Invertebrate diges sys are bags or tubes

● Single-celled org & sponges digest their food intracellularly

*others: extracellular (w/in a digestive cavity)
*In this case, the digestive enzymes are released into a cavity that is continuous w/ animal’s
external envi
● In cnidarian & flat worms (planarians): digestive cavity has only 1 opening that serves as
both mouth & anus
● No specialization w/in this type of digestive sys called gastrovascular cavity bc every cell
is exposed to all stages of food digestion
● Specialization occurs when the digestive tract or alimentary canal, has a separate mouth
& anus so that transport of food is one-way
● The most primitive diges tract is seen in nematodes (phylum: Nematoda), where it is
simply a tubular gut lined by an epith membrane
● Earthworms (phylum: Annelida) have a diges tract specialized in diff regions for the
ingestion, storage, fragmentation, digestion , & absorption of food
*All more complex animal grps including all vertebrates, show similar specializations

● The ingested food may be stored in a specialized region of diges tract or it may 1st be
subjected to physical fragmentation (may occur through):
Chewing action of Teeth, or grinding action of pebble (gizzard of earthworms & birds)
● Chem diges
-Then occurs, breaking down the larger food molec of polysaccharides & disaccharides, fats, &
proteins into their smallest subunits;
- involves hydrolysis reac that liberate the subunit molec--primarily monosaccharides, amino
acids, & fatty acids-from food
-products pass through the epith lining of the gut into the blood, in a process known as
absorption.

Vertebrate digestive systems include highly specialized struc molded by diet

● In humans & other vertebrates, diges sys consists of a tubular gastrointestinal tracy &
accessory diges organs
Overview of diges tract
Initial components: mouth & pharynx (common passageway of the oral & nasal cavities)
● Pharynx leads to esophagus (muscular tube) that delivers food to the stomach, where
some preliminary digestion occurs
● From the stomach, food passes to the small intestine, where a battery of digest enzymes
continues the diges process
● The produc of diges, together w/ minerals & water, are absorbed across the wall of the
small intestine into the bloodstream
● What remains is emptied into the large intestine, where some of the remaining water &
minerals are absorbed
 ● Cloaca: a cavity, where waste products emerge from the large intestine & receives the
products of urinary & reproductive systems in most vertebrates other than mammals
● In mammals, separate ang urogenital producs from fecal matl in large intestine
*fecal enters rectum & is expelled through the anus
● The accessory digestive organs:
>Liver : produces bile (a green sol that emulsifies fat)
>Gall Bladder: stores & concentrates the bile
>Pancreas: produce pancreatic juice w/c contains diges enzymes & bicarbonate buffer
*Both bile & pancreatic juice are secreted in the 1st region of the small intestine, the
duodenum where they aid in digestion
Tissues of the digestive tract
● Mucosa
-innermost layer
-epith that lines the interior, or lumen of the tract
● Submucosa
-next major tissue layer, made of connective tissue
● Muscularis
-outside submucosa
-consists of a double layer of smooth muscles
=nner layer have a circular orientation & serve to constrict the gut
=outer layer are arranged longitudinally & work to shorten it
● Serosa
-epith tissue layer, covers the external surface of the tract
-found here are the nerve networks, intertwined in plexuses between muscle layers & help
regulate the gastrointestinal activities

THE MOUTH & TEETH:FOOD CAPTURE & BULK PROCESSING

● Specialization of diges sys in diff kinds of vertebrates reflect the way these animals live
● Birds w/c lack teeth, break up food in their 2 chambered stomachs
-gizzard: small pebbles ingested by the bird are churned together w/ the food by muscular
action. (grinds up seed & other hard plant matl into smaller chunks)

Vertebrate teeth are adapted to diff types of food items

● Teeth used for chewing or mastication that break up food into small particles & mix it w/
fluid secretions
● CARNIVORES:
-pointed teeth that lack flat grinding surfaces (cutting & shearing)
-often tear pieces of their prey but have little need to chew them, bc diges enzymes can act
directly on animal cells
● HERBIVORES:
-pulverize cellulose cell walls of plant tissue b4 the bacteria in their rumens or cecae can digest
them
-have large flat teeth w/ complex ridges well suited to grinding
● OMNIVORES:
-Humans: carnivores in front of mouth & herbivores in back
-4 front teeth in upper & lower jaws are sharp, chisel-shaped incisors used for biting
-On each side of the incisors are sharp pointed teeth called cuspids/ canine w/c are used for
tearing food
-behind canines are 2 premolars & 3 molars, all w/ flattened, ridged surfaces for grinding &
crushing food

The mouth is a chamber for ingestion & initial processing

● Inside the mouth, the tongue mixes food w/ a mucous sol, saliva
● Humans-- 3pairs of salivary glands secrete saliva into the mouth through ducts in one’s
mouth’s mucosal lining
>saliva moistens & lubricates food so that it is easier to swallow & does not abrade the tissue of
the esophagus as it passes through
>saliva also contains hydrolytic enzyme salivary amylase, w/c initiates the breakdown of the
polysaccharide starch into the disaccharide maltose
*this diges is minimal in humans bc most dont chew their food very long

Stimulation of salivation
● Nervous sys controls secretion of salivary glands, w/c in human maintains a constant
flow of abt half a mililiter per min when the mouth is empty of food
*continuous secretion keeps the mouth moist
● Presence of food=increased rate of secretion
● Olfactory (smell), sight or sound= stimulate salivation
● The most potent stimuli are acidic solution
*lemon juice increase rate of salivation eightfold

Swallowing
● Initiated by voluntary action then is continued under involuntary control
*when food is ready to be swallowed, the tongue moves it to the back of the mouth
● In mammals, the process of swallowing begins when the soft palate elevates, pushing
against the back wall of the pharynx
● Elevation of the soft palate seals off the nasal cavity & prevents food from entering it
● Pressure against the pharynx triggers an automatic, involuntary response, the
swallowing reflec (cannot be stopped once initiated)

These actions keep food out of respiratory tract, directing it instead into the esophagus
● Neurons w/in the walls of the pharynx send impulses to the swallowing center of the
brain
● In response, electrical impulses in motor neuron stimulate muscles to contract & raise
the larynx (voice box)
● This pushes the glottis, the opening from the larynx into the trachea (windpipe), against
a flap of tissue called the epiglottis
In short:
1. As food moves to the back of the mouth, the soft palate seals of nasal cavity
 2. During swallowing, larynx rises & is sealed off by epiglottis (seal of trachea). This forces
the bolus (processed lump of food) into the esophagus & prevents entry into the trachea.
As the bolus moves into the esophagus, the larynx relaxes

ESOPHAGUS & STOMACH: EARLY STAGES OF DIGESTION

● Swallowed food enters a muscular tube called esophagus, w/c connects the pharynx to
the stomach
● The esophagus actively moves a processed lump of food called a bolus through the
action of muscles

Muscular contractions of esophagus move food to the stomach

● Esophagus in adult humans: abt 25 cm long
-upper 3rd is enveloped in skeletal muscle for voluntary control of swallowing
-lower 2-3rds is surrounded by involuntary smooth muscle
● Swallowing center stimulates successive one-directional waves of contraction in these
muscles that move food along the esophagus to the stomach
● These rhythmic waves of muscular contraction are called peristalsis (enable humans &
other verteb to swallow even if they are upside down)
● In many verteb, movement of food from the esophagus into the stomach is controlled by
a ring of circular smooth muscle or sphincter , that opens in response to the pressure
exerted by the food
*contraction of this sphincter prevents food in stomach from moving back into the esophagus
-rodents & horses have true sphincter & as a result they cannot regurgitate= bring (swallowed
food) up again to the mouth.
*normally, esophagus is closed off except during swallowing

Stomach is a “holding station” involved in acidic breakdown of food

● Stomach contains a 3rd layer of smooth muscle for churning food & mixing it w/ gastric
juice (acidic secretion of the tubular gastric glands of the mucosa)
These exocrine gastric glands contain 3 kinds of secretory cells:
1. Mucus-secreting cells
2. Parietal cells
-secrete hydrochloric acid (HCl)
3. Chief cells
-secrete pepsinogen, the inactive form of the protease (protein digesting enzyme) pepsin

Pepsinogen has44 additional amino acids that block its active site
● HCl causes pepsinogen to unfold, exposing the active site, w/c then acts to remove the
44 amino acids
● This yields the active protease, pepsin
● This process of secreting an inactive form that is then converted into an active enzyme
outside the cell prevents the chief cells from digesting themselves
● In the stomach, muscus produced by muscus-secreting cells serves the same purpose,
covering the interior walls & preventing them from being digested
 ● In addition to producing HCl, the parietal cells of stomach also secrete intrinsic factor,
a polypeptide needed for the intestinal absorption of vit B12
-bc this vitamin is req for the production of red blood cells, people who lack sufficient intrinsic
factor develop a type of anemia (low rbc) called pernicious anemia

Action of acid
● Human stomach prod abt 2L of HCl & other gastric secretions every day, creating a very
acidic solution
*concentration of HCl-abt 10 millimolar (mM), equal to a pH of 2.
=Thus gastric juice is abt 250k times more acidic than blood, whose normal pH is 7.4
● The low pH in stomach helps denature (take away or alter the natural qualities) food
proteins, making them easier to digest, & keeps pepsin maximally active
*active pepsin hydrolyzes food proteins into shorter chains of polypeptides that are not fully
digested until the mixture enters the small intestine
● Mixture of partially digested food & gastric juice is called chyme
*in adult humans, only proteins are partially digested in the stomach-no significant digestion of
carbohydrates or fats occurs there
● The acidic sol in stomach also kills most of bacteria that are ingested w/ the food
*few bacteria that survive enter the intestine intact are able to grow & multiply (large intestine)
*bacteria are a major component of feces

Ulcers
● Overprod of gastric acid can occasionally eat a hole through the wall of stomach or
duodenum, causing a peptic ulcer
*common cause: infection w/ bacterium Heliocobacter pylori
-can grow on the lining of human stomach, surviving acid pH by secreting substances
that buffer the pH of its immediate surroundings
-common in US (abt 20% of people younger than 40 & 50% older than 60), most people
are asymptomatic
● Infection by H. pylori can reduce or weaken the mucosal layer in stomach or duodenum,
allowing acidic secretions to attack the underlying epith
● Antibiotic treatment of the infection can reduce symptoms & often even cure the ulcer

Leaving the stomach

● Chyme leaves stomach through the pyloric sphincter to enter small intestine
>where all terminal digestion of carb, lipids, & proteins occurs
>where products of digestion- amino acids, glucose, and so on- are absorbed into the blood
> only some of the h20 in chyme & a few substances (aspirin & alcohol) are absorbed through
wall of stomach
INTESTINES: BREAKDOWN, ABSORPTION & ELIMINATION
● Capacity of small intes is limited & its digestive processes take time
● Efficient diges req that only relatively amts of chyme be introduced from the stomach into
the small intes at any one time
*coordination between gastric and intestinal activities is regulated by neural & hormonal signals
Structure of small intestine is specialized for digestion & nutrient uptake
● Small intestine
-approx 4.5m long (living) but 6m long at autopsy (muscles relaxed)
>1st 25cm= duodenum
> remainder= jejunum & ileum
● Duodenum receives acidic chyme from stomach, diges enzymes & bicarbonate from the
pancreas, & bile from liver & gallbladder
*enzymes in pancreatic juice digest larger food molec into smaller fragments
-this digestion occurs primarily in the duodenum & jejunum
● The epith wall of small intestine is covered w/ a tiny fingerlike projections called villi
● In turn, each epith cell lining the villi is covered on its apical surface (side facing lumen)
by many foldings of the plasma membrane that form cytoplasmic extensions called
microvilli
-also participate in digestion bc a # of diges enzymes are embedded w/in the epith cell’s plasma
membranes, w/ their active sites exposed to the chyme
-these brush border enzymes include those that hydrolyze the disaccharides lactose & sucrose,
among others
● Epith wall of small intestine is also called a brush border
● Many adult humans lose the ability to produce the brush border enzymes lactase &
therefore cannot digest lactose (milk sugar), a rather common condition called lactose
intolerance
● Brush border enzymes complete the digestive process that started w/ the action of
salivary amylase in mouth

Accessory organs secrete enzymes into small intestine

● Main organs that aid in digestion: pancreas, liver & gallbladder
● They empty their secretions, primarily enzymes through ducts directly into the small
intestine

Secretions of the pancreas

Pancreas
● large gland situated near the junction of stomach & small intestine,
● secretes pancreatic fluid into the duodenum through the pancreatic duct; thus, the
pancreatic functions as an exocrine gland (contains a host of enzymes)
>trypsin & chymotrypsin, w/c digest proteins;
>pancreatic amylase w/c digests starch;
>lipase, which digests fat
*like pepsin in the stomach, these enzymes are released into the duodenum primarily as
inactive enzymes & are then activated by trypsin, w/c is 1st activated by a brush border enzyme
of the intestine
● Pancreatic enzymes digest proteins into smaller polypeptides, polysaccharides into
shorter chains of sugars, & fats into free fatty acids & monoglycerides
● Digestion of proteins & carb is then completed by brush border enzymes
 ● Pancreatic fluid also contains bicarbonate, w/c neutralizes the HCl from the stomah &
gives the chyme in the duodenum a slightly alkaline pH
● The digestive enzymes & bicarbonate are prod by clusters of secretory cells known as
acini
● Endocrine: secreting several hormones into blood that control blood level of glucose &
other nutrients
> these hormones are prod in islets of Langerhans, clusters of endocrine cells scattered
throughout the pancreas
> 2 most imp pancreatic hormones: insulin & glucagon

Liver & gallbladder

Liver
● Largest internal organ of body (weigh 1.5 kg, size of a football)
● Detoxification of harmful substances, protein synthesis, & glycogen storage
*Chronic abuse of alcohol can overtax the liver’s detoxification capabilities, leading to cirrhosis &
eventually to liver failure
● Main exocrine secretion is bile, a fluid mixture consisting of bile pigments & bile salts that
is delivered into the duodenum during the digestion of a meal
>Bile pigments do not participate in digestion; they are waste products resulting from the liver’s
destruction of old red blood cells & are ultimately eliminated w/ the feces
>if the excretion of bile pigments by the liver is blocked, the pigments can accumulate in the
blood & cause a yellow staining of the tissue known as jaundice

>bile salts: prepare fats for subsequent enzymatic digestion bc fats are insoluble in water, they
enter the intestine as drops w/in the watery chyme
>bile salts w/c are partly lipid-soluble & partly water-soluble, work like detergents, dispersing the
large drops of fat into a fine suspension of smaller droplets
*this emulsification action prod a greater surface area of fat for the action of the lipase
enzymes, & thus allows the digestion of fat to proceed more rapidly

● After bile is prod in liver, it is stored & concentrated in gallbladder

● The arrival of fatty food in duodenum triggers a neural & endocrine reflex that stimulates
the gallbladder to contract, causing bile to be transported through the common bile duct
& injected into the duodenum
● Gallstones are hardened precipitates of cholesterol that form in some indiv
>if these stones block the bile duct, contraction of the gallbladder causes intense pain, often felt
in the back
>in severe cases of blockage, surgical removal of gallbladder may be performed

Absorbed nutrients move into blood or lymph capillaries

● After enzymatic breakdown, proteins & carb are absorbed as amino acids &
monosaccharides
*they are transported across the brush border into the epith cells that line the intestine by a
combination of active transport & facilitated diffusion
 ● Glucose is transported by coupled transport w/ Na+ions (a.k.a secondary active
transport)
*fructose found in most fruits is transported by facilitated diffusion
*most amino acids are transported by active transport using a variety of diff transporters
*Some of these carrier proteins use cotransport w/ Na+ions; others transport only amino acids
● Once they have entered epith cells across the apical membrane, these
monosaccharides & amino acids move through the cytoplasm & are transported across
the basolateral membrane & into the blood capillaries w/in the billi
● The blood carries these products of digestion from the intestine to the liver via the
hepatic portal vein
> a portal vein connects 2 beds of capillaries instead of returning to the heart
>intestine is connected to the liver by the hepatic portal vein, thus the liver receives blood-borne
molec from the intestine
>bc of the hepatic portal vein, the liver is the 1st organ to receive most of the products of
digestion (except for fat)

● Products of fat digestion are absorbed by a diff mechanism

>fats are hydrolyzed into fatty acids & monoglycerides by digestion
> fatty acids & monoglycerides are nonpolar & can thus enter epith cells by simple diffusion
*once inside the intestinal epith cells they are reassembled into triglycerides
● triglycerides then combine w/ proteins to form small particles called chylomicrons, w/c
are too bulky to enter blood capillaries in the intestine
*instead of entering the hepatic portal circulation, the chylomicrons are absorbed into lymphatic
capillaries, w/c empty their contents into the blood in veins near the neck
*chylomicrons can make the blood plasma appear cloudy if a sample of blood is drawn after a
fatty meal

● The amt of fluid passing through small intestine in a day is startlingly large: approx 9L
-almost all of this fluid is absorbed into the body rather than eliminated in the feces
-almost 8.5L is absorbed in small intestine & an additional 350mL in large intestine
-only abt 50g of solid & 100mL of liquid leaves the body as feces
-normal fluid absorption efficiency of human diges tract approaches 99% w/c is very high indeed

Large intestine eliminates waste material

Large Intestine/Colon
● Much shorter, occupying the last meter of digestive tract, large bc of its diameter
● The small intestine empties directly into large intestine at a junction where 2 vestigial
structures: cecum & appendix remain
● No digestion takes place & only abt 4% of absorption of fluids occur
● Not as convoluted as small intestine, & its inner surface has no villi
● Less than 1/30 the absorptive surface of small intestine
● Function
-absorb water, remaining electrolytes, & products of bacterial metabolism (including vitK). Large
intestine prepares waste matl to be expelled from the body
-prepares waste matl to be expelled from the body
● Many bacteria live & reproduce w/in large intestine, & excess bacteria are incorporated
into the refuse matl called feces
>bacterial fermentation produces gas w/in colon at a rate of abt 500mL per day
>this rate increases greatly after the consumption of beans or other types of vegetables bc the
passage of undigested plant matl (fiber) into the large intestine provides substrates for bacterial
fermentation

● Human colon evolved to process food w/ relatively high fiber content

● Diet in low fiber (common in US) results in a slower passage of food through the colon
● Low dietary fiber content is thought to be assoc w/ the level of colon cancer in US, w/c is
among the highest in the world

● Compacted feces, driven by peristaltic contractions of large intestine, pass from the
large intestine into a short tube called the rectum
● From rectum, feces exit body through the anus
● 2 sphincters control passage through anus
1. Smooth muscle & opens involuntarily in response to pressure inside the rectum
2. Striated muscle can be controlled voluntarily by brain, thus permitting a conscious decision to
delay defacation

VARIATIONS IN VERTEBRATE DIGESTIVE SYSTEMS

● Some animals contain bacteria & protists that convert cellulose into substances the host
can absorb
● Gastrointestinal microorganism plays a relatively small role in human nutrition, it is an
essential elem in nutrition of many other kinds of animals, including insects such as
termites & cockroaches, & a few grps of herbivorous mammals
● The relationships between these microorganisms & their animal hosts are mutually
beneficial & provide an excellent ex of symbiosis
● Plant cellulose is partic resistant to digestion
>herbivores tend to have much longer digestive tracts, allowing greater time for digestion to
occur

Ruminants rechew regurgitated food

● Ruminants have 4 chambered stomach
>reticulum
>rumen
-may hold up to 50 gallons, serves as a fermentation vat where bacteria & protists
convert cellulose & other molec into a variety of simpler compounds
-at the front of the 4 chambers allows the animal to regurgitate (bring (swallowed food)
up again to the mouth.) & rechew the contents of the rumen, an activity called rumination or
“chewing the cud”
 -this breaks tougher fiber in the diet into smaller particles, increasing the surface area for
microbial attachment
*after chewing, the cud is swallowed for further microbial digestion in rumen then pases to the
omasum and then to the abomasum (where it is finally mixed w/ gastric juice)
-this process leads to far more efficient digestion of cellulose in ruminant than in
mammalian herbivores that lack a rumen (horse)
>omasum
>true stomach
>abomasum

1. Rumen
>contains bacteria that break down cellulose from the plant cell walls
> b4 moving to 2nd chamber, food may be rechewed or regurgitated (bring (swallowed food) up
again to the mouth.)
2. Reticulum
3. -4. Omasum &Abomasum
*omasum secretes gastric juice as in the human stomach

Foregut fermentation has evolved convergently many times

● Hippopotamuses, langur monkeys, sloths, kangaroos, & hoatzins (type of bird) have
evolved large stomachs to enhance microbial fermentation
*these species have evolved a variety of other anatomical struc that serve to slow down the
passage of food through the stomach, leading to increased time for fermentation
● A remarkable case of convergent evol at the molecular level is exhibited by ruminants &
the langur monkey w/c subsist primarily on leave s
● In most mammals, lysozymes are enzymes found in saliva & tears, w/c attack invading
bacteria
● However, in ruminants & langurs, lysozymes have been modified for digesting bacteria
in stomach
● In both cases, 5 identical amino acid changes have evolved; the result is that the
lysozyme molec of ruminants & langurs are more similar to each other than they are to
lysozymes in more closely related species
*this ex illus that convergent evol has occurred in distantly related species by the exact same
evolutionary changes
Other herbivores have alternative strategies for digestion
● Some animals such as rodents, horses, deer & lagomorphs (rabbits & hares), the
digestion of cellulose by microorganism takes place in the cecum, w/c is greatly enlarged
● Bc the cecum is located beyond the stomach, regurgitation of its contents is impossible
● Rodents & lagomorphs have evolved another way to capture nutrients from cellulose
that achieves a degree of efficiency similar to ruminant digestion
● They do this by producing some feces packed w/ nutrients, w/c they then eat, a practice
known as coprophagy--thus passing the food through their digestive tract a 2nd time
● The 2nd passage allows the animal to absorb nutrients produced by the microorganism
in its cecum
*coprophagic animals cannot remain healthy if they are prevented from eating their feces
*animals w/ diets that dont include cellulose dont have a cecum, or if they do, it is greatly
reduced (insectivores or carnivores)

Vitamin K
● Another ex of intestinal microorganisms function in metabolism of animal hosts is
provided by synthesis of vit K
● All mammals rely on intestinal bacteria to synthesize this vitamin, w/c is necessary for
the clotting of blood
● Birds, w/c lack these bacteria, must consume the req quantities of vit K in their food
● In humans, prolonged treatment w/ antibiotics greatly reduces populations of bacteria in
intestine
● Restoring normal flora of digestive tract w/ beneficial bacteria may also help replace vit K

NEURAL & HORMONAL REGULATION OF THE DIGES TRACT

● Activities of gastrointestinal tract are coordinated by nervous & endocrine sys
>Nervous sys: stimulates salivary & gastric secretions in response to sight, smell &
consumption of food
>Endocrine sys: when food arrives in stomach, proteins in food stimulate the secretion of a
stomach hormone called gastrin w/c in turn stimulates secretion of pepsinogen & HCl from
gastric glands
=the secreted HCl then lowers the pH of gastric juice, w/c acts as to inhibit further
secretion of gastrin in negative feedback loop
=secretion of gastric acid is kept under tight control
● The passage of chyme from the stomach into the duodenum of the small intestine
inhibits the contractions of the stomach, so that no additional chyme can enter the
duodenum until the prev amt can be processed
This stomach or gastric inhibitor is mediated by a neural reflex & by duodenal hormones
secreted into the blood (collectively known as enterogastrones)
● The major enterogastrones include cholecystokinin (CCK), secretin, & gastric
inhibitory peptide (GIP)
● The chyme w/ the high fat content is strongest stimulus for CCK & GIP secretions; the
result is that fatty meals remain in the stomach longer than nonfatty meals, allowing
more time for digestion of complex fat molec

● In addition to gastric inhibition, CCK & secretin have other imp regulatory functions
CCK:
-stimulates increased pancreatic secretions of diges enzymes & gallbladder contractions
*gallbladder contractions inject more bile into the duodenum, w/c enhances the emulsification &
efficient diges of fats
Secretin:
-stimulate pancreas to release more bicarbonate, w/c neutralize the acidity of chyme
-1st hormone ever discovered
ACCESSORY ORGAN FUNCTION
● Liver is a key organ in the breakdown of toxins, & the pancreas secrete hormones that
regulate the blood glucose level in part through actions on liver cells

Liver modifies chemicals to maintain homeostasis

● Bc the hepatic portal vein carries blood from stomach & intestine directly to the liver, the
liver is in a position to chemically modify the substances absorbed in the gastrointestinal
tract b4 they reach the rest of the body
*ingested alcohol & other drugs are taken into liver cells & metabolized (liver often damaged)
● Liver also removes toxins, pesticides, carcinogens, & other poisons, converting them
into less toxic forms
*liver converts the toxic ammonia produced by intestinal bacteria into urea, a compound that
can be contained safely & carried by the blood at higher concentrations
● Liver regulates the levels of many compounds produced w/in the body
*steroid hormones are converted into less active & more water-soluble forms by the liver. These
molec are then included in the bile & eliminated from the body in the feces or are carried by the
blood to the kidneys & excreted in the urine
● Liver also produces most of the proteins found in blood plasma
-total concentration of plasma protein is significant bc it must be kept w/ certain limits to
maintain osmotic balance between blood & interstitial (tissue fluid)
-if the concentration of plasma protein drops too low=liver disease (cirrhosis), fluid accumulates
in the tissues, a condition called edema

Blood glucose concentration is maintained by the actions of insulin & glucagon

● Neurons in the brain obtain energy primarily from the aerobic respi of glucose obtained
from the blood plasma
*vitally imp that the blood glucose concentration not fall too low, as might happen during fasting
or prolonged exercise
*imp that the blood glucose concentration not stay at too high a level as it does in people w/
untreated diabetes mellitus bc too high a level can lead to tissue damage
● After a carb-rich meal, the liver & skeletal muscles remove excess glucose from the
blood & store it as a polysaccharide glycogen
*stimulated by the hormone insulin, secreted by the beta cells in pancreatic islets of Langerhans
● Blood glucose level decrease (between meals, during fasting & during exercise)
=liver secretes glucose into the blood
*this glucose is obtained in part from the breakdown of liver glycogen to glucose-6-
phosphate, a process called glycogenolysis
=phosphate grp is then removed, & free glucose is secreted into the blood
=skeletal muscles lack the enzyme needed to remove the phosphate grp & so even though they
have glycogen stores, they cannot secrete glucose into the blood
>muscle cells can use this glucose directly for energy metabolism bc glucose-6-
phosphate is actually the product of the 1st reaction in glycolysis
 =breakdown of liver glycogen is stimulated by another hormone, glucagon , w/ is secreted by
alpha cells of islets of Langerhans in pancreas
● Gluconeogenesis: new formation of glucose occurs when fasting or exercise continues,
the liver begins to convert other molec such as amino acids & lactic acid into glucose
-amino acids used for gluconeogenesis are obtained from muscle protein, w/c explains severe
muscle wasting that occurs during prolonged fasting

FOOD ENERGY EXPENDITURE & ESSENTIAL NUTRIENTS

● Ingestion of food (primary functions):
-provides a source of energy
-provides raw matl the animal is unable to manufacture for itself
● Even an animal completely at rest req energy consumption under defined resting
conditions is called basal metabolic rate
-BMR is relatively constant for a given indiv (depending primarily on the person’s age, sex, &
body size)

Exertion increases metabolic rate

● Physical exertion raises the metabolic rate above the basal levels so the amt of the
energy the body consumes per day is determined not only be the BMR but also by the
level of physical activity
● If food energy taken in is greater than the energy consumed per day, the excess energy
will be stored in glycogen & fat
-bc glycogen reserves are limited, however, continued ingestion of excess food energy primarily
in the accumulation of fat
● Kilocalories:intake of food energy is measured
(1kilocalorie= 1000calories; nutritionists use calorie w/ a capital C instead of kilocalorie)
-measurement of kilocalories in food is “burned”, either literally in a testing device called a
calorimeter, or in the body when food is digested & later oxidized during cellular respiration
● Daily energy expenditure (metabolic rates) of humans vary between 1300 & 5000
kilocalories per day, depending on the person’s BMR & level of physical activity
● When the total kilocalories ingested exceeds the metabolic rate for a sustained period, a
person accumulates an amt of fat that is deleterious to health, a condition called obesity

Food intake is under neuroendocrine control

● Experiments w/ fasting & overfeeding in rats showed an increase in food intake when
fasting ends
>The increase restores lost body weight to baseline values & food intake then drops
=indicated the existence of control mechanisms to link food intake to energy balance
● Presence of hormonal satiety factor produced by adipose tissue was hypothesized to
explain these observations
● It has also been shown that regions of the hypothalamus are involved in feeding
behavior
● Other studies in rodent models had identified a # of genes that can lead to obesity
-as modern molecular genetics has allowed the cloning of many of these genes, the outlines of
a model to link dietary intake to energy balance have emerged
>afferent signalling from adipose tissue & feeding behavior into CNS
>efferent signaling outward from the CNS tied to energy expenditure, storage,
reproduction & feeding behavior

Leptin
● Animals homozygous for the recessive mutant allele become obese compared w/ wild-
type mice
● When the gene responsible for this dramatic phenotype was isolated, it proved to
encode a peptide hormone named leptin, leading to the hypothesis that the lack of leptin
production in mutant indiv is responsible for obesity
● When ob (obeses)/ob animals are injected w/ leptin, they stop overeating & lose weight
● Leptin
-main satiety factor & key to control of apetite
-gene for leptin receptor has also been isolated & it is expressed in brain neuron in the
hypothalamus involved in energy intake
-main signaling molec in the afferent portion of the control circuit for energy sensing, food
intake, & energy expenditure
-produced by adipose tissue in response to feeding, & leptin levels correlate w/ feeding behavior
& amt of body fat
● Dietary restrictions reduces leptin levels, signaling the brain that food intake is necessary
● whereas refeeding after fasting leads to rapid increase in leptin levels & a loss of apetite
-efferent part of this control circuit is complex & includes control of energy expenditure, energy
storage, & feeding behavior by the CNS
*reproduction is even affected by this sys as reproduction is inhibited under starvation
conditions
● Leptin gene has also been isolated in humans & leptin appears to function in humans
much as it does in mice
● However recent studies in humans show that the activity of the ob gene & the blood
concentrations of leptin are actually higher in obese than in lean people & that the leptin
produced by obese people appears to be normal
● It has been suggested that, in contrast w/ mutant mice, most cases of human obesity
may result from a reduced sensitivity to the actions of leptin in the brain, rather than from
reduced leptin production

Insulin
● Although the extreme obesity associated w/ the loss-of-function mutations in the ob gene
indicate that other hormonal signals cannot substitute for leptin signaling, other
hormones are also involved
● Insulin has been implicated in signaling satiety as well, & insulin levels also fall w/ fasting
& rise w/ obesity
● Insulin’s primary role is homeostasis of blood glucose
*its role in the control circuit of energy balance is complex
Gut hormones (enterogastrones)
● The gut produces a # of hormones that control the physiology of digestion
*Several of these have also been implicated in the regulation of food intake
=produced directly in response to feeding, necessary for their role in digestion
● The hormone GIP & CCK have receptors in the hypothalamus & seem to send the same
kind of inhibitory signals to the brain as leptin & insulin
● The levels of these gut hormones also vary w/ feeding behavior in a pattern similar to
leptin & insulin
● Ghrelin (gut hormone)
-also has receptors in the hypothalamus
Appears to stimulate food intake
-in rats: chronic administration of ghrelin leads to obesity
-appears to rise b4 feeding & may be involved in initiating feeding behavior
● One of the treatments for severe obesity , gastric bypass surgery, leads to reduced
levels of ghrelin

Neuropeptides
● Efferent control over feeding & energy balance is less clear than the afferent control
● Central regulator is the hypothalamus & 2 brain neuropeptides have been implicated:
-neuropeptide Y (NPY)
-alpha-melanocyte-stimulating hormone
*these peptide are antagonistic w/ NPY inducing feeding activity & alpha-melanocyte-stimulating
hormone suppressing it
● Evidence comes from expirements that show that the production & release of alpha-
melanocyte-stimulating hormone is stimulated by leptin & that the administration of
alpha-melanocyte-stimulating hormone suppresses feeding
● Loss of function for alpha-melanocyte-stimulating hormone= obesity
● Expression of NPY is negatively regulated by leptin & administration of NPY stimulates
feeding behavior

Model for energy balance

● Role for both leptin & insulin is long-term regulation of the afferent portion of this
signaling network
● Leptin & insulin are produced by adipose tissue & the pancreas, respectively, in
response to the effect of feeding behavior, not as direct response to feeding itself
=leads to circulating levels of leptin that correlate w/ amt of adipose tissue
● extreme example of this is the very high level of leptin seen in obese individuals
>high levels of leptin & insulin then act on the hypothalamus to increase levels of alpha-
melanocyte-stimulating hormone & reduce levels of NPY
>this causes a reduction in appetite & increased energy expenditure & allows the reproduction
& growth
● low levels of these hormones
>act on the hypothalamus to reduce alpha-melanocyte-stimulating hormone & increase NPY
>if very low levels of leptin persist, this can inhibit reproduction & growth
● Gut hormones CCK & GIP are produced in response to feeding & rep short-term
regulators of the afferent portion of the energy balance control circuit
*action is same w/ leptin & insulin
● Gut hormones ghrelin is also a short-term regulator that stimulates feeding

Essential nutrients are those that body cannot manufacture

● Essential nutrients:substances that an animal cannot manufacture for itself but that are
necessary for its health & must be obtained in the diet
1. Vitamins
-vit C (humans, monkeys, apes & guinea pigs cannot synthesize abscorbic acis (vitc) )
-vit c defficient: develop scurvy, a potentially fatal disease that results in degeneration of
connective tissues
-humans req at least 13 diff vit
2. Amino acids
-must be obtained from food they eat
-humans req 9amino acids
Strict vegetarians must choose their food so that the essential amino acids in 1 food
complement those in another; vegetarians may also need supplements to provide certain vit not
found in large amts of plants (B vits)
3. long chain unsaturated fatty acids
-obtain them in food
● Some essential nutrients that vertebrates can synthesize cannot be manufactured by
others
● Food also supplies essential nutrients
● Animals obtain trace elements (zinc & molybdenum) either directly prom plants or animal
that have eaten plants