Comparative Digestive on Vertebrates

Objective
1. After this study, the students can distinguish and determine the digestivus anatomy among

vertebrates.

Why Do Animals Digest?

Food not ingested in suitable state

Physical nature of food determined by:

gathering apparatus for uptake type of digestive system

Primary Functions of the Digestive Tract

Transport food peristaltic contractions Digestion

Mechanical breakdown Chemical breakdown Passive diffusion and active transport

Absorption

Synthesis - true protein, FA, starch, vitamins Excretion elimination of waste products

Via bile (toxins, microbes etc) Via rectum (Ca, Mg, P)

1. Regions of Alimentary Canal

Foregut functions

Ingestion and storage of feeds Mechanical, chemical & enzymatic digestion of feed Nutrient absorption

Midgut functions

Hindgut functions

Water & ion re-absorption Formation, storage, excretion of feces

2. Associated Structures

Pancreas Contribute to small Liver intestinal digestion Gallbladder Salivary glands

Primitive Gastrointestinal Tract

Found in monotremes (egg-laying mammals), insectivores (bats, shrews, moles), and dermopterans (colugos)

Simple stomach, little or no division between small intestines and large intestines, large intestine simple, presence of cecum, non-sacculated colon

Species-Dependent Nutritional Adaptations

Includes involvement of:

Teeth Jaws and jaw musculature Alimentary canal

Stomach - May be simple or become sacculated to compartmentalize functions for prolonged storage of feed and utilization of bacterial fermentation (langurs and ruminants)

May also become voluminous for storage of large amounts of feed (vampire bats)

Large intestine - varies substantially in length, compartmentalization, and complexity among species

Ruminants

2.8 billion domesticated ruminants

Cattle, sheep, deer, elk, bison Ability to chew cud at frequent intervals distinguishes true ruminant from other foregut fermenters

Pregastric fermentation

Kangaroo, colobine monkey are not true ruminants

Four compartment stomach

Reticulum Rumen Omasum Abomasum

Ruminants vary in size and habitat

Classification of Ruminants by Feeding Preference

Classes of ruminants Concentrate selectors Intermediate feeders Roughage grazers

Concentrate Selecting Species

Properties

Evolved early Small rumens Poorly developed omasums Large livers Limited ability to digest fiber Fruit and forage selectors

Classes

Very selective feeders Duikers, sunis Eat highly lignified plant tissues to extract cell solubles Deer, giraffes, kudus

Tree and shrub browsers

Intermediate Feeding Species

Properties

Seasonally adaptive

Feeding preference

Prefer browsing

Moose, goats, elands Sheep, impalas

Prefer grazing

Roughage Grazing Species

Properties

Most recently evolved Larger rumens and longer retention times Less selective Digests fermentable cell wall carbohydrates Fresh grass grazers

Classes

Buffalo, cattle, gnus Hartebeests, topis Camels, antelope, oryxes

Roughage grazers

Dry region grazers

Structures in Mouth

Lips Teeth Tongue Salivary glands

Mouth

Functions

Grasp food Taste Masticate food Mix with saliva

Digestion in the Mouth

Prehension

Bringing the food to the mouth

Mastication or chewing

Upper limbs, head, beak, claws, mouth, teeth and lips

To crush the food, increase surface area and allow enzymes to act on molecules

Carnivores only to reduce the size of the particle to a size small enough to swallow Herbivores must chew continuously (4050,000 times a day) to increase surface area

Prehension

Seizing and conveying feed to mouth

Mechanisms vary with behavior and diet Forelimbs

Primates, raccoon Elephant, tapir Anteater, cow, sheep Horse, sheep, rhinoceros

Snout

Tongue

Lips

Prehension

Domestic mammals use lips, teeth and tongue

Relative importance varies by species Horses

lips when eating from manger teeth when grazing Use long rough tongue to grasp forage

Cows and sheep have limited use of lips

Pigs use snout to root in ground and pointed lower lip to convey feed into mouth

Birds use beak and tongue Drinking varies as well

Most mammals use suction Dogs and cats use tongue to form ladle

The Importance of Prehension in Diet Formulation

White Rhino (wijd = wide)

Black Rhino

Squared off upper lip used to crop grass Grazes on savannah

Prehensile upper lip for browsing Consumes bushes and shrubs in forest

Mastication

Physical reduction of feed Especially important in non-ruminant herbivores

Adaptations with teeth

Carnivores Herbivores Edentates (sloths, armadilloes, anteater)

Relative toothlessness

Morphological Adaptations for Herbivory

All related to finding, ingesting, masticating, and digesting plant cell walls Dental adaptations for herbivory include changes to incisors, molar occlusal surfaces, & masseter Solution for digestive problems is to provide a place in digestive tract for anaerobic bacteria & protozoans (microflora) to colonize

Monogastric Teeth

Function:

Mechanically reduce particle size Increase surface area Incisors are used for cutting Canine (fangs, eye teeth, tusks) are tearing teeth Premolars and molars (cheek teeth) grind the food

Four types:

Ruminant Mouth - Teeth

Function:

Reduce particle size Upper dental pad Lower incisors Premolars Molars

Anatomy:

Teeth Specializations

Carnivores

Canine teeth highly developed and used for tearing Molars are pointed for bone crushing

Teeth Specializations

Omnivores

Grinding teeth patterns on posterior teeth (molars) Piercing and ripping cusps on anterior teeth (incisors) Tongue - used to move feed to teeth

Jaw & Teeth Specializations

Non-ruminant herbivores (horse)

Incisors for nipping, molars slightly angled, jaws move circularly (vertical and lateral) No upper incisors, have dental pad, molars allow only lateral movements Different classes - roughage eaters, transition types, selective eaters all differ in tongue mobility and cleft palate

Ruminants

Ruminant Mouth

Lips range from short, relatively immobile in nonselective grazing species to very mobile (prehensile) in selective grazing or concentrate selecting species Chew in a lateral (grinding) motion on one side of mouth at a time

Needed to increase surface area of feed particles Feed chewed primarily during rumination in grazing species

Jaw Muscles and Mastication

Temporalis muscle - develops maximum force on anterior portion of jaw (largest muscle in carnivores and smallest muscle in herbivores) Masseter and medial pterogoid - maximum force for crushing and grinding Lateral pterogoid - allows lateral movement which is important for grinding (highly important in herbivores, but carnivores and many omnivores have almost no lateral movement of jaws)

Monogastric Tongue
Function:

Comprised of three muscles Maneuvers food in the mouth

Moves feed to teeth for grinding and to the back of the mouth for swallowing

Can distinguish between feed and toxins by papillae or taste buds

Ruminant Mouth - Tongue

Drinking, chewing and forming boluses Prehension of feed

Covered with rough, hook-like papillae that assist in grasping feed Important in nonselective grazing species More numerous than monogastric species More numerous on nonselective grazing species Believed that taste is primarily used for food avoidance by grazing species while concentrate selecting species select on the basis of smell

Taste buds

Monogastric Salivary Glands

Types of Glands:
Zygomatic Parotid Sublingual Mandibular

Functions of Saliva

Moisten feed (salt and water) Lubrication (aids swallowing) Starch and(or) lipid digestion (amylase and(or) lipase)

Salivary Glands
Gland Parotid Submaxillary Type of secretion Serous Mucous or mixed Main constituents Water, enzymes, ions Mucin (mucous), mucin plus enzymes (mixed), water

Sublingual

Mucous or mixed

Mucin (mucous), mucin plus enzymes (mixed), water

Monogastric Salivary Glands

Flow rate affected by:

Parasympathetic nervous system

Increased tone = Increased flow Increased flow = Increased dilution Increased tone = Decreased flow Decreased flow = Increased concentration

Sympathetic nervous system

Ruminant Mouth - Saliva

From at least three paired glands

Submaxillary, sublingual, parotid (50% of secretions) No digestive enzymes in the saliva of mature

Aids in mastication, swallowing, forming bolus

ruminants

Provides N, P, S and Na for rumen microoganisms Buffering compounds to maintain rumen pH and mucin to prevent bloat

Salivation

Quantity and composition of saliva varies considerably between species

Quantity related to level of chewing activity Dogs Sheep Horse Cattle minimal (lubrication, no enzymes) 3-10 liters/d 10-12 liters/d 130-180 liters/d

Amount of secretion

Deglutition (Swallowing)

Reflex initiated by presence of food in pharnyx Propulsion of food to stomach by esophageal peristalsis

Monogastric Esophagus

Horse/Pig:

Striated muscles for first 2/3 Smooth muscles for last 1/3 In horse, esophagus joins stomach at an oblique angle and cardiac sphincter (the valve between the stomach and esophagus) only allows one-way flow

MOST horses cannot belch out gas or vomit

Dog:

Striated muscles throughout allow GREAT control of digesta movement both directions

Ruminant Esophagus

Involved in rumination Different from monogastric esophagus

Striated muscle along the entire length

Provides greater strength Allows some voluntary control

Funnel shaped

Contains three sphincters active in rumination and eructation

Esophagus

Species adaptations

Ability to control peristaltic contractions

Reverse peristalsis Amount and location of skeletal muscle Regurgitation vs. vomiting

Foregut in Birds

Crop

Bottom of the esophagus forms a sac called crop

Stores undigested food.

Birds

with crop gorge when food is available, store it in crop, and slowly digest it later

Stomach

Monogastric

One compartment

Varies in size by species

Ruminant

Four compartments

Reticulum Rumen Omasum Abomasum

Gastric Digestion

Functions

Reservoir for controlled release of digesta to small intestine

Horse has small capacity requires increased number of smaller sized meals

Mixing food Mechanical breakdown of feed Hydrolytic digestion by acid and enzymes

Mainly protein

Kill bacteria Secrete intrinsic factor: needed for vitamin B12 absorption Hormone production

Stomach Regions

Esophageal

Non-glandular Secretes mucus Parietal cells Chief cells Mucus

Cardiac

Fundic

Pyloric

Gastric Pits

Formed by numerous folds in the epithelium Glands empty into the gastric pit Many types of glands may empty into one gastric pit

Gastric Glands
Gland Cardia Pylorus or Antrum Fundus Chief cells Parietal cells Type of secretion Mucous Mucous Enzyme Acid Main constituents Mucin Mucin Pepsinogen Pepsin HCl, intrinsic factor
acid

Stomach Secretions

HCl

Decreases pH (~2-3) Denatures protein Kills bacteria Activates pepsinogen

Pepsinogen

Activated form is pepsin Hydrolyzes protein Clots milk

Mucus
Protects lining from acid and enzymes No autodigestion Lubricant

Rennin (abomasum)

Lipase

Some species

Gastric Motility and Emptying

Motility aids mixing, mechanical and hydrolytic reduction of feed to chyme

acid pulp

Emptying is stimulated by distension of antral wall and presence of liquid chyme

Control of Gastric Secretions and Gastric Motility

Cephalic phase Gastric phase Intestinal phase

Cephalic Phase

Vagal reflex

Parasympathetic innervation Increases gastric motility, enzyme secretion Small increase in HCl secretion

Gastric Phase

Local reflex, depends on presence of feed in stomach Mainly mediated by gastrin

Increases HCl secretion

Intestinal Phase

Stimulated by duodenal distension, pH, osmolarity, nutrients (fat) Cholecystokinin (CCK) is released by the small intestine

Decreases HCl secretion and gastric motility

Gastrointestinal Hormones

Gastrin

Origin: Stomach, Abomasum Stimulus: Food in stomach Function: Stimulates HCl & pepsinogen secretion, increases stomach motility Origin: Duodenum Stimulus: Acid Function: Stimulates pancreatic secretions. Slows stomach motility and acid production

Secretin

Gastrointestinal Hormones

Cholecystokinin (CCK)

Origin: Duodenum Stimulus: Fat & protein in duodenum Function: Stimulates bile and pancreatic secretions

Also regulates appetite and feed intake

Gastric Inhibitory Protein (GIP)

Origin: Duodenum Stimulus: Fats and bile Function: Inhibit stomach motility and secretion of acid and enzymes

Ruminant Stomach
Anatomy:

Reticulum Rumen Omasum Abomasum

Reticulorumen
Although

structurally they

appear as a single continuous compartment, functionally they are distinctly different

Reticulum

Honeycomb lining

No secretions

Formation of food bolus Regurgitation initiated here Collects hardware (nails, wire)

Rumen

Digestion and fermentation vat

40-50 gallons No secretions

Contains anaerobic microbes (25-50 billion bacteria/mL fluid)

Also protozoa, fungi Produce VFA, protein Increase surface area Passive diffusion

Papillae lining

Absorption of VFA

Omasum

Laminae/manyply lining

Muscular folds No secretions

Reduces particle size Absorption of water

~60% removed ~2/3 of VFAs entering or 10% of total produced Prevents buffering of the abomasum

Absorption of VFAs

Abomasum

True gastric stomach - four gallons in a cow

Three regions (cardiac, fundic, and pyloric) Digestive secretions

Proteolytic enzymes and HCl

pH decreases from 6 to 2.5

Denatures proteins Kills bacteria and pathogens Dissolves minerals Gastric digestion

Small Intestine

Composed of 3 segments (proximal to distal)

Duodenum

Releases bile and pancreatic secretions Active site of digestion Active site of nutrient absorption Active site of nutrient absorption Most water, vitamins & minerals Some bacterial presence Fermentation

Jejunum

Ileum

The pH of the small intestine increases towards 7.0 as

food moves from the duodenum to the ileum

Intestinal Epithelial Cell

Brush border

Specialized Cells Lining Villi

Nutrients Mucus

Absorptive epithelial cell

Contain brush border on lumen/apical side Brush border:

Enzymes Nutrient transport molecules

Goblet cell

Secretes mucus

Specialized Cells Lining Villi

Anti-microbial compounds

Endocrine cell

Secrete hormones into bloodstream or local cells Secretory granules with anti-microbial properties

Paneth cell

CCK, Secretin, etc.

Small Intestine Absorptive Surface

Villi Enterocyte Brush border Cell migration from crypts to tips of villus

2-3 days

Small Intestine - Structure

Lumen Mucosa Villi Crypts Lacteal Enterocyte Brush border

Intestinal Wall
Villi
Mucosa

Enhanced Surface Area for Increased Nutrient Absorption

Intestinal villi

Increased Surface Area in Small Intestine for Absorption

Structure Plicae circularis Description Regular ridges in small intestine Finger-like projections on mucosal (inner) surface 1 um projections on surface of epithelium Increase in surface area 3x

Villi

10x

Microvilli Brush Border

20x

Nutrient Absorption in the Small Intestine

Principal site of absorption of amino acids, vitamins, minerals and lipids

Glucose and other sugars in monogastrics

Generally, most absorption occurs in the proximal (upper) part of the small intestine but some absorption occurs in all segments

Duodenum, jejunum and ileum Within 30 minutes of entering SI

Digestion and absorption within SI is rapid

Nutrient Absorption

Variety of mechanisms

Diffusion Facilitated diffusion Active transport Pinocytosis or endocytosis Solubility of the nutrient (fat vs. water) Concentration or electrical gradient Size of the molecule to be absorbed

Dependent upon

Diffusion

Water and small lipid molecules pass freely through membrane Move down concentration gradient to equalize concentrations

Facilitated Diffusion
1) Carrier loads particle on outside of cell 2) Carrier releases particle on inside of cell 3) Reverse
Allows equalization of concentrations across membrane

Active Transport
1) Carrier loads particle on outside of cell 2) Carrier releases particle on inside of cell 3) Carrier returns to outside to pick up another particle

Active Transport

Unidirectional movement Transports nutrients against concentration gradient

Pinocytosis or Endocytosis

Substance contacts cell membrane Membrane wraps around or engulfs substance into sac Sac formed separates from the membrane and moves into cell

Transporters

Secretions Entering SI

Intestinal mucus Brush border enzymes Pancreatic juices

Secreted from within SI

Enters from ducts into SI

Produced & stored in pancreas Produced in liver Stored in gallbladder Horse has no gallbladder

Bile

Direct bile secretion into duodenum Cannot store bilecontinuous intake of food

Intestinal Mucus

Secreted by glands in wall of duodenum

Brunners glands

Acts as lubricant and buffer to protect duodenal wall

Primary Enzymes for Carbohydrates

Nutrient Starch, glycogen, dextrin Maltose Lactose Enzyme Amylase Origin Saliva & pancreas SI SI Product Maltose & Glucose Glucose Glucose & galactose Glucose & fructose

Maltase Lactase

Sucrose

Sucrase

SI

Primary Enzymes for Proteins

Nutrient Milk protein Enzyme Rennin Origin Gastric mucosa Product Curd

Proteins
Polypeptides

Pepsin
Trypsin Chymotrypsin Carboxypeptidase Aminopeptidase

Gastric mucosa
Pancreas Pancreas Pancreas Small intestine

Polypeptide
Peptides Peptides Peptides & amino acids

Peptides

Primary Enzymes for Lipids

Nutrient Lipids Enzyme Lipase & colipase Origin Pancreas Product Monoglycerides & free fatty acids

Bile

Green, viscous liquid

Alkaline ph (neutralize acidic chyme) Stored in gall bladder (except in horses)

Secreted by liver via bile duct to duodenum

Functions to emulsify fats Composition

Bile salts (glycocholic and taurocholic acids) Bile pigments (bilirubin and biliverdin) Cholesterol

95% reabsorbed and returned to liver NOT AN ENZYME

Nutrient Digestion - Lipids

Large Lipid Droplet

Small

Action of bile salts Lipid emulsion

Bile salts & pancreatic lipase and colipase Water soluble micelles

Pancreatic Juice

Clear, watery juice Enters duodenum via pancreatic duct Aids in fat, starch, and protein digestion

Contains

HCO3Trypsinogen ProChymotrypsinogen enzymes Procarboxypeptidase Amylase Lipase Nuclease

Importance of Pancreas for Digestion

Produces enzymes responsible for

50% of carbohydrate digestion 50% of protein digestion 90% of lipid digestion

Produces sodium bicarbonate for neutralization of chyme in duodenum

Activation of Pancreatic Enzymes

Enterokinase

Secreted from crypts in duodenum Trypsinogen trypsin Trypsinogen trypsin Chymotrypsinogen chymotrypsin Procarboxypeptidase carboxypeptidase

Trypsin then converts:

Overview of Digestive Enzymes

Stomach

Pepsinogen Chymosin (rennin)

Trypsinogen Chymotrypsinogen Procarboxypeptidase Amylase Lipase Nuclease

Brush Border (SI)

Pancreas

Sucrase Maltase Lactase Aminopeptidase Dipeptidase Enterokinase

Ruminant Small Intestine

Similar in structure and function to monogastric Differences are subtle but important

Limited ability to digest starches and sugars

Little to none presented except in exceptional circumstances (high-grain feeding)

Small Intestine
Digesta pH Functions Enzymes pH change Flow rate regulation Enzymes Absorption Absorption Limited fermentation

Duodenum

2.7 - 4

Jejunum
Ileum

47
7-8

Rate of pH increase through small intestine is slower than monogastrics

Better for peptic activity May limit pancreatic protease and amylolytic activity

Pancreatic Secretions

Secretion pH is 7.2-7.8 Enzymes

Amylase Lipase Proteases

Trypsinogen converted to trypsin Chymotrypsinogen converted to chymotrypsin Procarboxypeptidase converted to carboxypeptidase

Nucleases

Activity of Pancreatic Enzymes

Concentration of enzymes in pancreatic juice comparable to monogastrics Activity is lower and may be affected by:

Less juice secreted/kg BW Low digesta pH High rate of passage

Limited activity particularly a problem for intestinal digestion of starch escaping ruminal digestion

For ruminants fed high grain diets, less than 50% of starch reaching small intestine is digested

Bile

Secreted with pancreatic juice in the common bile duct of sheep Secreted in the bile duct of cattle

Large Intestine

Composed of three segments

Cecum Colon Rectum Fermentative digestion

Function

No enzyme secretion Relies on microbes or secretions washed out of the SI

Absorption of remaining water, volatile fatty acids (VFAs) from microbial fermentation and minerals Digesta storage

Degree of development is species dependent

Monogastric Cecum

Located at junction of small and large intestine Function similar to rumen in ruminants

Microbial activity and digestion of feeds

Since cecum is located AFTER major site of nutrient absorption (small intestine), then microbial cell proteins are not available to the animal

Contains a microbial population similar to the rumen Cellulolytic & hemicelluloytic bacteria

Fecal loss

Monogastric Large Intestine

Function:

Absorption of liquid Mass movements move fecal matter to anus Usually only a few times a day

Associated with defecation

Bacteria

Cellulolytic digest cellulose (forages) Amylolytic digest starches and sugars (concentrates or grains) Other types:

Proteolytic

Clostridium

Organic acid utilizers Methanogens

Produce CO2, H2, formate, CH4

Ruminant Large Intestine

Fermentative digestion

Bacteria similar to rumen, but no protozoa Digestion in colon may account for as much as:

27% of cellulose digestion 40% of hemicellulose digestion 10% of starch digestion

Only important in conditions that increase the amount of fermentative carbohydrate entering the large intestine

Increased rate of passage of forages High grain diets

May account for as much as 17% of total VFA absorption VFAs are efficiently absorbed, but primarily used as energy source for large intestinal mucosa cells

Ruminant Large Intestine

Absorption of ammonia-N

May account for as much as 30 to 40% of the net transport of N into body fluid Absorbed N may be used for:

Synthesis of nonessential amino acids Recycling of N to the rumen

Important on low protein diets

Regulated by:

Increased by increasing N concentration of diet Decreased by increasing the amount of carbohydrate fermented in the large intestine

Mineral absorption Water absorption

90% of water entering the LI is absorbed

Rectum

Muscular area of large intestine used for storage of feces and ultimately for defecation

Feces includes sloughed cells, undigested food and microbial matter

Digestive Adaptations to Varying Feed Sources

Gastric capacity and structure

Capacity is greatest in pregastric fermentors

Stomach acts as reservoir

Small stomach in carnivores is related to high nutrient density of the diet Distribution and composition of epithelial lining varies between species and dietary adaptations

Digestive Adaptations to Varying Feed Sources

Intestinal length and functions

Small intestine

Less variable among species than stomach and hind gut, but generally shorter in carnivores than in herbivores Importance of hind gut fermentation dictates variation in structure and size Some hind gut fermentation occurs in most species

Large intestine

Adaptations of the Digestive Tracts

Stomach Small Intestine Cecum

Large Intestine
Rule Size

= Function

Adaptations of Digestive Enzymes

Young animals produce little sucrase, maltase, amylase Ruminants produce no sucrase Adult pigs lack lactase Activity changes with age

Lactase Sucrase, maltase

Utilizing Cellulose

Advantages

Ultra-abundant in the environment Easily obtained no need to hunt plants Plant cell walls & fiber high in energy

Disadvantages

Indigestible by mammalian digestive enzymes Cellulase is found only in bacteria & some protozoans

Fermentative Digestion

All mammals have some fermentative capacity that allows for utilization of ingested fiber The comparative importance of fermentation is related to the fraction of total digesta contained in fermentative compartments of the gastrointestinal (GI) tract

Advantages of Pregastric Fermentation

Make better use of alternative nutrients

Cellulose Nonprotein nitrogen

Ability to detoxify some poisonous compounds

Oxalates, cyanide, alkaloids

More effective use of fermentation endproducts

Volatile fatty acids, microbial protein, B vitamins

Allows wild animals to eat and run

Disadvantages of Pregastric Fermentation

Fermentation is inefficient

Energy

Loss % of total caloric value Methane 5-8 Heat of fermentation 5-6 Relative efficiency is dependent on the diet NDF

Protein Some ammonia resulting from microbial degradation will be absorbed and excreted 20% of the nitrogen in microbes is in the form of nucleic acids

Disadvantages of Pregastric Fermentation

Ruminants are susceptible to ketosis Ruminants are susceptible to toxins produced by rumen microbes

Nitrates to nitrites Urea to ammonia Nonstructural carbohydrates to lactic acid Tryptophan to 3-methyl indole Isoflavonoid estrogens to estrogen coumestans

Pregastric Fermenters
Class Ruminants Species Cattle, sheep Deer, antelope, camel Nonruminants Colobine monkey, hamster, vole Dietary habit Grazing herbivores Selective herbivores, including folivores and frugivores Selective herbivores

Kangaroo, hippo
Hoatzin

Grazing and selective herbivores

Folivore

Postgastric Fermentors

Cecal fermentors

Mainly rodents, rabbits and other small herbivores Often associated with coprophagy

Colonic fermentors

Includes true herbivores (e.g., horse), omnivores (e.g., pig and human), and carnivores (e.g., cat and dog)

Horse has some expanded cecal fermentation in addition to greatly expanded colonic fermentation

Degree of colonic sacculation is related to importance of fiber digestion and fermentative capacity

Postgastric Fermenters
Class Cecal digesters Species Capybara Rabbit Rat, mouse Dietary habit Grazer Selective herbivores Omnivores Grazers Folivores Omnivores Herbivores Carnivore

Colonic digesters Sacculated Elephant, horse, zebra New World monkeys Pig, human Unsacculated Panda Dog, cat

Foregut vs. Hindgut Fermentation

Foregut

More efficient per unit volume of food Slower digestive process Animal may starve with a full belly Size restricted More efficient relative to time Faster turnover

Hindgut

Feeding Habits of Mammals & Taxonomic Distribution

Feeding Habit

Number of Orders
10 5 4 2 10 7

Percent of Species
40 4 12 <1 33 10

Herbivory Frugivory, Graniv., Nectivory Carnivory Planktonivory? (Krill feeders) Insectivory Omnivory

Classification of Animals by Preferred Ingested Feedstuffs

Carnivore consume animal products

dogs, cats komodo dragon, tigers, eagles, sharks, polar bear cattle, sheep, goats, horses giant panda, gorilla, elephant, ostrich, green iguana, giraffe, American bison

Herbivore consume plant products

Omnivore plant and(or) animal products

pigs opossum, raccoon, blue jays, black bear, human

Specialized Carnivores
Aphidivore - feeds on aphids Apivore - feeds on bees Erucivore - feeds on caterpillars Insectivore - feeds on insects Larvivore - feeds on larvae Myrmevore - feeds on ants Mucivore - feeds on flies Piscivore - feeds on fish Pupivore - feeds on pupae Ranivore - feeds on frogs Sanguivore - feeds on blood Zoosuccivore - feeds on liquid animal secretions of decaying animal matter

Specialized Herbivores
Ambivore - feeds on grasses and broad leaf plants Exudativore - feeds on gums, resins, and sap Folivore - feeds on foliage (leaves or trees) Forbivore - feeds on forbs (i.e. flowering plants) Frugivore - feeds on fruit Graminivore - feeds on grasses Granivore - feeds on grains Gumivore - feeds on gums secreted by some plants Lignivore - feeds on wood Mellivore - feeds on honey Nectarivore - feeds on nectar Nucivore - feeds on nuts (agouti) Phytisuccivore - feeds on (tree) sap Radicivore - feeds on roots

Classification by Type of Digestion or Site of Digestion

Monogastrics or non-ruminants Ruminants

OR

Pre-gastric fermentation (cranial) Post-gastric fermentation (caudal)

Monogastric Animals

Single, simple stomach structure Mostly carnivores and omnivores

Very simple - mink, cat and dog Cecal digestion - horse, rabbit, elephant or rat Sacculated stomach - kangaroo

Ruminant Animals

Ruminant herbivores possessing multiple digestive tract compartments for feed breakdown before feed reaches the true stomach

True ruminants - cattle, sheep, goats Pseudo-ruminants - camels, llamas, alpacas, vicunas

GIT Capacity - Volume

Carnivores

stomach (70%) > SI = LI (15%) GIT surface/body surface: 0.6:1

Omnivores

stomach = SI = LI (33%) GIT surface/body surface: intermediate

Ruminants

Herbivores

stomach (70%) > SI (20%) > LI (10%) GIT surface/body surface: 3:1
stomach (10%) < SI (30%) < LI (60%) GIT surface/body surface: 2:1

Non-ruminants

GIT Classifications - Dog

Monogastric carnivore with limited post-gastric fermentation

Simple stomach, not capable of effective utilization of forage-based (high fiber) diets Unable to digest some of the substances in grains, fruits and vegetables Similar to cat

GIT Classifications - Pig

Monogastric omnivore with limited post-gastric fermentation

Simple stomach, not capable of effective utilization of forage-based (high fiber) diets Unable to digest some of the substances in grains, fruits and vegetables Similar to human

Pig
_________________________________________
Stomach (2 gal)

Large Intestine (16, 2 gal) Esophagus

Mouth
Cecum (10, 0.5 gal) Small intestine (60, 2.5 gal)

Human Digestive Tract

Rat

GIT Classifications - Kangaroo

Monogastric omnivore with limited pre-gastric fermentation

Sacculated stomach,capable of utilization of foragebased (high fiber) diets Able to digest some of the substances in grains, fruits and vegetables

GIT Classifications - Horse

Monogastric herbivore with extensive postgastric fermentation

Simple stomach incapable of utilization of foragebased (high fiber) diets Extensive fermentation after primary sites of digestion and absorption

Horse
_________________________________________
Small Colon (12, 3 gal) Small intestine (70, 12 gal)

Esophagus

Large Colon (12, 19 gal) Cecum (4, 8 gal)

Mouth

Stomach (3.5 gal)

GIT Classifications - Sheep

Ruminant herbivore with extensive pre-gastric fermentation

Highly developed sacculated stomach capable of extensive and effective utilization of forage-based (high fiber) diets Extensive fermentation before primary sites of digestion and absorption Similar to cattle and goats

Cow
_________________________________________
Large intestine (33, 8 gal) Cecum (3, 3 gal) Esophagus Rumen (paunch) (43 gal)

Mouth Abomasum (glandular) (5 gal) Omasum (4 gal) Reticulum (honeycomb) (2.5 gal)

Small intestine (150, 16 gal)

GIT Classifications

Avian is modified monogastric

Beaks replace lips and teeth Crop (enlarged area of esophagus) stores and softens feed prior to entering stomach Proventriculus glandular stomach Gizzard muscular part of stomach Branched cecum postgastric fermentation Cloaca both fecal and urinary waste Uric acid rather than urea (insoluble)

Avians (Poultry)
Mouth

No teeth, rigid tongue Poorly developed salivary glands

Saliva contains amylase

Beak is adapted for prehension and mastication

Avians (Poultry)
Esophagus

Enlarged area called crop

Ingesta holding and moistening Location for breakdown of carbohydrate by amylase Fermentation Release of HCl and pepsin (gastric juices) Ingesta passes through very quickly (14 seconds)

Proventriculus (stomach)

Avians (Poultry)
Gizzard (ventriculus)

Muscular area with a hardened lining reduces particle size

Muscular contractions every 20-30 seconds Includes action of grit HCl and pepsin secreted in proventriculus

Small intestine

Similar to other monogastrics No Lacteals

Avians (Poultry)
Ceca and large intestine

Contain two ceca instead of one as in other monogastrics Large intestine is very short (2-4 in) and empties into cloaca where fecal material will be voided via the vent

Water resorption Fiber fermentation by bacteria H2O soluble vitamin synthesis by bacteria

Chicken

Beak

Esophagus Crop (2) Small Intestine (55) Proventriculus Gizzard (2) Pancreas

Ceca (7)
Large Intestine (4) Cloaca

Feeding Behaviors

Impact feed choices

Neophobia (avoidance of new feed sources)

Contact testing (based on taste and other sensory information collected in mouth) prior to swallowing Early introduction of a variety of feeds limits this problem

Chimpanzees select feeds based on easily digestible carbohydrate content (sugars and starches) rather than fat or protein content Grazers and browser select early growth grasses and plants vs. mature growth In confinement feeding situations, grazers consume concentrates first and then forages in ration based on particle size (basis for creating total mixed rations)

Feeding Behaviors

Impact feed intake (avoiding under- and over-consumption of feed)

Hiding feed in bear exhibit encourages search and gather behaviors, limiting intake and reducing the stress of captivity Predator behavior towards prey meals vs. bowl meals Grazing animals prefer to eat forage at ground level rather than in elevated feed bunks