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Objective
1. After this study, the students can distinguish and determine the digestivus anatomy among
vertebrates.
Absorption
Synthesis - true protein, FA, starch, vitamins Excretion elimination of waste products
Foregut functions
Ingestion and storage of feeds Mechanical, chemical & enzymatic digestion of feed Nutrient absorption
Midgut functions
Hindgut functions
2. Associated Structures
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
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
Cattle, sheep, deer, elk, bison Ability to chew cud at frequent intervals distinguishes true ruminant from other foregut fermenters
Pregastric fermentation
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
Properties
Seasonally adaptive
Feeding preference
Prefer browsing
Prefer grazing
Properties
Most recently evolved Larger rumens and longer retention times Less selective Digests fermentable cell wall carbohydrates Fresh grass grazers
Classes
Roughage grazers
Structures in Mouth
Mouth
Functions
Prehension
Mastication or chewing
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
Primates, raccoon Elephant, tapir Anteater, cow, sheep Horse, sheep, rhinoceros
Snout
Tongue
Lips
Prehension
lips when eating from manger teeth when grazing Use long rough tongue to grasp forage
Pigs use snout to root in ground and pointed lower lip to convey feed into mouth
Most mammals use suction Dogs and cats use tongue to form ladle
Black Rhino
Prehensile upper lip for browsing Consumes bushes and shrubs in forest
Mastication
Relative toothlessness
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:
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
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
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:
Moves feed to teeth for grinding and to the back of the mouth for swallowing
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
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
Increased tone = Increased flow Increased flow = Increased dilution Increased tone = Decreased flow Decreased flow = Increased concentration
Submaxillary, sublingual, parotid (50% of secretions) No digestive enzymes in the saliva of mature
ruminants
Provides N, P, S and Na for rumen microoganisms Buffering compounds to maintain rumen pH and mucin to prevent bloat
Salivation
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
Dog:
Striated muscles throughout allow GREAT control of digesta movement both directions
Ruminant Esophagus
Funnel shaped
Esophagus
Species adaptations
Reverse peristalsis Amount and location of skeletal muscle Regurgitation vs. vomiting
Foregut in Birds
Crop
Birds
with crop gorge when food is available, store it in crop, and slowly digest it later
Stomach
Monogastric
One compartment
Ruminant
Four compartments
Gastric Digestion
Functions
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
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
Pepsinogen
Mucus
Protects lining from acid and enzymes No autodigestion Lubricant
Rennin (abomasum)
Lipase
Some species
acid pulp
Cephalic Phase
Vagal reflex
Parasympathetic innervation Increases gastric motility, enzyme secretion Small increase in HCl secretion
Gastric Phase
Intestinal Phase
Stimulated by duodenal distension, pH, osmolarity, nutrients (fat) Cholecystokinin (CCK) is released by the small intestine
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
Origin: Duodenum Stimulus: Fats and bile Function: Inhibit stomach motility and secretion of acid and enzymes
Ruminant Stomach
Anatomy:
Reticulorumen
Although
structurally they
Reticulum
Honeycomb lining
No secretions
Formation of food bolus Regurgitation initiated here Collects hardware (nails, wire)
Rumen
Also protozoa, fungi Produce VFA, protein Increase surface area Passive diffusion
Papillae lining
Absorption of VFA
Omasum
Laminae/manyply lining
~60% removed ~2/3 of VFAs entering or 10% of total produced Prevents buffering of the abomasum
Absorption of VFAs
Abomasum
Denatures proteins Kills bacteria and pathogens Dissolves minerals Gastric digestion
Small Intestine
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
Goblet cell
Secretes mucus
Endocrine cell
Secrete hormones into bloodstream or local cells Secretory granules with anti-microbial properties
Paneth cell
Villi Enterocyte Brush border Cell migration from crypts to tips of villus
2-3 days
Intestinal Wall
Villi
Mucosa
Villi
10x
20x
Generally, most absorption occurs in the proximal (upper) part of the small intestine but some absorption occurs in all segments
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
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
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
Brunners glands
Maltase Lactase
Sucrose
Sucrase
SI
Proteins
Polypeptides
Pepsin
Trypsin Chymotrypsin Carboxypeptidase Aminopeptidase
Gastric mucosa
Pancreas Pancreas Pancreas Small intestine
Polypeptide
Peptides Peptides Peptides & amino acids
Peptides
Bile
Bile salts (glycocholic and taurocholic acids) Bile pigments (bilirubin and biliverdin) Cholesterol
Small
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
Enterokinase
Secreted from crypts in duodenum Trypsinogen trypsin Trypsinogen trypsin Chymotrypsinogen chymotrypsin Procarboxypeptidase carboxypeptidase
Stomach
Pancreas
Similar in structure and function to monogastric Differences are subtle but important
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
Better for peptic activity May limit pancreatic protease and amylolytic activity
Pancreatic Secretions
Nucleases
Concentration of enzymes in pancreatic juice comparable to monogastrics Activity is lower and may be affected by:
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
Function
Absorption of remaining water, volatile fatty acids (VFAs) from microbial fermentation and minerals Digesta storage
Monogastric Cecum
Located at junction of small and large intestine Function similar to rumen in ruminants
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
Function:
Absorption of liquid Mass movements move fecal matter to anus Usually only a few times a day
Bacteria
Cellulolytic digest cellulose (forages) Amylolytic digest starches and sugars (concentrates or grains) Other types:
Proteolytic
Clostridium
Fermentative digestion
Bacteria similar to rumen, but no protozoa Digestion in colon may account for as much as:
Only important in conditions that increase the amount of fermentative carbohydrate entering the large intestine
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
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:
Regulated by:
Increased by increasing N concentration of diet Decreased by increasing the amount of carbohydrate fermented in the large intestine
Rectum
Muscular area of large intestine used for storage of feces and ultimately for defecation
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
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
Large Intestine
Rule Size
= Function
Young animals produce little sucrase, maltase, amylase Ruminants produce no sucrase Adult pigs lack lactase Activity changes with age
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
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
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
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
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
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
dogs, cats komodo dragon, tigers, eagles, sharks, polar bear cattle, sheep, goats, horses giant panda, gorilla, elephant, ostrich, green iguana, giraffe, American bison
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
OR
Monogastric Animals
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
Carnivores
Omnivores
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
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
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
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Stomach (2 gal)
Mouth
Cecum (10, 0.5 gal) Small intestine (60, 2.5 gal)
Rat
Sacculated stomach,capable of utilization of foragebased (high fiber) diets Able to digest some of the substances in grains, fruits and vegetables
Simple stomach incapable of utilization of foragebased (high fiber) diets Extensive fermentation after primary sites of digestion and absorption
Horse
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Small Colon (12, 3 gal) Small intestine (70, 12 gal)
Esophagus
Mouth
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)
GIT Classifications
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
Avians (Poultry)
Esophagus
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 contractions every 20-30 seconds Includes action of grit HCl and pepsin secreted in proventriculus
Small intestine
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
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
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