Comparative Anatomy of the Vertebrates:

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Digestive System]
B. ACCESSORY ORGANS 1. PANCREAS, LIVER and GALLBLADDER evaginations from the embryonic digestive tract Digestive system = accessory organs + digestive tract C. EMBRYONIC DIGESTIVE TRACT 1. FOREGUT elongates to form part of the oral cavity, pharynx, esophagus, stomach and small intestine 2. MIDGUT part containing the yolk or to which the yolk sac is attached little remains in adults 3. HINDGUT become the remainder of the intestine and the cloaca D. PLEUROPERITONIAL CAVITY 1. PLEURAL CAVITIES subdivision of the coelom enclosing the lungs 2. PERITONEAL/ABDOMINAL CAVITY coelom occupied by the digestive organs beyond the esophagus in mammals, birds and a few other reptiles E. MESENTERY 1. DORSAL MESENTERY remains intact throughout life, conducting blood vessels and nerves from the roof of the coelom to the digestive organs 2. VENTRAL MESENTERY eventually disappears except at the level of the liver and urinary bladder F. PERITONEUM 1. VISCERAL PERITONEUM outer covering of the digestive tract 2. PARIETAL PERITONEUM continuous with the visceral peritoneum and lines the body wall G. RELATED STRUCTURES/TERMS 1. CECA other evaginations from the embryonic digestive tract

OUTLINE I. The Digestive Tract: An Overview A. Major Subdivisions of the Digestive Tract B. Accessory Organs C. Embryonic Digestive Tract D. Pleuroperitoneal Cavity E. Mesentery F. Peritoneum G. Related Structures/Terms II. Mouth and Oral Cavity A. Related Structures B. Tongue C. Oral Glands D. Teeth III. Pharynx A. Most Constant Features of a Tetrapod Larynx B. Additional Features of a Mammalian Pharynx IV. Gut Wall Morphology A. Four Histological Layers V. Esophagus A. Structure B. Comparative Anatomy of Esophagus VI. Stomach A. Parts of the Stomach B. Comparative Anatomy of Stomach C. Functions of the Stomach VII. Intestine A. Fishes B. Tetrapods VIII.Lives and Gallbladder A. Liver B. Gallbladder IX. Pancreas A. Exocrine Pancreas B. Endocrine Pancreas X. Cloaca

I. THE DIGESTIVE TRACT: AN OVERVIEW A. MAJOR SUBDIVISIONS OF THE DIGESTIVE TRACT 1. ORAL CAVITY and PHARYNX (oropharyngeal cavity in gnathostome fishes) 2. ESOPHAGUS 3. STOMACH 4. INTESTINE subdivided in tetrapods into: (a) small intestine (b) large intestine

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3. PALATAL FISSURE deep cleft in the roof of the oral cavity due to incomplete 2 palate channelizes respiratory air between the choanae and pharynx found in most reptiles 4. VOCAL SACS reverberating chambers beneath the pharyngeal floor in anurans, open into the oral cavity near the angle of the jaws 5. VESTIBULE a trench that separates the gums from the cheeks and lips in mammals 6. ALVEOLAR RIDGES gums 7. CHEEK POUCH used by rodents to transport grain from the fields to the burrow for storage as SEED POUCHES in birds B. TONGUE 1. PRIMARY TONGUE no glands and musculature (cannot be independently manipulated) but may assist the jaw in holding the prey forerunner of tetrapod tongue derived from mesenchyme of III branchial arch 2. GLANDULAR FIELD secretes a sticky mucus as TUBERCULUM IMPAR in amniotes 3. LATERAL LINGUAL SWELLINGS derived from mesenchyme of II branchial arch 4. FUNCTIONS OF THE VERTEBRATE TONGUE gathering or capturing food for sense of taste manipulation of food in the oral cavity swallowing/deglutition thermoregulation grooming human speech streognosis (prognosis of the shape, weight and texture of food; made possible by mucosa of tongue; found in amniotes)

2. PERISTALSIS process of propelling food from the pharynx to the vent/anus by smooth muscle tissue in the walls of the tract 3. STOMODEUM embryonic ectodermal invagination that gives rise to the anterior oral cavity or oropharynx of fishes 4. PROCTODEUM embryonic ectodermal invagination at the future exit of the hindgut 5. ORAL PLATE a thin membrane that temporarily separates the early embryonic foregut from the exterior; ruptures eventually to provide an anterior entrance to the digestive tract 6. CLOACAL PLATE ruptures to provide an exit from the hindgut The coelom arises by the delamination, or splitting, of the early lateral-plate mesoderm into two sheets, somatic and splanchnic mesoderm. The space between the two sheets becomes the coelom. Most of the embryonic digestive tract other than the endodermal lining arises in cephalochordates and craniates from splanchnic mesoderm. Early in embryonic life, the parietal and visceral peritonea are continuous via dorsal and ventral mesenteries. Like all chordates, craniates, including humans, are deuterostomous. II. MOUTH AND ORAL CAVITY A. RELATED STRUCTURES 1. OROPHARYNGEAL CAVITY where mouth opens into in gnathostome fishes walls perforated by gills; terminates at a short esophagus The roof of the oropharyngeal cavity of fishes and oral cavity of amphibians is a primary palate, pierced anteriorly by internal nares in lungfishes and amphibians. 2. ORAL (BUCCAL) CAVITY where mouth opens into in tetrapods houses teeth and tongue leads to the pharynx

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2. POISON-SECRETING GLANDS PALATAL GLANDS open onto the palate; large venom glands of venomous snakes 3. ANTICOAGULANT GLANDS secretions inhibit blood clotting of the prey found in SANGUINIVORES (animals that feed on blood) 4. OTHER GLANDS (named according to location) LABIAL GLANDS open into the oral vestibule at the base of the lips MOLAR GLANDS lie near the molar tooth INFRAORBITAL GLANDS in the floor of the orbit INTERMAXILLARY / INTERNASAL GLANDS lie near the premaxillary bone 5. RELATED STRUCTURES GOBLET CELLS mucus-producing cells that are about the only source of oral secretions in aquatic craniates BROOD POUCHES temporary folds or crypts in male catfishes used to carry fertilized eggs; produce a copious nutritious secretion for nourishment of the hatchlings Birds do not have a copious supply of saliva. Fishes have no saliva at all. Moisture is also essential for tastes buds to function because the stimulant for taste must be in solution to evoke a gustatory response.

5. COMPARATIVE ANATOMY OF TONGUE Vertebrate Group Jawed fishes and primitive amphibian Most amphibians Reptiles & mammals Description Primary tongue Primary tongue + glandular field = definitive tongue Primary tongue + glandular field (tuberculum impar) + a pair of lateral lingual swellings = definitive tongue Immobile tongue Lateral lingual swellings are suppressed; intrinsic muscles are absent; with ENTOGLOSSUS (a bone that supports the tongue) Tongue is sometimes long and may move in and out of the oral cavity Attached to the floor of the oral cavity via the FRENULUM

Turtles, crocodilians, whales & some birds Most birds

Snakes, insectivorous lizards & amphibians, and some birds Most mammals

The roof of the tongue of terrestrial urodeles becomes anchored to the basihyal and ceratohyal cartilages in the floor of the pharynx; that of anurans, unlike other tetrapods, becomes anchored to the floor of the oral cavity immediately behind the mandibular symphysis. The tongue of agnathans is not homologous with any component of the tongue of gnathostomes.

C. ORAL GLANDS 1. SALIVARY GLANDS empty SALIVA (mucus + serous fluids) into the mouth principal salivary glands: (a) PAROTID GLAND o largest tetrapod salivary gland o PTYALIN (amylase) initiates starch digestion o mumps (b) SUBLINGUAL GLAND (c) SUBMAXILLARY GLAND o together with sublingual, opens via a common papillae under the tongue Functions: (a) lubrication and binding (BOLUS) (b) solubilizes dry food (c) oral hygiene (d) initiates starch digestion (e) provides alkaline for buffering (f) evaporative cooling

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Comparative Anatomy of the Vertebrates: D. TEETH

[CHAPTER 12:

Digestive System]
Based on morphology: (a) HOMODONT o all teeth are essentially similar as in fishes (b) HETERODONT o vary morphologically from front to rear o arose in late synapsids o found in all but a very few mammals Teeth Incisors Description for cutting; have one horizontal cutting edge and a single root; best developed in herbivorous mammals; grow throughout life; lacking on upper jaw in bovines, on the lower jaw in vampire bats, and completely in sloths; modified as tusks in elephant and mastodon for piercing and tearing; modified as tusks of walruses; absent in lagomorphs forming a toothless interval, or DIASTEMA BICUSPIDS in most mammals other than ungulates; with one or two roots; absent in rodents TRICUSPIDS; usually have three roots; not replaced by a second set but late arrivals of the first set

Teeth are descendants of the denticles of the dermal armor. 1. COMPONENTS OF THE TEETH Component Dentin Description deposited by odontoblasts which remain alive throughout the life of the tooth deposited by enamel organ, an ectoderm of the dental lamina longitudinal ingrowth of ectoderm along the jaw; site of tooth formation; earliest indication of teeth development a type of acellular bone; anchors the tooth to the bone by means of collagenous fibers

Enamel or enameloid Dental lamina

Cementun

An enamel organ is present but functionless in armadillos and a few other vertebrates, hence, their teeth have no enamel. 2. TYPES OF DENTITION Based on attachment: (a) ACRODONT o attached to the summit of the jawbone as in many teleosts (b) PLEURODONT o attached to the inner side of the jawbone as in anurans, urodeles and many lizards (c) THECODONT o occupy bone sockets or alveoli as in many fishes, in crocodilians, extinct toothed birds and mammals Based on no. of replacement sets: (a) POLYPHYODONT o numerous as in most gnathostomes through amniotes (b) DIPHYODONT o two sets of teeth: DECIDUOUS TEETH (milk teeth) PERMANENT TEETH o in most mammals (c) MONOPHYODONT o one set of teeth

Canines

Cheek or Molariform teeth

Premolars

Molars

In platypus, the deciduous set is replaced by HORNY EPIDERMAL TEETH. Most marine mammals have reverted to homodont dentition, but their Cretaceous ancestors exhibited heterodonty. 3. TEETH IN DIFFERENT VERTEBRATE GROUPS SECONDONT and CARNASSIAL carnivores modified molars LOPHODONT proboscidians specialized grinding tooth enamel and the dentin are intricately interfolded, and the enamel is disposed on ridges (loops) on enormous plateaus of naked dentin BURNODONT with low cusps higher primates, rhinos, hogs, some primitive ruminants and some rodents Page 4 of 10

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(c) LINGUAL TONSIL o develop on the tongue near its attachment to the hyoid bone 4. ISTHMUS OF FAUCES narrow passageway between oral and pharyngeal cavities in mammals pillars of fauces: (a) anterior GLOSSOPALATINE ARCH (b) posterior PHARYNGOPALATINE ARCH 5. UVULA fleshy process hanging from the caudal border of the soft palate in humans and as a derived condition in some other primates 6. OPENINGS OF THE PAIRED AUDITORY / EUSTACHIAN TUBES 7. OPENING INTO THE ESOPHAGUS B. ADDITIONAL FEATURES OF A MAMMALIAN PHARYNX 1. NASAL PHARYNX above the soft palate OF THE 2. ORAL PHARYNX between the oral cavity and glottis 3. LARYNGEAL PHARYNX portion of pharynx adjacent to the larynx in those mammals where the opening to the esophagus is caudal to the glottis 4. SUPRABRANCHIAL ORGAN paired muscular tube extending from the esophagus anteriorly above the pharynx in some teleosts; may function to trap plankton from the incoming respiratory water stream IV. MORPHOLOGY OF THE GUT WALL

TRICONODONT crown has three conelike prominences arranged in a straight line cheek teeth of early protherians TRITUBERCULATE early therian tooth with three cusps arranged in a triangle forerunner of todays tricuspids EGG TOOTH keratinized and temporary used by turtles, crocodilians, Sphenodon, birds and monotremes before hatching for cracking the shell 4. RELATED STRUCTURES ROOT CUSPS GINGIVA (gum line) CROWN part above the gum line covered with enamel APICAL FORAMEN where blood vessels and nerves enter TEETH PRIMORDIUM enamel + dermal papilla III. PHARYNX A. MOST CONSTANT FEATURES TETRAPOD PHARYNX 1. GLOTTIS slit leading to the larynx 2. EPIGLOTTIS a firbrocartilaginous flap in mammals that covers the glottis in swallowing prevents food or liquids from entering the trachea during deglutition 3. TONSIL bodys first line of defense vertebrate tonsils: (a) PALATINE TONSIL o located between the pillars of the fauces o develops in the walls of the embryonic second pharyngeal pouches (b) PHARYNGEAL TONSIL/ADENOIDS o develop in the mucosa of the nasal pharynx

A. FOUR HISTOLOGICAL LAYERS 1. MUCOSA mucous glands are ubiquitous, providing a lubricant that facilitates passage of the contents of the tract during peristalsis consists chiefly of: EPITHELIUM o one-celled layer (cuboidal or columnar) o endodermal origin Page 5 of 10

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4. The esophagus is lined by a stratified squamous epithelium, which, in terrestrial turtles, birds and few mammals, is cornified, enabling the lining to withstand abrasions caused by roughage in the diet. B. COMPARATIVE ANATOMY OF ESOPHAGUS 1. The esophagus is shortest in fishes and neckless tetrapods. 2. In fishes, it serves as a sphincter that closes the passageway to the stomach during the phase of respiration when the water is being forced across the gills. 3. Its only function otherwise in fishes and tetrapods is to conduct foodstuffs to the stomach. 4. The esophagus of marine turtles is lined by horny papillae that are directed backward, thereby preventing regurgitation while making it easy to swallow slippery seaweed. 5. CROP a paired or unpaired membranous diverticulum, or sac, found chiefly in grain eaters, who use it for hoarding seeds and grain until there is no room for them in the stomach VI. STOMACH A. PARTS OF THE STOMACH 1. CARDIA and CARDIAC REGION no zymogenic cells glands are compound tubular with many goblet cells a few parietal cells are also present 2. FUNDUS characterized by a specific array of simple tubular gastric glands 3. GREATER CURVATURE in mammals having been the embryonic dorsal border connected to the coelomic roof by the dorsal mesentery (MESOGASTER) 4. LESSER CURVATURE

LAMINA PROPIA o loose/areolar conn. tiss. o abundant in epithelial glands, lymph nodules, and blood & lymph capillaries MUSCULARIS MUCOSAE o a thin coat of smooth muscle fibers 2. SUBMUCOSA a thicker layer of connective tissue supports the base of compound alveolar glands and a rich plexus of arterioles, venules and lymphatics 3. MUSCULARIS EXTERNA consists of two layers of smooth muscle tissue: (a) INNER CIRCULAR LAYER (b) OUTER LONGITUDINAL LAYER The combined action of the two produces the macerating, peristaltic and segmenting actions of the gut. 4. SEROSA consists of loose connective tissue (adventitia) and a covering of visceral peritoneum exudes serous fluid that lubricates the surface of the viscera, reducing friction generated when the organs rub against each other PERITONITIS inflammation of the serosa that results in exudation of excessive quantities of fluids The esophagus and the caudal portion of the intestine may be covered by serosa on only one surface, the one that bulges into the coelom. V. ESOPHAGUS A. STRUCTURE 1. The glands in its lining secrete only mucus. 2. Striated muscle at the cephalic end of a long esophagus is gradually replaced farther down by smooth muscle, except in ruminants. 3. The lining of the rumen is similar to that of the esophagus, and it produces no digestive enzymes.

5. BODY OF THE STOMACH 6. RUGAE increases surface area for absorption Page 6 of 10

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B. COMPARATIVE ANATOMY OF STOMACH

7. PYLORUS and PYLORIC REGION terminal opening of the stomach leading to the duodenum no zymogenic cells with simple branched tubular glands, many goblet cells and relatively few parietal cells 8. PYLORIC SPHINCTER/VALVE ring of smooth muscle regulates the entry of CHYME from the stomach into the small intestine

Vertebrate Agnathans, chimaeras & lungfishes Jawed fishes

Description No definitive stomach

Epithelium is sometimes ciliated; gar stomach is almost straight; sharks exhibit the more common J shape; entire stomach of some teleosts is one large cecum Not distinguishable grossly from the esophagus Divided into two parts: 1. PROVENTRICULUS bears gastric glands 2. GIZZARD with thick muscular wall ad hard cornified lining; grinding mill; contains gravel

Amphibians 9. LESSER OMENTUM liver to lesser curvature 10. GREATER OMENTUM portion of the dorsal mesentery attached to the greater curvature of the stomach covers the abdominal viscera like an apron in humans 11. LESSER PERITONEAL CAVITY portion of the peritoneal cavity enclosed within the greater omentum 12. EPIPLOIC FORAMEN opening between peritoneal cavities Crocodilians and birds

Ruminants

1. RUMEN fermentation chamber; site of anaerobic bacteria adding cellulose 2. RETICULUM digests food material via peristalsis; chewing the cud 3. OMASUM temporary holding chamber 4. ABOMASUM true glandular stomach

lesser

and

greater

13. CHIEF CELLS zymogenic cells that synthesize and release the preenzyme PEPSINOGEN 14. PARIETAL CELLS secrete HCl which splits the active enzyme PEPSIN (stomach protease that initiates the process of protein digestion that eventually yields absorbable amino acids) C. FUNCTIONS OF THE STOMACH 1. Absorption of H2O and X-OH 2. Bolus Chyme 3. Serves as storage and macerating site for ingested solids via vertical peristalsis (churning) 4. Secretes digestive enzymes

VII. INTESTINE A. FISHES 1. COMPARATIVE ANATOMY the intestine of living agnathans, chondrichthyes and bony basal fishes is quite straight 2. RELATED STRUCTURES SPIRAL INTESTINE present in many fishes other than in teleosts so named because a SPIRAL VALVE, or TYPHLOSOLE, is suspended within its lumen increases the epithelial area available for absorption

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(b) PANCREATIC JUICES o amylase that acts on carbohydrates o lipase that digests lipids yielding absorbable fatty acids and glycerol o proteolytic enzymes that continue the digestion initiated by pepsin The small intestine is the chief site of digestion and absorption of nutrients in the presence of intestinal juice and pancreatic enzymes. 2. LARGE INTESTINE (COLON) 3 segments: (a) ASCENDING (b) TRANSVERSE (c) DESCENDING o ends in a sigmoid flexure in humans RECTUM a straight terminal portion in the pelvic cavity ILEOCOLIC CECA ceca beyond the ileocolic sphincter two in birds may exceed the large intestine in capacity in herbivorous mammals abbreviated in carnivores and humans APPENDIX terminates the cecum in anthropoids, rodents and many mammals no digestive function Most of the water is absorbed in the colon. The large intestine is rarely coiled, but ceca are common. VIII. LIVER AND GALLBLADDER A. LIVER 1. LIVER BUD where the liver arises becomes anchored to the embryonic transverse septum by a CORONARY LIGAMENT 2. DUCTS COMMON BILE DUCT empties into the duodenum composed of: o HEPATIC DUCTS liver o CYSTIC DUCT gallbladder Page 8 of 10

POSTVALVULAR INTESTINE portion of the intestine posterior to the spiral intestine in fishes PYLORIC CECUM intestinal diverticulum near the pylorus in teleosts RECTAL GLAND chloride-secreting (extracts and excretes excess NaCl from the blood) gland emptying into the caudal intestine of elasmobranchs no digestive function Intestinal ceca are the major adaptations for increasing the absorptive area of the teleost intestine. B. TETRAPODS 1. SMALL INTESTINE 3 segments: (a) DUODENUM o 50% absorption o where the pancreatic empties (b) JEJUNUM (c) ILEUM o with PEYERS PATCHES, aggregation of lymphoid masses VILLI fingerlike projections of the small intestine increase the absorptive surface of the intestine LACTEAL dead-end lymph vessels draining lipid laden lymph from the intestine CHYLE lymphatic fluid milky in color due to lipid content ILEOCOLIC SPHINCTER small intestine of amniotes terminate in it; where the colon commences regulates the ejection of the contents of the ileum into the large intestine DIGESTIVE JUICES (a) INTESTINAL JUICES o secreted by glands in the epithelial lining of the crypts and by compound alveolar glands near the pylorus

duct

Comparative Anatomy of the Vertebrates:

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Digestive System]
The liver removes dietary amino acids from the hepatic portal system and deaminates them. Several blood proteins are manufactured in the liver, including FIBRINOGEN and PROTHROMBIN, which are essential for blood clotting. The liver is elongated in eel-like bodies. It is short and wide in short trunks as in humans. B. GALLBLADDER 1. COMPARATIVE ANATOMY a gallbladder develops in most craniates. None develop in lampreys, a few teleosts, many birds, perissodactyls, whales and some rodents, inclusing rats craniates lacking a gallbladder are among those that have a little fat in their diet 2. FUNCTIONS OF THE GALLBLADDER stores and concentrates (by removal of H2O) bile human beings can live without a gallbladder, but they must be careful about the amount of lipids they consume IX. PANCREAS Its location is referred to as retroperitoneal. It varies from diffuse to compact. When diffuse, as in teleosts and many other craniates, pancreatic tissue is distributed along the blood vessels in the ventral mesentery of the stomach and duodenum. When compact, it may consist of several discrete lobes. A. EXOCRINE PANCREAS produces digestive enzymes in ALVEOLI (ACINI) The enzymes are transported via PANCREATIC DUCTS to the duodenum. B. ENDOCRINE PANCREAS bears PANCREATIC ISLETS (ISLETS OF LANGERHANS) lacks ducts secretes 2 hormonal products (into the bloodstream): (a) INSULIN o decreases blood glucose levels o necessary for the use of glucose by the body cells

AMPULLA (PAPILLA) OF VATER terminal segment of the bile duct in the duodenum 3. LIGAMENTS LESSER OMENTUM serves as a bridge that conducts the common bile duct to the duodenum and hepatic artery and hepatic portal vein to the liver composed of: o HEPATODUODENAL LIGAMENT remnant of the ventral mesentery connecting the liver and the duodenum o GASTROHEPATIC LIGAMENT remnant of the ventral mesentery connecting the pyloric region of the stomach and liver CORONARY LIGAMENT attaches liver to the central tendon of the diaphragm FALCIFORM LIGAMENT remnant of the embryonic mesentery ventral to the liver in adults extends from between the two median lobes of the liver to the median ventral line 4. FUNCTIONS OF THE LIVER It produces BILE which emulsifies lipids in the small intestine and imparts to them an alkalinity necessary for digestion. Some liver cells phagocytose aging red blood cells, splitting the hemoglobin molecule and freeing the iron. The remainder of the molecule is converted into BILIRUBIN (red pigment) and BILIVERDIN (green pigment) that are excreted as part of the bile. The fetal liver is a source of red blood cells. The liver removes from the circulation glucose in excess of immediate tissue needs. The glucose is converted to glucose-6-phosphate and then stored in the liver as GLYCOGEN.

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(b) GLUCAGON o antagonist of insulin by breaking down glycogen in the liver In living agnathans, there is no definitive pancreas. C. RELATED STRUCTURES/TERMS 1. PANCREATIC BUDS where pancreas arises 2. TAIL dorsal lobe from the dorsal bud 3. BODY OF THE PANCREAS ventral lobe from the ventral buds 4. ACCESSORY PANCREATIC DUCT smaller of two pancreatic ducts seen in cats and humans In sharks, the entire pancreas develops from the dorsal bud. In most mammals, the pancreas develops from one ventral and one dorsal bud. X. CLOACA common passageway for: (a) gametes (b) urine and other nitrogenous wastes (c) fecal components ANUS separate tracts In many fishes and in most tetrapods other than therian mammals, the digestive tract terminates in a common chamber, the cloaca, into which the urinary and genital tracts empty. In therian mammals, the cloaca becomes partitioned during embryonic development into two or three separate passageways, one of which becomes the rectum leading to an anus.

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