Chapter 37 Notes
Animal Nutrition
Lecture Notes:
Animals are heterotrophic
and, like all heterotrophs, must rely on organic compounds in their food to
supply energy and the raw materials for growth and repair.
A variety of nutritional
adaptations have resulted from natural selection during evolutionary
development of animals.
I. Diets and feeding mechanisms vary
extensively among animals
Animals usually ingest other
organisms.
The food organism may be either dead of alive.
The food organism may be ingested whole or in pieces.
Some parasitic animals (i.e. tapeworm) are exceptions.
Animals are categorized based
on the kinds of food they usually eat and their adaptations for obtaining and
processing food items.
Herbivores
eat autotrophic organisms (plants, algae, and autotrophic bacteria).
Carnivores
eat other animals.
Omnivores
eat other animals and autotrophs.
The varied diets exhibited by
animals are accompanied by a variety of mechanisms used to obtain food.
Suspension-feeders sift small food particles from the water.
̃ Many are aquatic animals such as clams and oysters
(trap food on gills) and baleen whales (strain food from water forced through
the screen-like plates on their jaws).
Substrate-feeders live on or in their food source and eat their way
through food.
̃ Leaf miners (larvae of various insects) tunnel through
the interior of leaves.
Deposit-feeders are a type of substrate-feeder that ingests partially
decayed
organic materials along with their
substrate.
̃ Earthworms ingest soil and their digestive systems
extract the organic materials.
Fluid-feeders suck nutrient-rich fluids from a living host.
̃ Aphids ingest the phloem sap from plants; leeches and
mosquitoes suck blood from animals; hummingbirds and bees ingest nectar from
flowers.
Bulk-feeders eat relatively large pieces of food.
̃ Most animals; they possess various adaptations to kill
prey or tear off pieces of meat or vegetation.
II. Ingestion, digestion, absorption, and
elimination are the four main stages of food processing
Food processing involves four
stages: ingestion, digestion, absorption, and elimination.
Ingestion is the first stage and is the act of eating.
Digestion is the second stage and is the process of breaking
down food into small molecules the body can absorb.
Organic food material is composed of macromolecules
(proteins, fats, carbohydrates) that are too large to cross the membranes and
enter an animal’s cells.
Digestion enzymatically cleaves these macromolecules
into component monomers that can be used by the animal (polysaccharides to
simple sugars; proteins to amino acids; fats to glycerol and fatty acids).
Digestion uses enzymatic hydrolysis to
break bonds in macromolecules.
̃ Hydrolytic enzymes catalyze the digestion of each
class of macromolecules by adding water.
̃ The chemical digestion is usually preceded by
mechanical fragmentation (e.g. chewing) that increases the surface area exposed
to digestive juices.
̃ Occurs in a specialized compartment where the enzymes
are contained so they don’t damage the animal’s own cells.
Absorption is the third stage of the process and involves the
uptake of the small molecules resulting from digestion.
Elimination is the fourth, and the last, stage where undigested
material passes out of the digestive compartment.
III. Digestion occurs in
food vacuoles, gastrovascular cavities, and alimentary canals
A. Intracellular Digestion in Food Vacuoles
Food vacuoles are the simplest digestive compartments.
They are organelles where a single cell digests its
food without hydrolytic enzymes mixing with the cell’s cytoplasm.
Protozoa have food vacuoles that form around food by
endocytosis.
Hydrolytic enzymes are secreted into the food vacuole
and digestion occurs; this is referred to as intracellular digestion. (See Campbell, Figure 37.5).
Sponges differ from other animals in that all
digestion is by the intracellular mechanism. (See Campbell, Figure 37.6).
Extracellular digestion occurs within compartments that are continuous via
passages with the outside of the body.
At least some hydrolysis occurs in most animals by
this mechanism.
B. Digestion in Gastrovascular Cavities
Most animals that have simple
body plans possess a gastrovascular cavity.
Gastrovascular cavity = Digestive sac with a
single opening.
Function in both digestion
and nutrient distribution.
Digestion is Hydra
involves both extracellular and intracellular digestion (See Campbell, Figure
37.7a)
Hydra is a carnivore which
captures prey.
Food items are immobilized by
stings from nemacysts on the tentacles.
Tentacles then force prey
through the mouth into the gastrovascular cavity.
Specialized gastrodermal
cells secrete digestive enzymes that fragment the soft tissues of the prey into
tiny pieces.
Some gastrodermal cells also
possess flagella whose movement prevents settling of food particles and
distributes them through the cavity.
The small pieces are
phagocytized by nutritive gastrodermal cells and surrounded by food vacuoles.
Hydrolysis is completed by
intracellular digestion.
Undigested materials are
expelled from the gastrovascular cavity though the single opening.
The combination of
extracellular and intracellular digestion that occurs in most animals permits
these organisms to feed on larger prey items.
Phagocytosis is limited to microscopic food.
Extracellular digestion begins the digestive process
by breaking-down large food items into smaller particles.
Extracellular digestion may
even begin outside of the animal’s body as in planarians.
Planarians possess a gastrovascular cavity that is
extensively branched.
Associated with the cavity is a muscular, tubular pharynx
that can be everted ventrally though the mouth.
The pharynx penetrates the prey (small invertebrates
and dead organic matter) and releases digestive juices.
The partially digested food is sucked into the
gastrovascular cavity where digestion continues.
Digestion is completed by intracellular digestion in
cells that phagocytize the tiny particles.
C. Digestion in Alimentary
Canals
Animals with body plans more complex cnidarians and
platyhelminths have alimentary canals.
Alimentary canal = Digestive tube running between two openings: the mouth (where
food is initially ingested) and the anus (where undigested wastes are
egested). (See Campbell, Figure 37.8)
Since food moves in one
direction along the tube, the tube can be organized into specialized regions
that carry out digestion and absorption of nutrients in a stepwise fashion.
The unidirectional passage of
food and the specialization of function for different regions make the
alimentary canal more efficient.
IV. A tour of the mammalian digestive system
The digestive system in
mammals include the alimentary canal and accessory glands that secrete
digestive juices into the canal through ducts.
The digestive tract has a four-layered wall: the lumen
is lined by a mucous membrane (mucosa),
then a connective tissue layer followed by a layer of smooth muscle, and outermost is a connective
tissue layer attached to the membrane of the body
cavity.
Peristalsis
(rhythmic smooth muscle contractions) pushes food along the tract.
Sphincters
(modifications of the muscle layer into ring-like valves) occur at some junctions between compartments and
regulate passage of materials though the system.
The accessory glands are: three pairs of salivary
glands, the pancreas, the liver, and the gall bladder.
Refer to Figure 37.9 for
orientation of the specialized compartments in the human system.
A. The Oral Cavity
Physical and chemical
digestion begin in the oral cavity.
Chewing breaks down large pieces of food into smaller
pieces.
This makes food easier to swallow and increases the
surface area available for enzyme action.
The presence of food in the
oral cavity stimulates the salivary glands to secrete saliva into the
oral cavity.
Saliva contain mucin (protects mouth from
abrasion and lubricates food), buffers that
neutralize acids, antibacterial agents, and salivary amylase - an enzyme
that hydrolyzes starch and glycogen
to the disaccharide maltose or small polysaccharides.
The tongue manipulates food
during chewing and forms it into a bolus which is swallowed.
The tongue also tastes the food and pushes the bolus
to the pharynx.
B. The Pharynx
The pharynx is an
intersection for both digestive and respiratory systems.
The movement of swallowing
moves the epiglottis to block the entrance of the windpipe (the glottis).
This directs food through the pharynx
and into the esophagus. (See Campbell, Figure 37.10)
C. The Esophagus
The esophagus is a muscular
tube which conducts food from the pharynx to the stomach.
Peristalis moves the bolus
along the esophagus to the stomach.
The initial entrance of the
bolus into the esophagus in voluntary (swallowing); once in, the peristalsis
results from involuntary contraction of the smooth muscles.
Salivary amylase remains
active as the bolus moves though the esophagus.
D. The Stomach
The stomach is a large,
saclike structure located just below the diaphragm on the left side of the
abdominal cavity. It functions in:
Food storage.
̃ The stomach has an elastic wall with rugae -
folds that can expand to accommodate up to 2 liters of food.
̃ Storage capacity permits periodic feeding (meals).
Churning.
̃ The stomach has longitudinal, vertical and diagonal
muscles that contract the stomach in churning movements that mix the food.
̃ Stomach contents are mixed every 20 minutes.
̃ Churning, mixing, and addition of stomach acid convert
food to a nutrient broth called acid chyme.
§ The passage of acid chyme into
the small intestine is regulated by the pyloric sphincter at the bottom
of the stomach.
§ The pyloric sphincter relaxes
at intervals and permits small quantities of chyme to pass.
Secretion. Gastric secretion is controlled by nerve
impulses and the hormone gastrin.
The stomach epithelium contains three types of secretory cells.
̃ Mucous cells secrete:
§ mucin - a thin mucus that protects the stomach lining from
being digested.
§ gastrin - a hormone produced
by the stomach. Gastrin is released
into the bloodstream and its action is to stimulate further secretion of
gastric juice (HCl and pepsin).
̃ Cheif cells secrete:
§ pepsinogen, an inactive protease or zymogen that is the
precursor to pepsin. Zymogen =
An inactive form of a protein-digesting enzyme.
̃ Parietal cells
secrete:
§ HCl
Protein digestion. Both components of gastric juice,
HCl, and pepsin, are involved with protein digestion:
̃ HCl - provides acidity (pH 1-4) which:
§ kills bacteria.
§ denatures protein.
§ starts the conversion of
pepsinogen to pepsin. Newly formed
pepsin can also catalyze this reaction:
Pepsinogen HCl ® Pepsin
̃ Pepsin - splits peptide bonds next to some amino
acids.
§ Does not hydrolyze protein
completely
§ Is an endopeptidase that
splits peptide bonds located within the polypeptide chain.
E. The Small Intestine
NOTE: A discussion of hormonal control of
digestion includes several examples of positive and negative feedback. This is a good place to reinforce the
concepts learned in Chapter 36 and to make a transition to the topic of the
endocrine system (Chapter 41).
The
human small intestine is about 6 m long in length and is the site of most
enzymatic hydrolysis of food and absorption of nutrients.
Remember, only limited
digestion of carbohydrates occurs in the oral cavity and esophagus (by salivary
amylase) and of proteins in the stomach (by pepsin).
The pancreas,
liver, gall bladder, and small intestine all contribute to the digestion
that occurs in the small intestine.
Their products are released into the duodenum, the first 25 cm of
the small intestine.
The pancreas produces:
̃ hydrolytic enzymes that break down all major classes
of macromolecules - carbohydrates, lipids, proteins, and nucleic acids.
̃ bicabonate buffer that helps neutralize the acid chyme
coming from the stomach.
The liver performs many
functions including the production of bile, which:
̃ is stored in the gall bladder.
̃ does not contain digestive enzymes.
̃ contains bile salts which emulsify fat.
̃ contains pigments that are byproducts of destroyed red
blood cells.
Hormonal
control of digestion involves the following regulatory hormones:
Gastrin - Released from the stomach in response to presence of
food; stimulates the stomach to release gastric juice (HCl and pepsin).
Secretin - Released from the duodenum in response to acid
chyme entering from the stomach; signals the pancreas to release bicarbonate
buffer to neutralize acid chyme.
Cholecystokinin (CKK) - Released from the duodenum in response to
chyme entering from the stomach; signals the gall bladder to release bile and
the pancreas to release pancreatic enzymes into the duodenum.
Enterogastrone - Released from the duodenum in response to the
presence of fat in the chyme; inhibits peristalsis in the stomach and slows
digestion.
Carbohydrate
digestion:
Begins with the action of
salivary amylase in the mouth.
Begins again in the duodenum
where pancreatic amylase hydrolyze starch and glycogen into
disaccharides.
Disaccharides attached to the surface of the duodenal epithelium
hydrolyze disaccharides into monosaccharides.
̃ Each disaccharide has its own disaccharidase. For example, maltose is hydrolyzed by maltase,
sucrose by sucrase, and lactose by lactase, etc.
̃ Since the disaccharidases are on the surface of the
epithelium, the final breakdown of carbohydrates occurs where the sugars will
be absorbed.
Protein
digestion involves the efforts of teams of enzymes:
Pepsin, and endopeptidase,
begins protein digestion in the stomach.
The pancreas secretes
proteases in the form of zymogens that will be activated only in the lumen of
the duodenum by the intestinal enzyme, enteropeptidase.
̃ Enteropeptidase converts trypsinogen to trypsin
Trypsinogen
enteropeptidase > Trypsin
̃ Trypsin catalyzes the conversion of more trypsinogen
to trypsin.
̃ Trypsin catalyzes the conversion of the other
zymogens.
Procarboxypeptidase trypsin
> Carboxypeptidase
Chymotrypinogen trypsin > Chymotrypsin
Trypsin and chymotrypsin
(endopeptidases) digest large polypeptides into shorter chains by breaking
internal peptide bonds adjacent to certain amino acids.
Carboxypeptidase
(exopeptidase) splits amino acids, one at a time, off the end of a polypeptide
that has a free carboxyl group.
The lining of the small
intestine also secretes protein-digesting enzymes, aminopeptidasei and
dipeptidases.
̃ Aminopeptidase beings at the end of a polypeptide that
has a free amino acid group and splits off one amino acid at a time.
̃ Dipeptides attached to the intestinal lining split
small polypeptides.
Since the protein-digesting
enzymes from the pancreas and small intestine break bond in specific areas of
the polypeptide, protein digestion to amino acids is a combined effort from all
of these enzymes.
Nucleic
acid digestion also involves teams of enzymes.
Nucleases hydrolyze DNA and RNA into nucleotides.
Other hydrolytic enzymes
(nucleotidases and nucleosidases) break nucleotides into nucleosides and
nitrogenous bases, sugars, and phosphates.
Fat
digestion occurs only in the small intestine, so most fat in food is undigested
when it reaches the duodenum. If fat is
present in chyme,
The duodenum secretes entergastrone,
a hormone that inhibits peristalsis in the stomach, slowing the entry of food
into the duodenum.
In response to CCK, the gall
bladder releases bile into the upper duodenum.
Bile salts emulsify the insoluble fat by coating fat droplets and thus
preventing them from coalescing.
Emulsification produces many small fat droplets that collectively
have a large surface area exposed for digestion.
Pancreatic lipase,
secreted into the duodenum, hydrolyzes fats into the building blocks, glycerol,
and fatty acids.
Macromolecules
are completely hydrolyzed as peristalsis moves the digestive juice-chyme
mixture through the duodenum. See Table
37.1 for a summary of digestion.
The remaining areas of the
small intestine, the jejunum and ileum, are specialized for
absorption of nutrients.
Nutrients
resulting from digestion must cross the digestive tract lining to enter the
body. While a small number of nutrients
are absorbed by the stomach and large and intestine, most absorption occurs in
the small intestine.
Large folds in the walls are
covered with projections called villi, which in turn have many
microscopic microvilli; this results in a surface area for absorption of
about 300 m2. (See Campbell,
Figure 37.14)
̃ This brush border (microvilli surface) is
exposed to the lumen of the intestine.
Penetrating the hollow core
of each villus are capillaries and a tiny lymph vessel called a lacteal.
Nutrients area absorbed by
diffusion or active transport across the 2 cell-thick epithelium and into the
capillaries pr lacteals.
̃ Amino acids and sugars enter the capillaries and are
transported by the blood.
̃ Absorbed glycerol and fatty acids are recombined in
epithelial cells to form fats; most are coated with proteins to form chylomicrons
which enter the lacteals.
Capillaries and veins
draining nutrients away from the villi converge into the hepatic portal vein,
which leads directly to the liver.
̃ Here various organic molecules are used, stored, or
converted to different forms.
̃ Blood flows at a rate of about 1 L per minute through
the hepatic vessel.
F. The Large Intestine
The
large intestine, or colon, connects to the small intestine at a T-shaped
junction containing a sphincter; the blind end of the T is called the cecum.
The appendix is a
fingerlike extension of the cecum and is composed of lymphoid tissue.
The colon is about 1.5
m long and is the shape of an inverted “U”.
Its major function is water re-absorption.
Feces (wastes of the digestive tract) are moved through the
colon by peristalsis.
̃ Intestinal bacteria live on organic material in the
feces and some produce vitamin K which is absorbed by the host.
̃ An abundance of salts in excreted by the colon lining.
̃ Feces are stored in the rectum and pass through
the two sphincter (one involuntary, one voluntary) to the anus for
elimination.
V. Vertebrate digestive systems exhibit many
evolutionary adaptations associated with diet
The
digestive systems of vertebrates are variations on a common plan, and many
adaptations associated with diet are found.
Variation
in the dentition (assortment of teeth) of mammals reflects the animal’s diet. See Campbell, Figure 37.15.
Carnivores (i.e. dogs and
cats) generally have pointed canines and incisors.
̃ These teeth are used to kill prey and rip away pieces
of flesh.
̃ Premolars
and molars are jagged and used to crush and shred the food.
Herbivores (i.e. cows) have
teeth with broad, ridged surfaces that are used for grinding vegetation.
̃ Incisors and canines are modified for biting off
pieces of vegetation.
Omnivores (i.e. humans) have
relatively unspecialized dentition.
̃ They are adapted to eat both vegetation and meat and
teeth similar to those of both herbivores and carnivores are found.
Non-mammalian
vertebrates typically have less specialized dentition, although there are
exceptions.
Poisonous snakes have fangs
which are teeth modified to inject venom into prey.
̃ Some fangs are hollow, others are grooved.
̃ Snakes can also swallow very large prey items due to
loosely hinged lower jaw-skull articulation.
A
correlation is also found between length of the vertebrate digestive system and
diet.
Herbivores and omnivores
have longer alimentary canals than carnivores relative to size.
The cell walls in vegetation
make it more difficult to digest than meat, and nutrients are less
concentrated.
The longer tract allows for
more time for digestion and provides a greater surface area for absorption.
The functional length
of an alimentary canal may be increased by increased folding as with the spiral
valve structure in sharks.
Special
fermentation chambers are present in the alimentary canals of many herbivores.
Symbiotic bacteria and
protozoa present in these chambers produce cellulase which can digest the
cellulose. (Animals do not produce
cellulase.)
The microorganisms digest
cellulose to simple sugars and convert the sugars to nutrients essential to the
animal.
The microorganisms may be
housed in the cecum (i.e. horses), cecum and colon (i.e. rabbit), or the much
more elaborate structure found in ruminants.
VI. An adequate diet provides
carbon skeletons for biosynthesis and essential nutrients
A
nutritionally adequate diet provides an animal with: fuel (chemical energy) for
cellular respiration, raw organic materials for biosynthesis, and essential
nutrients which must be obtained in prefabricated form.
A. Food as Fuel
Chemical
energy is obtained from the oxidation of complex organic molecules.
Monomers from any of the
complex organic molecules can be used to produce energy, although those from
carbohydrates and fats are used first.
Oxidation of a gram of fat
liberates 9.5 kcal, twice that of a gram of carbohydrate or protein.
The
basal energy requirements of an animal must be met to sustain their metabolic
functions.
When an animal takes in more
calories than are consumed, the liver and muscles store the excess in the form
of glycogen; further excess is stored in adipose tissue in the form of fat.
When the diet is deficient in
calories, glycogen stored in the liver and muscles is utilized first and fat is
then withdrawn from adipose tissues.
An undernourished
person or animal is one whose diet is deficient in calories.
̃ If starvation persists, the body begins to breakdown
its own proteins as a source of energy.
̃ The breakdown of the body’s own proteins can cause
muscles to atrophy and can result in the consumption of the brain’s proteins.
Obesity (overnourishment) is
a greater problem in the United States and other developed
countries
than undernourishment.
It increases the risk of heart
attack, diabetes, and other disorders.
B. Food for Biosynthesis
Heterotrophs
cannot use inorganic materials to make organic molecules; they must obtain
organic precursors for these molecules from the food they ingest.
Given
a source of carbon and nitrogen, heterotrophs can fabricate a great variety of
organic molecules by using enzymes to rearrange the molecular skeletons of
precursors acquired from food.
A single type of amino acid
can supply the nitrogen necessary to build other amino acids.
Fats can be synthesized from
carbohydrates.
The liver is responsible for
most of the conversion of nutrients from one type of organic molecule to
another.
C. Essential Nutrients
An
animal’s diet must include essential nutrients in addition to providing
fuel and carbon skeletons.
Essential nutrients =
Chemicals an animal requires but cannot synthesize.
Vary from species to species.
An animal is malnourished
if its diet is missing one or more essential nutrients.
Includes essential amino
acids, essential fatty acids, vitamins, and minerals.
Essential amino acids are
those that must be obtained in the diet in a prefabricated form.
Most animals can synthesize
about half of the 20 kinds of amino acids needed to make proteins.
Human adults can produce 12,
leaving 8 as essential in the diet.
(Human infants can only produce 11.)
Protein deficiency results when the diet lacks one or more essential
amino acids.
̃ The syndrome known as kwashiorkor is a form of
protein deficiency in some parts of Africa.
The human body cannot store
essential amino acids, thus a deficiency retards protein synthesis.
̃ This is most frequent in individuals, who for economic
or other reasons, have unbalanced diets.
Essential fatty acids are
those unsaturated fatty acids that cannot be produced by the body.
An example in humans in
linoleic acid which is required to produce some of the phospholipids necessary
for membranes.
Fatty acid deficiencies are
rare as most diets include sufficient quantities.
Vitamins are organic
molecules required in the diet in much smaller quantities (0.01 - 100 mg/day)
than essential amino acids or fatty acids.
Many serve a catalytic
function as coenzymes are parts of coenzymes.
Vitamin deficiencies can cause
very severe effects as shown in Table 37.2.
Water-soluble vitamins are not
stock-piled in the body tissues; amounts ingested in excess of body needs are
excreted in the urine.
Fat-soluble vitamins (vitamins
A, D, E, and K) can be held in the body; excess amounts are stored in body fat
and may accumulate over time to toxic levels.
If the body of an animal can
synthesize a certain compound, it is not a vitamin.
̃ A compound such as ascorbic acid is a vitamin for
humans (vitamin C) and must be included in our diets; it is not a vitamin in
rabbits where the normal intestinal bacteria produce all that is needed.
Minerals are inorganic
nutrients required in the diet in a small quantities ranging from 1 mg to 2500
mg per day, depending on the mineral.
Some minerals serve structural
and maintenance roles in the body (calcium, phosphorous) while others serve as
part of enzymes (copper) or other molecules (iron).
Refer to Table 37.3 for the
mineral requirements of humans.