Lecture 13, 14

Coelomic Cavities, Mouth and Digestive System

Kardong Chapter 13, Hildebrand Chapter 12

KK 13.1

Origin of the Coelom and Gut

Organs of the gut develop as evaginations of the gut into the mesenteries. The coelomic space is a split in the hypomere.

KK 13.2, H&G 12.1

The coelom or peritoneal cavity is undivided in hagfish, some sharks.

In most fishes and Amphibia there is a separate pericardial cavity just behind the gills. It is separated from the peritoneal cavity by the transverse septum.

In reptiles, which have a neck, the heart moves back under the lungs.
KK 5.34, H&G 11.2

In mammals, the lungs are in a separate pleural cavity. The new membrane separating the pleural and peritoneal cavities houses the diaphram. The lungs surround the heart in the adult and the membrane separating the pleural and pericardial cavities is the mediastinum.

KK 5.34, H&G 11.3

Origin of the Mouth

KK 13.4

The mouth develops de novo in deuterostomes. The invagination that meets the gut and creates the mouth is called the stomodeum. The nasal epithelium and the hypophyseal (Rathkes) pouch develop on the head. But, are drawn in by the deepening stomodeum (a). In cyclostomes (b) both remain on top of the head adjacent to the shallow stomodeum. In most fishes (c) only the hypophyseal pouch is drawn into the stomodeum. In vertebrates with choanae or internal nares (a,d) the mouth is deeper, and both the hypophyseal pouch and the nasal epithelium are drawn into the stomodeum. The nasal placode deepens to emerge on the top of the head creating the external nares.

Mouths in Sagittal Section KK 13.3, 13.37, H&G 12.2&12.3

Fish have no neck, their heart is in their pharyngeal region, and the opening to lungs (if present) is dorsal.

Tetrapods changes include salivary glands, and, especially in mammals, a much larger and more mobile tongue.

Specialization of Homodont Dentition

Teeth vary greatly among fish and ectothermic tetrapods, but generally all are the same within a species. Many have sharply pointed teeth to grasp prey. Chewing is minimal.

KK 13.13, H&G 7.5

Heterodont Dentition of Mammals

KK 13.7, H&G 7.7, 30.17

Most mammals and synapsids had differentiated teeth: incisors, canines, premolars and molars These teeth are usually replaced once. Premolars and molars are very similar, the difference being that molars are not replaced. They can be collectively referred to as cheek teeth. Carnassial teeth = molars specialized for shearing.

Adaptations for Herbivory

Herbivores may have fewer or no incisors and canines, but high and deeply folded cheek teeth. Why?

KK 13.14, H&G 30.22

Adaptations for tooth wear

Tooth wear limits the lifespan of many herbivores.
Rodents (e.g., beaver) have opening-rooted incisors that keep growing. Elephants use their 6 immense cheek teeth one at a time.

KK 13.9, H&G 30.20, 30.21

Gut Functions
Transportation (via peristalsis) and storage Physical treatment: mixing and churning Chemical treatment: breakdown of
macromolecules into subunits by acids and enzymes prior to assimilation. May also be biological treatment by microbes.

Absorption Production of feces, reclamation of H2O

Structure of the Gut

KK 13.22, H&G Fig. 12.4

The muscularis externa and the serosa are from the splanchnic hypomere. The mucosa and submucosa are derived from endoderm. Note that the ventral mesentery mostly disappears.

Structure of the small intestine

In the absorbtive area of the gut, i.e., the small intestine, the internal surface of the gut is intensely folded to increase the surface area. There is a rich blood supply. Note also lymph vessels (lacteals).
KK 13.25, H&G 12.4

Fish Guts KK 13.29, H&G 12.6

Fish (ectothermic predators) generally have short and simple guts. The esophagus is short and starts right behind the mouth. The large intestine is not distinct. The shark and some Osteichthyes have increased the area for absorbtion via a spiral valve in the intestine. Teleosts like the perch and trout have pyloric ceca to increase absorbtive area instead of a spiral valve. The small intestine may loop back on itself.

Note that the liver and pancreas (where present) are not shown.

Tetrapod Guts 1
Tetrapods have a more distinct large intestine, and most have a urinary bladder. They never have pyloric ceca or a spiral valve. Instead, the small intestine is long and coiled rather than a single loop.

Tetrapod Guts 2
Birds and many reptiles lack a urinary bladder. They both have a gizzard. Birds also have a crop.

KK 13.32, H&G 12.7

Tetrapod Guts 3

KK 12.27

Tetrapods increase absorbtive area by lengthening the small intestine.

Amphibia have a short esophagus, like fish.

Birds have a crop and a gizzard. They also have colic ceca that are enlarged in herbivorous species.

Functional variation in mammal guts

KK 13.28, H&G 12.8

Simple guts of carnivores and invertivores.

Complex guts of herbivores.

Foregut vs. Hindgut Fermentation

Foregut fermentation (artiodactyls, kangaroos, sloth, hippopotamus, ostrich) is ahead of the small intestine so absorbtion is favoured, and allows for re-chewing or chewing the cud in some species. But it is slow. Hindgut fermentation (horses, rodents, rabbits, rhinoceros) is below the small intestine involving the large intestine or the cecum, so absorbtion of products is limited but it allows faster throughput. Hindgut fermenters cannot regurgitate and re-chew, but many are coprophagous.

The Stomach
Vertebrate stomachs are rather similar. In birds (and some diapsids) the pylorus is particularly muscular, and called the gizzard. It often contains stones.
In ruminants, e.g., the cow, the foregut is expanded as an microbial fermentation chamber, allowing them to extract nutrition from coarse plant material. G

KK13.31, H&G 12.5

The Rumen
KK 12.34, H&G 12.9 The rumen is actually a part of the esophagus upstream from the true stomach. The rumen allows ruminant animals to extract nutrition from poor quality food via microbial fermentation, and to minimize their exposure to predators. They can consume large amounts of food quickly, move to cover, then regurgitate and re-chew it at their leisure. Methane gas production from ruminant animals is a significant contribution to atmospheric methane, a potent greenhouse gas.

Liver Functions
The liver is the largest organ in the body, and universal (and very similar) in all vertebrates.
Digestive: - produces bile acids, which emulsify fats in the digestive tract. Bile acids are stored in the gall bladder. Metabolic: - Carbohydrate metabolism regulates blood glucose by synthesizing and de-synthesizing glycogen. - Stores fat to various degrees (e.g., shark, cod). - Protein metabolism De-aminates amino acids and synthesizes urea with the waste N. Also: - stores iron and some vitamins - detoxifies toxins - phagocytosis of old blood cells, foreign cells - blood cell formation in fish and embryos

The Liver in Relation to the Circulatory System

Mesenteric arteries Absorption in Gut Capillaries Hepatic Portal Vein

Livers Metabolic Functions

Interlobular veins Capillaries Hepatic Artery Central Veins

Hepatic Vein

Heart

Liver Lobule in Transverse Section

KK 13.39, H&G 12.11

Liver functions occur as the blood from the hepatic portal vein passes through the liver lobules to collect in central veins and ultimately the hepatic veins.

Pancreas Functions
As an exocrine digestive gland (secretions to the outside, ie, the gut lumen) it produces proteolytic enzymes as zymogens or pro-enzymes. These are in an inactive state until worked on by proteolytic enzymes already in the gut.
As an endocrine gland (secretions into the body, i.e., the blood) it secretes insulin (stimulates deposition of glycogen) and glucagon (stimulates release of glycogen) While the liver is universal in vertebrates, the pancreas is not a discrete organ in most fish. Instead, tissue with pancreatic function is scattered in the mesentery adjacent to the liver.

The pancreas develops from separate structures in the dorsal and ventral mesenteries that fuse during development in many groups.

Development of the Liver and Pancreas

KK 13.38, H&G 12.1

The Pancreas
In many vertebrates, the dorsal and ventral pancreas are more or less separate and two ducts may persist, as in the panda. The ducts carry digestive enzymes to the gut.
KK 13.40

The Pancreas
In other vertebrates, including humans, the dorsal and ventral pancreas are intimately fused, and endocrine functions of the dorsal pancreas are carried out in the pancreatic islets. KK 15.11, H&G 12.12