breeding and hybrid formation is
termed ethological isolation.

15.5.2 Sex Differences in Courtship Behaviour

The behavioural pattern9 of males to woo females are diverse and often strarige. No less
strange is the choosiness that females exhibit. Why males take the initiative to court
females and why females sometimesreject males following his courtship may be
attributed to the difference in the investnient of either parent in the offspring in ien,\s of
time and energy. Such differences in the reproductive tactics of male and female r n ~ y
have helped to create a kind of natural selection termed sexual selection. Sexual sslection
 occurs when individuals vary in their ability to (a) compete with others for mates or (b) to
attract members of the opposite sex. Sexual selection involves.

1. competition to gain access to mates, often but not always among males. (i~trasexual
competition) Males compete with other males of the species for a mate. This
probably led to the evolution of aggressive. behaviour which males at times show.
2. competition to choose the best possible mates mostly among females (intersexual
selection)
Mate preference exhibited by females probably resulted from practical value of:
i) what kind of genes the male has to offer, and
ii) if the chosen mate is of any other value.

The theory of sexual selection was first suggested by Charles Darwin (1871). He
recognised that the reproductive success depended on the ability of individuals of one sex
to locate the op~ositesex, fight with other conspecific members of its own sex and win
over a mate Competition for mates among the individuals of one sex was thought to be
responsible for the evolution of the characteristic features of that sex, including the
beha1,loural mechanisms involved in selection of mate.

It is now known that reproductive success .or ability to produce large number of viable
offspring is more variable with males than females. This is due to difference in
contribution of reproductive resources by male and female to the zygote and hence to the
developing embryo i.e., parental investment of the male in the offspring is less than that
of the female.
I
Jn case of females parental investment is more because:

I
(i) the female gamete, egg, is larger than sperm, the male gamete,
(ii) generally only a few eggs are liberated by the ovary at a time,
(iii) egg contains nutritive substance such as yolk for the developing embryo,
(iv) females, in many cases, spend extra time after mating for parental care.

All these activities take time energy and materials and consequently limit the
number of young that can be produced. On the contrary:
(i) males release, larger number of sperms,
(ii) sperms are much smaller than ova,
(iii) males contribute only their genes through sperms, but no nutrients,
(iv) in most invertebrate species, males do not do aaything beyond transfer of sperms to
the female.

Sexual selection should produce different degrees of variation in reproductive success of

males and females. Some males may mate with several females while some males may
fail altogether. Females need only to mate once or twice for they can receive all the
sperms from one or two males to fertilise the relative!^ few eggs they produce. Thus
reproductive output of males will depend on how many females it can inseminate and
hence vary among males. Females, on the other hand will produce more or less the same
number of offspring.

Females, the choosy sex, prefer certain attributes in males which leads to mate selection.
The preference differs in different species. Some of these attributes are:
large size,
visual, mechanical and chemical signals or display by the male,
a offer of nuptial gifts from the male,
a superior fighting ability and aggressive behaviour of male,
e ability of the male to guard the mate,
a attempt of a inale to become socially dominant.

In certain species, males force the female into copulation.

Let us now examine some examples of male courtship behaviour patterns used for
attracting conspecifio females.
Adaptation and Behavioural
Pattern 15.5.3 Visual, Mechanical and Chemical Displays
The crab Uca waves an enlarged chela (Fig. 15.15) to attract the female. Horns and
snouts of certain male beetles have a similar effect. Male crickets and locusts stridulate to
produce species specific sounds. The sounds differ in frequency and may mean mating
-- (see subsection 15.4.2 - for auditory signals).
receptivity, victory- etc., i

In scorpions, both male and female, indulge in a courtship dance during the monsoon
(Fig. 15.16). The male and female face each other, raise the post abdominal part high into
air, then move in circles. The male holds the pedipalpi (clawed appendages on the
cephalothorax) ofthe female with his own pedipalpi and both intertwine with post
abdomen above them. Then they walk forward and backward and in rings. While moving
in a ring, male retreats and drags the female along. The courtship play may last for hours
or even days. The male digs a burrow. Both enter and mate. The male scorpion deposits a
spermatophore and moves the female so that her genital area is above the spermatophore.
Sperms enter through the female genital opening. The female eats up the male after
mating!

Fig. 15.16: Courtship dance of scorpions.

In some spiders, the female sits at the centre of its web. The male approaches the web and
I
plucks a thread of the web at a specific frequency. The female reduces her natural - ,

i
aggressive form. But if the male, by chance, tries to attract the female of another species,
it is killed.

Pheromones also act as sex attractants (see section 15.4.3).The pheromone is released by
eversible glands at the tip of the abdomen of unfertilised adult female moth. Antennae of
male moths have olfactory receptors for these pheromones, and they fly towards the
female in a definite species specific pattern and mate. Male of butterfly Danaus gilippus
brushes, pheromone laden 'hairpencils' everted from the tip of his abdomen on the
female. The female is stimulated to mate.

15.5.4 Nuptial Gifts

Gifts offered by males to court females may either be in the form of (a) food or (b)
spermatophore. The female black-tipped hangingfly (Hylobiitacusapicalis) chooses for
mating, the male which offers a large edible nutritional gift. The gift is a dead insect (Fig.
15.17). If unpalatable lady bird beetles are offered as gifts, the female rejects the male.
Duration of mating depends on size and quality of the prey which is gifted by the male. If
the gift is small, female unhooks itself fmm the male to give up mating without accepting
sperms.

Among katydids, relatives of grasshoppers the male offers "spermatophore" (a packet of

sperms). The spermatophore is rich in protein and after copulation, the female feeds on it.
If the male offers a small spermatophore, the female does not permit entry of sperm into
her spermatheca (sperm storage organ of female).
 lebavioural Patterns

FigIi5Tl'I: ale hanging fly carrying captured prey as nuptial glR for the mate.
In the Mormon cricket, Anabrus simplex the male attaches a two-part spermatophore to
the mate's genital opening during copulation (Fig. 15.18). The male separates after
copulation and the female feeds on one part of the spermatophore the spermatophylax
leaving the other part containing the sperms in place. Larger the spermatophore, longer
she feeds and more time elapses with more sperms entering the female spermatheca from
the sperm ampulla. When the first part is eaten she eats up whatever is left of the second
part the ampulla.

Fig. 15.18: Diagram of the posterior part of a female katydid sbowing male spermatphort after mating.
The spermatophore is made up of the ampulla containing the sperm and the spermatophylax
containing nutritional material.

The male balloon fly (Hilara sartor), an empid fly constructs silken balloons as nuptial
gifts (Fig. 15.19). A group of balloon canying males display collectively to visiting
females. The female selects one balloon canying male and accepts the balloon as a
precondition to mating.

Fig. 15.19: Males of emphids. Carrying silken balloons for mates.

Adaptation and Behavioura!
Pattern 15.5.5 Sperm Competition and Mate Guarding
Sperm competition and mate guarding is particularly important in insects, because female
insects can store sperms in an organ called spermatheca and use the sperm later.
Copulation by one male does not exclude successful copulation by other rival males. An
interesting example of sperm competition is seen in damsel fly Calopteryx maculata.
Some males defend a territory of one to three meter long stretch of the bank of a stream,
from which they drive away all other males. Females come to lay eggs on underwater
vegetation, the male courts her and mates with her before she oviposits. The male has
specialised claspers at the tip of his abdomen to grasp her. He bends the abdomen to form
a loop so that sperms are transferred to the penis. The receptive female swings her
abdomen and places her genitals on the penis. The penis is inserted by the rhythmic
movement of the abdomen. If there are sperms of another male stored in the spermatheca,
they are brushed out of the spermatheca before fresh sperms are deposited by the penis.
He then releases her but guards her while she lays eggs in his territory. If the male cannot
guard her, she flies away to another territory to mate with another male who repeats the
performance.

15.5.6 Alternative Tactics of Mate Competition

Not all males of a species compete for mates in the same way. In many species the males
often adopt alternative means of mating. For example, the scorpionfly Panorpa employs
three tactics for mating. (1) Some males defend dead insects which attract receptive
females that feed upon them. (2) Others secrete salivary material on leaves and wait for
females to come and consume them. (3) Still others, the smaller males, do not give any
gifts to the female but force her into copulation.

Not only scorpionflies but males of other insect species also force females to copulate.
The assaulting males especially pounce on females while they are laying eggs. Some ,
species may have the potential to behave in any of the available ways depending on the
prevailing conditions.

Mate choice by males

Uptill now we have examined examples where mate competition occurs between males
and mate choices by females. K and when parental investment is made by males rather
than females, the role of the sexes can get reversed. In high density population of
Mormon crickets, where the food becomes scarce, when a male gathers enough resources
to,produce a spermatophore, he climbs up a bush and begins to sing. Hungry females are
attracted and when more than one female arrives they compete to copulate. The male
accepts only that female that is significantly heavier than the rejected females. Such sex
role reversals are rare and show that when parental investment is greater by the male then
competition for mate arises amongst the females.

15.5.7 Rejection and Deceit in Courtship

. Females-ofcertain species have specific signals for rejection of the mate. The female
ground beetle Plerostichus lucublandus sprays the rejected male with toxic liquid and the
male may be in coma for a long time.

Female butterfly Colias is unreceptive during egg laying and Ecourzed, flies very high
when the males run away, she descends and oviposits in peace.

Deceit in courtship is shoh n by some males of the hangingfly Hylobittacus. A male

without a nuptial gift poses a female, takes a gift from another male which mistakes
L-:

him for a female and gives-thegift to a female.

SZAQ 4
1. WMt is courtship behaviour?
......................................................................................................
.......................................................................................................
. .

2. Whatis the major dif-ference between the courtship tactics of a male and female?
.......................................................................................................
3. What are some of the nuptial gifts offered to the females by males? Behrvioural $atterns

15.6 SOCIAL ORGANISATION IN INSECTS

The term social organisation refers to populations or groups and not to individuals and
defines how members of a species interact with each other. Social organisation among
non-chordates is best examplified in insects. Insects such as ants, wasps, honey bees and
termites survive through cooperative hiving. A number of individuals live as a society or
colony. Work such as, building the abode, procuring food, cleaning etc., are didded
among members of the society. Thus when a number of individuals of a species co-
operate and live together sharing the duties of life they are said to exhibit social
behaviour.

15.6.1 Advantages and Disadvantages of Social Behaviour

Social life has several benefits. A collective defence strategy can easily drive away
predators. Tlia young ones can be better looked after and protected in a colony of
individuals. Social life has its disadvantages too. The following table (Table 15.2) shows
some ma,jor advantages and disadvantages of sociality.

Table 15.2: Benefits and cost of social organisations

ADVANTAGES DISADVANTAGES

I. Defence from predators 1. Competition within the group

(i) One member detects predator, can warn Members of the society may compete amongst
and save other members. themselves for food, nest material, nest site and
(ii) A joint attack can easily repel the enemy. mate.
(iii) The sight of a colony of prey individuals
has a better chance of scaring the predator.
2. Collection of food 2. Risk of epidemic and parasites
(i) Food resources may be spotted by one Increased risk of infection by contagious
member and others are led to the food diser '5 and parasites.
source. This saves energy expended on
searching foraging grounds.
(ii) gathered Food is shared by the young, the
aged, the infirm apart from others.
3. Care of offspring
3. Exploitation of parental eare by conspecifics
:mproved care of offspring through comnlunal
feeding and protection from enemies. The lazy members may leave the care of their
* progeny to others.
4. Dcfenee ofoecupied territnry
'sk of loss of one's progeny
Intruders from other groups of the same species
looking for the same habitat and food are driven , ?,.eased risk of progeny being killed by
away througli a joint effort. conspeciflics.

15.6.2 Characteristics of Social Insects

You have already read about polymorphism in highly organised insect societies of
termites (Isoptera); ant and bees (Hymenoptera), in Block - I, Unit- 7 of this course. In
these societies we find such remarkable adaptation to social life that they are not called
just social but elisocial insects.

All truly eusocial insects possess three traits.

(1) many individuals of the same species cooperate in caring for the
(2) there is division of labour with sterile individuals working on hehalf of fecund
(fertile) individuals;
(3) there is an overlap of at least two generations in life stages capable of contributing to
colony labour, so that offspringsBssist parents during some period of life.

61
A d a p t i o n and Behavioural
Pa~,ni
You would recall that these societies are divided into castes, such as soldiers, workers and
reproductives which are morphologically different from one another and each has a
separate role to perform in the colony. Eusocial insects show cooperation.and altruism in
an incredible manner. They build elaborate nests where temperature and humidity are
kept constant. Group foraging allows sharing and harvesting of widely spread food
resources. Communication involved in foraging and maintaing the colony structure are
complex and the nest dr hive is fiercely protected as in the case of honeybees. When the
honeybee worker stings, she leaves her sting along with the poison gland and a set of
contracting muscles behind as she pulls away. As a result she continues to pump poison
in the victim even though she herself dies. Fig. 15.20 shows the sting apparatus which
consists of the muscles, pheromones that attract other guard bees and venom sac. The
evolution of nonreproductive castes showing a readiness for self sacrifice is intr.iguing. In
the following subsections we describe some interesting social organisations in insects.

15.6.3 Social Wasps

Protractor muscle
Among wasps eusocial behaviour is limited almost entirely to family Vespidae. The paper
Retractor muscle wasp Polistes, with its painful sting is a familiar example.

I
Sting
These wasps build small open nests beneath roofs of houses, barns, garages and other
Barbs
outhouses. They build annual colonies. The colony consists of three castes: the queens
Fig. 15.20: Sting apparatus of (fertile femates); the workers (nonfertile females) and drones (males). Reproductive
the honey bee. The females (queen) emerge from cells of their papery nest late in the breeding season, mate
barbs of the sting and hibernate in winter. In spring, these females emerge and construct either alone
get lodged in the (foundress) or with co-operation of other similar queens (auxiliary queens) a nest. The
victim's flesh. As a
result the sting faundress queen lays eggs and rears the first group of her larvae on macerated insects. All
apparatus tears out of these become workers. These workers thereafter, maintain the nest. They add new cells
of the bee's body to the nest, look after the queen henceforth, forage and bring food (Fig. 15.21). The
and she dies. The foundress remains in the nest thereafter, and'lays eggs. The colony now grows rapidly as
muscles left back
with the apparatus more and more workers emerge. At the end of summer, some new queens and a number
continue to contract. of males emerge. These leave the nest, mate and hibernate. After winter they build new
pumping more nests. The old queen and the old workers die and the original colony disappears.
venom into the
victim.

Fig. 15.21: A social wasp Polides. One of the adults shown is the queen the others are workers that aasist
in building the nest and caring for the young i.e. the larvae inside the cells.

15.6.4' Ants
The ants belong to the family Fonnicidae (Order Hymenoptera). Common ants are the
small black Monomorium and the large black Camponotus. Formica, Lasius are other
genera. They build nests in cavities in plants, in the soil, or under stones or logs. The
tropical ant Oecophylla smarsedina weaves leaves of trees into a nest with threads of silk.

All ants are social and polymorphic YOU have already learnt in unit-7, Block 1 of this
cou~sethat there are mainly three castes. (1) wingless and sterile females are either
soldiers having huge'head and well developed mandibles, or ordinary workem. (2) '
winged fertile females which lay eggs (queens) and (3) winged fertile males which are
short lived and die after mating.
Fig. 15.22: An underground harvester ant colony. The nest eonsistsof various chambers and connecting
I
tunnels. One chamber houses the queen. There are'severalnurseries where workers take earn
of the young. Harvester a& also bave storage chambers for seeds.

The underground nests of ants are called formicaria (Fig. 15.22). After mating-in air,
females dealate (drop their wings) build a new nest and lay eggs in cells or chambers.
Initially the queen performs all the work alse. But once the initial clutch of eggs hatch
into workers they take up the work of the colony. Then the queen is fiee to concentrate on
egg laying and growing.

Ants have glands for secreting trail laying-pheromones (Fig. 15.23). These pheromones
help in communication and troops of ants can be seen marching along the trails laid.
Certain ants also have beetles and other insects living in their nests as guests. Some ants
even cultivate h g u s garden.

Fig. 15.23: Fire ant worker laying a trail by releasing phtromone along its extended sting. The sling is
touehcd to the ground periodically.

15.6.4 Honey Bees

Honey bees belong to family Apidae (Order Hymenoptera). There are three common
species of honey bees in our country namely the giant bee Apis dorsata, the little bee Apis
jlorea and the Indian honey bee Apis indica. Honey bees are socially very well organised.
About 10000 to 16000 individuals may live in a honey comb. Recall from Unit 7 of this
-
course that each colony has only one queen the fertile female, about 500 to 1000 males
(drones) and the rest are sterile, female workers. The queen, the drones and workers are
all winged. The functions of the colony are divided among these three castes. The queen
is the mother of the colony. She remains in the live except during nuptial flight when she
mates with a drone in air.

The workers build the honey comb with wax which is secreted by wax glands on the
abdomen. They collect nectar from flowers, for which their mouth parts are suited. Pollen
is an important part of the food of the larvae and adult bees. The legs of the workers are
modified for collection of pollen. The hind legs (Fig. 15.24) have tibia fringed with long,
curved hairs and these forms the pollen brush. The spa& encloseddy the hairs form the
Adaptation and Behavioural
pollen baskets. The legs have spurs and bristles to clean antennae and pollen sticklng to
Pattern
body. The workers have a sting ;t the abdominal tip into which leads a poison gland (see
Fig. 15.20). The queen has a very well developed sting to drive away rival queens.
Drones do not have sting.

Fig. 15.24: Legs of honeybee workers that are especially modified for collecting pollen. The hind leg has
long hair forming a pollen basket. The joint between tibia a.nd metatarsus of front leg is
modified as an antenna cleaner.

15.6.5 Termites
All termites are social insects, and belong to the primitive order Isoptera. They are
popularly called white ants. Examples are KalotermesflavicoNis and Rettcrrliierme~
luc~fugus,Miacrocerotermies sp. The immense damage done to wooden components of
houses and fivniture by tennites is well known. Termites are also polymorphic (refer to
page 164-165 of unit 7 of this course for a detailed description of termite castes) and live
in communities leading a social life, in galleries. Most termite species live in soil inside
mounds of earth known as termitaria (Fig. 15.25). Termites feed on wood which they
digest with the help of symbiotic flagellate protoLoans in their intestine. Some tennites
Fig. 15.25 :Termite mounds are
have fungal beds or gardens in their nests specially prepared by workers.
elaborate constructions.
The nests ofAftrican
termites MacmIerm?s SAQ 5
bellicosus can reach
skyscraper proportions. i) Name the different castes found in
The nests have rock hard (a) honey bee
water tight walls of earth (b) tenites.
and plant material bound
together with the saliva
and excreta. The walls
help to insulate the nest
from extremes of
temperatures.
ii). What is meant by euiocial insects?
......................................................................................................

1 5 7 PARASITISM
- - - - - - -

In the earlier section you learnt how eusoeial insects show cooperative behaviour and
often go to suicidal lengths to protect their colony from invaders. Inthis section we shall
learn about another behavio~ar- parasitism - that is the other end of the spectrum. While
so,c!cialbehaviour is an intraspecific behaviour. parasitism is a regular and c!ose
interspecific association in which one of the animals, tl~eparasite is benefitted especially
nutritionally. The parasite in other words, lives at the Cxpense of the host. The parasite is
usually smaller and weaker. It lives within or on the surface of body of the larger and
stronger host. The parasite gets its food from the host.
Parasitism cvolved separately in many major groups of animals. Platyhelminthes, Behavioural Patterns
Nemathelminthes, and Arthropoda are the main non-chordate groups that show examples
of parasitic behaviour.

15.7.1 Types of Parasites

Parasites may be classified into the following types. Ectoparasites are those that attach to
the surface of the host (eg. leech on cattle).

Endoparasites live inside the body ofthe host. eg. Round worm (Ascaris) and tape worm
(Taenia) live inside the human intestine.

Facultative parasites may live without a host.

Obligatory parasites spend at least part of their life in the host to complete their life
cycle eg., Taenia. They cannot complete their life history without the host.

Apart from Platyhelminthes and Nemathetminthes, parasitic species are common among
Arthropoda (many among crustaceans, arachnids & insects). You would come across
more exarnples of arthropod parasites from relevant sections, in the next unit Harmful and
Beneficial Non-chordates.

15.7.2 Effects of Parasitism on Parasites

Majority of ~lat~helminthks and Nemathelminthes lead a parasitic life. The greater the
degree of parasitism, the more is their departure from normal morphology and
physi,ology. All these deviations, however, are adaptations to a parasitic life. You may
recall here the adaptations discussed in the relevant units in the course.
The main parasitic adaptations are:
1. Loss of surface epidermis and its replacement by a tegument which plays an
important role in the physiology of the parasite, is a common feature among parasitic
platyhelminths. In nematodes, the cuticle is impervious. There are shells around eggs
and cyst walls around larvae, for protection.
2 . Presence of organs of adhesion to fix to the host. Hooks, claws, various types of
suckers and in certain cases adhesive secretions. Consequent on the development of
adhesive organs there is loss of motility. Parasites remain firmly attached to hosts.
3 . Loss of sense organs: Progressive parasitism is accompanied by corresponding loss
of sense organs and a concomitant degeneration of nervous system.
4. Simplification and eventual loss of digestive system : Parasites depend on hosts for
food. In tapeworms there is a complete loss of digestive system but flukes possess a
mouth, a suctorial pharynx and a blindly ending intestine. Tapeworms absorb wholly
digested food of the host, throt ;h the tegument.
5. Reproductive system of parasites is highly complex. They show excessive capacity
for egg production. Tapeworms show multiplication of sex organs along the strobila
(proglottides of the body) for further augmentation of egg production. Many
parasites are hermaphrodites to ensure fertilisation.
6. Life cycle of parasites is complicated. Flukes have stages of life cycle passing
through one to three intermediate hosts. To suit this, the parasites often have a
number of larval stages. These often show polyembryony giving the animal a further
reproductive advantage, resulting in multiplicat~on.

SAQ 6
1. Define parasitism.
.....................................................................................................

2 . What are the usual organs of attachment of parasites?

.......................................................................................................
3. Why is the digestive system of parasites inco~npleteor lost?
........................................................................................................
Adaptation and Behmvioural ---- .--.----- - ----
Pattern 15.8 SUitIMARY

In this unit you have studied:

Taxis and kinesis involt. ijiovri~:clitsin response to environmental stimuli. Taxis is a
directional movement of thc organism. Taxis can be classified according to st~muli
into:
- phototaxis in response to light
- thermotaxis in response to temperature
- thigmotaxis in response to mechanical contact
- chemotaxis in response to chemicals
- rheotaxis in response to wind and water currents
- galvanotaxis in response to electric currents
- geotaxis in response to gravity.
Kinesis is a non-directional movement.
Biorhythms are behavioural activities performed at regular intervals. Daily rhythms
in which activities follow approximately twenty four hour cycle are circadian rythms.
Activit~esregulated by tides in marine animals are termed tidal rhythms while those
controlled by phases of moon are lunar rhythms. Rhythms shown once a year are
called circaannual rhythms. Biorhythms are exogenous when rhythmic activity is
controlled by external stimuli such as by photoperiod. Endogenous biorhythms are
regulated by an inbuilt biological clock.
Biological clocks are internal mechanisms which can measure time. They ars set to
cyclical events such as day and night. In case of change in the environmenta. -vcle
that is, such as when kept in continous darkness, the timing of biological clock
changes slightly and this is termed free running of the clock. If the organism moves
to adifferent photoperiod such as when animals travel from one part of theworld to .
another, the biological clock sets to new environmental conditions and this is termed
entrainment. Stimulus which sets the time of biological clock is called.Zeitgeber.
Animals communicate with each other through signals. Various signals are visual, ,
mechanical, auditory, chemical etc. and indicate availability of food, predator around
mate etc. When communication signals occur in regular stereotyped sequences they
are fixed action patterns.
Some visual signals such as changes in body posture or specific colour patterns lure prey
or mate, or frighten or warn the predator. Tactile signals are employed by ants where
touch makes the ant regurgitate food for consumption by the larva. Auditory signals are
common among insects. Pheromones play an important role in chemical communication.
Honey bees employ "dance language" for communication among the members of the
hive.
Courtship behaviour helps to attract the opposite sex of the same species for mating
and reproduction. Courtship behaviour brings about synchronisation of mating
activity and proper orientation of the mates. Courtship behaviour is usually initiated
by the male and the female has the privilege of choosing the mate. Courtship
behaviour patterns include visual displays, production of sex attracting pheromones,
nuptial gifts.
Ants, termites, wasps and bees are some examples of social insects. All these four
groups show polymorphic castes with division of labour. The castes are broadly
similar among all these. But there are some basic differences also between
hymenopteran groups (bees, ants and wasps) on the one hand and isopterans
(termites) on the other. The most important of these is that the worker caste in
hymenopteran consists of adult females only, whereas in termites (isopterans) the
workers consist of both adult males and females. The immatures also contribute to
work of the colony. The advantages of social life are:
- collective defence from intruders and predators;
- co-operative collection and sharing of food;
- ccirnmunity care of young and sick.
The disadvantages are competition within the social group; risk of epidemics and
parasites; and exploitation ofaparentalcare by conspecifies.
Parasitism is a regular and close interspecific association in which the parasite is
benefitted at the expense of the host. Advantages of parasitism are: parasite gets
ready supply of food; shelter and other metabolites without any effort. Disadvantage!
are that the parasite has to produce large number of eggs so that at least a few will
find a host. Once it finds a host it depends on the host for its life.
 Behnvioursl Pnlterns
Adaptations to parasitism are:
- presence of impervious cuticle, cyst or shell,
.
- presence of hooks or suckers for attachment to host; loss of motility,
- loss of sense organs and poorly developed nervous system,
- simple digestive system or its absence altogether, accompanied by development
of a special tegument for absorption of predigested food in the environment,
- well developed complex reproductive system,
- capable of producing enormous number of eggs, and
- hermaphroditism,
- life cycle complicated with many larval stages accompanied with polyembryony
in each stage.

15.9 TERMINAL QUESTIONS

1. Enumerate the different types of biorhythm.. and explain with examples.
......................................................................................................

2. Write a short paragraph on "biological clock".

......................................................................................................

3. With two examples each, explain the different types of signals used for
communication among invertebrates.

4. Elucida~ehow insects employ pheromones in communication.

......................................................................................................
5. What does mate selection mean?
......................................................................................................
6. What is the significance of courtship display in animals.
......................................................................................................
7. What is social behaviour? numerate its merits and demerits?
......................................................................................................

15.10 ANSWERS
Felf-Assessment Questions
.
i) Taxis is directional movement in response to a stimulus.
ii) Kinesis is non-directional movement in response to a stimulus.
iii) (i) positive phototxis; (ii) negative galvanotaxis; (iii) geotaxis.
2. i) Behavioural activities performed at regular intervals.
i i) i - e; ii - a; iii - d; iv - c; v - b.
iii) In exogenous rhythm, an external stimulus triggers the behavioural pattern of an
animal at regular inte~als.In endogenous rhythm, an internal biological clock
triggers activities of an organism at regular intervals.
iv) Setting the biological clock in phase with the environmental cycle.
v) Nervous system.
3. i) Actionlsignal of an individual which alters the pattern of behaviour of another
individual.
ii) Behaviour patterns for communication established through evolution.
iii) Stereotyped sequences occurring at regular intervals.
iv) a. frightening predatorlluring preylattracting mate.
b. courtshiplalarm.
Sound made by rubbing surfaces against one another.
antennae
67
Adaptation and Bchavloural
Pattern
vii) a. to inform the workers that source of food is close by (within 50 meters from
the hive).
b. to inform fellow worker bees that source of food is far b yond 50 meters
from the hive.
4. i) Specialised behavioural patterns in animals for attracting the opposite sex for
reproductive purposes.
ii) Male exhibits visual displays, offers nuptial gifts, or fights with conspecifics to .
. attract female. Female chooses to accept or reject him.
iii) Food/spermatophores/silkballoons.
5. a) Honey bee - Queen (fertile female), workers (sterile females), drones (fertile
males)
Termites
1) King and queen (Primary reproductives)
2) Supplementary reproductives (males and females)
3) Workers (sterile males and females)
4) Soldiers (sterile males and females)
b) Insects with highly developed and defined castes.
6. i) Close and regular interspecific association in which one of the organisms derives
nourishment from the other. '
ii) Hooks, suckers, adhesive secretions.
iii) parasites absorb food which is already digested by host.

Terminal Questions
1. Refer Sub-sections 15.3.1
2. Refer Sections 15.3
3. Refer Sub-sections 15.4.3
4. Refer Sub-sections 15.4.3
5. Refersub-section 15.5.2
6. Refer Sub-section ' 5.5.1
7. i) A number of individuals of a species living together co-operatively sharing the
duties of life.
ii) Collective means of defence from predatorslintruders; cooperative collection and
sharing of food; care of young.
iii) competition within group; risk of loss of progeny; proneness to epidemics1
parasites.

Credits
Figs. 15.3; 15.12 b from The Illustrated Encyclopaedia of Animals (Exeter Books).
Figs. 15.6a and b; 15.7; 15.11; 15.12a from Living Invertebrates, Pearse and Buchsbaum
(Blackwell Scientific Publication).
Fig. 15.8a from The Unii :nd Diversity of Life, Starr and Taggart (Wadsworth
Publishing Company Inc.).
Fig. 15.10a and b from Biology, Arms and Camp (CBS College Publishing) 3rdEdition.
Fig. 15.15 from Animal Behaviour (Time Life Series).
Fig. 15.21 from Animal Behaviour John Alcock.
UNIT 16 HARMFUL NON CHORDATES
Structure
16.1 Introduction
Objectives
16.2 Parasitic Platphelminthes
Class Monogenea
Class Trematoda
Class Cestoda
16.3 Parasitic Nemathelminthes
Class Nematoda
Plant Parasitic Nematodes
Animal Parasitic Nematodes
16.4 Injurius and Harmful Arthropods
Arachnids of Medical, Veterinary and Agricultural importance
Insects of Medical Importance
Illsects of House Hold Importance
lnsects of Veterinary Importance
Insects of Agricultural Importance
16.5 Summary
16.6 Terminal Questions
16.7 Answers

16.1 INTRODUCTION
In Blocks 1 and 2 you would have got a fairly good background about the diversity that
exists within the Animal Kingdom. YoB have learnt about various groups of protozoans
and metazoans and also the characteristic features that differentiate them. The units of
Block 3 have given you the basic idea about their structural organisation and the
functions of the various structures. Most of these animals have a free existence, whereas
some form various types of associations wjth other living beings. Their association or
dependence on other organisms can be for shelter, transport or for acquiring food.
Sometimes the dependence may be deeper or more intimate. Parasitism is one such
association. It may be defined as a regular and close association in which the parasite
lives on or inside the body of the host, deriving nourishment from it but not necessarily
destroying it. In Unit 4 Block 2 you have already learnt about some of these parasites.
Many of these parasites are unable to survive independently on their own in nature. These
parasites share or rob resources of their host, thereby debilitating it. Parasites may inflict
injuries to the host by causing destruction of the tissues, which may result in disease on
the host. In extreme cases even death of the host occurs due to release of toxic
metabolites by the parasite.

Thus some non-chordate parasites directly cause diseases in the host, which may be
plants and animals including man. Some non-chordates called vectors are however,
indirectly harmful as they transmit various parasites or pathogenic agents from one host
to the other. 'These vectors are thus responsible for the spread of several dreadful diseases.
Let us, in this unit study from the point of view of humans some animals among the non-
chordates which are harmful in one way or the other to them, and therefore are important
from medical, veterinary or agricultural point of view.

When we look into the various non-cllordate groups, it emerges that major species
harmful to human health and/or economy come under the protozoans (animal protists),
platyhelminths, nematodes and arthropods. Of these the protozoans have already been
dealt with in a previous U ~ I L In
. the present unit we will study some selected members
from the Phvla Platyhelniinthes, Nemathelminthes and Arthropoda. *e will also discuss
how t h ~ parasites
~e or vectors complete their life cycle and the way they affect their
hosts. This knowledge \ '"slso give us an idcs as to how their propagation and spread in
nature could be brought unaer control.

Objectives
After studying this unit you should be able to:
name a few important helminth parasites of medico - veterinary and agricultural
importance.
Adaptation and Behrvioural descriSe the ways in which these helminth parasites complete their life cycle and are
Pattern
transmitted from one host to another,
give examples of some important arthropods which act as vectors or as pathogens of
plants, animals and man,
identify the ways in which the propagation and trans~nissionof the parasites can be
checked.

16.2 PARASITIC PLATYHELMINTHES

Before attempting to study this unit, you will do well to go through the relevant sections
of this course which deal with classifications (Refer to Block 2 of this course- sections
4.6 and 4.7 of Unit 4 and section 5.3 of Unit 5). In Phylum Platyhelminthes the classes
Monogenea, Trematoda and Cestoide (Cestoda) are parasitic during the whole or part of
their life cycle. The turbellarians are free living. The trematode and cestode species are of
veterinary importance and are also cause of concern in human health. Monogeneans are
ectoparasitic usually on lower vertebrates, especially on fish.

16.2.1 Class Monogenea

Most Monogeneans are ectoparasites especially on fish. A few flukes are also found in
the urinary bladder of various aquatic vertebrates.

The monogenetic flukes are small, less than 1 mm to 2 mm. They are (Fig. 16.1) circular
to spindle shaped and dorsoventraly flattened. Their body is composed of a head, trunk
and haptor (opisthaptor). The head lacks an oral sucker but has adhesive glands (head
organs). A complex well developed opisthaptor is present at the posterior end which helps
the flukes to cling tenaciously to their host and to withstand the flow of water over the
gills and skin of the host.

The opisthaptor of Monogeneans vary in different species and may bear hooks, suckers
and clamps, often in combination with each other. They may also have adhesive
secretions.

I'oster~or haptor
t i:h hooks

(8)

Fig. 16.1; S,)n~ccctoprra~,;"- rnonclgeneans. (a) Cyrodacr)ltus,r fish parasir - is a serious pest in fish ponds
ant1 lirtchcries where fishes ;Ire crowtled. (h) Doctytogyrus vaslrrinr i s ectoparasitic on gills of
fresh water fishes and is a serious pest in fish hatcheries.

There is no intermediate host in monogeneans and one egg gives rise to only one adult
worm, hence the name monogenea - 'one generation'. The egg hatches into a free
swimming ciliated larva-the o:~comiracidium-thatattaches to the host with its opisthaptor,
and metamorphoses rapidly into an adult.
 Harmful Non Chordates
Damage: The monogeneans become serious pathogens only in the'case of fish farming
where their hosts the fish are crowded together. Dactylogyrus is an ectoparasite in gills of
various fresh water fishes. It can become a serious problem in fish hatcheries.

16.2.2 Class Trematoda

The class Trematoda consists of flukes. The majority of these flukes are a few millimeter
in length (Fig. 16.2). They are all parasites. Some are ectoparasitic living on the surface
of their hosts while others are endoparasitic, found within the host body. The class
Trematoda, includes the Digenea, a large, economically and medically important taxon.

Digenean parasites produces infection in higher animals, including man. They are
common parasites of all classes of vertebrates. Some species particularly those inhabiting
the liver, lungs or blood are serious pathogens of humans and livestock in which they
cause debilitating diseases.
Oral sucker
Pharynx
Oesophagus

Cirrus
Ventral sucker
Metatherm
Seminal vesicle
Vitelline glands (vitellaria)
Laurer's canal
Receptaculum seminis
Ootype

Intestinal cecum

Fig. 16.2: Diagram of a generalized trematode.

Life cycle I
Almost all digenetic trematodes are endoparasitic in the vertebrates. The vertebrates are Majority of the digenean trematodes
the primary or definitive hosts of the parasites as the sexual reproduction of the parasite are located in the digestive tract of
their final (definitive)hosts. Some
takes place in the vertebrate host. Digeneans usually have an invertebrat; as their first species however live in bile ducts, a
intermediate host. Thus digeneans have involved or complicated life histories with at least few in the lungs and gill others in the
) portal venous blood.
two generations and two hosts.Hence the name 'Digenea' meaning having two
generations. In such a complicated life cycle there may be two or more intermediate hosts
, that are infected by the larval stages of the parasite before the life cycle is completed. The
final larval stage locates in the definitive host in which the parasite undergoes sexual
reproduction.

A gastropodsnail (mollusc) usually acts as the first intermediate host for larval Definitive or prlrnary hosts are
development of the parasite. Some species may even need a second (fish, crab or other those in which the par:lsiti
crustaceans) or even a third (amphibian, reptile or mammal) intermediate host. The reproduces.
digeneans, after successf,tily invading the first intermediate host, undergo two rounds of
asexual division which gleatly increase their numbers and so also their chances to
complete the life cycle.
I
The life cycles of digeneans as mentioned before vary, depending upon the species. The
life cycle of the liver fluke Fasciola has been taken as an example in this unit.
Adaptation and Behavioural LIVER FLUKE ( ~ ~ s c l o l a )
Pattern
The cosmopolitan liver fluke of the ruminants sheep, goat and cattle is Fasciola hepatica
(Fig. 16.3). It causes in them a serious disease, known as liver rot. The Indian species is
Fascida indica. The adult liver fluke lives in the bile passages of the liver of the host
where in case of heavy infections it causes extensive damage and necrosis of liver. Heavy
infection also interferes with the normal flow of bile which results in obstructive jaundice
(Also refer to unit 4.6.2 of this course for the life cycle).

Fig. 16.3: The life cycle of F::ciola hepatica. a liver fluke of sheep. goats, and cattle.

The Chiniie liver fluke Clonorchis (Opisfhorchis) sinewis is the liver fluke commonly
found in China, Southern Asia and Japan.

LUNG FLUKES (Paragonimus)

Species of genus ~ a r a ~ o n i m (Fig.
u s 16.4) occur as lung parasite in humans and in many
species of carnivorous mammals. The.species P. westermani, a human parasite, is widely
distributed in the Far East Africa and South America. In India, it has been frequently
 Harmful Kon Chordates
reported from West Bmgal, Assam and South Indian States. Life cycle of P. westermani
has two intermediate hosts; a snail is the first intermediate host, while a fresh water
crayfish or crab is the second intermediate host.

Lung fluke infection can best be controlled by the eradication of the host species of snails
and by only eating crabs and cray fish after cooking them well.

LIFE CYCLE OF Paragonimus westeamni

Fig. 16.4: Life cycle of Pnrqgonimus weslermni'.
BLOOD FLUKES (fihisiosoma)
Blood flukes cause the human disease schistoson~iasis.The three important spccies of
this genus are Schistosoma haematobium, Schistosomcs vnamoni and Schistosoma
japanicum (Fig. 16.5) which are serials parasites of humans (Refer also to unit 4.6.2 of
this CCIPP~SB).

Schistosomes have a life span of 20-30 years. The adult Schistoroma lives in humans and
may be 2 cm long and 1 mm in width. The blood flakes differ fiom most other flukes in
beiilg dioscious (Fig. 16.5). The male and feinale flukes, though separate individuals, are
permanently paired. The male is shorter, broader and heavier. It has a large ventral
groove, called h e gynecophoric canal which is posterior to the ventral sucker into which
is lodged the more slender and longer female, Depending on the species the schistosomes
hirabit blood venules of different organs of the body and have specific intermediate snail
hosts.
Adaptation rod Bchrviourrl Oral sucker
Pattern Ventral
\ sucker Ventral sucker
Oral sucker Oral sucker
\

Ventral /
Oral
sucker

Fig. 16.5: important schistosomea of man. (A) Schktosomrr haemdobium. (B)S. mansoni. (C)S.jqmnic~~m.
Schistmomia mansini is widely distributed in Afiica, the Northern parts of South America and
the West Indies. S. mansoni adults live primarily in the venules draining the large intestine of
the human host. Their principal intermediate snail host. are species of Biomphalaria.
Schistosoma haematobium occurs in the Middle East and parts of Afiica. The adults of S.
haematobium are lodged in the venule of the urinary bladder of the human host. Their
principal intermediate snail hosts belong to the genera Bulinus and Physopsis.
S. japonicunr is mostly f o h d in Japan and other Far East countries. Adults of S.
japonicum mostly lodge in the venules of small intestine of the human host. Their main
intermediate snail hosts are species of Oncomelania.
The life cycle of blood flukes is generally similar in all sjxcies. In this unit we will study
the life cycle of Schistosoma mansoni.

Lye Qcle of Schistosomia mansoni

The adult male and female couple live in humans and feed on blood in the venules of the
large intestine. The female deposits the e'ggs in the small intestinal venules. Using its
spine the egg breaks through into the lumen of the intestine and is excreted outside. When
released from the host each egg contains a fully developed ciliated miracidium (miracidia
- plural). The eggs on comhg in contact with water hatch and the miracidia escape into
the water. The miracidia seek out and penetrate the specific intermediate host, a fresh
water snail. Within the intermediate host the miracidiatransforms into a sporocyst. This
sporocysts produces, a brood of second-generation sporocysts. These daughter sporocysts
directly produce cercariae, without going through redia stage. The cercariae come gut of
the snail host and swim about in water with their forked tail. On coming in contact with
the human host they penetrate the skin,enter the cutaneous blood vessels, and knsform
into a juvenile or sehistosomula. The schistosomulae migrate via blood circulation
through lungs, into liver and fmally to their final destination i.e. to the venules of the
large intestine.

During penetration through the skin, the cercariae cause local dermatitis reaction at the
site of entrance. During their growth phase in the portal system of the liver, the parasites
liberate toxic metabolites causing fever and enlargement of liver and spleen of the human
hosts. The eggs, while being extruded from the blood capillaries, cause extensive damage.
Tissue reactions occur around the eggs remaining in tissues like liver, resulting in the
formation of nodules which may gradually become irreversibly calcified, thus damaging
the organ. Blood circulation in liver may also be obstructed by these parasites which may
cause cirrhosis, interfering with the proper functioning of the organ.

SAQ 1
a) Fill in the blanks with appropriate words. '

i) The free swimming, ciliated larva of blood flukes is called ........................

 .-

Har Non
ii) The scientific name of the cosmopolitan liver fluke of ruminants is
..................................................... \ \

iii) The scientific name of Chinese liver fluke is ................................

........................................
iv) Paragominus a parasite of human lodges in their ..................................
v) The second intermediate host of lung fluke is either a ............................
o r a .......................................
vi) The principal intermediate snail host of Schistosoma haematobium belongs to
the genus ....................................
b) Indicate whether following statements are true (T) or false (F).
i) Digenean fluke have only one host.
ii) The lung fluke does not occur in India.
iii) Schistosoma mansoni is found mostly in the venules of large intestine.
iv) The intermediate host of Fasciola hepatica is a snail.

16.2.3 Class Cestoda

All cestodes are called tapeworms. Adults of all tape worms are endoparasites in the gut
of vertebrates. One of the intermediate hosts is usually a non-chordate. The young stages
live in various tissues of non-chordates and also of vertebrates. Several species of
cestodes occur in man and animals of economic importance of which the two.most
important economically are Taenia solium and Taenia (Taeniarhynchus)saginatus (Refer
to Unit 4.6.2 of this course also).

THE PIG TAPEWORM (Taenia Solium) and THE BEEF TAPE WORM
h
7beain Sagi natris -\
//-
Taenia solium and Taenia sagindrm are cosmopolitan species of tapeworms occurring in
man. The intermediate host for Taenia solium as you will recall is pig, as a result of
which it is also called pork or pig tape worm. The intermediate host for Taenia saginatus
however is cattle and hence it is called the beef-tape worm (Fig. 16.6).
- - - /f--"',
- ,

inverted scol~x
of future
tapewarm

" d
g.

--

Fig. 16.6: Life cycle of Taenia saginatus (beef tape worm)

Adaptation and Behavioural Heavy parasitic infections in humans causes digestive problems, diarrhoea, weight loss,
Pattern abdominal pain, often blockage of the intestinal lumen and reactions to toxic wastes of
the worm. Proper cooking of meat and better hygienic practices would prevent Taenia
infection.

DWARF TAPE WORM (Hymenolepis Nunu)

h'ymenolepis nana is another tape worm of humam. Its development is usually direct
without any intermediate host. The eggs pass out and when ingested by another human
host hatch .in the duodenum releasing ancospheres or the hexacanth embryos.

THE FISH TAPEWORM (Diphyllobothrium &turn)

The adult Diphyllobothrium latum is found in the intestine of man, dog, cat and other
mammals. The immature stages occur in copepods (crustacean) and fish which are its I
intermediate hosts.

THE DOG TAPEWORM (Echinococcus Grunrrlosus) .

One of the most serious larval cestode infection in man is caused by Echinococcus
granulosus (Refer Unit 4 of this course also). The adult parasite is minute being only 3 to
6 mm long. It lives in the small intestine of canines which are the main host. The parasite
has a scolex and strobila consisting of 3 or 4 segments. The larval stages occur in a
number of mammals such as sheep, cattle, horses or even humans, all of which serve as
intermediate hosts. Thus humans serve as an intermediate host in this case and are
infected from dogs due to their association with them.

The eggs ingested by intermediate host lodge in the intestine of the host and liberate the
onchospheres or hexacanths. The hexacanth leave the intestine and makes its way to
various organs such as lungs, liver, kidney, heart, brain or even bone via circulation. In
these organs the onchosphere develops in@ a larval form called hydatid cyst. The
hydatid cyst which is the bladder worm, contains a single or unilocular chamber within
which numerous daughteicyst or bladder worms develop or bud off. Each of these in turn
has many scolices within it (Each scolex will produce a worm when eaten by a canine
host). In course of time the hydatid cyst may grow to a very large size, of more than 15
cm in diameter. Depending on its size and location, the cyst in the intermediate host may
cause varying degree of damage. Rupture of a cyst in human may cause toxic reactions
(fever, vomiting, shock, even sudden death). If the hydatid grows in a critical site such a
heart or central nervous system, serious symptoms occur. Hydatid cysts may remain alive
and viable for many years after the initiation of the infection and in humans their surgical
removal is the only way out. Adequate personal hygiene of man and also of the pet dogs
along with their proper care are the suggested control measures against hydatid infection.
-
SAQ 2
Fi!l in the blanks with appropriate words.
i. Cestodes are commonly called ................................
ii. .......................................................
.... ........................................................isisthe beef tape worm.
111. the pork tape worm.
iv. ........................ ..is.aihuman tape worm usually with direct development.
v. ............................is a fish tape worm that you have studied in this unit.
vi. In humans, the onchosphere of the dog tape worm develops into a larva1 form called
............................... cyst.

16.3 PARASITIC NEMATHEMELMINTHES

16.3.1 Class Nernatoda
Many nematodes are parasitic in plants or animals (food crops. domestic animals and
humans). You may go through the earlier unit 4 subsection 4.7 of this course for an
account of this group, which gives you an idea about their classification and organisation.
The size of nematodes varies greatly, from being microscopic to even a metre or so long.
Most species of plant parasitic nematodes range from 0.3 rnrn - 5 mm in length. Many are
 Harmful Non Chordates
even smaller. Most free-living nematodes are much larger. Animal parasites show a vast
range from less than 5 mrn to a metre:

The buccal cavity has usually teeth or cutting plates. The buccal cavity of plant parasitic
(phytoparsites) nematodes have usually a spear or stylet, instead (Fig. 16.7). The sexes in
nematodes are nearly always separate (i.e. dioecious) and readily distinguishable
externally (sexual dimorphism).

Fig. 16.7: hiematode buccal cavities (a) Free living bacteriovore, Rhabditb (b) the plant root parasite,
Criconemides. (c) The intestinal parasite, Ancylostom.

16.3.2 plait Parasitic Nematodes

Plant parasitic nematodes or phytoparasitic nematodes cause extensive damage to
cultivated plants; resulting in heavy losses. Most cultivated plants are affected. The
damage may be caused directly, or indirectly when the parasitic phytonematodes transmit
plant viruses or allow other pathogens to gain entry into the plant through damaged areas
created by these nematodes. All the plant parasitic nematodes possess a s h w , pointed,
prbtrusible buccal stylet or spear (Fig. 16.7 b). This is used to puncture plant cells. ?be
parasitic nematode suck the cell sap from the punctured cells. The nematode also injects
saliva into plant cell while feeding. This saliva is toxic to plants and causes many
symptoms in the infected plants.

The life cycle of plant parasitic nematodes (Fig. 16.8) is simple. The adult female lay
200-500 eggs. The first stage larva or juvenile moults within the egg to form the second
stage larva or juvenile which comes out of the egg into the soil and then enters the root of
the host piant and starts feeding. Three more moults follow giving rise to the adult with
fully developed reproductive system. In some plant parasitic nematodes like Meloidogyne
,species and Heterodera species, only the female is parasitic, while the male develops from
the second stage juvenile into a free living adult that lives in the soil.

The phytoparasitic nematodes spread from one plot to another through percolating water
or through soil which is transported from place to place. Flood water also causes
extensive spreading. Contaminated seed material is also an important source of spread of
these nematodes.

Fig. 16.8: Generalized life cycle of a plant parasitic nematode showing the four larval stages
- L l , L 2 , L3,LA.
Symptoms on Plants due to attack by some Phytoparasitic Nematode
The nematode Aphelenchoides damages the buds and shoot tips and causes distortion of
stem and crinkling of foliage. The juveniles ofAnguina cause damage of leaves and
wheat grains. Di~lenchusis responsible for extensive rot of potato tuber and onion bulbs.
Pratylenchus and Radopholus cause root lesions as well as root rot in chillies, coffee,
corn, cotton, rice, pineapple, wheat, banana, coconut, sweetpotato, tomato etc. The female 77
Adaptation and Behavioural ' of Meloido~neis parasitic (Fig. 16.9 a) and causes formation of root galls or knots in
Pattern the roots of many plants specially the solenaceous plants. Meloidogyne species affect
chillies, tomato, brinjal, carrot, cotton and a number of other useful crops. Heterodera
species (Fig. 16.9 b) are the root cyst nematodes and occur in colder climates. The female
of Heterodera forms resistant cysts full of eggs. Heterodera species damage potato,
sugarbeet, oats, wheat, cabbage tobacco etc.

Some specific symptoms due to nematode infestation are poor crop growth, winter injury,
wilting of trees, loss of seedling, stunting of plant, patchiness in leaves and growth and
discoloured foliage in a crop.

Fig. 16.9: Some important plant parasitic nematodes and the damage they cause (a) Meloidogyne species
(b) Heterodera species.

16.3.3 Animal Parasitic Nematodes

Just like plant nearly all vertebrate and invertebrate animals are parasitized by nematodes.

ASCARIOD NEMATODES ,

A number of these nematodes are very important pathogens of humans and domestic
animals of which Ascaris Iumbricoides the common human round worm is an important
pest. It is found more commonly in children than in adults (Refer to unit 4 of course for
more details about this nematode).

THE PIN WORM (Entorobius vermicularis)

The pin worm, Enterobius vermicularis, is most common among children (see unit 4).
Adult worms are about 10 mm and live in the caecum of large intestine. The adult
female deposits eggs at the anal region. This causes intense itching in the perianal
region. While scratching, the finger tips of the children get contaminated with the eggs.
These eggs get into the mouth of children when they suck their finger and so are
swallowed. In the intestine the eggs hatch and develop into adult worms in 3-4 weeks.
The parasites however except for intense itching cause comparatively little disease
symptoms.

THE HOOK WORM (Ancylostoma duodenale and Necator arnericanus)

The common hookworms of man, Ancylostoma duodenale (common in the old world)
and Necator americanus, occur widely in the developing nations of the tropics and in
the temperate regions of the world. The mouth region of these parasites is usually
provided with cutting plates, hooks teeth or a,combination of these structures for
attaching to the gut wall of the host. Adult worms live attached to the mucosal wall of
the intestine.

Female lays several thousand eggs per day, which escape via faeces. The larvae on
hatching come out of the eggs in the soil. In about a week's time they become infective
juveniles and gain entry into the human host by penetrating the skin of their feet. These
larvae then travel through circulating blood to reach the lungs, into the air passages, the
trachea, to the epiglottis and when swallowed, enter the gut. In the intestine the juveniles
become finnly attached to the mucosal wall and feed on blood and abraded tissue, for
which their hooks, cutting plates and teeth are useful. Within a week's time they develop
into full adults reaching about 10 mm in length. Hookworm infections cause mechanical
damage to the intestinal wall and loss of blood resulting in anaemia. Infected persons
show stunted growth.
 Harmful Non Chordates

Fig. 16.10: A generalized life cycle of hookworm.

THE WHIPWORM (Trichuris trichiura)
Whip worms or Trichuris are parasites in the alimentary tracts of vertebrates, especially
human beings, dogs, cats, cattle and other mammals. They are relatively small in size.
Trichuris trichiura is parasitic in humans. The adult is about 4 cm long and lives in the
large intestine.

The life cycle is similar to that of the pinworm. The eggs are ingested by the host along
with contaminated food and hatch into larvae which pass into the caecum of the large
intestine and develop into adult worms in about 3 months. In the case of heavy infection,
the worms cause diarrhoea and dysentery. (Refer also unit 4 of LSE-09).

THE TRICHINA WORM (Trichinella Spiralk)

Trichinellaspiralis is a minute (about 3 mm long) nematode found in rats, pigs and
human. In humans it causes the disease 'trichinosis' which is prevalent in most countries
of the world. Both larvae and adults are present in the same host. However, infection
requires another host. Adult worms,(Fig. 16.1 1) live in the mucosa of the small intestine.
They do not have much effect on the host then. The adult female bores through the
intestine into a spiral lymph space and gives birth to about a thousand juveniles at a time.
The juveniles enter the blood stream and are carried throughout the body where they may
be found in almost any tissue or body space. Eventually they penetrate the skeletal
muscles and encyst there. The cyst walls gradually calcify. The skeletal muscle cells of
the host undergo extensive changes and serve to nourish the juveniles. There is no further
development till these cysts are ingested by a new host. When humans ingest pig meat
containing the juveniles the cysts break down and the juveniles escape in the intestine
where they mature. Humans are usually infected due to eating pcsrly cooked pork.
I
Adaptation and Behavioural Heavy infection may be fatal leading to trichinosis and even death. Severe. pain and
Pattern inflammation as well as toxic damage occur when the juvenile encyst in muscles. They
cause degeneration of muscle tissue,

I.AI<VAli arc carricd by

/ \
Whcn the infictcd muscle is blood to skclctal muscle
eaten the larvae cxcyst in
,I \
\
\ wherc they enter the
libcrs ... ' /
\
the intestine an'd ... 1

Fig. 16.11: Life cycle of Trichinellaspiralis.

FILARIAL NEMATODES
Several species of filarial nematodes are parasites of humans and cause diseases. These
are thread-like worms and inhabit the lymph glands and lymphatic vessels. The female
gives birth to juveniles called microfilariae. The life cycle of these nematodes require an
arthropod intermediate host such as certain species of fleas, flies or mosquitoes.

FILARIASIS WORM (WuchereriaBancrojii and Brugia Malayi)

About 250 million people in tropical countries are infected with Wuchereria bancrofti
(40-90 mm long; 1-24 mm broad) or Brugia malayi both of which have certain species of
mosquitoes as their intermediate host. These nematodes are responsible for causing the
disease 'filariasis', and in extreme cases, 'elephantiasis'.

The life cycles of both Wuchereria bancrofti and Brugia malayi are similar. Adults live in the
lymphatic ducts. The larvae or microfilariae are liberated in the blood and lymphatic system
They show nocturnal periodicity i.e., they move to the peripheral blood vesskls at night.

Certain specific mosquito species (refer to subsection 16.4.2 of this unit) act as
intermediate hosts. When these bite the infected human host, the microfilariae enter the
mosquito,with the host's blood. Development within the intermediate host involves
migration of the microfilariae through the gut to the thoracic muscles and then into the
proboscis. From the proboscis the microfilariae are introduced back into the primary host,
when the mosquito bites a another human.

In severe filarial infections the blocking of the lymph vessels by large number of worms
causes serious lymphatic inflammation marked by pain and fever. In chronic (long term)
infection, increase of connective tissue (fibrosis of infected areas) may cause irrevesible
massive enlargement and deformites of lower extremities such as legs, arms and
I sometimes scrotum (vulva and breast are rarely affected). Such enlargement is called Harmful Non Chordates
'elephantiasis' and can only be corrected by surgery. Fortunately extreme cases of
elephantiasis are now rare.

In India this disease is very damaging and is prevalent in Eastern Uttar Pradesh, Bihar,
West Bengal, Orissa and Eastern coastal regions of India.
Mosquito takes second blood meal

Larvae deposited on skin,

enter bite wound

Microiiiariae
enter. bloodstream

PATHALOGY

Fig. 16.12: Life cycle of Wuchereriabancrofli.

RlVER BLINDNESS WORM (Onchucerca Volvulus)

Onchocerca ~~olvulus is another filarial nematode, which causes the disease known as
river blindness or onchocerciasis in humans. This disease affects more than 30 million
oeople in parts of Africa, Arabia, Central America and South America, It causes severe
itching and damage to the eye and ultimately leads to blindness. The intermediate host is
tbe black fly Simulium.

THE GUINEA WORM (Dracunculus medinensis)

Dmcunculus ~nedinensis,the 'Guinea Worm' is a thread like nematode. It is a common
parasite of humans throughout Africa, the Middle East, Pakistan, and parts of India,
particularly Rajasthan and Gu.jarat. The female is about 1 mm in diameter and upto 120
cm in length.

The gravid female, after a period of development in the body cavity and connective tissue
of the h~~lnan host migrates to the subcutaneous tissue and produces a pzinfbl blister that
bursts to form an ulcerated opening. If the ulcerated area of the host comes in contact
with water, the active first stage juvenile larvae are discharged into the water. The
juveniles are ingested by a fresh water copepod crustacean Cyclops which acts as the
vector. In the Cyclops the juvenile mouths twice to form the infective stage. The infective
juvenile re-enters the human host through infected drinking water. The nematode
juveniles are released from the ingested cyclops, in the human gut. These juveniles then
Adaptation and Behavioural MITES AND TICKS OF VETERINARY AND MEDICAL IMPORTANCE
Pattern
CHICKEN MITE (Dermanyssus gallinae) ,
I
fhese (Fig. 16.16) suck the,blood of fowls, through their sharp piercing mouth parts.
When the number of chicken mite increases, egg-laying as well as weight gain in fowls is
affected.

Fig. 16.16: The chicken mite (Dermunyssusgallinae) nymph before gorging on blood.
SHEEP SCAB MITE (Psoroptes)
'The sheep scab mite, Psoroptes (Fig. 16.17) infests the skin of sheep, goats and cattle.
The attack results in loss of wool.

Fig. 16.17: The female of sheep scab mite (Psoropiesequii ovis).

ITCH MITE (Sarcopres scabiei)
The human itch mite (Sarcoptes scabieg (Fig. 16.18) is responsible for human
scabies. It lives in cutaneous burrows of the skin, where eggs are laid and
development takes place. The burrowing and feeding of mites cause extreme itching
Fig. 16.18: ~ a n g or
e itch mite, which is the chief symptom of the disease. The mites spread through contact of the
Sarcoptes scabiei. patient or his clothings.

CHIGGERS
Chiggers are the parasitic larval stages of the mites of the family Trombiculidae eg.
Eutromhicula and Trombicula. They attack poultry, ground resting birds, and mammals,
.
including man. Their attack results in scattered red blotches on skin accompanied by
intense itching.

TICKS
All ticks are parasites of vertebrates. They feed for days or weeks through the
skin of their hosts. They cause local inflammatory damage and itching. They
serve also as vectors of diseases caused by many viruses, ricketsiae, bacteria and
protozoa.

CATTLE TICK (Boophilus)

Cattle Ticks (Fig. 16.19) feed on cattle, sucking their blood. They transmits 'Texas fever',
as they serve as the vector of the organism responsible for the fever which is the
protozoan parasite, Babesia.
 Harmful Non Chordates

Fig. 16.19: Cattle tick Bwplrilus microplus.

SAQ 4
a) State whether true or false.
(i) All spiders are predacesus and carnivorous.
(ii) Spiders belong to the Order Araneae.
(iii) The main distinguishing feature between ticks and mites is the
difference in the thickness of their cuticle.
(b) Match the items listed in Column A with those in Column B.
Column A Column B
i) Tetranychus (a) Attacks a number of important vegetables
ii) Sarcoptes scabiei (b) Texas fever
iii) Boophilus microplus (c) itch mites
iv) Psoroptes equi ovis (d) poultry mite
v) Dermanyssus (e) . sheep scab mite
(c) Fill in the blanks with appropriate choices.
(i) The cattle ticks belong to the genus ................................
(ii) The parasitic protozoan responsible for Texas fever is the ..............................
(iii) Tetranychid mites are known as ..............................mites.

16.4.2 Insects of Medical Importance

Many insects transmit diseases in humans and domesticated animals. Others cause much
damage to crops, stored products and forests. A number of them serve as vectors of
viruses and other microorganisms which produce diseases in plants.

Mosquitoes (Order Diptera) have a world wide distribution. They mostly affect warm
blooded animals like birds and mammals, including humans.

The most important mosquitoes which affect man belong to the genera Anopheles, Culex,
Aedes and Mansonia (Fig. 16.20). Anopheles species are vectors of a malaria parasite
Plasmodium. The main malarial vectors in India are Anopheles culicifacies, and
Anophelese stephemi. Culexfatigans is the chief intermediate host of the filarial
nematode (Wuchereria bancrofti) which causes filariasis. Encephalitis viruses .are
transmitted by Culex tarsalis. Mansonia transmits the filarial nematode Brugia malayi.
Certain species of genus Aedes spread yellow fever and dengue.

The male and female mosquitoes can easily be identified (Fig. 16.20). The males are
smaller than females and do not feed on blood and their palpi are more prominent. The
antennae of the females have whorls of short hairs but in males the antennae have many
long hairs giving them a feathery or plumose appearance. The females also have specially
elongated and modified houthparts suitable for piercing and sucking human blood. '

Mosquitoes develop in water (kefer to Fig, 16.20) which contains microscopie~ntsand

animals that serve as food for the larvae. The eggs are laid on water or in places where
water is likely to collect. The eggs of Culex are laid singly or in minute rafts or masses
Adaptation and Behavioural
Pattern
while those of anophiline mosquitoes are laid singly and have floats. Mosquitoes have
four larval instars ("wrigglers"). I
1

ANOPHELINES CULICINES j

ANOPHELES AEDES CULEX

ON DRY SURFAC ON WATER

SINGLE HAIRS ON

RESTS PARALLEL TO WATER RESTS AT ANGLE TO

I1 I I
SURFACE HEAD ROTATED 180°
WHEN FEEDING

I GREATER PROPORTION OF BODY I

I
WATER SURFACE
HEAD NOT ROTATED

SMALLER PROPORTION OF BODY

I E S T I N G POSITION EXCEPT WHEN ENGORGED OR HIBERNATING I

Fig. 16.20: Diagrammatic representation of the principal characters separating the various stages in the
life cycle of anophcline and culicine mosquitoes.
Most mosquito larvae must come to the surface to breathe atmospheric air through
spiracles which are often situated at the tip of an air tube called siphon. Mansonia larvae
have specialised siphon which is capable of being inserted ihto roots or stems of aquatic
plants from where oxygen for larval respiration is obtained. Larvae of anopheline
mosquitoes however do not have siphons though they posses spiracles. They have also
anal gills.
 The full grown larvae change to the non-feeding but actively swimming pupae or Harmful Non Chordates - a

"tumblers". They are comma shaped. Their cephalothorax has dorsally a pair of
respiratory "trumpets".

The principal characters enabling one to distinguish differences between the anopheline
and culcine mosquitoes and their developing stages are given in Table 16.1.

Table 16.1 : Principal characters distinguishing anopheline and culicine mosquitoes

Stage Anophelinae Culicinae
EBZs Laid singly, possess floats Laid singly or in egg rafts or masses.
Never possess floats
Lar+ae Never have a siphon. Lie parallel All larvae have a short or long siphon.
to water surface Subtend an angle from the water surface.
Pupae Breathing trump& short and Breathing trum+& long and tubular.
broad apically.
Adults (both sexes) Rest at an angle to the surface. Rest with the bodies more or less
parallel to the surface.

SAQ 5
Fill in the blanks.
i) The four most important mosquitoes that affect humans belong to the genera
and ....................
..........................................................................
ii) .......................................is the chief intermediate host of the filarial nematode
Wurcheria bancrofti.
iii) ...............................species are vectors of malarial parwite Plasmodium species.
.iv) The mosquito larvae are called .........................................
v) The Mansonia species transmit the filarial nematode .................................
vi) Male and female mosquitoes can be easily identified by their size, mouth parts,
.................................and ...................................
vii) The mosquito ................................... transmits encephalitis virus.
viii) Dengue virus and yellow fever ,wetransmitted by the mosquito species belonging to
the genus .....................................

THE COMMON HOUSE FLY (Order -Diptera)

House flies are cosmopolitan in distribution. Though Musca domestics (Fig. 16.21) is
world wide in distribution, in India, Musca vicina and Musca nebulo are more common.
Houseflies transmit a large number of diseases in man, through contamination offood.
Some of the diseases they cause are typhoid fever, cholera, poliomyelitis, diarrhoea and
dysentery. They also act as carriers of eggs of a number of worms (helminths).
Adult female lays eggs on decaying organic matter. The larva emerging from the egg is
called a maggot.

Fig. 16.21: Life stages of the house fly. Musea d o d c a .

BED BUGS (Order - Hemiptera)

mese x e Cimex species. Bed bugs (Fig. 16.22) are nocturnal in habit. m e y are flattened
--A ,,,inoc
A . - ~ ~ - . reduced to inconspicuous oads. The adults as well'a~
Adaptation and Behavioural nymphs suck blood of man, other mammals and poultry. The two common species that
Pattern parasitise humans are Cima lectularius and C. hemipterus.

FLEAS (Order - Siphonoptem)

The homan flea.Pulex irritum (Fig. 16.23) and the rat flea or the oriental flea Xenopsyla
cheopis are the common gpecies. They suck blood and act as transmitters.of a bacterium
which causes bubonic plague. .

Adult fleas are laterally compressed, wingless, with piecing and sucking type of mouth
parts. Their posterior pair of legs are adapted for jumping,

Fir. 16.22: Common bed bug.

Fig. 16.23: Female human flea, Pnlcxirrifacns.

-
Human Lice (Order Anoplura)
-
Three species of lice usually infect man. 1. Body louse Pediculus humanus corporis
(Fig. 16.24 a). 2. Head louse, Pediculus humanus capitis. 3. Pubic louse - or crab louse
Phthiwpubis (Fig, 16.24 b). The head and body lice are vectors of relapsing fever,
epidemic typhus fever and trench fever.

Fig. 16.24: Human lice (a) body louse is one of the most dangerous paragite. It is a carrier of epidemic
typhus. (b) pubic louse Phthlnrspubls lives on areas of the body with widely spaced coarse
hair. For this reason its main location Is the publc area.
-
Sand Fly (Order Diptera)
Sand fly (Fig. 16.25) are responsible for spreading several diseases. Phleboromus
argentipes is the Indian species, responsible for spreading 'Kala A m ' , oriental sore and
phlebotomus fever. They are vectors cif Leishmania donovani which causes Kala-azar and
Leishmania tropica which causes Oriental sore, as well as of a virus which is the
causative agent of phlebotomous fever.

Fig. 16.25: Sand fly.

 Harmful Non Chordates
SAQ 6
Match the following items listed in Column A with those listed in Column B.
Column A Column B
i. Musca domestics (a) Flea
ii. Cimex (b) Trench fever
iii. Pulex irritan; (c) Common house fly
iv. Pediculus humanus (d) Bed bug

16.4.3 Insects as Household Pests

ANTS (Order - Hymenoptera)
Ants are social insects (see unit 7 of this course). Some common Indian species are
Monomor 'um indicum, and Componotus compressus (the large black ant known as the
carpenter ant (Fig. 16.26 a and b) and Dorylus labiatus (the small red house ant). All
these are house hold pests as their worker ants feed on a variety of material including the
food we eat, seeds, fruits and nectar.

Fig. 16.26: Ants (a) Monomorirrm indicum (b) worker of the Canlponotus conpressus.

TERMITES (Order Isoptera)-

They are pests causing heavy damage to wood in all its forms, and to a number of house
hold goods (Refer to unit 7 of this course for greater detail).

SILVER FISH (Order Thysanura)-

These are small, primitive, wingless insects, commonly found among books. Examples
are: Lepisma (Fig. 16.27), and Ctenolepisma.These cause damage to a variety of
materials on which they feed such as starched clothes, binding of books, etc. Fig. 16.27: Silver fish.

COCKROACHES (Order - Orthoptera)

Cockroaches (Periplaneta americana (Fig. 16.28), and Blatta orientalis) are very
familiar. ~ o t h ~ a d u land
t s nymphs cause damage, feeding on many kirids of materials like
binding or leaves of books, and various food products in kitchens.

Fig. 16.28: Periplaneta americana (a) Egg case with two longitudinal rows of eggs. (b) Second instar
nymph. (c) Adult. .%.
Adaptation and Behavioural The fertilized female lays about 30 eggs in a capsule or case called ootheca which is
carried by the female for a few days before being deposited. The eggs hatch into nymphs
which mature into adults (Fig. 16.28).

16.4.4 Insects of Veterinary Importance

STABLE FLY (Order Diptera) -
The stable ffy (Stomoyxs calcihans) attacks mules, horses and other animals as well as
man. It (Fig. 16.29) resembles the house fly but is smaller. Both males and females suck
blook. Animal attacked by these flies lose weight, and their milk yield is reduced.

-
HORSE FLY (Order Diptera)
The horse flies (Tabanus rtriatus) resemble house flies but are larger and stouter. The
females attack horses, cattles, dogs, humans, deer etc. They pierce the skin and suck
blood. Blood continues to ooze fiom the wound even after the ffy leaves the animal.
I(

OX WARBLE FLY (Order Diptera) -

The common cattle grub is the larval stage of the fly Hypoderma lineatum (Fig. 16.30).
The maggots of these flies form tumour or cyst under the skin of the back of cattle. Each
tumour contains a maggot. It emerges when mature and falls to the ground and pupates.
Adult emerges fiom pupa. Eggs are glued to hair near hoof of the host and hatched
maggots penetrate skin.

Fig. 16.30: Adult female l y of the common raffle grub, Hjpmkrnm Pnmmr.

-
BLOW FLY (Order Diptera)
Lucilia serenissima is an example of blow fly. It is small, metallic blue or green in colour.
Other blow flies are Calliphora and Phormia. Only larvae feed on livestock. Wounds
attract female blow fly which lay eggs in them.

Fig. 1631: The shaft louse SHAFT LOUSE (Order Mallophaga) - I

Menopon gallinae.
Shaft louse (Menopon gallinae) (Fig. 16.3 1) mainly infests chickens. The shaft louse
feeds by nibbling or chewing dry s k i scales, feathers or scabs on the skin of the host.
The irritation results in a nervous condition of the bird like sleeplessness, loss of appetite. Harmful Non ~ 6 r d a t e s

CHICKEN FLEA (Order - Siphonoptera)

The chicken flea Echidnophaga gallinaceus attacks chickens, and other poultry. Young
fowls infested by chicken flea are often killed. Furthermore egg-laying and growth is
severely affected in infested poultry.

SAQ 7
Indicate which of the following statements are true (T) or False (F).
i) The common cattle grub forms tumours on the back skin of the host.
ii) Only larvae of blow flies feed on the necrotic tissue of the host.
iii) The horse flies (Tabanus) attack only horses.
iv) The milk yield of cattle attacked by stable fly is often reduced.
v) The common name of Lepisma and Ctenolepisma is ox warble fly.
vi) The female of cocltroach lays about 30 eggs in a capsule called ootheca.

16.4.5 Insects of Agricultural Importance

Insects attack a large number of trees, vegetable, fruit and ornamental plants. Different
parts of trees and plants may be attacked.

Types of Losses of Agriculture

Losses of agriculture by insects may be due to (1) direct losses to the plants (2) indirect
losses.

1. Direct losses are due to:

i) -
Leaf eaters or defoliators Such insects posses biting, and chewing type of mouth
parts for eating leaves and often cause serious damage to the crop. Examples include
grasshoppers, locusts, beetles, weevils and larval stages of caterpillars of moths and
butterflies.
ii) Leaf miners - Insects of this group lodge in between upper and lower epidermis of
leaves and devour their green parts eg. citrus leaf miner.
iii) Leaf rollers - Caterpillars of insects, feeding on leaves also cause the leaves to roll
up which later shrivel and fall off eg. cotton leaf roller.
iv) Stem and root borers - Caterpillars of insects bore though the stems and roots of
various plants, often seriously dimaging them. As a result affected plants dry up
and become stunted in growth - e.g. sugarcane stem borer and rice stem borer.
v) Sap suckers - Such insects pierce and suck cell sap of plants. They cause serious
injury to the plants when present in large numbers e.g. rice gandhy bug, mustard
aphis etc.
vi) Bark and wood feeders - This group consists primarily of caterpillars, beetles and
weevils which tunnel between bark and wood of shrubs and trees e.g. bark eating
caterpillar.
vii) -
Fruit destroyers Insects of this group attack fruits, making them inedible for human
consumption and unfit for seed purposes e.g. fniit flies.
viii) Seed feeders or storage insects - Much damage is caused to stored food material and
grains by certain insects like rice weevil.

2. Indirect losses to plants

Most leaf hoppers and aphids indirectly damage host plants by secreting honey dew on
their leaves whiph result in the development of sooty moulds. This retards the
development and growth of the plants. Several sap suckers also indirectly transmit fungal,
bacterial and viral diseases (called blight, mould, wilt) while feeding on plants. Beetles
transmit bacterial wilt on cucurbits, leaf hoppers, aphids and fulgorids transmit viral
diseases of tobacco, potato, peach erc.
Adaptation and Behavioural Nature of Insect Pest
Pattern
Some insects restrict their attack to one species of plants and are called monophagous.
Others attack and feed on a few related plant species and are called oligophagous. While
many others attack a considerable number of host species and are called polyphagous.

In this section we will discuss two or three agricultural pests each of (1) Paddy, (2)
Wheat, (3) Sugarcane, (4) Coconut, (5) Oilseeds, (6) Cotton, (7) Pulses, (8) Potato, (9)
Vegetables, (10) Fruit, (1 1) Stored grains, (12) Teak trees.

To make this study easier we will first briefly study the polyphagous insects like tenhites,
desert locusts and hairy caterpillars which attack a large number of plants. We will then
describe two or three important pests of the above listed plants which may be
monophagous, oligophagous or polyphagous.

A. SOME POLYPHAGOUS PESTS

-
Termites (Order Isoptera)
Termites have already been covered previously both in this unit (sub section 16.4.2) as
well as in unit 7 of this course. The important crop damaging subterranean termite species
are Microtermes obesi and Odontotermes obesus. Termites are polyphagous. They
destroy wood and attack a large number of crops likes paddy, wheat, sugarcane, barley,
cotton, groundnut,jowar, bajra, maize, chillies, brinjal, guava, lemon, mango, coconut
stored grains, fruit trees, forest trees etc. The attacked plants show drying up of leaves
and damaged buds.

-
LOCUSTS (Order Orthoptera)
Locusts are grasshopper species which multiply rapidly when conditions are favourable.
They form swarms and migrat~from place to place travelling long distances and
destroying vegetation on their way.
There are three species of locusts that occur in India.
I . Schistocera gregaria (desert locust)
2. Locusta migratoria (migratory locust)
3. Patanga succinata (Bombay locust)
Locusts are polyphagous and attack most plants. The locust have mainly two phases (1)
Solitary (2) gregarious (Fig. 16.32). &A

Pig. 16.32: Locusts of the two phases are of different colours.

In solitary phase the locusts are dispersed, leading solitary and sedentary life. They breed
in desert area which in India are in Rajasthan, parts of Sa~rashtraand Bafoda, Kutch and
Hissar and some districts in Punjab.

In the gregarious phase, the locusts are more L :tive and crowded. They form swarms and
migrate long distances. The desert locust is the most destructive. It has a wide range of
activity extending from Southern Portugal, Gib alter, North West, East and North East -
Africa to Arabia, Israel, Northern Russia, Iraq, A-an,Turkey, Afghanistan, Pakistan and
India.
 - - - - -

The desert locusts copulate just after attaining maturity. The female lays 50-100 eggs in Harmlul Non Chordates
sandy soil in a compact cluster (Fig. 16.33).

Fig. 16.33: Scl~isfocarcrtgregana- (r Sr b) nymphs of solitary phase.; (c & d) Nymphs of grrgarious

phase. (e) Female locust laying eggs.

CATTER PILLARS O F MOTHS (Order - Lepidoptera)

Larvae of moths and butterflies are called caterpillars. Many of these are polyphagous
Especially, so the hairy caterpillars.

The more important hairy caterpillar pests are:

1. Amsacta nzoorei - Red Hairy caterpillar
2. Diacrisia obliqua - Bihar Hairy caterpillar

All these caterpillars are polyphagous attacking a variety of fooci piants. They are
sporadic pests and occur in large numbersduring certain seasons.

THE RED HAIRY CATERPILLAR (Amsacta moore0

The red hairy caterpillar (Fig. 16.34) is a voracious feeder of leaves devouring almost any
green matter. The pest occurs regularly but assume epidemic proportion in certain years.
Its main host plants are groundnut, cowpea, other pulses, cotton, maize, bajra, sun hemp
etc.

The adult moth is medium sized and iays 800-1000 eggs in batches in her lifetime, on the
underside of the leaf and even on the soil. The caterpillars feed on whatever green plant
material is available and move from plant to plant devouring all that is their path. The
fully grown red caterpillar pupates in the soil into a cocoon made of larval hairs. The
moth emerges from the cocoon to start the next generation.
Adaptation and Behavioural
Pattern

(a) .w

Fig. 16.34: Liie history oiRed hairy caterpillar (Amsada moore0 - (a) Egg (b) Larva (c) Pupa (d) Adult.
BIHAR HAIRY CATERPILLAR (Diacrisia obliqua)
The Bihar hairy caterpillar attacks sesamum, linseed, mustard, groundnut, castor, jute,
safflower, sunflower, potato, tomato, cauliflower and cabbage. The caterpillars feed on
the lower surface of young and tender leaves. When infestation is severe, only the mid-rib
is left hanging. Crops which are severely affected become completely defoliated.
The moth, (Fi4. 16.35) is medium sized. The female lays eggs in hundreds in clusters
mostly on the bnderside of the leaves. The young caterpillars feed on the lower epidermis
of the leaves. They gradually disperse, spread over the whole area, feedingvoraciously.
The cateaillars march from field to field in large numbers defoliating and destroying the
crops. Pupation occurs in the soil and a cocoon of silk and larval hairs is formed. The
adult moth emerges from the cocoon to start the next generation.

Fig. 1635: Liie history of Bihar hairy caterpillar Diacrisia obliqua. (a) adult (b) egg (c) larva (d) pupa.

B. SOME ECONOMICALLY IMPORTANT SPECIFIC PESTS

OF PLANTS
, .
PESTS OF RICE (PADDY)
There are many important pests that cause enormous damage to the paddy crop.

Some Pests of Paddy (Rice)

Leaf eating pests
1. -
Hieroglyphus nigrorepletw, H Banian Rice grass hopper
 Harmful Non Chordates
2. -
Dickadispa armigera Rice hispa
-
3. Spodoptera mauritia swarming caterpillar or army worm
- -
4 . Amsacta moorei Red hairy caterpillar Refer to poly phagous pests of this unit.

Leaf and stem sucking pests

5. Nephotettix viresc-, N. apicalis - Green rice leaf hopper

Stem borer pests

6. Sgiropophaga (Tryporyza)incertulas - the rice stem borer

Grain sucking pests .

7. Leptocorisa varicornis - Rice gundhy bug (rice earhead bug).

Leaf cutting pest

8. Mythimna separata - Paddy army worm

-
RICE GRASS HOPPER (Hierogbphus nigrorepletus, H. bunion) Order -
Orthoptera
Both adults and nymph of Hieroglyphys nigropletus (Fig. 16.36) damage paddy crop by
feeding on leaves and shoots. In severe infestations, they move from field to field over
large areas. Due to this the plants are completly defoliated resulting in low yield.

Fig. 16.36: Life history of Hieroglyphus nigrorepletus

RICE HISPA (Dicladispa armigera) Order Coleoptera -
The rice hispa (Fig. 16.37) beetle feeds on green matter of leaves. The'grubs of this beetle
mine (burrow into) the rice leaves feeding on tissues between the two epidermal layers.
Crops remain stunted and tips dry up.

Fig. 16.37: ~ d u lof

t Dicladispa armigera
GREEN RICE LEAF HOPPER (Nepliotettk virescens) Order - Hemiptera -
N. virescence, (Fig. 16.38) also called green jassids, is a green leaf hopper. Males have
two spots in the forewing which are absent in females. Both adults and nymphs suck the
sap from leaves, thus causing the plants to have a sickly appearance with the leaves
becoming yellow. N. Virescence transmit viral diseases which threaten rice cultivation in
certain areas.
Adaptation and Behavioural
Pattern

Fig. 16.38: Nepi~otefiirvirrscens. ,(a) Egg, (b) Eggs arranged in a row; (c) Nymph (d) Adult-

RICE SWARMING ~ A T E R P I L L A R(Spodoptera mauritia) Order Lepidoptera -

The caterpillars of Spodoptera mauritia (Fig. 16.39) when presert in large numbers,
cause severe damage to crop. They move in bands (and so are called swarming
caterpillars), feeding voraciously and leaving behind leafless plants. The fully grown
caterpillar pupates in the soil. The adult emerges from the pupa.

Fig. 16.39: Life history of Spodoptera mourilirr (a) adult (b) egg- (c) larva (d) pupa.

THE RICE STEM BORER (Sciropophaga incertulas), Order- Lepidopterr.

The cater pillar of the rice stem borer Sciropophaga (Tryporyza) incertulas (Fig. 16.4C
bores into the stem near the roots causing 'dead heart' or drying of central shoot which
comes off easily when pulled. When plants are attacked at flowering stage, the panicles
(ear head) dry up causing white ear heads.

Fig. 16.40: Life cycle of the rice stem borer Sciropophaga incertulas. (a) a infested plant with egg; (b)
larva; (c) pupa; (d) adult.
PADDY ARMY WORM (Mytlrimna separata), Ordkr L.e&doptera . - Harmful Non Chordates

The caterpillar of Mythimna separata (Fig. 16.41) is the destructive stage. It feeds
voraciously on the leaves and cuts the stem and unripe ears. The fmal larval stage cuts off
rice panicle from peduncle, hence the term 'ear cutting'. Iqis this stage of caterpillar
which causes maximum loss to crops. The fully grown caterpillar enters the soil,
constructs an earthern chamber and pupates within it.

Fig, 16.41: Life Stages of Mythimna separata (a) eggs; (b) caterpillar; (c) pupa; (d) npult.
RICE EARHEAD BUG (Leptocorka varicoinis), Order Hemiptera -
Both adult and nymph of Leptocorisa varicornis (Fig. 16.42) suck the sap of peduncle, tender
stem and grains which are at the milky stage as a result of which they become chaffy.

Fig. 16.42: Life history of Leptoeorisa vancorn&. ( a ) eggs; (b) nymph; (c) adult.
PESTS O F WHEAT
'Wheat is also attacked by a large number of pests.
Some important pests include:
-
1 . Agrostis ipsilon Cutworm
2. Sesamia inferens - Pink stem borer

CUT WORM (Agrostis ipsilon), Order Lepidoptera -

The larval stage of Agrostis ipsilon (Fig. 16.435 damages the standing crops by cutting the
tender plants at just above the soil surface. These larvae make earthern chambers
underground and pupate within them. The moth,emerges from the pupa at night.
Adaptation and Behavioural
Pattern

Fig. 16.43: Life stages ofAgrostir ipsilon. (a) adult (b) eggs (c) larva (d) pupa.
PINK STEM BORER (Sesomio infkrens), Order - Lepidoptera
b

Caterpillars of Sesamia inferem (Fig. 16.44) produce dead hearts similar to the rice stem borer.

Fig. 16.44: Sesatnia inferem (a) Adult (b) Larva.

PESTS O F SUGARCANE
The main destructive pests of sugarcane are:
1. -
Chilo infuscatellus The Shoot borer
2. Pyrilla perpusilla - Leaf hopper
3. Sesamia inferem - Refer to pests of wheat of this unit
4. Hieroglyphus nigrorepletus - Grass hoppers - Refer to pest of paddy in this unit.
5. Mythimarr seporoto - Paddy army worm - Refer to pests of paddy in this unit.

Fig. 16.45: Life history of Chilo infuscafeIIus. (a) infested plant (b) adult (c) eggs (d) larva (caterpillar) (e) pupa.
 - -- - -

Non Chordates
SUGARCAN B SHOOT BORER (Chilo infuscatellus), Order-Lepidoptera
The caterpillars of Chrlo rnfuscatellus (Fig. 16.45) bore into the stem and feed inside the
soft tissues causing "dead heart". The adult moth lays eggs in clusters of 8-10 in rows on
underside of leaves. The hatched caterpillars enter the stem just above ground level. Fully
grown caterpillar pupate within the shoot or stem from which fully grown moth emerges.

SUGAR CANE LEAF HOPPER (Pyrillaperpusilla), Order - Hemiptera

Both adult and nymph of Pyrilla perpusilla (Fig. 16.46) suck the cell sap of the leaves of
sugar cane. Consequently the leaves turn yellow and dry up. The adult and nymph also
secrete honey dew on the foliage. The honey dew attracts harmful fungi resulting in
growth of black sooty mould due to which the rate of photosynthesis in the anacked
plants is retarded, causing loss in sugar content of sugarcane.

The head of the adult insect is prolonged anteriorly into a snout-like structure called
rostrum. The female has a pair of anal tufts which produces waxy filaments.

Fig. 16.46: Life history of E!vriCIaperpusillo.(a) infested sugarcane plant (b) eggs (c) nympK(d) male (e)female.

PESTS OF COTTON
Cotton suffers severely due to many insect pests. Some of these are:
1. -
Earias vitella (Eariusfabia) Spotted boll worms
2. Dysdcrcus kocnigii - Red cotton bug
3. Amsacta spp. - Red hairy caterpillars - Refer to polyphagous pests of this unit.
- -
4. Diacrisia obliqua Bihar hairy caterpillar Refer to polyphagous pests of this
unit.
5. Mythimna separata - Cut worm - Refer to paddy pests of this unit.

SPOTTED BOLL WORM (Earius vitella), Order - Lepidoptera

The caterpillar of spotted boll worm (Earias vitella = E.fabiu) (Fig. 16.47) bores into the
growing shoots of cotton plants when the plants are 6 weeks old, causing the plants to
droop and wither. Later when the flower buds and cotton bolls appear they are also
attacked by the caterpillars and are consequently damaged. Due to this the bolls are shed.

Fig. 16.417: Life history of Ear& vilrlln (a) e@s (b) larva (c) pupa (d) adult.
 -

.
I Adaptation and Behavioural RED COTTON BUG ( ~ ~ s d e r c &
koenigi&
s o r d e r - Hemiptera
I Pattern .
Both the adults and nymphs of Dysdercus koenigii (Fig. 16.48) damage the host plant as
well as the bolls' by suckidg their sap.

Fig. 16.48;Life history of Dysdemus koenigii. (a) ggs (b)nymph (c) adult.

PESTS O F OIL SEEDS

Oil is excracted from the seeds of several types of plants like castor, ground nut and
coconut palm. All these are often attached by insect pests.
PESTS O F CASTOR PLANT
The following are some important pests of castor.
1 . Achaea janata - Castor semilooper
2. Amsncta moorei- Red hairy caterpillar - Refer to polyphagous pests in this unit.
CASTOR SEMILOOPER (Acltaea janata), Order Lepidoptera -
The caterpillars of Achaea Janata (Fig. 16.49) feed voraciously on leaves and when they
attain full growth, they pupate in the soil. Adult emerges from the pupa.

(:I) (1))
Fig. 16.49: Life history ofAchaeajanata (a) full grown semi looper caterpillar (b) Adult

PESTS O F GROUNDNUT CROP

Some important pests of Groundnut are:
1. Aphis craccivora - Groundnut aphid
2. Amsactn nioorei - Red hairy caterpillar - Refer to polyphagous pests in this unit.
-
3. Diacrisin obliqun - Bihar hairy caterpillar Refer to polyphagous pests in this unit.
-
4. Agrotis spp. - Cut worm Refer to wheat pests in this unit
-
5. Heliotlrb armigern - Gram pod borer Refer to pests of pulses later in this unit.

GROUNDNUT APHID (Aplth crnccivora), Order Hemipteaa -

Large number of adults (Fig. 16.50) and nymphs of Aphis craccivora may be observed on the
tender shoots sucking the cell sap. Consequently the shoots dry up. These aphids act also as
vector for "rosette disease" in groundnut. Population of these pests on plants increase very
rapidly. Infestation on the groundnut crop usually occurs 4-6 weeks after sowing.

Fig. 16.50: Adult of Aphis craccivora

PESTS O F COCONUT PALM

Some pests of Coconut Palm are:
1. Rhynchophorusferrugineus - Red palm weevil
 Harmful Non Chordates
2. Oryctes rhinoceros - Rhinoceros beetle '
3. Opisina arenosella (=hkphantis serinopa) - The black headed caterpillar

THE RE9 PALM YEEVIL (Rhyncophorusferrugineus), Order Coleoptera -

The adult weevil (Fig. 16.51) is about 35 mm and is cylindrical and reddish brown. The
damage is caused by the grubs of the weevil which bore into soft tissues of the trunk of
the palm and feed inside. When attack is severe the central shoot wilts and the palm dies.
The female makes small holes in the soft tissues of the host and lays eggs in the hole. The
apodous, (without eggs) soft whitish grubs with red head, feed on soft tissues resulting in
the formation of tunnels inside tree trunk.

Fig. 16.51: Rhynchophorusferrugineus - (a) adult (b) grub (c) cocoon.

THE RHINOCEROS BEETLE (Oryctes rhinoceros), Order Coleoptera -
The adult rhinocerous beetle (Fig. 16.52) is responsible for most of the damage in the
plants. They bore into unopened, folded tender leaves and petioles, chewing up tissues,
leaving fibrous mass in the burrow. Attacked trees becomes stunted and the growing
points are destroyed. This beetle is one of the major pests of coconut palms.

Thk adult beetle measures about 5 cm and has a stout cylindrical, reddish brown black Fig. 16.52: Adult beetle of drydcs
rhinooeros.
body. Its head is provided with a pointed horn projecting dorsally. T4e horn is longer in
the females.

The female lays 100-150 eggs in decaying organic matter or in manure pits. The grubs
feed on decaying matter. The grub pupates in a cocoon in the soil at a depth of 30 cm.
Adult beetles emerge from the cocoon and fly towards host tree and start feeding on it
thus damaging it. Life span of adult is about'200 days.

THE BLACK HEADED CATERPILLAR (Opisina arenosella), Order LePidoptera -

The black headed caterpillar of Opisina arenosella (=Nephantis serinopa) (Fig. 16.53)
feeds on the undersurface of leaflets scraping off the green matter. The leaves
consequently dry up. In case of severe attack the whole plantatiojl gives a burnt up
appearance.

Fig. 16.53: Adult of Opisina arenoseUa (=Nephtmls serinopa)

PESTS OF PULSES
The pulses attacked by the pests are green gram (moong), black gram (urd), Bengal gram
(chana), Pea (matar) and pigeon pea (arhar) etc.

Some important pests of pulses include:

-
1. Helicoverpa (Heliothis) armigera Gram pod borer.
- -
2. Aphis craccivora Aphid Refer to pests of groundnut crop in this unit.
- -
3. Amsacta moorel Red hairy caterpillar Refer to polyphagous pests in this unit.
Adaptation and Behavioural -
4.' Diacrisia obliqua - Bihar hairy caterpillar Refer to'polyphagous pests in this unit.
Pattern
GRAM POD BORER (Helicoverpa armigera), Order Lepidoptera -
The catter pillar of gram pod borer, Helicoverpa (=Heliothis) armigera (Fig. 16.54) is a
polyphagous pest of pulses attacki~gmany pulses (red gram, Bengal gram, soybean,
green gram, black gram and pea) and other important crpps including safflower, chillies,
sorghum, groundnut, tomato, cotton etc.

The caterpillars are voracious feeders and damage young pods, foliage and developing
grains by boring into them and feeding on them thus reducing yield. In America it causes
maximum destruction to cotton, hence it is called "American cotton boll worm".

The adult females lay eggs singly on tender parts of plants. These eggs release young
caterpillars which start feeding voraciously, initially attacking the leaves and then the
pods. The caterpillars when fully grown pupate in the soil. The adult moth emerges from
the pupa and lays eggs on'leaves.

Fig. 16.54: Life history uf Gram pod borer Helicoverpa (=Heliothis) arwgera (a) eggs (b)infested plant (c)
pupa below soil (d) adult.
PESTS OF VEGETABLES
Pests of Potato and Brinjal
The fruits leaves and often entire plants of vegetables like brinjal, potato, okra, cabbage,
cauliflower etc. are attacked by a large number of insect pests some of which are given
here.

Some pests of Potato and Brinjal

1.
2.
,Leucinodes orbonalis - The shoot and fruit borer
Epilancha vigintioctopwctata - Spotted leaf beetle
3. - -
Agrostis ipsilon Cut worm Refer to pests of wheat in this un:t.
4. -
Helicoverpa armigera Gram pod borer - Refer pests of pulses in this unit.
5. Myzusperisicae - Refer pests of mustard in the unit.

THE SHOOT AND FRUIT BORER (Leucinodes orbonalis), Order - Lepidoptera

Leucinodes orbonalis (Fig. 16.55) is a polyphagous pest on a large number of
solanaceous plants including potato and brinjal. The caterpillar produces damage by
boring into the tender shoots of young plants. The growing point wilts, drooping shcots
appear, which ultimately wither and die. The caterpillar also bores into flower and fruits.

Fig. 16.55: Life cycle of~eucitiodkorhnulis. (a) adult (b) egg (c) larva (d) pupa in brinjal fruit (e)
infested plant
 SPOTTED LEAF BEETLE (Epilaclrna vigin{ioctopunctata), Order Coleoptera - Harmful Non Chordates .

Epilachna vigintioctopunctata is also a polyphagous pest of solanceous plants. The adult

and the grubs both feed on potato, brinjal, tomato and other solanaceous plants. It is
however a very serious pest of potato and brinjal.

The adults and grub both are destructive stages which scrape and feed on the green
chlorophyll and sketelonize the leaves which then get gives a characterisitic lace like
appearance. These leaves dry up later. The adult of E. vigintioctopunctata (Fig. 16.56) is
dotted at several places on the elytra. . -

PESTS O F LADY'S FINGER (OKRA)

Some pests of lady finger o r okra are:
-
I. Enfiris vitclla Spotted boll worms - Refer to pests of cotton in this unit.
- -
2. Hclicovcrpcr rrrmigertr Gram pod borer Refer to pests of pulses in this unit.
3. Dystlcrcus koenigii - ?'he red cotton bug - Refer to pests of cotton in this unit.

PEST O F CUCURBITACEOUS VEGETABLES

Cucurhitaceous vegetables include the various gourds.

One important pest of cucui-bitaceous vegetables is:

I. Dacus cucurbitae - Fruit fly

FRUIT FLY (Drrcus cucurbitae), Order - Diptera

The fruit fly (Ducus cucurbitae) (Fig. 16.57) is the most important pest of cucurbitaceous
vegetables. The attacked plants include bottle gourd, bitter gourd, pumpkin, water melon,
musk melon, cucumber etc. Damage is caused by larvae called maggots which feed on
the pulp of fruits. The infested fruit starts rotting. The female lay eggs into flowers and
tender fruits. The maggots which hatch from the eggs feed on the pulp of fruits. The fully
grown maggots pupate in soil. Adult flies emerge from the pupa.

cucurbitae (Melonfly),
PESTS O F
Cruciferous vegetables include cabbage, cauliflower, knol-khol, beet root, radish, turnip etc.
Some of their pests are:
I
I. Plutella xylo~tella- The diamond back moth
2 . Trichoplusia ni - The cabbage green semilooper
3. Lipaphis erysimi and Myzus persica; - Aphids
-
4. Agrostis ipsilon - The cutworm Refer to wheat pests in this unit.

DIAMOND BACK MOTH (Plutella xylostclla), Order Lepidoptera -

The damage is caused by the small caterpillar of the moth Plutellaxylostella (Fig.
16.58). The cater pillar feeds on the underside of the leaves cutting several holes.
Caterpillars hinder healthy growth of plants due to which yield is considerably
reduced.
Adaptation and Befiavioural
Patters

Fig. 16.58: The diamondback moth, Plutella xylostelln.

CABBAGE GREEN SEMI LOOPER (Triclroplusia no, Order Lepidoptera -
The caterpillar of Trichoplusia ni (Fig. 16.59) is the pest. This pest in addition to
attacking cabbage also attacks lettuce, spinach, beet, pea, celery, parsley, potato, tomato,
carnation, nastursium etc.

Fig. 16.59: Trichoplusia ni (a) adult (b) cater pillar.

MUSTARD APHID (Liaphis erysimi) and POTATO APHID (Myiwspersicae),
-
O'rder Hemiptera
The adults and nymphs of both (Liaphis erysimi) (Fig. 16.60 a) and Myzus perisicae
(Fig. 16.60 b) suck the sap from the host plant. The main plants attacked by mustard
aphids include mustard, cabbage, raddish and cruciferous plants. The potato aphid (Fig.
16.55 b) attacks mustard as well as potato, bean, lady's finger. tomato etr

Fig. 16.60: Aphids. (a) Mustard aphid (Liaphus erysinu) (b) Potato aphid Myzuspersiae.
PESTS OF FRLIIT CROPS
Fruit trees are also attached by a large'variety of insect pests of which only a few are .
given here.
PESTS OF MANGO
Some important pests of Mango are:
1.. Amritodus (Ideocerus) atkinsoni - Mango hoppers
2. Dacus dorsalis - Fruit fly
MANGO HOPPERS (Amritodus atkinsono, Order Hemiptera
4
-
Both adults and nymphs of the hoppers (Fig. 16.61) are serious pests of mango. They
 Harmful Non Chordates
suck cell sap from new shoots, inflorescence, buds and flowers.'When infestation is
severe the entire inflorescence and even tiny fruits wither. Tiny fruits and flower buds are
shed resulting in heavy crop loss due to poor fruit setting. The hoppers also secrete
honey-dew on which a black sooty mould develops.

Fig. 16.61: Amrirodus (Idiocerus) atkinsoni. (a) adult (b) nymph.

FRUIT FLY (Dacns dorsnlis) Order - Diptera
Dacus dorsalis (Fig. 16.62)It is a polyphagous pest on mango, guava, banana, citrus, apricot,
apple, pomegranate, locat, plum, peach, pear and a number of other fruits and vegetables too.
The damage is mainly caused by maggots of the adult fruit fly which feed on the pulp of
fruits, causing the fruits to rot and fall off.
-

Fig. 16.62: Adult of fruit fly Dacus dorsalis.

PEST OF CITRUS
An important pest of citrus is:
Papilio demoleus - The lemon butterfly
LEMON OR CITRUS BUTTERFLY (Papilio demoleus) Order - Lepidoptera
The Lemon butterfly (Fig. 16.63) is one of the important pests of citrus. Damage is
caused by the caterpillar which feeds on fresh leaves and terminal shoots of the citrus
plant. When infestation is severe the plant becomes unfit for fruit bearing.

Fig. 16.63: Citrus butterfly (Papillo dcmoleus) (a) adult (b) lama
Adaptation and Oehaviourrl PESTS OF STORED GRAINS
Pattern
Stored grains are damaged by a large number of insect pests.

PESTS,OF gRDER COLEOPTERA

Some pests belonging to order Coleoptera which attack stored grains are only listed here
alongwith their figures (Fig. 16.64): '

I. Sitophilus otyae - Rice weevil, (Fig. 16.64 a)

2. -
Tribolilcm castaneum Red flour beetle, (Fig. 16.64 b) !
?
3. -
Cal/osobruchrcs chinensis The pulse beetle, (Fig. 16.64 c)
4. -
Rhizopertha dominica The gram borer or paddy borer beetle, (Fig. 16.64 d)
-5. -
Trogodmla granarium The khapra beetle, (Fig. 16.64 e)

(a)

Fig. 16.64: Some Coleopterrn pests of stored grains. (a) Rice weevil (silophilus oryzae) (b) Red flour
beetle (Tribolium castaneurn) (c) Pulse beetle (Callosobrucltus chinemis) (d) Cram
borer or- paddy borer beetle (Rltizoprrtlm dominien) (e) Khapra beetle (Trogoderm
grarmri~ml).

PESTS OF ORDER LEPIDOPTERA

The stored grains are also damaged by several lepidoptera species. A few major pests-.
belonging to the order lepidoptera are also only listed here along with their figures (Fig
16.65).
I . Sitotroga cerealella - Angoumois gram moth, (Fig. 16.65 a)
2. Plodia interpunctella - Indian meal moth, (Fig. 16.65 b)
3. Corcyra cephalonica - The rice moth, (Fig. 16.65 C)
 Ha~mfulNon Chordates

Fig. 16.65: Some Lepidopteran pests of stored grains. (a) Angoumois grain moth (Sirotrogo cerealella).(b)
Indian meal moth (Plodio inlerpunclello).(c) Rice moth (Corcyracephdonica).
PESTS O F TREES
A number of insects are pests on trees. One of the most commercially important of these
tree, is teak, which yields much valuable timber:
PESTS O F TEAK TREE
An most important pest of teak tree is the:
1. Hyblaea puera - Teak defoliator

TEAK DEFOLIATOR (Hybfaeapuera) Order - Lepidoptera

The caterpillars of the adult moth Hyblaea puera (Fig. 16.66) feed on the tender leaves $

and skeletonize the older ones, causing extensive defoliation.

(a) (b)
Fig. 16.66: Hyblaeapuera. a) larva; (b) adult moth.
-- --

SAQ 8
Match the icsect in Column A with the crop it damages listed in Column B.
Column A Column B
i) Nephantis serinopa Cotton
' ii) Chilo infuscatellus Wheat
iii) Agrostis ipsilon Castor
iv) Helicoverpa armigera Cabbage
v) Epilachna vigintioc topuanctata Pulse
vi) Earias vitella Ground nut
vii) Nephotettix virescens Coconut
viii) Achaea janata Brinjal
ix) Trichoplusiani Rice
x) Aphis craccivora
xi) Dacus dorsalis
- Sugarcane
Citrus
xii) Papilio demoleus Mango
xiii) Tribolium castaneum Teak
xiv) Hyblaea puera Wheat grain
Adaptation and Bel~avioural PREVENTION AND CONTROL OF ARTHROPOD PESTS
Pattern
In modem times, large number of insecticides are available to bring down the populatioq
of arthropod pests to reasonable levels. However, none of them totally eliminate any pest.
So there will be pest build up again, and insecticides will have to be thus repeatedly
employed. Most of these insecticides are poisonous not only to man and domestic
animals, but to beneficial arthropods like honey bee as well. So pesticide application has
to be avoided as far as possible or at least kept to the minimum level. This can be done by
resorting to suitable preventive measures which prevent build up of pest.

There are several preventive methods which we can employ against many arthropod ,
pests. This however, depends upon a thorough knowledge of the biology~ofthe pest. We
will here discuss some :>fthese preventive measures.
Most of the house-hold pests and those of veterinary and medical importance, breed
and multiply in unhygienic conditions. So keeping our home and the surroundings
clean and hygienic is important to prevent buildup of pests like mosquitoes,
houseflies and other related flies, bedbugs, cockroaches, lice, fleas etc. It is to be
especially noted that many species of niosquitoes breed in stagnant and foul water or
where there are chances of such water accumulating. This should be avoided.
Larivorous fish like Gambusia and Aplochilus should be grown in stagnant water
bodies like wells, tanks, ponds etc. as they will devour the mosquito larvae.
Application of crude oil (spraying) on foultstagnant water bodies will also result in
elimination of many types of mosquito larvae, as this process will prevent them
from breathing. Similarly, house flies and many related flies breed in decaying
matter, which therefore should be properly disposed off and prevented from
accumulating.
Sanitation methods can qlso be used irr agriculture to prevent build up of pests in the
field. Pest-free seeds should be employed for sowing. After harvesting the stubble
should be burned so that the different stages of the pests in the stubble will be
destroyed. The field should also be ploughed thoroughly so that egg masses and
pupae of those pests in the soil are brought above soil in the open where birds can
easily eat and destroy. Since many pests are polyphagous they will migrate after
harvesting to weeds in the field which are also their hosts. So keeping the field free
of weeds, removing the weeds from time to time, will facilitate prevention of pest
build up. L
Some pests (either adults or other stages) are fairly large, as is the case with the red
palm weevil and rhinoceros beetle. These can be hand picked, or mechanjcally
scooped out from their holes or burrows in tender palm shoot by inserting sharp
hooked or barbed wire for pulling them out and destroying.
The build up stored products pests can be prevented by drying grain and similar
products before storage, as this will kill their sensitive stage. Periodical drying may
be needed to prevent further build up of storage pests.
Borer attack can be brought down by removing the "drooping" and "wilted" shoots
or stem (which are tell-tale symptoms of the borer inside) from plants and burning
them promptly.
Many night flying insects, especiakly moths and beetles, are attracted to light during
night. So light traps can be set in fields and the pests trapped and destroyed.
Many insects are attracted to food and so mixing insecticides with their food (poison
baits) can be used to attract and kill them. Cockroaches and many moths can thus be
easily attracted and killed by this method.

These are some of the many methods available to us, and by judiciously resorting to such
tactics, pest build up can lie prevented to a certain extent.
- - -

16.5 SUMMARY
In this unit you have studied some harmful non-chordates. They may be harmful in many
ways. They may cause direct or indirect injury to man, his domestic animals or to
economically important plants or to their products. They may be parasites and cause
diseases in plants or animals including man, or they may serve only as tools in
transmitting some disease agents. Platyhelminthes, Nemathelminthes and Arthropods are
the major non-chordate groups which have many harmful species.
 Harmful Non Chordates
~la~helminthes include animal parasites belonging to the classes ~onogknea,
Trematoda and Cestoda. Many of these are serious pathogens causing disease in man
and animals. Monogeneans are ectoparasites of lower vertebrates, especially of fish.
They are often quite damaging in fish hatcheries. ~arasiiictrematodes are exclusively
parasitic on or in vertebrates. Among them, Fasciola and Clonorchis are important
liver parasites. Paragonimus is a lung fluke and the species of Schistosoma occur as
blood parasites. All trematodes complete their life c$le in two or more hosts. Their
larval stages develop in a snail which acts as a intermediate host. Additional
intermediate hosts like fish or crab may be involved in the life cycle of some .
trematodes. Cercarial or the metacercarial stage is the infective stage though which
the final host gets infected.
Cestades, like trematodes, are exclusively parasitic and have usually an indirect life
cycle. Tapeworms of the genera Taenia, Hymenolopis and Diphyllobothrivm are
common intestinal parasites of man. while it is the adult which is the main parasite
and causes injury to the hosts, larval stages of some species may also be seriously
harmful. Infection by the bladder worm of cysticercus of Taenia solium and hydatid
cyst of Echinococcus granulosus (the dog tapeworm) may be dangerous.
The animals of class Nemathelminthes are called round worms or nematodes. A large
number of nematode species (roundworms) are parasitic on either plants or animals.
They all have the same basic pattern of life cycle - adult worm, the egg and 4 '
successive larval stages each following moulting. Nematode pests of plants are called
phytopa:asites. Some nematodes act both as vectors of viruses and bacteria as well as
pathogens of plants. Meloidog~nethe root knot nematode which produces galls in
root and Heterodera, the cyst nematode, are major parasites of plants.
Animal nematode parasites lodge in the intestine or other systems in the body of
their host. The common roundworm (Ascaris), the trichina worm (Trichinella) the
whipworm (Trichuris), the pinworm (Enterobius), the filarial worms (Wurchereria
and Brugia), the river blindness worm (Onchocerca); the Guinea worm
(Dracunculus) and the hookworm (Ancyclostoma and Necator) are important
nematode parasites of man. Ascaris, whipworm, pinworm and hookworm occur as
parasites of the digestive tract of humans who contract the infection by ingesting
contaminated food or water'containing infective eggs or through penetration of skin
by infective larvae present in the soil. The filarial worms (Wuchereria and Brugia)
have in their development, a microfilaria stage which occurs in the blood stream.
Mosquitoes spread the filaria infection. The Medina worm or Guinea worm
(Dracumlus) occurs in the subcutaneous tissue of man and produces blisters which
burst releasing the infective juveniles. Cyclops infected with the larvae ofthe worm
.. spread the infection.
Many species of arthropods are harmful as they serve as carriers of disease agents for
man, e.g., typhoid, plague, cholera, yellow fever, etc. Arthropods serve as biological
vectors for many helminth parasites, and are essential as intermediate host for the
completion of the parasite's life cycle. These include mosquitoes, flies, bugs, beetles,
copepods and ticks. Many arthropods are directly harmful to inan as they themselves
cause injury or disease. Species of scorpions, spiders, ticks, mites, beetles, lice, flea
and flies cause localised injuries to man while termites, cockroaches, silverfish and
ants cause indirect damage to man. Several species of flies, lice and flea are
important parasites of domestic animals.
Many insect pests are found on a wide variety of plants. Several of these cause
damage to the plant directly, while others are indirectly damaging as they transmit
pathogenic micro- organisms. Mites, termites, locusts, several species of.beetles and
moths are important insect pests of various economically important plants, trees w d
grains.

16.6 TERMINAL QUESTIONS

1. List the three important species of Schistosomes of man and describe the life cycle of
any one.
Adaptation and Behavioural
Pattern

2. Describe the life cycle of the dog tape worm. Explain how the larval forms of this
tapeworms could be harmful to man.

3. Mention the nematodes parasitising different organs in the human body. Describe the
diseases caused by any three important nematodes and thir mode of their infection.

......................................................................................... I""""""

4. Write a brief account of arthropods as pathogens and as vectors of human diseases.

......................................................................................................
 .

. .

Harmful Non Chordates

16.7 ANSWERS
Self-Assessment Questions
1. a (i) miracidium; (ii) Fasciola hepatica; (iii) Chlornorchis (Opisthorchis) sinensis;
(iv) lung; (v) Cray fish, Crab; (vi) Bulinus, Physopsis
b (i) False; (ii) False; (iii) True; (iv) True.

2. (i) tapeworm; (ii) Taenia saginatus; (iii) Taenia solium; (iv) Hymenolepis nana
(v) Diphyllobothrium latum; (vi) hydatid.

4. (a) (i) True; (ii) True; (iii) True; (iv) False

(b) i a; ii c; iii b; iv e; v d;
(c) (i) Boophilus (ii) Babesia (iii) red spider;

5. (i)' Anophelese, Culex, Aedes, Mansonia (ii) Culexfatigans (iii) Anophelese (iv)
wrigglers, (v) Brugia malayi vi) antennae and palpi (vii) Culex tarsalis (viii) Aedes

6. (i) c; (ii) d; (iii) g; (iv) a; (v) b;

7. (i) ~ i e(ii)
; True; (iii) False; (iv) True; (v) False; (vi) False;

8. (i) g; (ii) j; (iii) b; (iv) e; (v) h; (vi) a; (vii) i; (viii) c; (ix) d; (x) fi (xi) I;
(xii) h; (xiii) n; (xiv) m.

Terminal Questions
1. Refer to Blood Flukes in subsection 16.2.2.
2. Refer to Echinococcus granulosus in subsection 16.2.2.
3. Ascaris, Enterobius, Ancyclostoma, Necator and Trichuris occur in the digestive
tract; larvae of Trichinella remain encysted in the skeletal muscles; Wuchereria
inhabits the lymph ducts while its microfilariae occur in the blood; Dracunculus
occurs in the subcutaneous tissue in the gravid stage. Trichinosis is caused by
Trichinella larvae. This infection results in severe muscular pain. Infection occurs
Adaptation and Behavioural when man ingests infected pork. Wuchereriabancrofri is responsible for
Pattern elephantiasis, a disease in which the limbs (arms and legs) become abnormally
swollen and deformed. Iylosquitoes serve as the vector and transmit the infection.
Dracunculus medinensis causes blisters with acute pain. Larvae of the worm get into
Cyclops present in water bodies and man acquires the infection on drinking cyclops-
contaminated water.
4. Refer to section 16.4 for giving examples of disease causing arthropods and also of
those which serve as vectors by referring to section 16.4.
L
 UNIT 17 BENEFICIAL NON-CHORDATES
structure
Introduction
Objectives
Broad Categorisation of beneficial nature of non-chordates
on-chordates used as food
Phylum Arthropods as source of food
Phylum Mollusca.as source of food
Non-chordates yielding food-Honey
Composition of honey
Kinds of honey bees
How is honey produced
Non-chordates yielding Industrial Products
Silk
Lac
Beeswax
Shells
Pearls
Precious corals
Sponges
Dyes and pigments
Medicinal uses of Non-chordates
Non-chordates useful in agriculture
Non-chordates improve fertility of soil
Non-chordates as pollinators
Non-chordates as destroyers of pests
Non-chordates as components in food chains and as scavengers
Summary
~erminalquestions
Answers

17.1 INTRODUCTION
'In Unit- 16 you have studied the harmful non-chordates and have seen that some of the
worst disease-producers and food-destroyers are non-chordates. In this Unit you pill
realise a a t some non-chordates are beneficial to man.

It is not the intention here to point out how they constitute significant links in the natural
cycles including the food chains, or how they serve as research material in the hands of
scientists but we will make a sqrvey as to how some of these we find extremely useful in
a variety of other.ways.

This unit will familiarise you with the variety of ways in which non-chordates are of
benefit to man, directly and indirectly.

Objectives
After studying this unit you will be able to :
categorise the ways in which the non-chordates are beneficial to humans,
i describe the usefulness of non-chordates as sources of food,
list the various industrial products of non-chordates and briefly describe the manner
I in which these are produced by them,
I cite examples of non-chordates that have been used as sources of medicines and as
1 objects of decoration,
I
I
point out how they are useful iii agriculture.

17.2 BROAD CATEGORISATION OF BENEFICIAL

NATWRE OF NON-CHORDATES
I
Many non-chordates are beneficial to humans in numerous ways This beneficial nature
can be either direct or indirect. The direct usefulness may include their utilization as food
1 or as a source of prodi.e+s thzt fulfil human needs. The indirect benefits are the result of
I their activites whirh in turn contribute to human welfare. The list below aims to classify
broadly how the non-chordates are usehl to humans.
Adaptation and Behavioural
Pattern
A. Directly beneficial
1. Used as food
2. The food they gather is used as food for human consumption.
3 . They produce useful substances
4. Used in medicine
5. Used as.objects of beauty and decoration

B. Indirectly beneficial
1. Useful in agriculture
2. They clean the environment and thus serve as scavengers.

' We shall now examine each of the above points in detail and survey the contributions of
different non-chordates to human welfare.

17.3 NON-CHORDATES USED AS FOOD

- - - - - - - -

Among non-chordates only arthropods and molluscs contain edible species.

17.3.1 Phylum Arthropoda as Source of Food

Though some primithe tribes are known to eat insects like termites and grasshoppers,
well-recognised edible species are limited to crustaceans.

CRUSTACEANS
Though many crustaceans are edible, only the larger ones are of commercial significance.
Most shrimps are small and are hence not of much commercial value. But prawns,
lobsters and crabs are larger and are hence exploited commercially. All these come under
order Decapoda. These are of great food value and are an important source of earning
especially foreign exchange. The main export products are frozen prawns, canned '
prawns, dried prawns, prawn pickles, frozen crab meat and canned crab meat.

1. Prawns and Shrimps

Prawns and shrimps have cylindrical or compressed body and the abdomen is large with q .
peculiar bend (Fig. 17.1). The carapace has a well developed rostrum. Thoracic legs are
slender, anterior ones being chelate. These swim in water.

Prawns are considered a delicacy throughout the world. There are various species of
prawns that inhabit the sea, freshwater reservoirs, rivers and estuaries. The following are
some important species of prawns that are commercially exploited:

Penaeus indicus (Indian prawn) found in coastal regions only; Penaeus monodon and
Metapenaeus monoceros (Giant tiger prawn) found in fresh as well as salt watets;
Macrobrachium malcomsonii : inhabit fresh and brackish water; Macrobrachium
scabriculum : seen in fresh water only; and Macrobrachium rosenbergii: The giant fresh
water prawn.

Carapace

Fig. 17.1: A Shrimp Crangon, side view

Penaeus indicus is the commoneit Indian prawn which constitutes the bulk of the catch.
It grows upto 20 cm in length. P. monodon grows even longer upto 30 cm. Metapenaeus
monoceros is one of the most suitable species to be cultured in India and other tropical
countries. For fresh water culture the giant prawn Macrobrachium rosenbergii is the most
suitable.

Prawn culture or farming is rapidly growing in various countries like Philippines, Japan,
U.S.A., Taiwan, as well as India where ponds, tanks, paddy fields, estuaries and coastal
sea waters are being.converted into prawn raising areas.

2. Lobsters
Lobsters are large heavy bodied marine crustaceans. They have straight dorsoventrally
flattened body, with heavier legs. Their first pair of walking legs are strongly chelate
(chelipeds) for capturing prey. They are adapted for creeping or crawling. Homarus
americanus (Fig. 17.2) the American lobster, may grow to a length of 60 cm and weigh
over 20 kg. The European lobster Homarus gammarus is smaller. Scyllarus is the Spanish
lobster. Panulirus is the Indian lobster.

Fig. 17.2: A lobsterHomarusamericanu.~.

3. Crabs
Crabs are chiefly marine crustaceans with short body and flattened broad carapace
(cephalothorax) usually as wide as It is long. They have a small abdomen that curls
forward and is held tightly beneath the cephalothorax. The chelipeds are modified as
grasping pincers (Fig. 17.3). Crabs crawl on the surface when crawling rapidly they move
sideways. Many crabs are treated as delicacy in mahy parts of the world. Porrunus
sanguinolentus, P. pelagicus and Chatybdis cruciata are some edible Indian crabs.

Fig. 17.3: A cr.b

17.3.2 Phylum Mollusca as Source of Food

Many molluscs are edible, Scallops, mussels, clams, oysters, snails, squids and octopuses
are examples. Ostrea is perhaps the best known edible oyster Crassostrea madrapemis
(the Indian backwater oyster), Crassostrea cucullata (the rock oyster) and C. discoidea
(disc oyster) are Indian edible oysters. They are marine bivalves. Several fresh water
mussels such as Unio and Anodonta are also eaten. The common edible European mussel
are Mytilus edulis and the closely related M. galloprovincialis. They are also cultivated in
the sea either on poles, or on the seabed or from fixed frames or from freely floating
structures in the sea. h@tilus viridis and M. edulis are the Indian species. The important
molluscan beds in India are along the west coast and the Palk strait and the Gulf of
Adaptation and Eehaviourrl
Pattern Mannar. The gastropods are not of much significance as a source of food, though some
species are eaten by poorpeople. So are cephalopods also in India, thou@ they are
caught from the east coast for export. Sepioteuthis arctipinnis is such an Indiaq species.

SAQ 1
i. Which two non-chordate phyla provide the largest variety of edible forms ?
......................................................................................................

ii. List any four crustaceans that are commonly eaten.

......................................................................................................

,iii. Mention if the following statements are true (T) or false (F).
(a) Prawn culture is a growing industry.
(b) Shrimps are chiefly marine forms.
(c) No insects are eaten in any part of the world.
(d) All fiesh water mussels are avoided as food.
(e) Primitive human races often include insects like beetles, grubs and caterpillars
in their food.
( f ) Shrimps can grow to a size of 30 cm in length.

17.4 NON-CHORDATES YIELDING FOOD HONEY

- - - -- - - - - -
-
We have all tasted honey at some time or the other in our lives and are familiar with the
honey bees. The honey bees are insects (order Hymenoptera). Social life has attained a
high degree of perfection in these insects and their social organisation and behaviour will
be dealt with separately under the unit Behavioural Patterns. These honey bees produce
honey and store it in their hive and use it for themselves and for their young ones. Man
discovered it and started gathering honey long back. In the early days they squeezed out
the honey from the hives of the bees that lived in wild state. Even today many tribal
people collect honey like this fiom'forests. Later, they devised vaious types of apiaries
and domesticated the honey bees. The practice of bee keeping is now very common
almost throughout the world and is designated as apiculture. In India, the Khadi and
Village Industries Commission is encouraging bee keeping on a large scale. Tamil Nadu,
Kerala, Karanataka, Jammu and Kashmir and Himachal Pradesh are major honey
producing states in India.

17.4.1 Composition of Honey

Honey is an aromatic viscid product. It is sweet a d a highlynutritious food. It is a
product modified by the honey bees from plant nectar, and is consumed in many ways, as
a spread, and is used in making drinks, candies, cakes, and is also used as a medicine.

Honey is an instant source of'energy. Its solid constituents are mostly monosaccharides
which are readily absorbed into the blood. The rich vitamin and mineral contents further
enhance the nutritive value of honey.

Chemical composition of honey: The composition of honey varies and the composition
of an average sample of honey shows the following table:

Sugars
Fructose - 40-50%
-
Glucose 32-37%
Sucrose 2% -
Maltose - traces
Polysaccharides
-
Dextrins 1- 12%
 Beneficin\ Non-ckordates
Vitamins - Abundant A,B and C vitamins
-
Minerals Traces of iron, copper, manganese, magnesium, sodium, potassium, calcium,
"
si,lica and phosphates. Acids like acetic, butyric, citric, formic, lactic, malic and succinic
are also presed as also amino acids. Enzymes like invertase, diastase and phosphatase
also occur.
-
Water 13-20%

The colour and odour of honey usually depend on the flowers from which the nectar has
; been collected.

17.4.2 Kinds of Honey Bees

.
The honeybees are insects that belong to the order Hymenoptera. There are four species
of honey bees. .
1. Apis dorsata (The rock-bee): This is the largest honey bee. It constructs its huge,
single open comb or hive on high branches of trees, on tall deserted buildings or on
rocks. 'This species cannot be domesticated, it is ferocious and has a poisonous sting
that can cause fever.
2. Apis indica (The Indian bee): This is a medium-sized bee. It builds hives consisting
of several parallel combs side by Side in dark places like the cavities of tree trunks,
mud walls, earthen pots, etc. This is not so ferocious and can be easily domesticated.
This is the species used in apiculture (Fig. 17.4).
3. ApisJlorea (The little bee): This bee is even smaller. It builds single small combs in
bushes, hedges, etc. Its honey yield is poor and therefore not economical. .
4. Apis mellijiera (The European bee): It is primarily found in Europe but it has also
been introduced in many parts of the world including India, because of the greater
quantity of honey that can be gathered from its hive.

Apis dorsata

Apis florea

Drone

Fig. 17.4: Three lndian bees: Apis indica 1. Worker. 2. Queen, 3. Drone, and other two common Indian
bees A. dorsata and A./lorea.

17.4.3 How is Honey Produced .

The worker bees visit flowers and obtain nectar from them.

Mouth parts of hon~y-bee:Adult-honeybees have mouth parts that are modified in

-
order to utilize liquid food i.e. nectar and honey. Mouth parts are chewing lapping type.
The major feeding apparatus consists of a maxillo - labial complex. Surrounding the
central 'tongue' the glossae of the labium, is a tube formed from the galeal part of the
maxillae. With the combined action of both the sucking pump and 'tongue' moving up
and down, nectar is drawn up into the body. The mandibles usually do not function
Adaptation and Behavioural
Pattern
directly in feeding but may be used not only for cutting flowers that have long corolla to
gain access to the nectar but also for defense and for molding into combs for storing
honey in the hive.

Nectar contains disaccharides (mainly sucrose). While sucking nectar the bees mix their
own saliva with it and swallow it into a honey stomach (crop). After flying back to the
hive the workers regurgitate the mixture into their mouth where they again masticate
thoroughly with saliva. The saliva is rich in the enzyme invertase which hydrolyses
sucrose into glucose and fructose. This processed nectar (honey) is deposited in the
storage cells of the hive. This rather watery honey is then thickened by evaporating a
large portion of water content by rapid beating of the wings of the worker bees. When a
cell is filled with "ripened" honey it is capped over with wax.

SAQ 2
1) List any two uses of honey.
i) .............................................................................................
ii) ..............................................................................................

2) Column I below lists some of the constituents of honey. Rearrange these in column
II according to their approximate percentage given alongside.
Column I Column I1 Percentage (Aprox.)
Water
Fructose
Sucrose

3) Give the scientific names of the following bees :

i) The domesticated Indian bee - .................................
ii) The rock bees - ...............................

4) How does saliva of bees contribute in changing nectar into honey?

......................................................................................................
......................................................................................................

5) Name any three principal Indian states which produce large quantities of honey.
i) .....................................
ii) .....................................
iii) .....................................

NON-CHORDATES YIELDING INDUSTRIAL

PRODUCTS
We have seen that many non-chordates, serve as human food or indirectly their products
like honey serve as food. We shall describe here some other uses of non-chordates.

The following industrial products are obtained from some non-chordates:

Silk
Shellac
Beeswax
She1Is
Pearls
Corals
Sponges
Dyes and pigments

17.5.1 Silk
Though various species of insects produce silk, which is'a secretion of their salivary
glands, silk of cothmercial quality is produced only by a very limited number of species.
The silk produced by the mulberrysilk worm ofthe species Bombyx mori is the most
 Beneficial Non-chordates
valued of these. This species belongsto the family Bombycidae. This silk is one of the
most valuable and widely used products. It is the secretion of the paired salivary glands of
the full grown last larval stage of the silk moth Bombyx mori (Fig. 17.5). The worm feeds
on mulbeny leaves (Morus aha).

rd&*
"p$; \
Newly born
catetpillars
I

\t
Different stages(
of larva \,
',
,

cut open Mature caterpillar

Fig. 17.5: Life-cycle of Bombyx mori.

The adult moth is creamy white about 5 cm across with fully spread wings. From head to
the tip of the abdomen, it measures about 3 cm. The wings are however feeble and the
insect scarcely flies. As adult it takes no food and lives for only 2-3 days during which
the female lays 300-500 eggs. The eggs hatch into tiny larvae by 8-12 days. These feed
on mulberry leaves and grow in size. These moult four times to ,reach full grown size of
about 8 cm in length. It is grayish and has a hump behind the head and a spine-like horn
at the hind end dorsally. Full growth is reached by 28-30 days. When full grown it
becomes restless and spins a cocoon.

Spinning of cocoon takes about 3 days. During this operation the head is constantly
moved from side to side swiftly at the rate of about 65 times per minute. As this larva
does so, the secretion of the salivary gland is continuously poured out through the
common opening at the tip of a median cylindrical spinneret on the lower lip or labium.
This is a clear viscous fluid, but as it is exposed to the air it hardens into the fine silk
fibre. This fibre forming the cocoon is continuous and ranges in length from 700 to 1100
metres. Silk consists of two proteins: fibroin and sericin. The silk thread is elastic,
resistant and non-conductor of heat and electricity. It has also good tensile strength,
comparable to steel. The cocoons are oval. The colour of the silk varies from.white to a
beautiful golden yellow.

Pupation: The larva pupates within the cocoon. The cocoon is made up of a single
reelable thread. The pupa is the inactive stage undergoing metamorphosis to become
adult moth. The adult moth emerges from the cocoon after 10-12 days. While coming
out it softens one end of the cocoon by an alkaline secretion which enables it to break
Adaptation and &bavioural through thestrands of silk. Such cocoons through which the moths emerge are called
Pattern
pierced cocoons. These are of low value because they cannot be reeled.

Reeling: For reeling (winding the thread on a wheel), the cocoons are gathered about 8
days after spinning begins, and the pupae are killed usually by heat from steam or hot air
or even by fumigation and then thoroughly dried. Next, the cocoons are immersed in
warm water. This loosens the fibre. The fibres from four or five cocoons are caught up
together and twisted into a thread that is wound on the reel.

History of Silk: Usefulness of silk was first discovered by Empreqs Lotzu in China about
2700 B.C. Since then silkworms were reared and silk was produced in that country as a
monoply for about next 2000 years. In the year 550 A.D. two European monks smuggled
out eggs of silk moth and thus introduced silk culture in Europe.

Today, sericulture, or the commercial production of silk, is an important industry in

several countries including China, Japan, India, France, Spain and Italy. In India the
~ulberrysilk is produced extensively in Karnataka, Tamil Nadu, Assam, West-Bengal,
Jammu-Kashmir and Punjab.

Non-mulberry silk industry.

You have studied above how mulberry silk is produced by the silkworm Bombyx mori.
This moth no more exists in wild state and is completely domesticated. ow ever, other
moths still exist in the wild condition. They and their products are described below.

1. Tasar Silk
Tasar silk is produced by three species of Antheraea, a moth belonging to the family
Saturnidae. The commonest of this genus is A. paphia (Fig. 17.6). Its larvae feed on
some trees like Terminalia,Dalbergia, Shorea, Zizyphus, Ficus etc. in the thick
jungles of Bihar, Madhya Pradesh and 0-rissa. The silk of its cocoons is also reelable
and is brown or coppery in colour.

Fig. 17.6: Tasar silk moth Anrheraeapaphia: Adult, egg, larva, pupa and cocoon. I

2. Muga SiIk
This is produced by Antheraea assamia (Fig. 17.7) confmed to Assam along
Brahrnaputra Valley of Jndia.
 Fig. 17.7: M u p silk moth Anthgraea m.samia.Adult, eggs, cocoon (enclosed inside leaf) and pupa.
3. Eri Silk
Eri silk is the product of the eri silkworm Phylosamia ricini (Fig. 17.8) that feeds on
Ricinus communis (Castor plants). This species also is a moth belonging to the
family Saturnidae. It is raised in Orissa and Assam commercially, and,can be reared
on castor leaves. Its silk is white or brick red in colour, though not as glossy as
mulberry silk. It is not in one single strand. So it is not reelable, but has to be spun
like cotton. So the moths can be allowed to emerge from the cocoon.

Fig. 17.8: Eri silk moth Phylosanu'a ricini, Adult, eggs, larva, cocoon and pupa.
;'Adaptation and Behariourai
Pattern 17.5.2 Lac
Lacand the related shellac are familiar substances used for making french polish and
floor polish, gramophone records, bangles, printing ink, electrical insulators and sealing
wax. It is also employed for a host of other purposes. However, now-a-days substitute
synthetic substances are being used for some of the above items.
I

Lac is the product of a tiny insect Laccij2r facca, popularly called the lac insect: The
insect lives on a number of forest trees, especially Buteafrondosa, (Palas), Zizyphus
ji!jrrha (Ber) and Schkichera frijuga (Kusum) . Besides India, lac insect also grows on
certain forest trees in Pakistan, Myanpar (formerly Burma), Malaysia, Sri Lanka, China,
Thailand and IndoChina.

Life History: Minute young lac insects called crawlers (larvae) hatching out of the eggs
find a suitable place or a twig. Then they insert their proboscis or beak into the succulent
plant tissues. and start sucking the plant sap. Thus they grow and secrete a resinous
material that ulti~natelycovers it. The resinous encrustation increases in size with growth
of the insect. As thousands of crawlers settle side by side and the resinous encrustation
builds up around them, it ultimately completely encloses the twig. After three months,
most crawlers develop into females which cannot escape from the resinous mass. The
males, which may be winged or wingless forms, emerge and fertilise the females through
minute openings, that extend to the surface of encrustation, thereafter the males die. As
tlie eggs develop inside the body of the female, she grows fast and assumes a sac-like
bright red appearance. This red pigment is a source of lac dye. The female dies inside its
little chamber, the eggs hatch into crawlers which escape through the narrow opening,
move to a nearby uninfested part of the twig and the process is repeated (Fig. 17.9).

Wingless male

k .8 Male and female lac

Lac encrustate on ,

a twig

Fig. 17.9: Lac insect. I.Larva,.2. Wingless male, 3. Winged male, 4. Adult female, 5. mala and females on
a twig. 6. Resinous encrustation encloses twig.

, Stick lac and Seed lac - The encrusted twigs are called the stick lac. Such twigs are
removed and the encrustation is then scraped, ground, soaked and washed, now it is
 Beneficial Non-chordates
called seed lac. It is then processed with certain other materials to make shellac in the
form of thin orange or yellow flakes.

17.5.3 Beeswax
Beeswax is the material that makes the beehive. This material is a secretion of the worker
bees that is poured out from the wax glands that open on the underside of the abdomen
forming thin delicate scales. The worker bee removes these wax flakes with its legs, picks
them up into mouth, manipulates by the mandibles and builds the combs of the hive. Wild
bees especially Apis dorsata is the main source of beeswax in India. Beeswax is used in
manufacturing candles, shaving creams, cosmetics (various creams, lotions, lipsticks,
eyebrow pencils, rouges, pomades etc.), polishes, floor waxes, models, crayons, etc.
However, now-a-days, in many cases the use of beeswax has been replaced by paraffin
wax and other products.

17.5.4 Shells
"Shell" is applicable to the hard skeletal structures, chiefly of the two phyla Protozoa and
Mollusca. In Protozoa the foraminiferans possess a hard calcareous shell and in Mollusca
most have either a single valved shell (gastropods and cephalopods) or a double valved
shell (bivalves). The protozoan shells have mainly contributed to the formation of chalk
as you have already learnt in Unit-:! of this course.

Chalk - Chalk is a soft white, gray or buff limestone chiefly composed of the shells of
foraminiferans such as Elphidium (Polystomella), and Globigerina. Shells of these dead
organisms cover 30 to 40 per cent of the ocean floor as "foraminiferan ooze". In course of
time the products of these bottom depbsits have become rocky limestone and chalk.
When uplifted geologically these form the.limestone mountains and the chalk cliffs.
Chalk finds use in several ways.

Molluscan shells
Some of the well known commercial or industrial uses of molluscan shells are the
following: Shells of a variety of molluscs have been used for making quick lime used for
construction work. Shells of fresh water mussels in particular have been used for making
buttons almost throughout the world. These are lustrous and shiny and are not affected by
washing. However, these days buttons are mostly made from synthetic material.

Cowries - Cowries are exquisite shells of Cypraea and related species (Cypraeacea).
Their shells have been used in making ornaments and other decorative items. Cowries are
also used in certain ind~orgames.

Conches - These are the shells of certain large spiral shelled marine gastropods (such as
the genera Strombus and Cassis). These shells have been used for making sculptures or
carving on them. The sacred shankh, Xancus pyrum is a conch. This is used in temples,
for blowing. A number of other shells are beautiful items for collectors. Several
gastropods like Triton and Terebra arid a cephalopod Nautilus are specially very
attractive. The sectional view of Nautilus reveals the lovely internal design. The inner
surface of the shell is very smooth with pearl-like shine and hence this mollusc is also
called pearly Nautilus.

-
Cuttle bone It is the calcareous internal shell of cuttle fish Sepia. Sometimes it reaches
a fairly large size of about 20 cm length.

Cuttle bones are light due to minute air spaces inside. As the animal dies and finally
decomposes, the clean cuttle bones float on the surface of the sea and are often thrown on
the sea shore. The cuttle bone has been widely used for polishing and as a food
,supplementfor pet birds including poultry.

17.5.5 Pearls
Pearls have been highly valued by man since ancient times. These beautiful and costly
objects, have been used in making jewellery. Pearls are produced by two kinds of
molluscs:
Adaptation and Ikhauioural
Pattern
i. Pearl oystm: Several species of P inctada, with common species P. vulgaris and P.
margaritifera. Pearl pysters (Fig. 17.10 A) are all marine and these produce oriental
pearls of good quality.
ii. Freshwater mussels (Unio and ~nodonta):Pearls produced by musSels are of
inferior quality.

Fig. 17.10: A. Pearl oyster, complete. 8. Pearl oyster openedYo showprltrk in it.

Sites of pearl production: Main sites of pearl production are Persian Gulf, Gulf of
Mannar, Palk Bay, Gulf of Kutch, Panama Bay, and Gulf of California, But now Japan is
the largest pearl producer through their pearl culture technique.

Formation of Pearl
Before we study formation ofpearl, let us see the structure of the shell of the mollusc.
The shell is secreted by the underlying mantle. Cross section through a shell valve of the
mussel or oyster shows three distinct layers that constitute the shell (Fig. 17.1 I).
i. . Periostracum: Outermost horny layer. This protects the shell from dissolution.
ii. Prismatic layer: It is calcareous and is made up of minute crystals of calcium
carbonate, separated by thin layers of conchiolin.
iii. Nacreous layer: Innermost smooth layer of the shell. This is also called the "mother
of pearl". It is composed of calcium carbonate crystals aligned parallel to the mantle
surface. This is the lovely shining and smooth inner surface of the shell valves.

The mantle has also three layers.

i. Columnar epithelium: This is immediately next to the nacreous layer of the shell.
It consists of cells that secrete the "Mother of Pearl".
ii. Connective tissue layer: This is the middle layer.
iii. Ciliated epithelium: This is the layer farthest from the nacreous layer (Mother of
pearl) of the shell and it secretes mucus.

Pearl is produced when a minute foreign pbject like a sand grain or a parasite larva lodges
itself between the mantle and the shell valve. As a protective measure against the
irritating foreign object, the epithelium ofthe mantle slowly encloses the object forming
an epithelial sac. The epithelium secretes concentric layers of nacreous substance. In
about 3-4 years, the pearl is completed (Fig. 17.1 1).

Fishing of pearl oysters is carried out by divers. These are left exposed to air for a few
days for the organic matter to decompose. Finally they are washed and the pearls are
picked up by hand. '

Pearl culture is a new technique chiefly developed and practised in Japan. Small artificial
objects are introduced between mantle and the shell of the pearl oyster. This
"impregnated" oyster is allowed to live in its normal habitat and in course of time it
produces pearl.
 :ial Non-chordates

parasite \ - Pearl
Mantle
epithelium

*
Fig. 17.11: P a r 1 formation. A. A foreign object caught between mantle ahd shell. B. the mantle
epithelium encloses the object and starts secreting nacre i n concentric layers arqund it. C. ,
Completely enclosed sae showing a pearl inside it. D. A pearl attached to the shell valve.

17.5.6 Precious Corals

Coral is a general term employed to denbte the calcareous or the horny skeletal deposits
produced by certain cnidarians (anthozoans and rarely hydrozoans.). One of these, the red

+Pinnate tentacles

Fig. 17.12: The red coral of commerce Corallium rubrum.

Septa

Fig. 17.13: Fungus coral, Fungia.

Adaptation and Behavioural
coral of commerce is highly valued and is precious. It is used in making jewellery. This
Pattern
coral is the skeleton produced by Corallium rubrum which grows as a branched colony
(Fig. 17.12). 'The red coral of commerce naturally occurs ig the sea around Japan and in
the Mediterranean sea.

Several corals (Cnidaria) are appreciated for their beauty. The fungus coral (Fungia),
(Fig. 17.13) rose coral are some such examples that lend beauty to any collection of
natural objects.

Fig. 17.14:Venus's flower basket, Euplectella aspergillurn.

17.5.7 Sponges
Sponges have been in use since ancient times. The Greeks, and later the Moghuls in
particular, used the dried fibrous skeletons of the sponges for various purposes - bathing,
washing, scrubbing floors and padding shields and amour. The Romans used them for
painting. These were also being used in hospitals, infant nurseries, etc. All these are
actually the skeleton of the sponges. However, the synthetic "sponges" have virtually
replaced the natural ones.

The water-holding capacity of sponges, is on account of the minute capillary spaces

enclosed by the spongin fibres.

Commercial sponges are obtained exclusively frorh the sea. The common bath sponges
are Spongia and Hippospongia. They come under family Spongiidae. They are gathered
from the Gulf of Mexico, the Caribbean and the Mediterranean. Some of these are as
follows:
Source animal Place of occurrence
Euspongia offcinalis Mediterranean and Adriatic Sea
Spongia dura Bahamas
Hippospongia lachne Bahamas

After collection their living tissues are allowed to decay. These are then thoroughly
washed, cleansed, cut, trimmed, bleached and graded.

Venus flower basket Euplectella is a glassy sponge, its skeleton (Fig. 17.1) is an object of
beauty often used as a gift.

17.5.8 Dyes and Pigments

Best known,examples under this category are certain pigments obtained from insects. A
very comnion pigment widely used since olden times is that of cochineal dye. It is a
beautiful red dye that has been much used in textiles and is highly preferred as it imparts
permanent colour. It has also been used in cosmetics, as a colouring for beverages and in
decorating cakes and pastries. Its source is'a tiny insect Dactylopius coccus that lives on
the prickly pear Opuntia coccinellijera in Mexico. The related Dactylopius~omentosus
occurs in India, on Opuntia dillenii. These insects are close relative of lac insects. The
dye is obtained by grinding the dried body of these insects.
Dyes from insects galls: Galls are peculiar growths in plants caused by a number of
insects. These insects insert their eggs in plant tissues which produce the gall containing
the developing larval stages. Some galls yield dyes used for dyeing hair, wool, leather
and for preparing permanent inks. Some galls contain high percentage of tannic acid.

17.6 MEDICINAL USES OF NON-CHORDATES

Some non-chordates are of use in medicine especially in traditional, ancient medicine.
Leeches are still widely used in blood letting. People believe that the leeches suck the
"bad blood".

Honey bees have been used to extract a medicine from the excited worker bees used to
cure diphtheria. Bee venom is used for curing rheumatism and arthritis.

"Allantoin" is obtained from the maggot larvae. It is said to be effective in treating deep
wounds.

The Spanish fly Lytta (Cantharis)visicatoria and Mylabris species are an excellent
source of cantharidin, a pharmaceutical product.

SAQ 3.
1) Match the commercial products given in column I with the source animals in
column 11.
Column I Column I1
(Commercial Product) (Source animal)
a) Mulbeny silk i. Laccfer lacca
b) Pearl ii. Antheraea paphia
c) Lac iii. Bombyx mori
d) Tasar silk iv. Corallium rubrum
e) Honey v. Pinctada margaritifera
-f) Red coral of commerce vi. Euspongia
g) Bath sponge vi i . Apis indica

2) Why is it necessary to kill the pupa in silk cocoons to obtain silk fibre for realing?

3) ' Fill in the blanks:

.. The eri silkworm mainly feeds on the leaves of ................................tree.

i)
11) The adult ............................ of the lac insect cannot come out of its
resinous chamber, and finally dies in it.
iii) Chalk is a substance produced by certain members of the phylum
.................................
iv) The cochineal insect lives on ...................................
4) Name any four kinds of molluscan shells and their main commercial or industrial
use.

.5) What induces the pearl oyster to produce a pearl?

.......................................................................................................

6) ' Given below is a list of certain non-chordates:

Maggot larvae, honeybee, leech, Spanish fly
Adaptation and Behavioural
Pattern
Fill in the blanks by choosing appropriate non-chordate
.
i) Allantoin for treating deep wounds is obtained from ..............................
ii) .:.
cantharidin is obtained from. ...... .............................
iii) The bee venom used to treat rheumatism and arthritis is obtained from
. . ...............................
, .
i.......... .
. . iv) ............;..;,,'. ......:.........:......is sometimes used for blood letting.
7. Give one example each from the following phyla, which is used as decoration
object.
. t.
i) Mollusca ...............,.'.......................
.:. ......,.
ii) Cnidaria ........................
. ,
~

17.7 NON-CHORDATES - USEFUL IN AGRICULTURE

Though many non-chordates are harmful to the crops as you have already, seen in Unit 16,
there are several of this group that are remarkably useful. Three major areas of their
usefulness are:
1. In improving fertility of the soil
2. In bringing-aboutpollination
3. In destroying harmti1 pests

17.7.1 Noh-Chordates Improve Fertility of Soil

There are two major contributors which improve the physical condition and the fertility
of soil. These arc:
i. Earthworms ( Phylum Annelida)
ii. A variety of insects ( Phylum Arthropoda)

Earthworms: Earthworms have been regarded as excellent friends of farmers. They

,improvethe soil in many ways:
i. As they burrow through the soil they build channels and furrows which assist in
aeration and irrigation and improve drainage. It helps in mixing and churning of
the soil. This is perhaps the biggest use of earthworms.
ii. During feeding, earthworms take decomposing matter from surface leaves and
coarse soil particles into their digestive tracts, grind them up and after absorbing
organic matter from them, pass out the fine pulverised soil constituents as
castings. This is an excellent organic manure. Thus the soil is pulverised and
turned over from lower levels to upper surface.
iii. Nephridlal excretions add to the nitrogenous content of the soil.
iv. At death the earthworms enrich the soil with the organic remains of their
decomposed body. = ,

Insects: Many insects and their larval stages habitually live in soil. They make burrows
and make the soil porous thus contributing to what the earthworms also do. Some
common examples of such insects are ants, beetles and grubs (beetle larvae), and certain
species of wild b q s . Many insects like beetles, ants, termites and spring tails
(collembolans)break down the fallen leaves, twigs, etc. into minute pieces, eating them
up and return the nutrients back to the soil.
h

17.7.2 Nan-Chordates as Pollinators

Fruits rarely develop and seeds seldom set in without pollination. Moreover, cross
pollination b e e e n two plants of the same kind has definite advantages. Animal visitors
which go round from flower to flower in pursuit of nectar and pollen play an important
role in cross pollination. Insects are intimately associated with this activiv, and there are
'
thousands ofspecies of insects playing this role as pollinators.

Plants pollinated by insects

Some major crops that largely benefit from insect pollin~tionleading to hl $er
agricultural and horticultural yield are given below.
 Beneficial Non~hordates
Apples Oranges Brinjals
Pears Lemons Tomatoes
, Peaches Melons Soybeans
Plums ~ucum be& Sunflower
Cherries Beans Cotton
Strawberries Peas Tobacco
Figs Many ornamental flowers
'
A large number of forest trees and other wild plants owe their continuance of species
solely to insects.

Common insect Pollinators

Among the thousands of insect pollinators some are mentioned below.

Honey bees, wild bees, bumble bees, wasps, a variety of flies, butterflies, moths, some
beetles and ants. In fact, orcharders, especially in many western countries practise bee
keeping to enhance productivity in orzhards.

-
Some adaptations of insect pollinators and the flowers pollinated by them.
The insect pollinated flowers in general, are large, usually brightly coloured, scented, and
possess nectaries. Their pollen grains are relatively large and sticky.

Most insect pollinators have a long probing type oftongue (proboscis) which they thrust
into the deep nectaries of flowers. While they do so their bodies are rubbed against the
anthers to'get a powdery smear of the pollen. As the insect visits another flower of the
same kind, this pollen gets brushed against its stigma to accomplish pollination. The
honey bees are one of the most frequent visitors of flowers for nectar but they also collect
large quantities of pollen which they use as "bee-bread" for their developing larvae. For
collecting pollen the bees have a special "pollen basket" on the tibia of their hind legs
(Fig. 17.15). This pollen gathering habit of honey bees make them potent pollinators.

I Fig. 17.15: Hind leg of honey bee, showing pollen basket on tibia and wax sheen on tarsus.
In the so-called hawk-moth orchids, the nectar in their flqwers lies at theibottom of a long
narrow tube and is accessible only to the "long-tongued" hawk moths. While probing for
nectar the moth brings each eye againit a sticky disc to which a massgf pollen is
I attached. Then the moth flies away carrying the mass (pollinia) on. its eyes. When the
moth inserts its proboscis into another flower the pollinia fit perfectly against the stigma
and adhere to it.
I
I
17.7.3 Non-Chordates as Destroyers of the Pest (Biological Control)
The three non-chordate groups which play important role in destroying various pests
come under Protozoa, Nematoda and Arthropods respectively.
Adnptntiun and Behnviournl
Pattern
Protozoa
Several Protozoan species are parasites o f certain nematodes and insects that attack the
agricultural crops. This is happening in a natural way, and offers the possibility of a wider
application for achieving biological control o f harmfill nematodes. The protozoan
Mul/>igharnoehaIocilstae is known to be pathogenic to grasshoppers. Farinocystis tribolii
infects the grain beetle Triboliurn castaneurn in India. Many protozoans such as Noserna
hornbycis and Nosenla lyniantriae are used against the insect pest European corn borer
(Ilstrinia nuhilulis).

Nematoda
Many nematodes, especially the rhabditids, in conjunction with bacteria, form disease
complex affecting maly insect pests. The nematode Neoaplectana carpocapsae forming
a co~nplexcalled DD-136,along with the bacterium Achrornobacter nernatophilus is an
example. I t affects caterpillars o f the codling moth Cydiapomonella.

Arthropoda
Arthropoda includes both predators and parasites that destroy harmful pests. Predators
capture and devour their prey. Scorpions, spiders, centipedes and a very large number o f
insects are useful predators. A few exa~npleso f insect predators are given below:

~iagonflies(odonates) - Adults feed on some pests

Aphid-lions - These are larvae o f caddisflies that eat aphids (plant lice)
Lady bird beetles - Feed on aphids
(Coleopterans)
and their larvae
Ground beetles . - Adults feed on various pests
(Coleopterans)
Syrphid fly larvae - Feed chiefly on aphids
(dipterans)

There are several parasitic insects which lay their eggs in or on the body o f certain insect
pests (Fig. 17.16). These parasitic insects belong mainly to the order Hymenoptera
(Families: Ichneumonidae, Braconidae, Encyrtidae, Eulophidae, Aphelinidae,
Pteromalidae and Trichogrammatidae) and Diptera (Family: Tachinidae). The eggs hatch
and the larvae feed on the tissues o f the host, ultimately destroying it. The adult parasite
emerges out before the host insect dies continues egg laying, seeking out other host

-
insects. Certain parasites lay their eggs inside the eggs o f other insects. Trichograrnma
rninlrtlrrn is an example o f such an egg parasite (Fig. 17.27).

Fig. 17.16: A parasitic chnlrid wnsn i ~ ~ t r o d u rnn

i ~ egg
~ g inside nn aphid.

Fig. 17.17: Trichogramma dnutum lnjectillg its egg into the egg of n moth.
 -3cncfleirl Non-ehordrtcs
Many of the predators and parasites are being employed in a scientific way to control
pests. This method of controlling pests is known as biological control. Thus the pest
cottony cushion scale, Iceryapurchasi (coccid) was controlled in California (USA) by the
predatory beetle Rodolia cardinalis (a lady bird beetle of the family Coccinellidae).
Attempts are being made to control the coconut blackheaded caterpillar Opisina
I arenosella (Nephantis serinopa) by parasites Peresierola nephantidis, Trichospilus
pupivora and Bracon brevicornis. These are only some examples of organisms employed
in biological control. It may also be mentioned here that some insects keep weeds under
check and have been used as biological control agents. Thus, the prickly pear, Opuntia
inermis infesting extensive land mass in Australia was eradicated by introduction of an
'insect Cactoblastis cactorum.

SAQ 4
1. List any three ways in which non-chordates are useful to agriculture.
1.
11.
iii.

2. Name any four non-chordates that substantially increase the porosity of the soil.

ii.
...
111.
iv.
I 3. How do earthworms improve the fertility of soil?
i!

4. Match the items in column I with the animals in column 11.

Column I Column 11
a) Pollen basket i. Larvae of ladybird beetles
b) Predator of aphid ii. Trichogramma minutum ,

c) Parasitic insect iii. Honey bee

17.8 NON-CHORDATES AS COMPONRNTS IN FOOD

CHAINS AND AS SCAVANGERS
The great majority of non-chordates form integral parts of food chains. Many such food
chains ultimately produce organisms that may either be used as human food or in some
other way useful to human beings. Small fish feed on minute aquatic organisms
collectively called the plankton (a collective term for the p~ssivelq'floating or weakly
swimming very small organisms). The larger fish which devour the smaller ones
ultimately depend on this planktonic life since the small fish eat plankton. In this way,
most planktonic organisms in the ocean constitute important useful components in the
food chain, ultimately increasing the fish population in the ocean and inland reservoirs.

Scavengers : Any organism that feeds on dead material belongs to this category.
Practically all phyla have some examples of scavengers. Arthropoda in particular have
numerous such examples centipedes, millipedes, many crustaceans, common flies and a
number of insects are scavengers. Ants, dung beetles, flesh flies etc. are examples.

SAQ 5
1. Mention if the following statements are true (T) or false (F).
i) All food chains are beneficial to humans..
-
l
.l
ii) Marine zooplankton largely consists of some ciliates and flagellates and
a lot of small crllstaceans.
iii) Non-chordates are constituents of food chains leading ultimately only to
fishes.
Adaptation and Behavioural '

Pattern
2. "Centipedes and ants can be regarded as scavengers". How do you justify this
statement?

Many non-chordates are beneficial to man in a number of ways. Some are used as
1
food, some collect food that can be used by man, some yield industrial products, and
some have an important role in food chains.
The commonest edible non-chordates are prawns, shrimps, lobsters, crabs, etc. and a 1
few molluscs like the oysters and freshwater mussels. Honey is a main food item
I
collected by a non-chordate that is widely used by man. Honey is mainly composed
of predigested monosaccharides with rich vitamin content. In India apiculture
employs mainly the Indian honey bee Apis indica for obtaining honey. Apis melljfera
is the European species, gaining popularity in India also, due to greater quantity of
honey it produces. Silk is the salivary secretion of silkwohns, with which they spin
cocoons. There are four common silk moths in India Bombyx mori, the mulberry silk
moth, Antheraea assamia whicb gives muga silk; Antheraea paphia, the source of
tasar silk and Phylosamia ricini which produces eri silk. I

India is the largest lac producing country. The tiny insect Laccifer lacca thrives on
several forest trees. The minute male and female lac insects crowd on the twigs and
secrete resinous protective chambers which form encrustations, which is lac. Lac is
used extensively in several industries. Usehl body secretions of non-chordates also
include beeswax and a variety of protective mollusc shells. Chalk is the shell
material of foraminiferan protozoans. Molluscan shells find several uses such as in
making of buttons, and object of beauty and decoration. Pearls are objects of beauty
and are used in making jewellery. These are produced by pearl oysters and clams as a
protective measure by secreting pearly layers around some foreign object like a
parasite larva lodged between mantle and shell. Pearl culture and pearl fishing are
now well established industries.
Certain corals like the red coral of commerce have ornamental value. Some other
corals may be used as decoration objects. Several species of sponges are used for
bathing, washing or padding purposes.
A crimson red cochineal dye is obtained by grinding the dead bodies of the cochineal
insect Dactylopius cactus that lives on a prickly pear.
Medicinal uses of non-chordates are largely traditional. Leeches, honey bees, and
"spanish flies" have been used for treating certain ailments. Many non-chordates lend
beauty to the environment and serve as articles of decoration.
Earthworms and several insects improve the soil through their burrowing and other
activities. Pollination of crop plants by the bees and other insects results in fruit
setting and thus contributes towards human welfare. Similarly, many arthropods kill
agricultural pests either as predators or as parasites, and keep them under check.
Some insects feed on weeds and keep them under check. A great many non-
chordates are intermediate links in food chains that are ultimately beneficial to man.
Quite a few non-chordates are valuable scavengers keeping the environment clean.

TERMINAL QUESTIONS
1. Categorise the different ways in which non-chordates are beneficial to man giving .
one example of each.
2. Describe how the non-chordates serve as food. J
Beneficial No chordates

. How are honey bees useful to man?

.....................................................................................................

4. Describe the varieties of silk and how it is obtained.

.....................................................................................................
.....................................................................................................
.....................................................................................................
................
....................................................................................
i

5. What is lac? List any three trees on which lac is commonly produced.
.....................................................................................................

6. How are pearls produced?

17.11 ANSWERS
,
-- -

Self Assessment Questions

1. i. Arthropods, Mollusca
ii. Prawns (eg. Penaeus), shrimps (eg. Crangon), lobsters(eg. Homarus) crabs
(eg. Portunus).
iii. (a) T, (b) T, (4 F, (4 F, (el T, (0 F

2. i. (a) used as a foodlnutrient

(b) used in medicines ,

ii. (a) Fructose

(b) Water
(c) Sucrose
Adaptation and Behavioural
Pattern
iii. (a) Apis indica
(b) Apis dorsata
iv. saliva contains invertase which converts sucrose of nectar into glucose and
fructose.
v. Tamil Nadu, Karnataka, J,ammu and Kashmir.
,
3. i. (a) iii, (b) v, (c) ii, (d) vii, (e) iv, (f) vi
ii. If the pupa is allowed to grow into adult, it will break through the cocoon and
will not yield a continuous silk fibre.
iii. (a) Castor, (b) female, (c) Protozoa, (d) Prickly pearlopuntia coccinellifera,
- -
iv. Freshwater mussel button; cuttlefish cuttle bone; Marine gastropods1
-
Shombus - conch, Cyprea cowries, Oyster - quick lime a building material
v. Any foreign object such as tiny parasite which gets lodged between shell and
mantle.
vi. (a) Maggot larvae, (b) Spanish fly, (c) Honeybee, (d) Leech
vii. (a) Nautilus, (b) Fungus coral ( ~ u n ~ i a )

4. i. (a) Improve fertility of soillmake soil porous

(b) Bring about pollination
(c) Destroy harmful pests
ii. (a) Earthworms, (b) ants, (c) grubs, (d) termites
iii. (a) Burrow through soil making it porous
(b) Excrete out nitrogenous wastes which are thrown out mixed with soil as
castings
(c) Soil turned over from deeper layers to upper surface
(d) At death their organic remains decompose
iv. (a) iii, (b) i, (c) ii

5. i. (i) F, (ii) T, (iii) F

ii. Both can eat dead organisms, and thus clean the environment.

Terminal Questions
1. i) used as food prawns
ii) Food gatherer honey bee
iii) Secrete industrial products silk worm
vi) Yield medicine allantoin
v) Decoration corals
vi) Support agriculture honeybee
vii) Parts of food chains crustacean larvae
viii) Clean environment dung beetles .
2. Primarily two non-chordates phyla are providing food to humans, these are
Arthropoda and Molluscs. Among arthropods the prawns (several species of the
genera Penaeus, Metapenaeus, Macrobrachiurn). shrimps such as Crangon, lobsters
like Homarus and several crabs are delicacy. Among molluscs the edible oysters an(
mussels are a precious food item.
3. Honey bees pollinate flowers, contributing to higher yield from agricultural and
horticultural crops. Honey bees produce honey which is a rich source of energy and
nutrients. Honey is used in medicines. Honey bees secrete wax as the nest-building
material. Bees wax has numerous industrial uses.
4. There are four principal kinds of silk - mulberry silk obtained from Bombyx mori,
tasar silk from Antheraeapaphia, Muga silk from Antheraea assamia and Eri silk
from Phylosamia ric.ini. Silk is the secretion of the last larval stage, and it comes
from the salivary glands. It is a continuous fibre that is used by the larva to spin the
cocoon. Normally adult moth emerges from the cocoon through a hole produced
cocoon. But for industrial purpose the cocoons are heated to kill the moth inside and
a continuous silk fibre is obtained for reeling.
5. Lac is a resinous secretion of the insect Laccifer lacca. Lac commonly grows on
- "Palas", "Ber" and "Kusum" trees.
6. Pearls are produced by the pearl oyster and by some freshwater mussels, when any
foreign object like parasites or tiny inert materials lodge themselves between shell
valve and the mantle.. The mantle epithelium encloses the object and starts secreting
pearly material around it in concentric layers to produce the spherical pearl.
 Beneficial Non-chordates
GLOSSARY
Host Animal or plant which sustain a parasite.

Intermediate host: ordinarily the host in which live the l q a l or immature or asexual
stages of a parasite:

Parasite: an organism that depends on its host for some essential metabolite, i.e., for its
survival.

Periodicity: recurrence of a parastie at regular time intervals.

Vector: an essential intermediate host in which the parasite undergoes some development
and which transmits the parasite further to a newer host.

FURTHER MADINGS -.-

- - -

Animal Agents and vectors of Human diseases, E.C. Faust, P.C. Beaver and R.C.
Jung, Lea and Febiger, Philadelphia, 1975.
Parasitology, the Biology of Animal Parasites, E.R. Noble and G.A. Noble, Lea and
Febiger, Philadelphia, 1982.
Introduction to Parasitology, A.C. Chandler and C.P. Read, Chapman and Hall,
London, 1961.
Parasitic Diseases, M. Katz, D.D. Despommier and R. Gwadz, Springer Verlag, -
N.Y., London, 1988.
Ecology ofAnimal Parasites, Jean G. Bear, University of Illinois Press, 1952.
Applied Entomology, P.C. Finemore and Alka Prakash, Wiley Eastern Limited,
1992.
Elements ofEconomic Entomology, B. Vasantharaj David and T. Kumara Swami,
Popular Book Depot, Madras, 1975.
Text Book of Applied Entomology 4 K.P. Srivastava, Kalgani Publishers, 1993.
Invertebrate Structure and Function by Barrington, E.J.W. (1967). Thomas eli ion &
Sons, London.
The insects Structure and Function by Chapman, R.F. (1978), ELBS series.
Invertebrate Zoology (Vol I and 11) by Kaestner, A. (1967), Interscience Publishers
New York.
Zoology Phylum Series (Coelenterata, Annelida, Arthropoda and Mollusca) by ?
Kotpal, R.L. (1983), Rastogi Publication, Meerut.
General Entomology by Mani, M.S. (1988), Oxford and IBH Publsihing Co., New
Delhi. ,
The Procession by Life by Romer, A.S. (1968, The World Publishing Co., New
York.
NOTES