Source: http://www.mhhe.com/biosci/genbio/rjbiology/ELOs/ELO45.html

Chapter 45 Outline

INTRODUCTION

	Vertebrates Contain Many Cells with Specialized Functions

	No Vertebrate Cells Are Specialized for Photosynthesis
		Plants are self-sustaining autotrophs
		Animals are heterotrophs
			Cells must be nourished by food obtained from outside the body
			Many major organ systems are associated with acquisition of food energy

THE NATURE OF DIGESTION

	Animals Obtain Energy by Degrading Chemical Bonds of Organic Molecules
		Process acts on simple molecules:  amino acids, lipids and sugars
		Organisms rarely contain large amounts of simple molecules
			Eating an organism does not provide immediate energy
			Simple molecules incorporated into long macromolecular chains
			Macromolecules include proteins, fats and starches
		First must degrade macromolecules into simple constituent parts	fig 45.1

	Process Called Digestion

ORGANIZATION OF VERTEBRATE DIGESTIVE SYSTEMS

	General Organization of the Vertebrate Digestive System
		Consists of a tubular gastrointestinal tract and accessory digestive organs	fig 45.2
			Initial components are mouth and pharynx
				Common passage of oral and nasal cavities
				Pharynx leads to esophagus
			Esophagus is a muscular tube leading to stomach
			Preliminary digestion occurs in stomach
			Food passes into duodenum, upper part of small intestine
				Battery of digestive enzymes continue digestion
				Products pass across small intestine wall into bloodstream
		Tubular gastrointestinal tract has a layered structure	fig 45.3
			Mucosa is innermost layer
				Epithelial layer
				Separates interior (lumen) from blood vessels in next layer
			Submucosa is next layer of connective tissue
			Next outermost layer is the muscularis
				Double layer of smooth muscle
				Inner muscles have circular orientation
				Outer layer are arranged longitudinally
			Serosa connective tissue layer covers external surface
			Nerve plexuses regulate activities of gastrointestinal tract

	Specializations of Digestive Systems Indicate Different Ways of Living
		Fish have large pharynx with gill slits
		Air-breathing vertebrates have reduced pharynx
		Many vertebrates have teeth and chew food particles
		Birds lack teeth, break up food in  two-chambered stomach
			Gizzard grinds material with small pebbles
			Seeds and hard materials ground up for digestion in second chamber
		Carnivores have shorter intestines than herbivores
			Most animal macromolecules are readily digested
			Herbivores eat cellulose, have convoluted intestines to prolong digestion
			Herbivorous mammals have multiple-chambered stomachs with cellulose-degrading bacteria

FOOD ENTERS THE DIGESTIVE TRACT THROUGH THE MOUTH

	Teeth Are Important to Animal Digestion
		Capture food in different ways, teeth specialized for such capture
		Carnivores possess pointed teeth for capture, cutting and shearing
		Herbivores have large, flat teeth suited for grinding plant materials
		Omnivores have both types, front like carnivores, back like herbivores	fig 45.5
			Incisors:  four front teeth, used for biting
			Canines:  one on each side of incisors, used for tearing food
			Premolars:  two on either side behind canines, chewing teeth
			Molars:  three on either side behind canines, chewing teeth

	Food Is Moistened and Lubricated in the Mouth
		Tongue mixes food with saliva
			Saliva secreted by three pairs of salivary glands
			Empty through mucosal lining of mouth
			Contains salivary amylase to initiate breakdown of starch
		Secretion of saliva controlled by the nervous system
			Continuous secretion to keep the mouth moist
			Secretion stimulated by presence of food

FOOD PASSES TO THE STOMACH THROUGH THE ESOPHAGUS

	Food Passes Beyond the Teeth to the Back of the Mouth
		Palate elevates, pushes against back wall of pharynx	fig 45.6
			Seals off nasal cavity
			Prevents entry of food into nasal cavity
		Pressure on pharynx stimulates receptors to signal swallowing center
		Swallowing center signals respiratory tract
			Inhibits respiration
			Seals trachea by raising larynx and closing glottis with epiglottis

	Food Enters Esophagus Connecting Pharynx and Stomach
		Upper portion of esophagus enveloped in skeletal muscle
		Lower two-thirds enveloped in smooth muscle
		Food propelled to stomach by peristaltic waves
		Exit of food from esophagus to stomach controlled by a sphincter
			Muscular constriction at junction of two organs
			Prevents food in stomach from re-entering esophagus
			Rodents and horses have true sphincter cannot regurgitate, humans can

PRELIMINARY DIGESTION OCCURS IN THE STOMACH

	Stomach Is a Saclike Portion of the Digestive Tract	fig 45.7
		Interior of stomach is highly convoluted
			Folds up when empty, expands when full of food
			Carnivores that gorge sporadically can distend stomachs greatly
		Stomach has extra layer of smooth muscle to churn food

	Gastric Glands of Mucosa Are Exocrine Glands that Produce Secretions	fig 45.8
		Parietal cells secrete hydrochloric acid (HCl)
		Chief cells secrete pepsinogen, acid-loving ,weak protein-digesting enzyme
		Activated pepsinogen molecules cleave fragment from each other, make pepsin
			Pepsin is more active molecule
			Production of inactive molecule, converted to active enzyme outside
			Chemical generically called a zymogen
			Prevents chief cells from self-digestion
		Stomach produces 2 liters of acid and gastric secretions per day
			Produces pH of 2, compared to blood pH of 7.4
			Low pH helps denature proteins, keeps pepsin active
			Proteins denatured into polypeptides
			Digestion to amino acids occurs in small intestine
			No digestion of carbohydrates or fats in stomach
		Chyme:  mix of partly digested food and gastric juice
		Acid solution also kills bacteria ingested with food
		Overproduction of acids may occur
			In stomach, cause gastric ulcers
				Are rare due to protective alkaline mucus produced by mucosa
				Mucosal cells readily replaced when damaged
			Duodenal ulcers are more common
				Produced when excessive amount of acidic chyme delivered into duodenum
				Alkaline secretions of pancreas cannot neutralize chyme
		Parietal cells also produce intrinsic factor
			Polypeptide needed for intestinal absorption of vitamin B12
			Required for formation of red blood cells
			Deficiency causes pernicious anemia
		Little absorption occurs in stomach, all other absorption in intestine
			A little water
			Substances like aspirin and alcohol

TERMINAL DIGESTION AND ABSORPTION TAKE PLACE IN THE SMALL INTESTINE

	Food Passes From Stomach to Small Intestine
		Controlled by muscular pyloric sphincter	fig 45.7
		Capacity of small intestine limited, digestion takes time
			Relatively small amounts of chyme can enter at a time
			Coordination regulated by neural and hormonal signals

	Small Intestine Is the Primary Location of Digestion
		Length is approximately six meters
			Duodenum  comprises first 25 centimeters, or 4%
			Jejunum and ileum comprise rest of small intestine
		Duodenum receives chyme, pancreatic enzymes, bile from liver and gallbladder
		Absorption occurs in all three regions of small intestine

	Digestion of Food in the Intestine
		Epithelial wall covered with small projections called villi	fig 45.9
		Epithelium of villi covered with microvilli, cytoplasmic projections	fig 45.10
			Seen clearly with electron microscope
			Epithelial wall also called brush border
		Both increase the absorptive surface of the small intestine
		Microvilli also participate in digestion
			Digestive enzymes embedded in epithelial cell plasma membranes	fig 45.11
			Brush border enzymes hydrolyze lactose, sucrose and others	tbl 45.1
				Adult humans lose ability to produce lactase
				Condition called lactose intolerance

	Absorption of Food in the Intestine
		Components of protein and carbohydrate digestion transported across brush border
			Amino acids and monosaccharides cross to intestinal epithelial cells	fig 45.12
			Transported across intestinal epithelium to capillaries in villi
			Blood carries digestion products to liver
			Travel via hepatic portal vein	fig 45.13
		Products of fat digestion absorbed by different mechanism	fig 45.12
			Fats hydrolyzed into fatty acids and monoglycerides
			Absorbed by intestinal epithelium
			Reassembled into triglycerides
			Combine with proteins to form water-soluble chylomicrons
			Absorbed into lymphatic capillaries, not hepatic portal system
			Contents of lymphatic system enter blood stream in veins near neck
Total volume of food and water equals 2 liters (800 grams of solids)
		Body adds 7.0 liters of its own fluids making a total of 9.0 
			1.5 liters salivary enzymes
			2.0 liters of gastric secretions
			1.5 liters of pancreatic secretions
			0.5 liters of bile from the liver
			1.5 liters of intestinal secretions
		Nearly all fluids and solids are absorbed
			8.5 liters reabsorbed in the small intestine
			350 milliliters reabsorbed in the large intestine
		Only 50 grams of solids and 100 milliliters of liquid leave as feces
		Fluid absorption efficiency = 99%

THE PANCREAS SECRETES ENZYMES, BICARBONATE AND HORMONES

	The Pancreas Makes Digestive Enzymes
		Pancreas located at junction of stomach and small intestine	fig 45.2
		Fluid secreted into duodenum via pancreatic duct
			Pancreas is thus an exocrine organ
			Fluid contains
				Protein digesting trypsin and chymotrypsin
				Starch digesting pancreatic amylase
				Fat digesting lipase
			Enzymes released primarily as zymogens, activated by brush border
			Also contains bicarbonate to neutralize HCl from stomach
				Chyme in intestine is slightly alkaline
				Bicarbonate produced by acini, clusters of secretory cells

	Pancreas Also Serves as an Endocrine Gland
		Produces hormones that regulate levels of blood sugar and other nutrients
		Produced in islets of Langerhans clustered throughout pancreas
		Most important hormones are insulin and glucagon

THE LIVER PRODUCES BILE AND REGULATES BLOOD COMPOSITION

	Bile Production
		Liver is largest internal organ of body	fig 45.2
		Main secretion of liver is bile
			Mixture of bile pigments and bile salts delivered into duodenum
			Bile pigments do not participate in digestion
				Are waste products from liver's destruction of old red blood cells
				Eliminated with feces
				Accumulation of pigments result in jaundice
			Bile salts are lipid and water soluble
				Disperse fat droplets in chyme into emulsion of smaller droplets
				Emulsification increases surface area for lipase to work on
		Bile is stored and concentrated in gall bladder
		Fatty food in duodenum triggers contraction of gallbladder to release bile

	Regulation of Blood Composition
		Hepatic portal vein carries blood from stomach and intestine to liver	fig 45.13
		Liver absorbs or chemically modifies substances before they reach rest of body
			Ingested alcohol and drugs metabolized by liver cells
			Toxins, pesticides, carcinogens, poisons detoxified
			Ammonia from intestinal bacteria converted into urea
		Controls level of substances produced in body
			Steroid hormones converted into less active water-soluble forms
			Molecules included in bile, eliminated in feces or through kidneys
		Produces proteins found in blood plasma
			Includes most blood clotting factors
			Maintains blood protein concentration within narrow limits
			Imbalance can cause edema

	Regulation of Blood Glucose Levels
		Constant concentration of blood glucose must be maintained
			Brain cells totally dependent on blood for supply of glucose
			Brain cells store little glucose, cannot convert fat or amino acids into glucose
		Maintaining level requires active control by various body organs
			Vertebrates eat sporadically, ingestion followed by fasting
			Most food digested rapidly, metabolites enter blood stream
			Without control, level of metabolites would change drastically
		Liver removes glucose from blood, converts it into glycogen
			Glycogen stored in liver tissue and 
			Also stored in skeletal muscle fibers, but can only be used in muscles
			Process stimulated by pancreatic hormone, insulin
		If blood glucose level is low, liver secretes glucose into blood
			Occurs between meals, during fasting
			Glucose partly obtained from breakdown of glycogen
			Conversion stimulated by glucagon, other pancreatic hormone
			Only liver can secrete glucose into blood
			Liver stores enough glycogen for 10 hours of fasting
			For greater fasting liver converts amino acids, lactic acid into glucose
				Process called gluconeogenesis
				Amino acids come from muscle protein

THE LARGE INTESTINE CONCENTRATES SOLIDS

	Large Intestine or Colon Comprises Last Meter of Digestive Tract
		 Has no digestive function, absorbs 4% of fluids
		Shorter in length than the small intestine
			Lies in three relatively straight segments
			Surface is not convoluted
			Inner surface lacks villi
			Significantly less surface area over which to absorb
		Absorb sodium, vitamin K, other products of bacterial metabolism
		Primary function is a refuse dump
			Undigested material compacted and stored
			Bacteria live and reproduce and are incorporated into feces
			Bacterial fermentation produces gas within the colon
		Human colon evolved to process food with high fiber content
			Low fiber diets result in slower passage of food through colon
			May be associated with high level of colon cancer in U. S. 
		The rectum is the terminal portion of the large intestine
			Feces pass into rectum by peristaltic contractions
			Material exits anus through two sphincters
				First sphincter is smooth muscle, opens involuntarily
				Second sphincter is striated muscle, under voluntary control

	All Vertebrates Except Placental Mammals Possess a Cloaca
		A collective cavity for digestive, reproductive and urinary functions
		Placental mammals have separate exits for each tract

NEURAL AND HORMONAL REGULATION OF DIGESTION

	Coordination by Nervous System
		Stimulates salivary and gastric secretions in response to sight and smell of food
			Food in stomach stimulates secretion of gastrin by stomach	tbl 45.2
			In turn stimulates secretion of pepsinogen and HCl in stomach	fig 45.14
			Negative feedback loop controls HCl secretions
				Decreased stomach pH reduces secretion of gastrin
				Decrease in gastrin decreases HCl production
		Passage of chyme from stomach inhibits stomach contractions
			No more chyme enters intestine until previous amount processed
			Process mediated by neural impulses and gastric inhibitory peptide (GIP)
				GIP released by duodenum
				Production stimulated most strongly by fat in chyme
				Fatty meals take longer to process

	Coordination by Hormones
		Cholecystokinin (CCK) is secreted in response to fat in chyme
			Stimulates contraction of gallbladder to release bile
			Bile emulsifies fats, increases efficiency of digestion
			Stimulates secretion of pancreatic digestive enzymes
		Secretin released in response to acidity of chyme
			Stimulates pancreas to release bicarbonate
			Neutralizes acidity of chyme
			First hormone ever discovered

SYMBIOSIS WITHIN THE VERTEBRATE DIGESTIVE SYSTEM

	Bacterial Digestion of Cellulose Within Animals
		Vertebrates lack enzymes to digest plant material
		Some bacteria can do so and are harbored by animals	fig 45.15
			Plays relatively small role in human nutrition
			Essential nutrition for termites, cockroaches and some herbivores
		Cows and related ruminants possess two stomachs	fig 45.16
			First stomach has two chambers:  rumen and reticulum
			Second stomach has two chambers, omasum and abomasum
		Capacity of rumen is 50 gallons
			Provides a fermentation vat for bacteria and protozoa to process cellulose
			Allows cows to regurgitate and rechew their food (cud)
		Re-chewed food swallowed, goes into reticulum, omasum and abomasum
			Abomasum released gastric juices
			Is equivalent to human stomach
			Leads to a very efficient digestion of cellulose
		Horses, rodents and lagomorphs retain bacteria in the caecum
			Cannot regurgitate material from caecum
			Rats and rabbits redigest cellulose another way
			Eat feces and literally redigest them a second time
			Efficiency approaches that of ruminants

	Additional Digestive Activity of Intestinal Bacteria
		Wax digested by bacteria in gut of honey guide birds
		Intestinal bacteria provide mammals with vitamin K
			Birds lack bacteria and must consume vitamin K in food
			Prolonged antibiotic treatment depletes bacteria
			Must supplement vitamin K until bacteria are re-established

NUTRITION

	Ingestion of Food Has Dual Purpose
		Provides source of energy
		Provides raw materials the animal cannot manufacture for itself	fig 45.17
		Intake of food required to maintain glycogen stores in the liver
			Excess glucose metabolized by muscles or converted to fat
			Basal metabolic rate (BMR):  rate energy is consumed at complete rest
			Food energy -(energy used at rest+exercise energy) = energy in glycogen and fat
			BMR is relatively constant within an individual
			Balance between food energy and exercise energy determines energy storage in fat
		Wealthy countries exhibit obesity from overeating and imbalanced diet
			Obese = 20% more than normal weight for a certain height	fig 45.18
			Obesity correlated with coronary heart disease, other disorders
			Obesity in children related to increase in number of fat cells
			Adult obesity related to change in size of fat cells, number does not decrease

	Essential Nutrients
		Over time many vertebrates have lost ability to synthesize substances
		Substances that cannot be manufactured must be obtained from diet
		Vitamins:  essential organic substances required in trace amounts	tbl 45.3
			Humans, apes, monkeys, guinea pigs cannot make vitamin C
			Humans require at least thirteen vitamins
		Essential amino acids:  eight of the total twenty
			Lysine, tryptophan, threonine, methionine, phenylalanine, leucine, isoleucine valine
			Must be obtained from proteins in food	fig 45.19
		Vertebrates synthesize cholesterol, insects cannot
		Essential minerals:  calcium, phosphorus and other trace elements	tbl 2.1

Source: http://www.mhhe.com/biosci/genbio/rjbiology/ELOs/ELO45.html