A.

The Origin and Parts of the Coelom and the  (where lungs are situated); transverse septum 
Mesenteries  assumes an ​oblique​ position 
  ● In some reptiles (esp Crocodilia), birds, 
1. Origin  mammals: pleuroperitoneal cavity divided into 
● Coelom​ or ​body cavity​ ​– cavity of the  anterior and posterior parts by various fusions 
hypomere; never segmented  between the transverse septum and other 
● Parietal peritoneum​ ​– lining of the body wall  coelomic folds 
formed from outer wall of hypomere in contact  ○ In mammals: ​pleuroperitoneal​ or 
with inner surface of the layer of voluntary  nephric fold​ ​– descends from dorsal 
muscles  body wall and fuses with TS 
● Visceral peritoneum​ or ​serosa​ – inner walls of  ○ Partition so formed in birds: ​oblique 
the hypomere that become the covering layer  septum​ ​(slightly muscular); in mammals: 
of the intestine and other viscera  diaphragm​ (extensively muscularized 
● Mesentery​ – double-walled membrane formed  from the cervical myotomes) 
by the two walls of the hypomere that are in  ● Two​ ​pleural cavities​ o
​ r​ p
​ leural sacs​ – each 
contact above and below the intestine  inclose a lung (anterior to OS or diaphragm); 
● Dorsal mesentery​ – between the dorsal wall of  separated from each other 
the coelom and the intestine; intact for its entire  ● Peritoneal​ ​or ​abdominal cavity​ – part of 
length  pleuroperitoneal cavity posterior to OS or 
● Ventral mesentery​ – between the ventral wall  diaphragm; incloses greater part of the 
and the intestine; disappears very early except  digestive tract and urogenital system 
in certain regions  ● In birds and mammals: coelom divided into 4 
● In amphioxus: mesoderm pouches off from the  compartments (pericardial, 2 pleural, peritoneal) 
archenteron as a paired axial series of sacs  – increases efficiency of lung respiration 
whose cavities later combine to form the  ● Remember! Viscera are covered by the visceral 
definitive coelom (​enterocoelous​)  peritoneum and are NOT really inside the 
● Mesoderm of vertebrates no longer arises in  coelom 
this fashion; ancestral for vertebrates; links   
vertebrates to enterocoelomates  B. The Digestive Tract and Its Derivatives 
(chaetognatha, echinodermata, hemichordata)   
  1. The Origin of the Digestive Tract 
2. Divisions of the Coelom  ● Primitive intestine or archenteron is produced 
● At first, coelom is divided into 2 longitudinal  by invagination or other processes in embryo’s 
halves by the dorsal and ventral mesenteries  gastrula stage 
● Partial disappearance of ventral mesentery →  ● Archenteron at first is a simple tube of 
two halves of coelom are connected ventral to  endoderm with one opening, the blastopore, 
the intestine  situated at the future posterior end of the 
● Transverse septum​ – partition that divides  embryo 
coelom into (at least) two compartments in the  ● This endodermal tube persists as the lining of 
adults of all vertebrates  the adult digestive tract and all its derivatives 
● Pericardial cavity​ ​– small anterior  ● Adult digestive tract consists of a thick-walled 
compartment of the coelom; contains only the  tube composed of both endodermal and 
heart  mesodermal elements (connective tissue and 
● Pleuroperitoneal cavity​ – very large posterior  muscular layers from splanchnic mesoderm of 
compartment; contains all other viscera  hypomere); latter predominates 
● In fishes and urodeles: pericardial cavity  ● Anterior and posterior ends of digestive tube 
anterior to pleuroperitoneal cavity; transverse  are lined by ectoderm (result of invagination 
septum passes ​transversely​ across the body  processes) 
● In Anura and all vertebrates above: pericardial  ● Stomodaeum​ – anterior invagination 
cavity descended posteriorly thus lying ventral  ● Proctodaeum​ – posterior invagination 
to anterior part of the pleuroperitoneal cavity  ● In adults, no trace remains of the boundary of 
said invaginations 
  ● Backward-shoving of choanae: for separation of 
  food and respiratory passages 
2. Homology of the Mouth  ● Primary tongue​ – in fishes; part of buccal floor 
● In Amphioxus: mouth originates on the left side  demarcated anteriorly by a fold; sheaths 
in series with the gill slits; only later moves to a  anterior end of hypobranchial apparatus and 
median position  contains no muscles or glands 
● Said mode of origin lent support to idea that  ● Definitive tongue​ – in amphibians; crescentic 
vertebrate mouth arose by fusion of first pair of  gland field​ in front of primary tongue fused with 
gill slits  it 
● However, more probable that these are not of  ● In amniotes: same process, ​tuberculum impar 
phylogenetic significance  (gland field) joined by a pair of ​lateral lingual 
● Annelid-arthropod ancestry for vertebrates  swellings​ → amniote tongue now compounded 
○ Vertebrate is an annelid turned on its  of 4 swellings 
back  ○ invaded by voluntary musculature from 
○ Original annelid mouth and  hypobranchial musculature 
stomodaeum are represented by the  ○ innervated by 12th (hypoglossal) 
pituitary and its opening  cranial nerve 
○ Present vertebrate mouth is a new  ○ with glands and taste organs 
formation  ○ develops intrinsic muscle fibers 
● Now discredited in favor of an origin from the  ● Very long, mobile tongues in snakes, some 
echinoderm-hemichordate line – vertebrate  lizards, woodpeckers, and ant-eating mammals 
mouth is same as that of its immediate  (in WP, tongue armed with horny covering or 
invertebrate ancestors and NOT a new  with spines and thorns) 
formation  ● Teeth – in lower vertebrates: on various bones 
  of palate and jaws; in higher vertebrates: limited 
3. Parts and Outgrowths of the Digestive Tube  to a single row along jaw margin 
   
Oral or Buccal Cavity  Oral glands* 
● Bounded by jaws in front and on sides, palate  ● In tetrapods: numerous multicellular glands 
above, buccal floor bearing tongue below  imbedded in walls of oral cavity and opening 
● Lips and cheeks in higher forms  into it 
● Vestibule​ ​– space between lips and teeth  ● 2 kinds: ​mucous-secreting glands​ (slimy) and 
● In Amphioxus and Ammocoetes: oral cavity  serous glands​ (watery, secrete enzymes, 
consists of space inclosed by oral hood;  poisons, etc) 
definitely bounded from the pharynx by the  ● Labial glands (lips) and dental glands (teeth) – 
velum  modified skin glands 
● Position of velum corresponds to the boundary  ● Glands in more internal parts of oral cavity – 
between stomodaeum and pharynx of  new formations 
gnathostomes (no landmarks in adults)  ● Few oral glands in amphibians; increase in 
● In Dipnoi, Crossopterygii, tetrapods: nasal  reptiles – ​lingual​ ​(tongue), ​sublingual​, 
cavities open into roof of oral cavity by ​internal  palatine​, ​labial​ ​(lip) groups of glands occur 
nares ​or ​choanae​ ​(primitively anteriorly placed)  ● Poison glands of snakes are enlarged, modified 
● In Crocodilia, birds, mammals: internal nares  upper labial glands 
assume a far posterior position through  ● Association of glands with teeth is nothing new, 
formation of secondary palate  occuring in amphibians 
● Secondary palate imperfectly formed in birds;  ● Most birds equipped with oral glands; 
best developed in mammals (continued  fish-eating birds may lack them 
backward by fleshy fold – ​soft palate​)  ● In mammals:​ ​salivary glands​ ​– enlarged oral 
● Nasopharyngeal passages​ – prolongs nasal  glands opening into oral cavity by one or more 
cavities posteriorly to open by the choanae  long ducts 
now situated near the beginning of the pharynx  ● Principal salivary glands 
○ Parotid​ – enlarged cheek gland 
 ○ Sublingual​ – occurs either as one large  ● Above fishes: number of gill slits in embryo 
gland or as group of smaller glands  gradually decreases; often the openings fail to 
each with own duct and opening  break through 
○ Submaxillary  ● No gill slits in adult vertebrates above urodeles 
○ Infraorbital  (close during development) 
  ● In amniotes: gills fail to appear even in embryo 
Rathke’s Pouch*  ● Pharynx originally functioned as a 
● Epithelial evagination from roof of buccal cavity  food-catching device (tunicates and 
occurs in embryo → blind pouch  Amphioxus) 
● Extends toward, comes in contact with, and  ● Pharynx lost said function when vertebrates 
fuses with brain wall  increased in size and ate larger objects → 
● Hypophysis​ ​or ​pituitary body​ – compound  respiratory with aid of gills 
structure formed; important gland of internal  ● Gills​ ​or ​branchiae​ – thin-walled projections, 
secretion  richly supplied with blood vessels and 
● Adenohypophysis​ – part derived from oral  presenting large surface for exchange of 
epithelium (anterior pituitary)  respiratory gases; either ​lamellar​ (plate-like) or 
● VS ​Neurohypophysis​ – part derived from brain  filamentous 
(posterior pituitary)  ● 2 kinds in vertebrates (absent in amniotes) 
  ○ External gills​ ​– borne on outer surface 
Pharynx  of branchial bars; found only in larvae 
● Immediately caudad to oral cavity;  of Crossopterygii and Dipnoi and in 
characterized by gill slits through its wall in  larvae and some adults of amphibians; 
adult or embryonic chordates  more ancient 
● In Amphioxus and Ammocoetes: very exactly  ○ Internal gills​ ​– borne on side walls of 
delimited anteriorly by the velum  branchial bars (facing gill pouches) 
● In gnathostomes: anterior boundary is indefinite  ■ Demibranch​ o ​ r ​half-gill​ – gill 
but may be set just anterior to 1st gill slit or its  on one side of branchial bar 
representative (opening of auditory tube)  ■ Whole gill​ ​or ​holobranch​ – 
● Gill slits – passages from pharynx to exterior  two gills on two sides of the 
○ Internal gill slits​ – pharyngeal  bar 
openings  ○ Epithelium of external gills is 
○ External gill slits​ – exterior openings  ectodermal 
● Gill pouch​ ​ or ​visceral pouch​ – passage from  ○ Open question as to whether lining of 
one slit to the other in the pharyngeal wall  pharynx and gill pouches and 
● Visceral arch​ ​or ​branchial bar​ – tissue  epithelium of internal gills are 
between successive gill slits; incloses  entodermal or not 
○ skeletal gill arch  ○ If epithelium of IG is ectodermal, then 
○ a blood vessel termed an aortic arch  EG and IG are variants of same organ 
○ a cranial nerve   
● Visceral pouches​ – paired evaginations put out  Thyroid Glands* 
by pharyngeal wall (typically 6)  ● Epithelial evagination from buccal or 
● Visceral furrow​ ​– invagination of ectoderm  pharyngeal floor at the level of the hyoid arch, 
opposite each visceral pouch  between primary tongue and tuberculum impar 
● Pouches meet furrows and the fusion area  ● Separates from the floor, becomes ductless, 
breaks through as a​ ​gill slit​ or ​gill cleft​ (which  proliferates into a considerable mass of 
persists as the external gill slit)  follicles, usually becoming bilateral 
● Large number of gill slits in ancestral chordates  ● Vital endocrine gland; secretion necessary for 
● In present vertebrates, greatest number is in  normal growth and sexual development 
hagfishes (up to 14); in gnathostomes, 6 (but 1st   
one, between mandibular and hyoid arches, is  Pharyngeal Glands* 
generally modified or reduced, forming the 
spiracle​) 
 ● Branchial buds​ ​– epithelial masses that  ● In Polypterus and Dipnoi, these sacs (lungs) are 
proliferate from dorsal and ventral ends of gill  employed in air respiration (to let fishes live out 
pouches (most complete set found in lampreys)  of water) 
● True​ ​or ​palatine tonsils​ – lymphoid masses  ● In teleosts: single sac displaced dorsally 
proliferated from the second pouches  through a rotation process; in most, connecting 
● Thymus​ ​– lymph gland of uncertain function;  duct or ​pneumatic duct​ remains although also 
formed when 2 to 4 of the dorsal buds unite  shifts to dorsal side of pharynx; in others, duct 
● Parathyroids​ ​– small glands concerned in  degenerates 
calcium metabolism; from ventral buds  ● In teleosts, swim bladder serves to alter specific 
● Epithelial bodies​ – glandular masses of  gravity of fish as it changes level in water, to 
unknown function  keep it in equilibrium with external pressures 
● In mammals: thymus comes from ventral buds,  (accomplished by alterations in gas content) 
parathyroid from dorsal buds  ● In tetrapods: lungs arose embryologically in the 
● Postbranchial​ ​or ​ultimobranchial body​ –  same way as those of Polypterus (homologous) 
outgrowth which separates from the pharynx  ● Pair of sacs formed grows backward into 
and produces a number of follicles; arises on  pleuroperitoneal cavity; mesodermal tissue 
one or both sides close behind last gill slits; of  added to original endodermal lining 
unknown significance  ● Walls of lungs subdivided internally into air 
  spaces which become smaller and more 
Tympanic Cavity and External Auditory Meatus*  numerous in the tetrapod series until finally the 
● Beginning with amphibians, 1st gill pouch puts  lungs are: 
out toward the internal ear an outgrowth whose  ● Alveoli​ – spongy mass of minute air pockets in 
end expands into the ​tympanic cavity​ (cavity of  mammals 
the middle ear)  ● Common in tetrapods na left is smaller 
● Auditory​ ​or ​Eustachian tube​ – stalk of the  ● Mammalian lungs commonly divided into few 
outgrowth, connects middle ear to the pharynx  large lobes 
● Tympanic membrane​ ​or ​eardrum​ –  ● Snakes: one lung (right) 
double-walled membrane formed when wall of  ● Some salamanders: altogether lost 
tympanic cavity meets bottom of invagination  ● Crocodilia: lungs approach most nearly among 
corresponding to the position of the 1st external  reptiles to those of mammals 
gill slit (in some reptiles, in birds and mammals)  ● Theory: lungs homologous to pair of gill slits but 
● External auditory meatus​ – passage formed  lacks supporting evidence 
by the invagination; marks the position on the  ● Trachea​ o ​ r ​windpipe​ – duct connecting the 
side of the head of the first gill slit  lungs with the pharynx (very short in 
  amphibians, considerable length in amniotes 
Swim Bladder, Lungs, Larynx, and Trachea*  due to backward retreat of viscera); wall 
● Gills slits and gills disappeared in adult  stiffened by complete or incomplete 
vertebrates when they adopted land habitats  cartilaginous rings 
● Swim bladder​ – pharyngeal outgrowth that  ● Glottis​ – opening into the pharynx 
took over physiological role of gill slits and gills  ● Trachea → 2 ​bronchi​ (in mammals, subdivide 
(already existed in crossopterygian, dipnoan,  repeatedly) → ​bronchial tree​ → ​bronchioles 
and bony fishes)  (final twigs) → little chambers encircled by 
● In Polypterus: this arises as a ventral  alveoli 
evagination from pharynx behind gill region;  ● In birds: number of large, thin-walled ​air sacs 
evagination divides into 2 sacs which grow  scattered throughout the body, connected to 
backward and retain ventral position and  lungs by bronchial tubes of large caliber 
ventral connection with pharynx (left smaller  ○ Inside the lung there are anastomoses 
than right)  between the larger bronchial tubes and 
● In Dipnoi: similar origin, single or bilobed sac is  the terminal bronchioles also anastome 
displaced to dorsal side but retains ventral  with each other, forming a complicated 
opening into pharynx  network 
 ○ No closed alveoli at ends of  sac-shaped appendage; serves for storage or 
bronchioles; occur along their sides  preliminary digestion 
○ Air passes through lungs in and out of  ○ In pigeon tribe: food in crop is mixed 
air sacs which act like bellows (so lungs  with milk, a secretion produced by fatty 
completely ventilated at each breath)  degeneration of crop epithelium; 
○ Lessen specific gravity of the bird +  mixture is fed by both parents to the 
other flight advantages  young by regurgitation 
● Larynx​ – chamber into which the beginning of  ● In cud-chewing mammals: lower end of 
the trachea at the glottis is differentiated; walls  esophagus contributes crop-like sacs to the 
supported by cartilages representing reduced  complicated “stomach” 
gill arches   
● In some urodeles: there is a pair of lateral  Stomach or Ventriculus 
cartilages derived from the last gill arches  ● Follows the esophagus as a spindle-shaped to 
● In other amphibians: these subdivide into a pair  sacciform enlargement provided with enzyme- 
of dorsally placed cartilages – ​arytenoids​, and  and acid-secreting glands and a thick muscular 
a cartilaginous ring – ​cricoid  wall of smooth muscles 
● Aryteno-cricoid primordium​ – furnishes the  ● In lower vertebrates: little or no external 
cartilaginous rings of the trachea when it  demarcation between E and S; proximal limit of 
elongates  stomach determined only by presence of 
● This condition persists through reptiles and  gastric glands 
birds; in mammals:  ● Cardiac region​ – region of the stomach next to 
● Thyroid cartilage​ – large, ventrally placed  the esophagus 
cartilage added anterior to the cricoid; prolly  ● Pyloric region​ – end leading to intestine 
comes from 4th and 5th gill arches + their  ● Pylorus​ ​– strong sphincter muscle by which 
copula  stomach terminates; embraces entrance into 
● In birds and mammals: middorsal part of cricoid  intestine 
frequently separates as a ​procricoid  ● In Amphioxus and cyclostomes: definite 
● Larynx (situated behind basihyal) is very closely  stomach lacking 
associated with hyoid apparatus  ● In lower fishes: stomach is a straight or J- or 
● Muscles of the larynx are branchial muscles  V-shaped tube; teleosts frequently lack a 
since its cartilages are gill arches  stomach 
● Function of larynx: production of the voice  ● In amphibians and reptiles: tubular, not much 
● Syrinx​ ​– voice-producing mechanism at the  enlarged stomach 
forking of the bronchi in birds  ● In birds: stomach divided into ​proventriculus 
● Epiglottis​ ​– cartilage-supported projection  (bears gastric glands) and ​gizzard​ (with thick 
protecting the glottis (first in reptiles, found  muscular wall and hard cornified lining) 
throughout mammals); improbable that its  ○ Best developed in grain- and 
cartilage is a modified gill arch  seed-eating birds; contains gravel to 
  assist in grinding up hard food 
Esophagus   
● Posterior to glottis, the digestive tract narrows  Small Intestine 
to the esophagus  ● Follows the pylorus and usually consists of a 
● Generally short in fishes and amphibians,  more or less coiled tube of considerable length 
elongated in amniotes (due to posterior  ● Duodenum​ – beginning beyond the pylorus; 
descent of viscera)  received the ducts of liver and pancreas 
● Lined by stratified epithelium similar to that of  ● In cyclostomes: presence of low fold in 
buccal cavity; provided by striated musculature  intestinal wall, apparently the forerunner of the 
(all or part of its length)  spiral valve​ – wide, spirally coiled fold filling 
● Crop​ ​or ​ingluvies​ – any esophageal  the greater part of the small intestine in 
enlargement; occurs in many birds (esp  elasmobranchs, dipnoans, and many ganoids 
grain-eating) where it may form a large 
 ● Valvular intestine​ – part of intestine having the  ○ Production of urea and other 
spiral valve (device for increasing absorptive  nitrogenous wastes 
surface of intestine)  ○ Control of food and other substances in 
● Teleosts lack a spiral valve; instead have one to  the blood 
many diverticula, the ​pyloric caeca​, projecting   
from the proximal part of the small intestine  Pancreas* 
(has same histological structure as intestine and  ● Gland that proliferates from the intestinal 
presumably a device for increasing the  epithelium, shortly beyond the liver 
absorptive surface)  proliferations, usually as 1 dorsal and 2 ventral 
● Longer in herbivores than in carnivores  outgrowths that unite 
  ● Grows out into adjacent mesenteries and may 
Liver*  form a compact or more or less diffuse organ 
● Has no counterpart in invertebrates  ● All 3 stalks of the outgrowths may persist as 
● In Amphioxus: seen in simple form as a hollow  pancreatic ducts​ ​(but usually reduced to 1 or 2) 
diverticulum from the intestine  ● These ducts open into the duodenum 
● In vertebrates: large, lobed organ suspended  separately from or after joining the common 
from the transverse septum or its  bile duct 
representative; arises as a single or double  ● Amphioxus: no pancreas 
hollow or solid proliferation from the intestinal  ● Vertebrates: one 
epithelium at the level of the TS, into which it  ● Teleosts: so diffuse that it cannot be 
grows  recognized macroscopically 
● The proliferations soon branch extensively and  ● Consists of 2 kinds of cells: 
the branches usually anastomose → adult liver is  ○ Regular gland cells (​acinar cells​) – 
a complicated network of epithelial columns of  secrete into the duodenum the 
endodermal origin with blood spaces filling the  enzyme-containing pancreatic juice 
meshes of the net  (most important digestive fluid of 
● Stalk of the liver outgrowth becomes the ​bile  vertebrates) 
duct  ○ Islets of Langerhans​ – small scattered 
● Gallbladder​ – a sac that usually develops from  nests of cells which produce insulin 
the liver substance along the course of the bile  (necessary for metabolism of 
duct  carbohydrates) 
● Ammocoetes larva has both gallbladder and   
bile duct, but these degenerate during  Yolk Sac* 
metamorphosis  ● Baglike extension of the ventral wall of the 
● Fishes, amphibians, reptiles generally have a  small intestine found in the embryos of 
gallbladder; lacking in birds (including pigeons)  vertebrates which develop by meroblastic 
and in a number of mammals  cleavage 
● Many vital functions, most obvious is secretion  ● Filled with yolk, gradually utilized by embryo as 
of bile (assists in digestion of fat)  food (kaya yolk sac is provided with numerous 
● Hepatic ducts​ ​– larger ducts formed by union  blood vessels early) 
of minute ducts into which bile is poured upon  ● Diminished yolk sac is withdrawn into the body 
secretion by liver cells  and becomes part of the intestinal wall 
● Cystic duct​ ​– duct from gallbladder, joined by  ● Body wall closes over the aperture through 
hepatic ducts  which the yolk sac protruded 
● Common bile duct ​or ​choledochal duct​ –  ● Mammals possess a vestigial yolk sac during 
joined by hepatic ducts; empties into small  embryonic stages as a reminiscence of reptilian 
intestine not far beyond the pylorus, either with  ancestry (despite not having meroblastic 
or without joining the pancreatic duct  cleavage) 
● Other functions of the liver   
○ Storage of excess sugar in the  Caecum 
digested food as glycogen  ● Colic caecum​ – diverticulum at the junction of 
small and large intestine 
 ● First seen in a few amphibians; usually present 
throughout amniotes 
● In reptiles: small size 
● In birds: 2 colic caeca 
● In mammals: conspicuous caecum (very long 
and large in herbivores); few mammals have 2 
● Vermiform appendix​ – slender appendage into 
which the end of the caecum degenerates (in 
primates and some other mammals) 
● Uncertain function except in animals where it is 
large (assists in digestion and absorption) 
 
Large Intestine or Colon 
● Increased diameter compared to SI; presence 
of one or two caeca 
● In fishes: short and not distinctly marked off 
from SI 
● Digitiform​ ​(or ​rectal​) ​gland​ – slender projection 
that secretes mucus in elasmobranchs 
● In amphibians and birds: large intestine is called 
cloaca​ when it receives urinary and genital 
ducts (as in most vertebrates) 
● No cloaca in Holocephali, ganoids, teleosts, 
and mammals except monotremes 
● Rectum​ – end section of large intestine in the 
absence of a cloaca 
 
Urinary Bladder* 
● In most tetrapods: serves as a reservoir for the 
urine; present as a ventral outgrowth of the 
cloaca or urogenital canal (if cloaca is absent) 
● Allantois​ ​– enlarged urinary bladder in amniote 
embryos; projects far beyond the embryo and 
serves as a respiratory organ (adult bladder is a 
portion of the allantois) 
● No urinary bladder in adult state of birds but 
large allantois in their embryos 
 
Anus 
● Digestive tract opens to the exterior by the 
anus 
● In vertebrates: not at the posterior end of the 
body; shifted to the ventral surface at the base 
of the tail region (happens through overgrowth 
of the tail) 
 
Postanal Gut 
● In most vertebrate embryos: solid or hollow 
extension of gut into the tail, later vanishes 
● Occurrence indicates an originally more 
posterior position of the anus