Respiration in Arthropoda

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Respiration in Arthropoda
Bsc zoology (University of Calicut)
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Zoology
Lesson: Respiration in Arthropoda
Lesson Developer: Sarita Kumar
College/Dept: Zoology, Acharya Narendra Dev College
University of Delhi
Institute of Lifelong Learning, University of Delhi

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Table of Contents
 Introduction
 Book Gills –Organs of Respiration in Aquatic Arthropods
 Structure of a book gill
 Mechanism of respiration through book gills
 Book Lungs –Organs of Respiration in Terrestrial
Arthropods
 Structure of a book lung
 Mechanism of respiration through book lungs
 Tracheal System – Respiratory System in Insects
 Structure of tracheal system
 Gaseous exchange via tracheal system
 Respiration in Aquatic Insects
 Plastron respiration
 Tracheal gills
 Blood gills
 Spiracular gills
 Breathing tube
 Summary
 Exercise/ Practice
 Glossary
 References/ Bibliography/ Further Reading

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Introduction
Arthropoda is the largest phylum of the animal kingdom which comprises on 75-80% of
animals. They live in diverse kinds of habitats and have, thus, developed a variety of
structures and organs for breathing. Some of these organs include book gills, book lungs
and tracheal organs. In fact, a few arthropods species have very soft skin and they can
exchange gases with the external environment through skin.
Value Addition: Did you know!!
Heading Text: Evolution of respiratory organs in arthropods
adaptations in locomotion, breathing, reproduction, and mechanisms for food capture

Body Text: The evolutionary transition from aquatic to terrestrial life led to. The
authors studied the fate of ancestral gills during the evolution of chelicerates by
examining the expression of developmental genes, pdm/nubbin and apterous in
crustacean epipods/gills. They found that –
a) In the horseshoe crab, pdm/nubbin was specifically expressed in opisthosomal
appendages that give rise to book gills.
b) In spiders, pdm/nubbin and apterous were expressed in successive segmental
primordia that give rise to book lungs, lateral tubular tracheae, and spinnerets.
The authors suggested that fundamentally different new organs (wings, air-breathing
organs, and spinnerets) evolved from the same ancestral structure (gills) in parallel
instances of terrestrialization.
Source: Damen WGM, Saridaki T and Averof M (2002) Diverse adaptations of an
ancestral gill: A common evolutionary origin for wings, breathing organs and
spinnerets. Current Biology, 12:1711–1716.
Book Gills – Organs of Respiration in Aquatic Arthropods
Book gills are characteristic respiratory organs of aquatic arthropods, especially

crustaceans; though these can also be observed in some arachnids, such as Limulus. These
are present at different places of the body, based on which they can be of various kinds.
a) Podobranchs: These are also called foot gills as they are attached to the coxa of
an appendage.
b) Arthrobranchs: These gills are attached to the arthrodial membrane which joins the
leg to the body of a crustacean. These are also called joint gills.
c) Pleurobranchs: The side gills attached to the pleural membrane of the body
segment bearing the limb are termed as pleurobranchs.

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Fig. 1: Different Kinds of gills in a crayfish based on their location

A. 1. Walking legs, 2. Gills 3. Carapace 4. Cheliped 5. Maxilliped 6. Compound eye 7.
Antennule 8. Antenna
Source: A.
https://bentonzoology.wikispaces.com/Crawdad+and+Grasshopper+Dissection+-+Internal
B. ILLL in house

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Structure of a book gill
The book gills are often flap-like or crescentic in shape. They are called so because they
resemble a book. Each gill is made up of a number of thin structures, called gill lamellae
or gill plates. These are arranged like the leaves/pages of a book at right angle to the gill
base which is also called gill axis. The nerves and blood vessels enter and leave the gills
through the gill axis.
The gill lamellae of a book gill are of diverse shapes. Based on their structure, the book gills
in arthropods are of three types:
a) Phyllobranch: The gill lamellae are leaf-like, for example in Palaemon.

b) Trichobranch: These gills have hair-like lamellae, e.g. in Astacus
c) Dendrobranch: These are the gills with lamellae divided into arborescent bundles.
(a) (b)
Fig. 2: Location of book gills in horseshoe crab
Source: (a)
http://faculty.clintoncc.suny.edu/faculty/michael.gregory/files/bio%20102/Bio%20102%20l
ectures/Animal%20Diversity/Protostomes/Ecdysozoans/Image11.jpg
(b) https://stormhawks2013.wikispaces.com/JK

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Fig. 3: Location of book gills in a crayfish

Source:
http://faculty.clintoncc.suny.edu/faculty/michael.gregory/files/bio%20102/Bio%20102%20l
ectures/Animal%20Diversity/Protostomes/Ecdysozoans/Image13.jpg
Value Addition: Interesting..!!
Heading Text: Multiple functions of crustacean book gills
Body Text: The crustacean gill is a multi-functional organ and it is the site of a number
of physiological processes, including ion transport, which is the basis for hemolymph
osmoregulation; acid-base balance; and ammonia excretion. The gill is also the site by
which many toxic metals are taken up by aquatic crustaceans, and thus it plays an
important role in the toxicology of these species.
Source: Henry RP, Lucu C, Onken H and Weihrauch D (2012) Multiple functions of
the crustacean gill: osmotic/ionic regulation, acid-base balance, ammonia excretion, and
bioaccumulation of toxic metals. Frontiers in Physiology, 3:431.
Mechanism of respiration through book gills
Book gills, like fish gills are bathed in water and supplied with blood vessels. The exchange
of respiratory gases takes place between water and blood flowing through the gill lamellae.
 The deoxygenated blood is carried to the gill lamellae with the help of afferent
branchial vessel.

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 During the flow of blood, gaseous exchange takes place with water which enters the gill
chamber in the form of a current.
 The thin membranes of the gill lamellae act as an excellent surface for diffusion of
respiratory gases.
 Consequently, oxygen from the flowing water diffuses into the blood while carbon
dioxide from the deoxygenated blood diffuses in the water.
 The blood gets oxygenated which is carried by the efferent branchial vessel to the
heart and then to all body parts.
 The carbon dioxide-rich water flows out of the gill chamber.

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Fig. 4: Flow of water through gill chambers of a crustacean
Source: http://sallylightfootcrabresource.weebly.com/gas-exchange.html
Book Lungs – Organs of Respiration in Terrestrial

Arthropods
Book lungs are the characteristic respiratory organs of terrestrial arthropods; arachnids.
These are always in pairs and considered to be evolved from book gills as an adaptation of
terrestrial mode of life. Book lungs are often located in the abdominal segments of the
arachnids; for example, in scorpion, there are four pairs of book lungs located in 3-6th
abdominal segments.

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Fig. 5: Location of book lungs in a spider

Source:
http://commons.wikimedia.org/wiki/File:Spin_vent_diag.jpg#mediaviewer/File:Spin_vent_d
iag.jpg
Structure of a book lung
A book lung is consists of two parts:
a) Dorsal Chamber
 It is also called pulmonary chamber and is formed by the invagination of ventral
abdominal wall.
 It consists of approximately 150 leaf-like lamellae, each of which is lined with the
cuticle.
 The lamellae lie parallel to each other like the leaves of a book.
 The thin space between two lamellae is filled with the air.
b) Ventral Chamber
 The ventral chamber, also called atrial chamber is the non-folded part of the book
lung and is filled with the air.
 On one side, it is continuous with the interlamellar spaces of pulmonary chamber and
on the other side it opens to the outside through a slit-like opening, called stigma or
spiracles.

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Fig. 6: Structure of a book lung
Source: http://upload.wikimedia.org/wikipedia/commons/9/98/Comstock-book-
lungs.png
Fig. 7: Detailed section of a book lung
Source: http://lovelornphyla2011.wikispaces.com/Wecht+Arachnid

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Mechanism of respiration through book lungs
The movement of air in the book lungs is controlled by certain muscles attached to them.
These include; atrial muscles and dorso-ventral muscles.
 The contraction of the muscles results in the compression of book lungs. As a

result, the air is forced out from the interlamellar spaces to the atrial chamber from
where it is released outside through stigma.
 On the contrary, the book lungs regain their shape by the relaxation of the muscles
and the air enters the atrial chamber through stigma reaching interlamellar spaces.
 The lamellae contain deoxygenated blood which is collected from the body of
arachnids and drained through ventral sinus.
 The gaseous exchange takes place between blood and the air present in the
interlamellar spaces through the thin membranous walls of lamellae.
 The blood gets oxygenated while carbon dioxide is released into the air.
 The oxygenated blood is collected into a pulmonary vein which carries blood to the
pericardium from where it circulates into whole body.
Fig. 8: Cross-section of a book lung showing gaseous exchange

Source: http://www.britannica.com/EBchecked/topic/73354/book-lung

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Fig. 9: Exchange of gases in a book lung

Source: http://arthropods2011.wikispaces.com/arachnids
Value Addition: Must know!!
Heading Text: Homology between book lungs and book gills
Body Text: The authors reported and provided evidence regarding the homology;
differences and similarities; in the formation of book lung and book gill lamellae. They
showed that in scorpion embryos, there is ingression (inward migration) of atrial
hypodermal cells rather than invagination or infolding of the atrial hypodermal layer. The
features of cell alignment and apical-basal polarity described in the reports are common
among epithelial cells in animals, but the precursor cells for book gills and book lungs are
distinctive in organizing themselves into a compact mass of many parallel, non-branching
rows, i.e., the page-like pattern of alternating hemolymph and air or water channels.

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Source: Roger D Farley (2011) The ultrastructure of book lung development in the bark
scorpion Centruroides gracilis (Scorpiones: Buthidae). Frontiers in Zoology, 8:18;
doi:10.1186/1742-9994-8-18
Fig. 10: Comparison between book lungs and book gills
The book lungs of (a) arachnids are made up of alternating air pockets and hemocoel tissue
shaped like a stack of books. The book gills of (b) crustaceans are similar to book lungs but
are external so that gas exchange can occur with the surrounding water. (Credit a:
modification of work by Ryan Wilson based on original work by John Henry Comstock; credit
b: modification of work by Angel Schatz)
Source: http://cnx.org/contents/9ba68810-990a-472b-840a-2c6942e270bd@1.33:146
Tracheal System – Respiratory System in Insects
Insects, unlike other arthropods, do not have any respiratory pigment and thus have
inefficient circulatory system to meet their respiratory demands. Consequently, they have
evolved a tracheal system which comprises of a large number of small tubes that carry
oxygen to each and every body cell for oxidation of food.
Structure of tracheal system
The insect tracheal system consists of the following parts:

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(a) Tracheae
The tracheal system constitutes of special tubes called tracheae, formed by the
invagination of cuticle. These tubes form an extensive network which spread throughout the
insect body and supply oxygen. Generally, insects have a pair or more of longitudinal
tracheal trunks which are cross-connected to each other by horizontal tracheae.
Fig. 11: Outline of insect tracheal system
Source: http://cnx.org/contents/185cbf87-c72e-48f5-b51e-f14f21b5eabd@9.45:204
The tracheal tubes are lined internally by a chitinous layer, called intima. The intima forms
ring-like spiral thickenings, called taenidia, which provide them support and elasticity
preventing their collapse.
(b) Tracheoles
The tracheae branch repeatedly into smaller tubes, called tracheoles. These are devoid of
cuticular thickenings and are lined with a proteinaceous layer formed of trachein. The
tracheoles finally enter and terminate into each body cell. The distal ends of these
tracheoles are closed and are filled up with tracheole fluid. The fluid-filled parts of
tracheoles are termed as tracheole end cells.
(c) Spiracles
The prime tracheal trunks open to the exterior by segmentally arranged apertures present
in the lateral region of the body wall; spiracles or stigmata. There are often ten pairs of
spiracles in the insects; two pairs present in the thoracic region and eight pairs in the
abdominal region.

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Fig. 12: Diagrammatic view of insect tracheal system
Source: https://ec-2biol.wikispaces.com/Edens!!!!+page
Fig. 13: Spiracles in an insect tracheal system
Source:
http://en.wikipedia.org/wiki/Respiratory_system_of_insects#mediaviewer/File:Actias_selen
e_5th_instar_spiracles_sjh.jpg
A spiracle is surrounded by an annular sclerite, called peritreme. It also has a valve wich
regulate the opening and closure of the spiracle with the help of certain
sphincter/spiracular muscles. This helps to regulate the flow of air and prevents the loss
of moisture from the body.

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In certain insects, such as cockroaches, the spiracle leads to a chamber, atrium which has
a filtering apparatus in the form of hairs or bristles preventing the entry of dirt in the
tracheal system.
Fig. 14: Scanning electron micrograph of a cricket spiracle valve
Source:
http://en.wikipedia.org/wiki/Respiratory_system_of_insects#mediaviewer/File:Spiraclevalve
s.jpg
Value Addition: Video
Heading Text: Development of tracheal system

Body Text: This interesting video depicts the development of tracheal system in a
fruit fly, Drosophila melanogaster.
Source: http://en.wikipedia.org/wiki/Respiratory_system_of_insects
(d) Air Sacs
In many insects, such as bees, grasshoppers, butterflies, etc., the tracheae lack the spiral
thickenings and dilate at certain places. These are called air sacs which permit increased
oxygen supply leading to more efficient respiration and enhanced release of energy.

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(a)
(b)
Fig. 15: Air sacs in tracheal system of insects
Source: (a) http://lovelornphyla2011.wikispaces.com/Katie-+Insecta; (b)

http://arthropods012.wikispaces.com/INSECTS+1
Gaseous exchange via tracheal system
The gaseous exchange in the tracheal system is primarily by diffusion and ventilation. It
takes place by the following steps:

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 Alternate contractions and expansions of the insect abdomen lead to the changes in
the diameter of tracheae which results in movement of air in/out of the tracheae.
 During inspiration, air enters the tracheae through the spiracles and passes
through the tracheoles to the fluid-filled tips where it gets dissolved in the fluid.
 Now, the oxygen diffuses from the tracheoles into the body cells whereas CO2
diffuses from the cells into the tracheoles.
 During expiration, CO2-rich air is released outside the body through tracheae.
However, CO2 can also diffuse outside through the cuticular covering of the body.
During active metabolism, the tracheole fluid enters the surrounding tissues leading to
the exposure of more surface area of tracheoles to oxygen and more oxygen can enter
the tissues.
Fig. 16: Network of tracheoles terminating in each body cell
Source: http://ahsbiology3.wikispaces.com/Exchange+in+other+organisms

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Fig. 17: Inspiration and expiration during breathing via tracheal

system
Source: https://ec-2biol.wikispaces.com/Courtney+D%27s+Page
Value Addition: Interesting!!
Heading Text: Synchrotron X-ray Imaging for Visualizing Tracheal

Respiration
Body Text: Beetles, crickets and ants exhibit rapid cycles of tracheal compression
and expansion in their head and thorax. However, body movements and
hemolymph circulation cannot account for these cycles. Thus, a synchrotron beam
was used to obtain x-ray videos of living, breathing insects which demonstrated a
previously unknown mechanism of respiration in insects analogous to the inflation
and deflation of vertebrate lungs.
Source: Westneat MW, Betz O, Blob RW, Fezzaa K, Cooper WJ and Lee WK (2003)
Tracheal respiration in insects visualized with synchrotron x-ray imaging. Science, 299:558-
560.

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Fig. 18: Mechanism of breathing in tracheal system
Source:
http://honorsbiologyp6.wikispaces.com/P6+Insects+Gas+Exchange+and+Circulation
Value Addition: Videos
Heading Text: Respiratory Structure and Breathing Mechanism of

Insects
Body Text: The videos comprehensively show and explain the structure of tracheal
system in insects and the mechanism of breathing making all the concepts clear.
Source: http://www.youtube.com/watch?v=JwPsstFBCvI;
http://www.youtube.com/watch?v=quwhcgkVO3c

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Respiration in Aquatic Insects

Aquatic insects have developed diverse structures and mechanisms for respiration. A few of
such structures are discussed below.
A. Plastron Respiration
Certain aquatic insects with tracheal respiration carry an air bubble or a thin gaseous layer
restricted mostly to their ventral surface. This is called plastron which is actually a 8gas
store9 which communicates with the tracheal system.
 Plastron is held in position between the hydrofuge hairs or scales; the structures
present at the angle of contact between the water surface and the body region.
 It has a constant and negligible volume which never decreases in size.
 The surface film of plastron in contact with water acts as a diffusion membrane
through which exchange of respiratory gases takes place. The large surface area of
plastron helps in efficient gaseous exchange.
 It enables the insect to live submerged aquatic life in water nearly indefinitely,
provided there is adequate oxygen in water.
Fig. 19: A giant water bug, Abedus heberti, breathing using plastron
Source: http://thedragonflywoman.com/category/from-the-literature/

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Heading Text: How do insects breathe without lungs?

Body Text: This interesting video is an interesting complination of varios modes
of insect breathing.
<iframe width="560" height="315"

src="//www.youtube.com/embed/wMbi_NcI2JY" frameborder="0"
allowfullscreen></iframe>
Source: https://www.youtube.com/watch?v=wMbi_NcI2JY
Value Addition: First Reports..
Heading Text: Arachnids with plastron respiration
Body Text: Non-tracheated terrestrial arachnids; Phrynus marginemaculatus which

respire by book lungs were reported to be responsive when submerged in water for more
than 24 hours. The data suggested that these arthropods utilized dissolved oxygen from the
surrounding water by plastron respiration. The authors proved the occurrence of plastron
through scanning electron microscopy and light microscope sections. These are the first
reports which showed the presence of plastron respiration in arachnids without tracheal
system.
Source: Hebets EA and Chapman RF (2000) Surviving the flood: plastron respiration in
the non-tracheate arthropod Phrynus marginemaculatus (Amblypygi: Arachnida). Journal of
Insect Physiology; 46: 13-19.
B. Tracheal Gills
The immature stages of many aquatic insects have developed tracheal gills. These are thin
plates or filaments-like structures with abundant tracheal supply but only a very small blood
cavity. The tracheae present in these gills exchange gases with the water and help in
respiration.
The tracheal gills commonly occur on the abdomen, but can also be observed on the thorax.
Depending on the position of these gills, these can be of the following types:
a) Abdominal tracheal gills: The naiads of mayfly (Ephemeroptera) have 7 pairs of

leaf-like tracheal gills on the abdominal segments. The larvae of trichopterans,
caddis flies also possess leaf-like abdominal gills for respiration.
b) Thoracic tracheal gills: The larvae of a few species of stone fly (Plecoptera) have
tracheal gills on their thorax.

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c) Caudal tracheal gills: These are characteristics of the damselfly naiads which have
3 flattened or leaf-like respiratory gills at the end of abdomen. The abdominal end of
mosquito larvae also has four leaf-like tracheal gills. However, they are believed to
help in ionic balance instead of respiration.
d) Rectal tracheal gills: These are internal tracheal folds found in the rectum of
dragonfly naiads. Water enters in the rectum through anus and passes over these
folds resulting in exchange of gases.
e) Coxal tracheal gills: These can be found originating from the coxae certain
stoneflies. However, these probably function in osmoregulation rather than
respiration.
(a) (b)
Fig. 20: Tracheal gills in the sub-imago of May fly, Cloeon dipterum
Source: (a) http://commons.wikimedia.org/wiki/File:Cloeon_dipterum.jpg

(b) http://www.bioimages.org.uk/html/r154959.htm

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Fig. 21: Filamentous tracheal gills on the abdominal segments of

Hydropsyche (Trichoptera: Caddis fly)
Source: http://commons.wikimedia.org/wiki/File:Hydropsyche.jpg
(a)

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(b)
Fig. 22: Enlarged view of abdominal Tracheal Gills in the naiads of

Mayfly, Cloeon dipterum
Source: (a) http://www.bioimages.org.uk/html/r154959.htm
(b) http://commons.wikimedia.org/wiki/Cloeon_dipterum#mediaviewer/Fil
e:Cloeon_dipterum_larva_detail.jpg
(a)

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(b)
Fig. 23: Caudal Tracheal Gills in the naiads of a damselfly
Source:
(a)
https://naturalhistoryofnorthernindiana.wikispaces.com/Damselfly+nymph+~+Sara
+Yob
(b)
http://upload.wikimedia.org/wikipedia/commons/8/8b/Pond_10.18.2012%2C_dams
elfly_nymph_2%2C_side%2C_beltsville%2C_MD_2012-10-18-
14.46_%288100821203%29.jpg?uselang=en-gb
C. Blood Gills
A few aquatic insects contain gills which have a spacious lumen but are either
devoid of tracheae or tracheae are poorly developed in these gills. Their lumen is
filled with the blood and thus, these are called blood gills.
For example, the ventral and anal gills of Chironomus. They can also be observed
as white filaments projecting from the anus on the dorsal side of the abdominal

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segments of Black Fly, Simulium larva. These may or may not be branched,
depending on various species.
D. Spiracular Gills
These gills are ectodermal and filament-like which are covered by a thin layer of cuticle. The
cuticle has thin air spaces which are connected to the tracheae and help in breathing. These
gills can be observed in the pupae of black fly, Simulium.
Fig. 24: Spiracular Gills in the pupa of a black fly, Simulium aureum
Fries
Source: http://www.bioimages.org.uk/html/r161850.htm
E. Breathing Tube:
A few aquatic insects are unable to breathe oxygen dissolved in the water and are
dependent on the atmospheric air for respiration. They have developed certain structures
for taking in atmospheric air. A few of these are as follows:
a) Caudal tube:
Certain adult aquatic insects, such as water scorpion Nepa, have a pair of caudal breathing
tubes at the end of abdomen to form a siphon. The tube is thrusted above the water surface
by which air is drawn to the tracheae.
b) Respiratory siphon:

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Mosquito larvae have a tube-like respiratory siphon at the end of their abdomen which
serves as a breathing tube. The tip of the siphon opens to the outside by a pair of spiracles
through which atmospheric air is drawn inside the tracheal system.
The larvae frequently move towards the surface of water to breathe. At rest, The larvae
hang at an angle from the water surface, head downwards, in order to project the end of
siphon above the water surface. Different species of mosquito larvae have different size of
siphon. The larvae of Culex sp. have the longest siphon followed by that of Aedes while the
siphon in Anopheles larvae is exceedingly short. As a result, the larvae of Culex and Aedes
hang at an angle from the water surface while that of Anopheles lie parallel to the water
surface.
Fig. 25: Larvae of Culex sp. hanging from the water surface by
respiratory siphons
Source: http://commons.wikimedia.org/wiki/File:Culex_sp_larvae.png
c) Respiratory trumpets:
The pupae of mosquito and midges have a pair of breathing tubes, called respiratory
trumpets, which are located dorsally on the thoracic region. They are structurally and
functionally similar to the respiratory siphons present in the larvae and help in breathing.

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Fig. 26: Pupa of Culex pipiens hanging from the water surface by
respiratory trumpets
Source: http://influentialpoints.com/Gallery/Mosquitoes_and_Biting_Midges.htm
Heading Text: Mosquito larvae and pupae
Body Text: The video describes the characteristic upward movement in larvae and pupae
of Culex sp. to reach the water surface at frequent intervals for breathing.
Source: https://www.youtube.com/watch?v=_9zhLZpgrn4
Value Addition: Interesting to know
Heading Text: Amphibious caterpillars in Hawaii
Body Text:
Click on the link and about the amphibious caerpillars found in Hawaii. The article also ahs
supplementary videos showing the activity of these caterpillars.
http://www.pnas.org/content/107/13/5903.full?sid=b80c5bb3-0cef-4f4a-ac8e-
eaafda3ace84

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Source: http://www.pnas.org/content/107/13/5903.full?sid=b80c5bb3-0cef-4f4a-ac8e-
eaafda3ace84
Summary
 Arthropods have developed a variety of structures and organs for breathing, which
include book gills, book lungs, tracheae, etc.
 Book gills are characteristic respiratory organs of aquatic arthropods, especially

crustaceans.
 Book gills are of various kinds depending upon their location and structure.
 Each book gill is made up of a number of thin gill lamellae arranged like the
leaves/pages of a book at right angle to the gill base.
 In a book gill, the exchange of respiratory gases takes place between water and
blood flowing through the gill lamellae.
 Book lungs are the characteristic abdominal respiratory organs of terrestrial

arthropods; arachnids.
 A book lung is consists of a dorsal pulmonary chamber consisting of approximately

150 leaf-like lamellae arranged like the leaves of a book and a ventral atrial chamber
filled with the air.
 In a book lung, the gaseous exchange takes place between blood and the air present
in the interlamellar spaces through the thin membranous walls of lamellae.
 Insects are devoid of any respiratory pigment and thus have evolved a tracheal
system comprising of a large number of small tubes that carry oxygen to each and
every body cell for oxidation of food.
 The tracheae, formed by the invagination of cuticle, posses spirally thickened

taenidia and form an extensive network throughout the insect body.
 The tracheae branch into smaller tracheoles which are lined with proteinaceous
trachein. Their distal ends terminating into body cells are filled up with tracheole
fluid.
 The tracheal trunks open to the exterior by segementally arranged lateral spiracles
which helps to regulate the flow of air and prevents the loss of moisture from the
body.
 The tracheae of a few insects dilate at certain places into air sacs which permit
increased oxygen supply leading to more efficient respiration.

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 Air enters the tracheal system during inspiration and oxygen diffuses from the
tracheoles into the body cells whereas CO 2 diffuses from the cells into the tracheoles.
The CO2-rich air is released outside the body during expiration.
 Certain aquatic insects with tracheal respiration carry an air bubble, plastron on their
ventral surface which communicates with the tracheal system and helps in
respiration.
 The immature stages of many aquatic insects have thin tracheal gills with abundant
tracheal supply which exchange gases with the water and help in respiration. These
may be located at various places of the insect body.
 A few aquatic insects contain blood gills which have a spacious lumen but are either
devoid of tracheae or poorly developed tracheae the lumen of which is filled with the
blood.
 The spiracular gills are ectodermal, filament-like and are covered by a thin layer of
cuticle with has thin air spaces connected to the tracheae.
 A few aquatic insects are unable to breathe oxygen dissolved in the water and draw
in atmospheric air for respiration with the help of certain structures.
 Mosquito larvae have a tube-like respiratory siphon at the end of their abdomen
which serves as a breathing tube.

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Exercise/Practice
Q1. Differentiate between

a) Book lungs and book gills
b) Respiratory siphon and respiratory trumpet
c) Trachea and tracheoles
d) Blood gills and spiracular gills
Q2. Name the following
a) Respiratory organs of Palaemon
b) Respiratory organs of Palamnaeus
c) Breathing tube in mosquito larvae
d) Mosquito species with the longest breathing tube
e) An insect with air sacs in the tracheal system
f) Ventral gills of Chironomous
g) Gills of the dragon fly naiads
h) Gas store on the ventral side of an aquatic insect
Q3. Answer the following questions in short.
(a) List the different types of book gills based on their structure and function.
(b) Why do larvae of Culex sp. hang at an angle from the surface of water?
(c) Write the function of air sacs present in the respiratory system of a bee.
(d) How is opening and closing of a spiracle is regulated in the body of an insect?
(e) Why are book gills and book lungs called so?
(f) Name two chambers of a book lung and differentiate between them.
(g) How does plastron enable the insect to live submerged aquatic life in water nearly
indefinitely?
Q4. Answer the following questions in detail.
(a) Describe the structure of book gills and explain the mechanism of respiration through
them.
(b) Discuss the structure and function of a book lung. Write the similarities and
differences in their structure as compared to the book gills.
(c) Explain any three ways by which aquatic insects breathe.
(d) How does breathing between terrestrial and aquatic insects differ?
(e) Describe the tracheal respiration in detail. Support your answer with well-labelled
diagrams.

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Glossary
Book gills: Organs of aquatic respiration in crustacea made up of a number of thin gill
lamellae arranged like the leaves/pages of a book
Phyllobranch: Book gills with leaf-like gill lamellae
Trichobranch: Book gills with hair-like gill lamellae
Dendrobranch: Book gills with arborescent gill lamellae
Book lungs: Organs of terrestrial respiration in arachnida consisting of gill lamellae

arranged like the leaves/pages of a book in the pulmonary chamber
Taenidia: Ring-like spiral thickenings in the tracheae of insects
Trachein: Protein present in the inner wall of insect tracheoles
Spiracles: Segmentally arranged openings of the tracheal system present in the lateral
region of the insect body wall
Peritreme: Annular sclerite surrounding the insect spiracles
Air sacs: Dilated trachea without any spiral thickenings
Plastron: An air bubble or a thin gaseous layer restricted mostly to the ventral surface of
certain aquatic insects
Tracheal gills: Thin plates or filaments-like structures, in some aquatic insects, with
abundant tracheal supply but only a very small blood cavity
Blood gills: Gills with spacious lumen filled with the blood but either devoid of
tracheae or with poorly developed tracheae

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Spiracular gills: Ectodermal filament-like gills covered by a thin layer of cuticle with thin
air spaces connected to the tracheae
Respiratory siphon: Breathing tube at the end of abdomen of mosquito larvae
Respiratory trumpet: Breathing tube at the thoracic region of mosquito pupae
References/ Bibliography/ Further Reading
Suggested Readings
 Barnes, R.D. (1982). Invertebrate Zoology, V Edition. Holt Saunders International

Edition.
 Barnes, R.S.K., Calow, P., Olive, P.J.W., Golding, D.W. and Spicer, J.I. (2002). The
Invertebrates: A New Synthesis, III Edition, Blackwell Science
 Barrington, E.J.W. (1979). Invertebrate Structure and Functions. II Edition, E.L.B.S.
and Nelson
Useful Web Links
 https://stormhawks2013.wikispaces.com/JK
 http://sallylightfootcrabresource.weebly.com/gas-exchange.html
 http://lovelornphyla2011.wikispaces.com/Wecht+Arachnid
 http://www.britannica.com/EBchecked/topic/73354/book-lung
 http://arthropods2011.wikispaces.com/arachnids
 http://cnx.org/contents/185cbf87-c72e-48f5-b51e-f14f21b5eabd@9.45:204
 http://en.wikipedia.org/wiki/Respiratory_system_of_insects
 http://ahsbiology3.wikispaces.com/Exchange+in+other+organisms
 http://www.iaszoology.com/arthropod-respiration/
 http://rspp.weebly.com/arthropoda.html
 http://en.wikipedia.org/wiki/Arthropod

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Respiration in Arthropoda

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Bsc zoology (University of Calicut)

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Lesson: Respiration in Arthropoda

Lesson Developer: Sarita Kumar

College/Dept: Zoology, Acharya Narendra Dev College

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Heading Text: Evolution of respiratory organs in arthropods

adaptations in locomotion, breathing, reproduction, and mechanisms for food capture

Book Gills – Organs of Respiration in Aquatic Arthropods

Book gills are characteristic respiratory organs of aquatic arthropods, especially

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Fig. 1: Different Kinds of gills in a crayfish based on their location

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Structure of a book gill

a) Phyllobranch: The gill lamellae are leaf-like, for example in Palaemon.

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Fig. 3: Location of book gills in a crayfish

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Heading Text: Multiple functions of crustacean book gills

Mechanism of respiration through book gills

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 The carbon dioxide-rich water flows out of the gill chamber.

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Fig. 4: Flow of water through gill chambers of a crustacean

Book Lungs – Organs of Respiration in Terrestrial

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Fig. 5: Location of book lungs in a spider

Structure of a book lung

A book lung is consists of two parts:

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Fig. 6: Structure of a book lung

Fig. 7: Detailed section of a book lung

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Mechanism of respiration through book lungs

 The contraction of the muscles results in the compression of book lungs. As a

Fig. 8: Cross-section of a book lung showing gaseous exchange

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Fig. 9: Exchange of gases in a book lung

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Heading Text: Homology between book lungs and book gills

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Fig. 10: Comparison between book lungs and book gills

Tracheal System – Respiratory System in Insects

Structure of tracheal system