Chapter 12 Digestive System

PROCURING FOOD

Filter feeders: ancestral chordates (sea squirts, amphioxus), living larval agnathans (e.g. lampreys) o o This mode of feeding is primitive Can be employed only by aquatic organisms Passive filtration of organic matter from the incoming respiratory stream, propel particles to the rear of the pharynx for swallowing

More aggressive methods of obtaining food made possible by jaws & elaborate musculature of body walls (for locomotion & pursuit)

o o

Jaws were first invested with denticulated bony dermal armor In specialized predators, eventually furnished with sharp denticles (teeth)

Bite-tear-swallow modern sharks

technique,

as

in

More active version of filter feeding in some fishes (spoonbills, cluepids, basking sharks)

This required no tongue specializations of oral cavity

or

other

Plankton & small fish strained out of respiratory stream using long filamentous gill rakers hanging into pharyngeal chamber from gill arches

Less energy consuming procedure due to further adaptations of skull & hyoid arches o o Protrusible jaws, suction (as in goldfish) Lampreys, due to parasitic nature, rasp host tissues with spiny tongue (fleshy & cartilaginous rod armed w/ horny teeth) & suck the debris into pharynx

Whalebone/Baleen cacity

hangs

into

oral

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)

Carnivorous mammals: snap-bite-tear technique o Often involves piercing effect of saberlike tooth

Active predation/active acquisition of particulate food in adults of early fishes -> led to paired external sense organs & their concentration on the head

Food taking depends on food finding

Use chemical, mechanical, thermal, capsulated touch, visual, electroreceptors

THE DIGESTIVE TRACT: AN OVERVIEW

Digestive tract aka alimentary canal o 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:

Occupies the pleuroperitoneal cavity in fishes, amphibians, most reptiles 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)

o
o o o

Oral cavity & pharynx (in fishes: oropharyngeal cavity) Esophagus Stomach Intestine (divided tetrapods) into small & large in

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 Dorsal mesentery remains intact (conducts blood vessels & nerves from roof of coelom to digestive organs) Ventral mesentery disappears EXCEPT at the level of the liver & urinary bladder

Ducts from accessory organs also empty into the tract o o Pancreas, liver, gall bladder These organs arise as evaginations from the embryonic digestive tract

Ceca are also commonly present Digestive system = tract + accessory organs

Embryonic digestive tract has 3 regions:

Midgut contains the yolk (when present), where the yolk sac is attached; little remains in adults Hindgut becomes remainder of intestine and cloaca Foregut elongates to form part of oral cavity, pharynx, esophagus, stomach, much of small intestine

Oropharyngeal cavity short esophagus

terminates

at

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

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

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

Differences in anatomy of digestive tracts caudal to pharynx correlated with nature & abundance of food

MOUTH & ORAL CAVITY

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

Mouth entrance to digestive tract o opens into oropharyngeal cavity w/ teeth & walls perforated by gill slits (in gnathostome fishes)

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

Primary tongue develops from hyoid arch mesenchyme 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

Seed & grain eating birds: median sublingual seed pouch Lies upon caudal portion of mylohyoid muscle beneath oral cavity Retracted by homologue genioglossus muscle of mammals of

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

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

No tongue develops in toad family Pipidae

Reptiles & mammals: o 3 distinct features Pair of lateral lingual swellings From mesechyme mandibular arch; not outside of amniotes Suppressed in birds from found

Terrestrial urodeles, anurans: primary tongue + extension that can be flipped out of mouth

Primary component from hyoid arch that develops a glandular field

Mesenchyme from 3rd pharyngeal arch spreads forward over some of 2nd arch mesenchyme

Sensory epithelia thus formed from mesenchyme from pharyngeal arches 1, 2, 3; innervated by cranial nerves 5, 7, & 9 Hypobranchial musculature invades entire complex & receives somatic motor innervation from 12th cranial nerve Tongue of amniotes anchored to skeletal components of hyoid arch Turtles, crocs, alligators, some baleen whales: tongue also affixed to floor of oral cavity Garter snakes have no tongue 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 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 nail-like 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

Birds: tongue almost lacking in intrinsic muscles o Only movement comes from muscular operation of hyoid skeleton to which tongue is anchored w/ any

Tongue of agnathans NOT homologous component of gnathostome tongue o

Rodlike lingual cartilage of unknown homology capped by horny spines Operated by protractor & retractor muscles

Tongue widely used to capture or gather food

Encapsulated nerve endings enable insectivores to search for food in dark places using their tongue; also allows seed-eating 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 o o

Molar glands Infraorbital floor of orbit Palatal includes venom glands of 4 families of venomous snakes (venom exudes into groove/tube in the fang) Sublingual & submandibular common papillae under tongue open via

Oral Glands Secretes watery or viscuous fluids Chief ingredient: mucus chemical composition) o (varying viscosities & o

In Heloderma, sublingual glands secrete toxin

Intermaxillary/internasal near premaxilla

Moistens food to produce a bolus that can be manipulated by tongue Lubricates dry food for passage thru pharynx, down into esophagus Moisture also essential for taste buds to function (stimulant for taste must be in solution to evoke gustatory response) Other secretions include enzyme that digests starch (rare outside of mammals) Viscous secretions keep tongue sticky; venom tranquilizes prey

Saliva mixture of oral secretions; term usually reserved for oral secretions of mammals o Parotid largest tetrapod salivary gland Resembled histologically gland of reptiles by poison

Ptyalin (amylase) one of the secretions of the parotid Specific mix of Mucus + serous secretions + ptyalin correlated with dietary habits Birds do not have a copious supply of saliva

Usually named according to location o Labial glands open into oral vestibule at base of the lips

Aquatic craniates commonly have mucus-producing goblet cells o These are the only source of oral secretions

o o

Mucus lubricates esophagus 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

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 o 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

Rare instance of multicellular oral gland: anticoagulant gland of lampreys

Teeth Bony teeth: jawed fishes, amphibians, reptiles, most mammals, earliest birds o Achieved a peak in regional specialization in mammals 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 &

No teeth: sturgeons, numerous teleosts including sea horses, few amphibians, all turtles, modern birds, whalebone whales, South American & scaly anteaters, Echidna

Morphology of biting surface of dental plates correlated with putative function Flat crushing

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

Sharp shearing o 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 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 The cells at periphery of papilla become organized into a definite layer of odontoblasts that deposit dentin 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

Acanthodians: each denticle attached directly to endoskeleton of jaws, like modern teeth Jaws also denticles often flanked by additional

Some toothless or had lower teeth

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

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

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

Living remnants of dermal papilla remain within whatever is left of the pulp cavity (root canal) throughout the life of the tooth

Details of tooth development and emergence, time of initiation of different stages, ultimate fate of erupted teeth vary with the species

Types of dentition o Acrodont: as in many teleosts; teeth attached to outer surface or to summit of jawbone Pleurodont: as in anurans, urodeles, & many lizards; attached to inner side of jawbone Thecodont: occupy bone sockets or alveoli o Sockets are deepest in mammals

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 Blue sucker has 35-40 teeth on the LAST gill arch 0__0

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

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 o Least numerous ang teeth among mammals :D Teeth, like dermal armor, have tended towards a more restricted distribution with the passage of time

jaw during growth; teeth that are being replaced move beyond the edge & fall away o Diphyodont: 2 sets (deciduous/milk teeth & permanent teeth), definite sequence in which the teeth erupt Only in mammals is there number of teeth in a species a definite

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

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 an infants diet until the jaws are more stabilized structurally & have elongated sufficiently to accommodate large teeth o o

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

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

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)

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

Holocephalans & modern lungfishes: similar to early jawed fishes; few large plates of enameloid/enamel-covered 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

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

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

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 canines NOT incisors but o appearance in

Usually with 3 roots; occasionally 4 or 5 Not replaced by a 2nd set; late arrivals of the 1st

Canines: lie next to incisors Scarcely differ in generalized mammals

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 crushing bone flesh &

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

laterally compressed, 2 or 3 cusps interconnected by sharp ridges of enamel, long roots (secodont) sharp enamel ridges of crowns produce shearing effect because the cusps of upper jaw teeth fit between the cusps of lower jaw teeth; carnassial teeth (last upper premolar + first lower molar) larger and longer in order to deal with tough shearing problems

o Premolars

Most mammals EXCEPT prominent cusps (bicuspid)

ungulates:

1 or 2 roots; number of roots may differ on upper & lower jaw among different individuals of same population

Molars Tricuspid

Ungulates & some herbivores o for macerating vegetation

cheek teeth wider & longer, providing broad surfaces for grinding crowns are tall, allowing for plenty of wear crowns also consist of crescentic columns of dentin embedded in additional dentin devoid of enamel overlay; each column surrounded by enamel Selenodont

Cheek teeth exhibit a variety of molariform styles Omnivores & some herbivores: low rounded cusps instead of sharp edges and pointed cusps (bunodont teeth) Humans, rhinos, some hogs, primitive ruminants, rodents

o
o

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
o Bovines

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

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

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 o

Early prototherians Triconodont: crown has 3 conelike prominences arranged in a straight line

Early therians Trituberculate: crowns 3 conelike prominences arranged in a triangle Forerunners of todays tricuspids

Remaining mammals

Enamel crests connecting the cones is thought to account for selenodont & lophodont teeth

Opens into the esophagus Fishes: functional part of the respiratory system Most constant features: o o Glottis slit opening into larynx Openings of paired auditory tubes leads to the middle ear cavity Opening into the esophagus

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

In mammals, additional features: o Nasal pharynx above soft palate nasal passageways pharynx via choanae empty into nasal

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

auditory tubes derived from 1st pair of pharyngeal pouches open into its lateral walls pharyngeal tonsils (adenoids) develop in the mucosa

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

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)

PHARYNX Part of the digestive tract that had pharyngeal pouches in the embryo

Humans & some other primates: uvula hangs from caudal border of soft palate into oral pharynx Palatine tonsil in the hollow between the pillars

Other tetrapods: locations

fleshy valves

at appropriate

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

Develop in the walls of the 2 pharyngeal pouch

nd

Remnant of pouch remains as pocket-like crypt with tonsil at its wall

Laryngeal pharynx dorsal to the larynx in mammals in which the opening to the esophagus is caudal to the glottis Lingual tonsils develop on the tongue near its attachment to the hyoid bone Tonsils as lymphoid masses that serve as the bodys first line of defense against infective agents 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 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

MORPHOLOGY OF THE GUT WALL 4 histological layers (outward): muscularis externa, serosa mucosa, submucosa,

Differences lie in the thickness and the nature of glands 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) Mucous glands provides a lubricant that facilitates passage of contents during peristalsis Serosa o

longitudinal & circular muscle layers and between circular layer & submucosa

Mucosa o

Consists of loose connective tissue (adventitia) + covering of visceral peritoneum Exudes small amounts of serous fluid lubricates surface of viscera, reducing friction Peritonitis: inflammation of serosa, leads exudation of excessive quantities of fluid that

to

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

Esophagus & caudalmost portion of intestine: covered by serosa on one surface because they generally lie against the body wall as in protochordates: Entire

Many larval craniates, 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

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

Combined action of smooth muscles produces the macerating, peristaltic, & (in mammalian colon) segmenting actions of the gut Neural stimuli for contraction supplied from autonomic (visceral) nerve plexuses between the