Chapter 12 Digestive System

PROCURING FOOD

Filter feeders: ancestral chordates (sea squirts, amphioxus),

living larval agnathans (e.g. lampreys)
o This mode of feeding is primitive
o Can be employed only by aquatic organisms
o Passive filtration of organic matter from the incoming
respiratory stream, propel particles to the rear of the
pharynx for swallowing
More active version of filter feeding in some fishes
(spoonbills, cluepids, basking sharks)
o Plankton & small fish strained out of respiratory
stream using long filamentous gill rakers hanging
into pharyngeal chamber from gill arches
Whalebone/Baleen hangs into oral cacity
Note that whales take water into their oral
cavity for the sole purpose of feeding; they
DO NOT breathe with gills (water spills out
of the mouth to be sent back to sea)
Active predation/active acquisition of particulate food in
adults of early fishes -> led to paired external sense organs
& their concentration on the head
More aggressive methods of obtaining food made possible
by jaws & elaborate musculature of body walls (for
locomotion & pursuit)
o Jaws were first invested with denticulated bony
dermal armor
o In specialized predators, eventually furnished with
sharp denticles (teeth)
Bite-tear-swallow technique, as in modern
sharks

This required no tongue or other

specializations of oral cavity
Less energy consuming procedure due to further
adaptations of skull & hyoid arches
o Protrusible jaws, suction (as in goldfish)
o Lampreys, due to parasitic nature, rasp host tissues
with spiny tongue (fleshy & cartilaginous rod armed
w/ horny teeth) & suck the debris into pharynx
Carnivorous mammals: snap-bite-tear technique
o Often involves piercing effect of saberlike tooth
Food taking depends on food finding
o Use chemical, mechanical, thermal, capsulated
touch, visual, electroreceptors

THE DIGESTIVE TRACT: AN OVERVIEW

Digestive tract aka alimentary canal

From mouth to vented cloaca or anus (directly exterior)
Functions: digestion & absorption, elimination of undigested
wastes
Peristalsis: food propelled from pharynx to vent or anus due
to action of smooth muscle tissue in walls of the tract
Major subdivisions:
o Oral cavity & pharynx (in fishes: oropharyngeal
cavity)
o Esophagus
o Stomach
o Intestine (divided into small & large in tetrapods)
Ducts from accessory organs also empty into the tract
o Pancreas, liver, gall bladder
o These organs arise as evaginations from the
embryonic digestive tract
Ceca are also commonly present

Digestive system = tract + accessory organs

o Occupies the pleuroperitoneal cavity in fishes,
amphibians, most reptiles
o Lungs occupy separate pleural cavities; digestive
organs
beyond
esophagus
occupy
abdominal/peritoneal cavity in mammals, birds, &
few other reptiles
Coelom arises via delamination of early
lateral-plate mesoderm into somatic &
splanchnic mesoderm
Most of embryonic digestive tract EXCEPT
the endodermal lining comes from
splanchnic mesoderm (in cephalochordates
& craniates)
Visceral peritoneum outer covering of digestive tract,
continuous with parietal peritoneum
o Early in embryonic life, parietal & visceral petionia
continuous via dorsal & ventral mesenteries; coelom
is divided into separate right & left cavities
o Dorsal mesentery remains intact (conducts blood
vessels & nerves from roof of coelom to digestive
organs)
o Ventral mesentery disappears EXCEPT at the level
of the liver & urinary bladder
Embryonic digestive tract has 3 regions:
o Midgut contains the yolk (when present), where
the yolk sac is attached; little remains in adults
o Hindgut becomes remainder of intestine and
cloaca
o Foregut elongates to form part of oral cavity,
pharynx, esophagus, stomach, much of small
intestine
Stomodeum anterior portion of oral cavity, or in the case of
fishes, oropharynx; arises as midventral invagination of
ectoderm of the head

Oral plate temporarily separates early embryonic foregut

from exterior; ruptures to provide an anterior entrance to
digestive tract
Proctodeum counterpart of stomodeum in the hindut;
provides the exit when the cloacal plate ruptures
o Deuterostomous
Differences in anatomy of digestive tracts caudal to pharynx
correlated with nature & abundance of food

MOUTH & ORAL CAVITY

Mouth entrance to digestive tract

o
opens into oropharyngeal cavity w/ teeth & walls
perforated by gill slits (in gnathostome fishes)
Oropharyngeal cavity terminates at short
esophagus
o Opens into oral/buccal cavity w/ teeth & tongue (in
tetrapods)
Leads to the pharynx
Primary palate roof of oropharyngeal cavity of fishes &
amphibians
o Pierced anteriorly in lungfishes & amphibians by
internal nares
Most Reptiles have an incomplete secondary palate, leaving
in the roof of the oral cavity a palatal fissure that channelizes
respiratory air between the choanae & pharynx
Crocodilians & mammals: cleftless roof for oral cavity from
mouth to pharynx
Anurans: paired vocal sacs (reverberating chambers
beneath floor of pharynx); tetrapods: oral glands or their
ducts
o These open into the oral cavity :D
Oral vestibule trench; in mammals; separates the gums
(alveolar ridges) from the cheeks & lips
o Rodents: opening from vestibule leads into cheek
pouch, used for transport

Extend from 1st cheek teeth to a position

lateral to scapula
Lined with moderately low keratinized
squamous epithelium, for protection vs.
abrasion
Buccinators retractor; a slip of this muscle
inserts on the wall
Overlying skin is loose so that pouches can
expand
& grain eating birds: median sublingual seed

Seed
pouch
Lies upon caudal portion of mylohyoid
muscle beneath oral cavity
Retracted by homologue of genioglossus
muscle of mammals

Tongue

Elasmobranchs, bony fishes, perennibranchiate amphibians:

crescentic or angular elevation of floor of oropharyngeal
cavity shaped by basihyal & ceratohyal cartilages
o Primary tongue lean hyoid elevation; no
musculature, cannot be independently manipulated
May assist jaws in holding prey within
oropharyngeal cavity
Forerunner of tetrapod tongue
Terrestrial urodeles, anurans: primary tongue + extension
that can be flipped out of mouth
o Primary tongue develops from hyoid arch
mesenchyme
o Extension develops from embryonic glandular field
anterior to hyoid arch
Glandular field secretes sticky mucus that
entangles insects when tongue is suddenly
thrust out of mouth

Insectivorous amphibians: tip of tongue usually terminates in

an expansion that increases the probability that the prey will
be caught & delivered into oral cavity
Terrestrial urodeles: root of tongue anchored to basihyal &
ceratohyal cartilages of pharyngeal floor :: anurans: unique!
Floor of oral cavity immediately behind mandibular
symphysis
o No tongue develops in toad family Pipidae
Reptiles & mammals:
o 3 distinct features
Pair of lateral lingual swellings
From mesechyme from mandibular
arch; not found outside of amniotes
Suppressed in birds
Primary component from hyoid arch that
develops a glandular field
Mesenchyme from 3rd pharyngeal arch
spreads forward over some of 2nd arch
mesenchyme
o Sensory epithelia thus formed from mesenchyme
from pharyngeal arches 1, 2, 3; innervated by cranial
nerves 5, 7, & 9
o Hypobranchial musculature invades entire complex
& receives somatic motor innervation from 12 th
cranial nerve
o Tongue of amniotes anchored to skeletal
components of hyoid arch
o Turtles, crocs, alligators, some baleen whales:
tongue also affixed to floor of oral cavity
o Garter snakes have no tongue
Birds: tongue almost lacking in intrinsic muscles
o Only movement comes from muscular operation of
hyoid skeleton to which tongue is anchored
Tongue of agnathans NOT homologous w/ any component of
gnathostome tongue

Rodlike lingual cartilage of unknown homology

capped by horny spines
o Operated by protractor & retractor muscles
Tongue widely used to capture or gather food
o Flipping tongue (over mandibular symphysis) occurs
when long fibers of genioglossus medialis muscle
stiffen to form a complex of intrinsic rods & when
genioglossus basalis muscle swells; returns to
mouth via contraction of hypoglossal muscle
o Woodpeckers: barbed tongue; hummingbird tongue;
immobilized tongue of baleen whales; parrot tongue
armed w/ 2 flexible horny shields composed of
keratinized epithelial cells
These epithelial cells grow forward from naillike bed halfway back on tongue
Entoglossal bone embedded within tongue of birds &
lizards; anteriorly directed process of hyoid
Paraglossal bone attached to entoglossus in many birds
Tongues of most mammals protrusible though tied in floor of
oral cavity by the frenulum linguae
Stereognosis perception of shape, weight, & texture of
solid body
o Mucosa of amniote tongue contains receptors for
this
Encapsulated nerve endings enable insectivores to search
for food in dark places using their tongue; also allows seedeating birds to manipulate seed being husked in the seed
cup
Tongue manipulates fluids & solids; in most tetrapods,
participates in swallowing
Also functions in cooling the blood; lizards clean their
transparent eyelids/spectacles using their tongues; spiny
papillae on tongue surface may be used by carnivores to
rasp bones; may also be used by many mammals for
grooming (kaya may hairballs hihi :D); speech
o

Oral Glands

Secretes watery or viscuous fluids

Chief ingredient: mucus (varying viscosities & chemical
composition)
o Moistens food to produce a bolus that can be
manipulated by tongue
o Lubricates dry food for passage thru pharynx, down
into esophagus
o Moisture also essential for taste buds to function
(stimulant for taste must be in solution to evoke
gustatory response)
o Other secretions include enzyme that digests starch
(rare outside of mammals)
o Viscous secretions keep tongue sticky; venom
tranquilizes prey
Usually named according to location
o Labial glands open into oral vestibule at base of
the lips
o Molar glands
o Infraorbital floor of orbit
o Palatal includes venom glands of 4 families of
venomous snakes (venom exudes into groove/tube
in the fang)
o Sublingual & submandibular open via common
papillae under tongue
In Heloderma, sublingual glands secrete
toxin
o Intermaxillary/internasal near premaxilla
Saliva mixture of oral secretions; term usually reserved for
oral secretions of mammals
o Parotid largest tetrapod salivary gland
Resembled histologically by poison gland of
reptiles
o Ptyalin (amylase) one of the secretions of the
parotid

Specific mix of Mucus + serous secretions + ptyalin

correlated with dietary habits
o Birds do not have a copious supply of saliva
Aquatic craniates commonly have mucus-producing goblet
cells
o These are the only source of oral secretions
o Mucus lubricates esophagus
o Performs special function in male popn of few
species of catfishes
Males carry fertilized eggs in brood pouches
(crypts) in the mucosa of the palate
Goblet cells produce a copious nutritious
secretion that maintains a suitable envt for
the development of eggs; also nourishes the
hatchlings
Brood pouches atrophy after hatchlings
leave, in response to altered hormonal ratios
o Rare instance of multicellular oral gland:
anticoagulant gland of lampreys
o

Teeth

Bony teeth: jawed fishes, amphibians, reptiles, most

mammals, earliest birds
o Achieved a peak in regional specialization in
mammals
No teeth: sturgeons, numerous teleosts including sea
horses, few amphibians, all turtles, modern birds, whalebone
whales, South American & scaly anteaters, Echidna
o Many toothless species still develop an embryonic
set of teeth, but this doesnt disrupt, or it disappears
after disrupting
Descendant of denticles of dermal armor that covered the
head & extended into the oropharyngeal cavity of early
fishes

Gnathostomes with no teeth have lost the genetic

code necessary to induce or complete their
development
Early fishes: bony dental plates of dermal origin overlay
endoskeletal components of jaws
o Pointed, rounded, or jagged surface projections

These prevented the escape of live food

from oropharyngeal cavity

Used to crush shellfish

To bite flesh

To rasp vegetation
o Placoderms: pattern of dermal structures associated
with jaws incompletely known

Range from unossified Meckelian cartilage

associated with overlying denticles within
skin to fully ossified dermal plates overlying
or adjacent to Meckelian cartilage

Did NOT possess teeth

Surface of dental plates hardened & shaped

to perform toothlike functions

Morphology of biting surface of dental plates

correlated with putative function

Flat crushing

Sharp shearing

Spikes impaling

Cusps holding prey

Loss of specialized biting structures

in filter feeders

Single or paired upper dental plates

opposed the lower jaw on each side
o Acanthodians: each denticle attached directly to
endoskeleton of jaws, like modern teeth

Jaws also often flanked by additional

denticles

Some toothless or had lower teeth

o

Dermal denticles and placoid scales show a gradual

transition to teeth as they approach the cutting edges of jaws
o Proof that later vertebrate teeth are derived from
bony dermal armor
Teeth are like placoid scales in that it is also composed of
dentin (a variety of bone), surmounted by a crown of enamel
or enameloid
Development
o Earliest indication of devt of socketed teeth:
ingrowth into dermis of a longitudinal ridge of
ectoderm, called the dental lamina

Extends more or less the length of the jaws

o Dermal papillae, each designating the site of a future
tooth, may be present beneath or above it; these
papillae indent the lamina & organize blood vessels
necessary for further development of a tooth
primordium
o The cells at periphery of papilla become organized
into a definite layer of odontoblasts that deposit
dentin
o Odontoblasts slowly withdraw toward the center of
the primordium (pulp cavity) as deposition proceeds

Evidence of withdrawal of odontoblasts:

dentinal
tubules
that
contain
their
protoplasmic processes

Odontoblasts remain alive throughout the

life of the tooth
o Ectoderm of dental lamina organizes an enamel
organ (ameloblasts)

This deposits enamel on the surface of

dentin

Thin layer of cementum (acellular bone)

eventually anchors tooth to bone of jaw via
collagenous fibers

Living remnants of dermal papilla remain within

whatever is left of the pulp cavity (root canal)
throughout the life of the tooth
o Details of tooth development and emergence, time
of initiation of different stages, ultimate fate of
erupted teeth vary with the species
Armadillos & few other vertebrates: enamel organ present
but functionless
Mammals: enamel deposited by ameloblasts of ectodermal
origin
Fishes & other vertebrates: enameloids differ in physical
characteristics, & the ultimate source of scleroblasts that
elaborate them has been determined to be odontoblasts,
which form a compact dentin
Gnathostomes: teeth vary in number, distribution within oral
cavity, position with reference to summit of jaws, degree of
permanence, shape
o Develop on jaws, palatal bones, & even pharyngeal
skeleton in the oropharynx of living fishes
o Blue sucker has 35-40 teeth on the LAST gill arch
0__0
Early tetrapods: teeth widely distributed on the palate
o Most amphibians & many reptiles have teeth on the
vomer, palatine, & pterygoid bones; occasionally on
the parasphenoid
Crocodilians, fossil toothed birds, mammals: confined to the
jaws
o Least numerous ang teeth among mammals :D
o Teeth, like dermal armor, have tended towards a
more restricted distribution with the passage of time
Types of dentition
o Acrodont: as in many teleosts; teeth attached to
outer surface or to summit of jawbone
o Pleurodont: as in anurans, urodeles, & many lizards;
attached to inner side of jawbone
o Thecodont: occupy bone sockets or alveoli
o

Sockets are deepest in mammals

Polyphyodont: as in most gnathostomes through
amniotes; succession of teeth, number of
replacements during a lifetime indefinite but
numerous

Elderly crocodile: replace front tooth 50x

Crocodilians &
other
nonmammalian
gnathostomes replace teeth in waves that
sweep along the jaws eliminating &
replacing every other tooth

In one wave in tetrapods, evennumbered teeth are lost and oddnumbered ones in the next

Tooth germs for the next wave of

eruptions subsequently form

Waves sweep in different directions

in different species

Waves
ensure
a
balanced
distribution of teeth throughout life

Sharks: tooth germs form in dermis

on the oropharyngeal cavity side of
jaws; migrate onto cutting edge of
jaw during growth; teeth that are
being replaced move beyond the
edge & fall away
Diphyodont: 2 sets (deciduous/milk teeth &
permanent teeth), definite sequence in which the
teeth erupt

Only in mammals is there a definite number

of teeth in a species

Sequence of eruption in humans (1-8 from

front to rear): 6,1,2,4,5,3,7,8

Eruption of 8 is delayed in higher primates

(wisdom tooth)

First set provides constantly changing infant

jaw with small temporary teeth adequate for

o
o

an infants diet until the jaws are more

stabilized structurally & have elongated
sufficiently to accommodate large teeth
Monophyodont: 1st set only

Platypus: milk teeth replaced by horny

epidermal teeth

Toothless whales: 1st set forms within the

jawbone; these do not erupt or are shed if
they do

Freshwater manatee from the Amazon &

Australian Rock Wallaby: no sets, teeth
replaced throughout life by the forward
migration of new teeth formed at the rear of
the jaws

Manatee: migration rate is 1 or 2

mm/month

Thin bony sockets separate roots of

successive teeth, bony septa
resorbed under pressure from
migrating teeth

Grasses eaten by manatee contain

abrasives that appear to be
necessary for the teeth to move
forward

Proboscidians: slow but constant succession

of molar teeth that move forward from the
rear
Homodont: when all teeth are essentially similar
Heterodont: teeth vary morphologically from front to
rear

Applies to all but a very few mammals

(cetaceans, sirenians, & some marine
carnivores exhibit homodont dentition)

Incisors, canines, premolars, molars (last 2

are cheek teeth)

Arose in the late synapsids

Morphologic variants in fishes

o Most sharks: fish eaters,; numerous rows of jaw
teeth that are flat, sharp, or notched triangles used
to cut; single or multipointed tusks that curve toward
pharynx to hold struggling prey until it can be
swallowed

Each shark tooth has a broad basal plate of

bone embedded in dermis

Minority of sharks eat shellfish: teeth at

entrance have curved caudally directed
spines; rest are batteries of rounded
denticles to crush shells

Tiny stomodeal denticles line the pharynx in

some sharks (transitional shapes between
denticles & teeth pag near the jaws)
o Holocephalans & modern lungfishes: similar to early
jawed fishes; few large plates of enameloid/enamelcovered dentin that bear rows of various-sized
rounded mound-like denticles which become sharp
spines at the entrance to the oropharyngeal cavity

Chimaera: one large anterior + one small

posterior dental plate on each side of upper
jaw to cover the entire upper jaw; single
large plate on each side below

Modern lungfish: plates restricted to palate &

medial aspects of jaw
o Actinopterygii, amphibians, most reptiles: simple
pointed cones attached to one or more membrane
bones

Small teeth may be interspersed among

larger ones

Those in front sometimes larger & curved

slightly to rear

Specialized shapes sometimes appear on

one jaw or the other

Gars: few fanglike teeth shaped at

their ends like arrows

Venomous snakes: fangs borne on

maxillae; curved, bladelike, or
tubular for injecting venom
Morphologic variants in mammals
o Incisors: located on either side of mandibular
symphysis

One horizontal cutting edge

Single root

Best developed in herbivorous mammals

(used to hold, crop, or gnaw)

Rodents & large front pair of lagomorphs:

enamel on anterior surface ONLY

Lagomorphs have the 2nd pair of

incisors BEHIND the 1st

Gnawing wears away dentin faster;

this keeps cutting edges of incisors
sharp

Incisors grow throughout life

Bovines: lacking on the upper jaw

Vampire bats: lacking on the lower jaw

Sloths: no incisors

Elephant & mastodon tusks are modified

incisors that grow throughout life

Walrus tusks NOT incisors but

canines
o Canines: lie next to incisors

Scarcely differ in appearance in generalized

mammals

Carnivores: spearlike, used to pierce flesh

Absent in lagomorphs, so there is a

diastema between incisors & first cheek
tooth

Rodents: premolars are also missing so the

diastema is longer

Attained greatest length on the upper jaw of

saber-toothed tigers

20 cm below lower jaw with the

mouth closed

Lower canines reduced

Premolars

Most mammals EXCEPT ungulates: 2

prominent cusps (bicuspid)

1 or 2 roots; number of roots may differ on

upper & lower jaw among different
individuals of same population
Molars

Tricuspid

Usually with 3 roots; occasionally 4 or 5

Not replaced by a 2nd set; late arrivals of the

1st
Crown: part of the tooth above the gum line

Covered with enamel

Crown of cheek teeth of carnivores &

herbivorous ungulates show extreme
morphological differences

Carnivores
o
for tearing flesh & crushing
bone
o laterally compressed, 2 or 3
cusps interconnected by
sharp ridges of enamel,
long roots (secodont)
o sharp enamel ridges of
crowns produce shearing
effect because the cusps of
upper jaw teeth fit between
the cusps of lower jaw
teeth;
o carnassial teeth (last upper
premolar + first lower molar)

Bovines
selenodont cheek teeth
no teeth anterior to them in the upper jaw;
cheek teeth employed in chewing cud
Proboscidians

Adaptation for grinding exaggerated

Lophodont teeth: enamel & dentin intricately

interfolded & enamel disposed on ridges on
enormous plateaus of naked dentin

Reach a foot or more in length and a third of

a foot in width in the largest elephants
Remaining mammals

Cheek teeth exhibit a variety of molariform

styles

Omnivores & some herbivores: low rounded

cusps instead of sharp edges and pointed
cusps (bunodont teeth)

larger and longer in order to

deal with tough shearing
problems
Ungulates & some herbivores
o for macerating vegetation
o
cheek teeth wider & longer,
providing broad surfaces for
grinding
o crowns are tall, allowing for
plenty of wear
o
crowns also consist of
crescentic
columns
of
dentin
embedded
in
additional dentin devoid of
enamel
overlay;
each
column
surrounded
by
enamel
o Selenodont

hogs,

Rodents

Largest mammalian order, largest variety of

diets, largest variety of teeth

Squirrels: low crowned with long roots

Wood rats: high crowned with short roots

o Crabeater seal

Among the most unusual mammalian teeth

Employed to strain small crustaceans &

other plankton from mouthfuls of seawater
as it spills back into the sea
o Early prototherians

Triconodont: crown has 3 conelike

prominences arranged in a straight line
o Early therians

Trituberculate:
crowns
3
conelike
prominences arranged in a triangle

Forerunners of todays tricuspids

Enamel crests connecting the cones
is thought to account for selenodont
& lophodont teeth
1st therians: total of 44 teeth (3-1-4-3)
See book for other formulas of a few
selected mammals :p (p. 279)
Teeth, along with the tongue & hyoid, form a functional triad
that procures, manipulates, & (in mammals) masticates
foodstuffs at the entrance to the digestive tract, then starts a
bolus of food

Humans, rhinos, some

primitive ruminants, rodents

Epidermal Teeth
Keratinized (horny) teeth in living agnathans buccal cavity
and tongue, for rasping
Anuran tadpoles: temporary lips perched on poorly
developed jaws, for rasping algae and other vegetation;
shed and replaced by bony ones at metamorphosis

Turtles, crocodilians, Sphenodon, birds, monotremes:

temporary horny egg tooth used to crack egg shell
Platypus: horny teeth replace bony teeth and remain
throughout life
Horny beaks of turtles and modern birds have serrations that
perform some of the functions of teeth

PHARYNX
Part of the digestive tract that had pharyngeal pouches in the
embryo
Opens into the esophagus
Fishes: functional part of the respiratory system
Most constant features:
o Glottis slit opening into larynx
o Openings of paired auditory tubes leads to the
middle ear cavity
o Opening into the esophagus
In mammals, additional features:
o Nasal pharynx above soft palate
nasal passageways empty into nasal
pharynx via choanae
auditory tubes derived from 1 st pair of
pharyngeal pouches open into its lateral
walls
pharyngeal tonsils (adenoids) develop in the
mucosa
o Oral pharynx between oral cavity and glottis
Isthmus of the fauces: narrow passageway
marking transition from oral cavity to oral
pharynx
Lateral walls of isthmus exhibit 2 pillars of
the fauces, which are muscular folds that
arch upward from the side of the tongue to
the soft palate (glossopalatine arch) and
from the pharyngeal wall to the soft palate
(pharyngopalatine arch)

Humans & some other primates: uvula

hangs from caudal border of soft palate into
oral pharynx
Palatine tonsil in the hollow between the
pillars
Develop in the walls of the 2nd
pharyngeal pouch
Remnant of pouch remains as
pocket-like crypt with tonsil at its
wall
o Laryngeal pharynx dorsal to the larynx in mammals
in which the opening to the esophagus is caudal to
the glottis
o Lingual tonsils develop on the tongue near its
attachment to the hyoid bone
o Tonsils as lymphoid masses that serve as the bodys
first line of defense against infective agents
o Epiglottis: fibrocartilaginous flap that lies in the floor
of the pharynx ventral to pharyngeal chiasma;
attached to hyoid bone
Regulation of air & food traffic
o In many mammals, swallowing draws the larynx
forward (upward in humans) against the epiglottis,
closing the glottis
o In other mammals, the epiglottis and part of the
larynx can be drawn into the nasopharynx to provide
an uninterrupted air pathway to the lungs; foodstuffs
detour around the larynx & enter the esophagus
o Other tetrapods: fleshy valves at appropriate
locations
E.g. valves that open/close the entrance to
external nares in aquatic tetrapods
Some teleosts:
o Suprabranchial organs (muscular tubes) evaginate
from roof of pharynx, terminate as blind sacs

Elongated gill rakers from last 2 gill arches

form funnel-shaped baskets that extend into
the entrance of suprabranchial organs
Each tube surrounded by cartilaginous
capsules that furnishes attachment for the
striated muscle of the tube
Epithelium at blind ends have many goblet
cells
Sacs contain quantities of plankton
sometimes compressed into a bolus
Therefore, possible fxn: trap plankton from
incoming respiratory water stream &
concentrate it into mucus-rich masses that
are swallowed
Air-gulping teleost: cavity filled with air &
highly vascularized epithelial lining serves
as accessory respiratory membrane

MORPHOLOGY OF THE GUT WALL

4 histological layers (outward): mucosa, submucosa,
muscularis externa, serosa
o Differences lie in the thickness and the nature of glands
Mucosa
o Consists chiefly of glandular epithelial lining of endodermal
origin + layer of not very dense (areolar) connective tissue
supporting base of cryptlike epithelial glands, lymph nodes, & blood
and lymph capillaries + thin coat of smooth muscle fibers (muscularis
mucosae, may be absent in some regions)
o Mucous glands provides a lubricant that facilitates passage
of contents during peristalsis
Submucosa
o Thicker layer of connective tissue supporting base of
compound alveolar glands & rich plexus of arterioles, venules, &
lymphatics that service capillary beds of the mucosa
Muscularis externa
o Smooth muscle tissue arranged into 2 layers

 Inner circular layer smooth muscles, constricts lumen on

neural demand
Outer longitudinal layer contracts short segments of gut
o Combined action of smooth muscles produces the
macerating, peristaltic, & (in mammalian colon) segmenting actions
of the gut
o Neural stimuli for contraction supplied from autonomic
(visceral) nerve plexuses between the longitudinal & circular muscle
layers and between circular layer & submucosa
Serosa
o Consists of loose connective tissue (adventitia) + covering of
visceral peritoneum
o Exudes small amounts of serous fluid that lubricates surface
of viscera, reducing friction
o Peritonitis: inflammation of serosa, leads to exudation of
excessive quantities of fluid
o Esophagus & caudalmost portion of intestine: covered by
serosa on one surface because they generally lie against the body
wall
Many larval craniates, as in protochordates: Entire digestive
tract ciliated
Many Teleosts: cilia in the stomach
Some adult amphibians: cilia in the oral cavity, pharynx,
esophagus, & stomach
Cilia are present for a time in the stomach of human fetus
Peristalsis chiefly responsible for moving foodstuffs along
alimentary canal