 Nutrition : includes all of those processes by which an animal

takes in, digests, absorbs, stores, and uses food (nutrients) to

meet its metabolic needs
 Digestion: is the chemical and/or mechanical breakdown of food
into particles that individual cells of an animal can absorb.
 Intracellular digestion: In some of the simplest forms of life (the
protists and sponges), some cells take in whole food particles
directly from the environment by diffusion, active transport,
and/or endocytosis and break them down with enzymes to obtain
nutrients.
 Extracellular digestion: the enzymatic breakdown of larger pieces
of food into constituent molecules, usuallynin a special organ or
cavity . Nutrients from the food then pass into body cells lining the
organ or cavity and can take part in energy metabolism or
biosynthesis.
 Intracellular digestion occurs in some invertebrates (e.g., in
sponges); others (e.g., some Cnidarians and molluscs) utilize both
intracellular and extracellular digestion; and most higher
invertebrates (e.g., insects) have evolved variations in
extracellular digestion that allow them to exploit different food
sources .
 In primitive, multicellular animals, such as cnidarians, the gut is a
blind (closed) sac called a gastrovascular cavity. Its one opening is
 both entrance and exit; thus, it is an incomplete digestive tract.
Some specialized cells in the cavity secrete digestive enzymes that
begin the process of extracellular digestion. Other phagocytic cells
that line the cavity engulf food material and continue intracellular
digestion inside food vacuoles. Some flatworms have similar
digestive patterns
 The development of an anus and complete digestive tract in the
aschelmenthis was an evolutionary breakthrough. A complete
digestive tract permits the one-way flow of ingested food without
mixing it with progressive digestive processing in specialized
regions along the system.
Protozoa : Protozoa may be autotrophic, saprozoic or heterotrophic
(ingest food particles).Ciliated protozoa are good examples of
protists that utilize heterotrophic nutrition. Ciliary action directs food
from the environment into the buccal cavity and cytostome. The
cytostome opens into the cytopharynx, which enlarges as food and
pinches off a food-containing vacuole.
 The detached food vacuole then moves through the cytoplasm.
During this movement, excess water is removed from the vacuole,
the contents are acidified and then made alkaline and a lysosome
adds digestive enzymes. The food particles are then digested
within the vacuole and the nutrients absorbed into the cytoplasm.
The residual vacuole then excretes its waste products via the
cytophyge.

Porifera:
 Pore bearing animals
 Ex. Sponges
 Asymmetrical or radially symmetrical, three types of cells
pinacocytes, mesenchyme cells and choanocytes, central cavity or
series of branching chambers through which water circulates
during filter feeding, no tissues or organs.
 Thin flat cells called pinacocytes line the outer surface of a
sponge. Pinacocytes may be mildly contractile and their contraction
may change the shape of some sponges. In a number of sponges
some pinacocytes are specialised into tube like contractile
porocytes, which can regulate water circulation. Openings through
parasites are pathways for water moving through body wall. Just
below the pinacocytes layer of sponge is a Jelly like layer called
mesohyl . Ameboid cells called mesenchyme cells move about in
mesohyl and are specialized for reproduction, secreting skeletal
elements, transporting and storing food and forming contractile
rings around openings in the sponge wall. Below mesohyl and lining
of inner chambers are choanocytes or collar cells. Choanocytes are
flagellated cells that have collar like ring of microvilli surrounding a
flagellum. Microfilaments connect the microvilli forming a net like
mesh within the collar. The flagellum creates water currents
through the sponge and collar filters microscopic food particles
from the water. Water currents bring food and oxygen to sponge
and carry away metabolic and digestive wastes.
 Ascon sponges are vaselike. Ostia are outer openings of
 porocytes and lead directly to a chamber called spongocoel.
Choanocytes line the spongocoel and their flagellar movements
draw water into the spongocoel through ostia. Water exits the
sponge through osculum which is a single large opening at top of
sponge.
 In the sycon body form sponge wall appears folded. Water enters
a sycon sponge through openings called dermal pores. Dermal
pores are the openings of invaginations of the body wall called
incurrent canals. Pores in the body wall connect incurrent canals
to the radial canals and the radial canals lead to spongocoel.
Choanocytes line radial canals and the beating of choanocyte
flagella moves water from ostia through incurrent and radial
canals to spongocoel and out of osculem.
 Choanocytes filter small suspended food particles. Water passes
through their collar near the base of cell and then moves into sponge
chamber at the open end of collar. Suspended food is trapped on the
collar and moved along microvilli to the base of collar where it is
incorporated into a food vacuole. Digestion begins in food vacuole by
lysosomal enzymes and pH changes. Partially digested food is passed
to amoeboid cells which distribute it to other cells. Filtration is not
the only way that sponge feed. Pinacocytes lining incurrent canals
may phagocytize large food particles . Sponges may absorb by active
transport nutrients dissolved in seawater. Because of extensive canal
system and the circulation of large volumes of water through
sponges, all sponge cells are in close contact with water. Thus
nitrogenous waste removal and gas exchange occur by diffusion.

Cnideria:
 Radial or bilateral symmetry, diploblastic or tissue level
organization, gelatinous mesoglea between epidermal and gastro
dermal tissue layers, gastrovascular cavity, specialized cells called
cnidocytes used in defence feeding and attachment.
 The gastrodermis of all cnidrians lines a blind ending
gastrovascular cavity. This cavity functions in digestion, the
exchange of respiratory gases and metabolic wastes and discharge
 of gametes. Food, digestive wastes and reproductive stages enter
and leave the gastrovascular cavity through mouth
 The food of most cniderians consist of very small crustaceans,
although some cniderians feed on small fish. Nematocysts
entangle and paralyze prey and contractile cells in the tentacles
cause the tentacles to shorten which draws food toward the
mouth. As food enters the gastrovascular cavity, gastrodermal
glands cell secrete lubricating mucus and enzymes, which reduce
food to a soupy broth.
 Certain gastro dermal cells called nutritive muscular cells
phagocytize partially digested food and incorporate it into food
vacuole where digestion is completed. Nutritive muscular cells
also have circularly oriented contractile fibers that help move
materials into and out of the gastrovascular cavity by peristaltic
contractions
 During peristalsis, ringlike contractions move along the body wall
pushing contents of the gastrovascular cavity ahead of them,
expelling undigested material through the mouth. In cniderians,
the water filled gastrovascular cavity acts as a hydrostatic
skeleton. Cniderians have large surface area to volume ratio, all
cells are a short distance from the body surface and oxygen,
carbon dioxide and nitrogenous wastes exchange by diffusion.
Platyhelmenthis:
 Usually flattened dorsoventrally, triploblastic, acoelomate,
bilaterally symmetrical, unsegmented worms, incomplete gut
usually present but absent in cestoidea, highly branched
digested systems are an advancement that result in more
gastrodermis closer to the sites of digestion and absorption,
reducing the distance nutrients must diffuse.
 Both type of digestion is present but it is more expanded than
cniderians. Extracellular digestion takes place in gastrovascular
cavity by enzymatic secretions while intracellular digestion
takes place by phagocytic cells present in inside lining of
platyhelminthes.
 The turbellarian pharynx functions as an ingestive organ.It varies
in structure from a simple ciliated tube to a complex organ
developed from the folding of muscle layers. Most turbellarians
 such as common planarian are carnivores and feed on small live
invertebrates or scavenge on larger dead animals; some are
herbivores and feed on algae that they scrape from rocks.
Sensory cells (chemoreceptors) on their heads help them detect
food from a considerable distance. Food digestion is partially
extracellular. Pharyngeal glands secrete enzymes that help
breakdown food into smaller units that can be taken into the
pharynx . In the digestive cavity digestion is completely in
intracellular vesicles.
Mollusca:
 Mantle cavity functions in excretion, gas exchange, elimination of
digestive wastes and release of reproductive products, bilateral
symmetry. Most gastropods feed by scrapping algae or other
small attached organisms from their substrate. Others are
herbivores that feed on larger plants, scavengers parasites or
predators.
 The anterior portion of digestive tract may be modified into an
extensible proboscis, which contains radula. This structure is
important for some predatory snails that must extract animal
flesh from hard to reach areas. The digested tract of gastropods
like that of most molluscs is ciliated. Food is trapped in the mucus
strings and incorporated into a mucoid mass called protostyle
which extends to the stomach and is rotated by cilia. A digestive
gland in the viseral mass releases enzymes and acid into the
 stomach and food trapped on the protostyle is freed and digested.
Wastes from fecal pellets in the intestine.

 Cilia covering leaflike labial palps on either side of mouth also

sort filtered food particles. Cilia carry small particles into the
mouth and move larger particle to the edges of the palps and
gills. This rejected material called pseudofeces falls or is thrown
onto the mantle and the ciliary tract on the mentle transports
the pseudo feces posteriorly. Water rushes out when valves are
forcefully closed washes pseudofeces from the mantle cavity
 The digestive tract of bivalves is similar to that of other molluscs.
Food entering the esophagus entangles in a mucoid food string,
 which extends to the stomach and is rotated by cilia lining the
digestive tract. A consolidated mucoid mass,the crystalline style
project into the stomach from a diverticula called style sac.
Enzymes for the carbohydrate and fat digestion are incorporated
into the crystalline style . Cilia of style sac rotate the style against
chitinized gastric shield. This abrasion and acidic condition in
stomach dislodge enzymes.
 The mucoid food string winds around the crystalline style as it
rotates, which pulls the food string farther into the stomach from
esophagus. This action and acidic pH in the stomach dislodge food
particles in food string. Further sorting separates fine particles
from undigestible course material. The latter are sent to the
intestine. Partially digested food from stomach enters a digested
gland for intracellular digestion. Cilia carry undigested wastes in
the digestive gland back to the stomach and then to intestine.
The intestine empties through the anus near the excrunt opening
and excurrent water carries feces away.
 All cephalopods have Jaws and a radula. The jaws are powerful
beak like structures for tearing food and radula rasps food, forcing
it into mouth cavity.
 Many bivalve molluscs suspension feed and ingest small food
particles. The digestive tract has a short esophagus opening into a
stomach, midgut, hindgut, and rectum. The stomach contains a
crystalline style, gastric shield, and diverticulated region. These
 diverticulae are blind-ending sacs that increase the surface area
for absorption and intracellular digestion. The midgut, hindgut,
and rectum function in extracellular digestion and absorption.
 Digestion is a coordination of three cycles: (1) feeding,
(2)Extracellular digestion, and (3) intracellular digestion. The
resting phase is preparative for extracellular digestion. The
mechanical and enzymatic breakdown of food during feeding
provides the small particles for intracellular digestion. Intracellular
digestion releases the nutrients into the blood and produces the
fragmentation spherules that both excrete wastes and lower the
pH for optimal extracellular digestion. These three cycles are
linked to tidal immersion and emersion of the mollusc.

Aschelmenthis:

 The aschelmenthis are the first assemblage of animals to possess

a distinct body cavity, but they lack the peritoneal linings and
membranes called mesenteries found in more advanced animals.
As a result the various internal organs lie free in the cavity. Such a
cavity is called pseudocoel and the animals are called
pseudocoelomate . The pseudocoelom is often fluid filled or may
contain a gelatinous substance with mesenchyme cells, serves as a
cavity for circulation, aids in digestion and acts as an internal
hydrostatic skeleton that functions in lolocomotion
 Most aschelmenthis have complete tubular digestive tract that
extend from an anterior mouth to posterior anus. This complete
digestive tract was first encountered in nemerteans and is
characteristic of almost all other higher animals. It permits for the
first time, the mechanical breakdown of food, digestion,
absorption and feces formation to proceed sequentially and
continually from an anterior to posterior direction and
evolutionary advancement over blind ending digestive system.
Most also have specialized muscular pharynx that is adapted for
feeding.
 Most rotifers feed on small microorganisms and suspended
organic material. The coronal cilia create a current of water that
brings food particles to the mouth. The pharynx contain a unique
structure called Mastax(jaws) which is a muscular organ that
grinds food. The inner wall of mastax contain several sets of jaws
called trophi. The trophi very in morphological detail and
taxonomists use them to distinguish species.
  From the mastax food passes through a short ciliated esophagus
to ciliated stomach. Salivary and digestive glands secrete
digestive enzymes into the pharynx and stomach. Complete
extracellular digestion and absorption of food occurs in the
stomach
 . In some species a short ciliated intestine extends posteriorly and

becomes a cloacal bladder which receives water from the

protonephridia and eggs from the ovaries, as well as digestive
waste. The cloacal bladder opens to the outside via an anus at
junction of foot with the trunk.
 Nematodes have a complete digestive system consisting of a
mouth, which may have teeth, jaws or stylets, buccal cavity,
 muscular pharynx, long tubular intestine where digestion and
absorption occur, short rectum and anus. Hydrostatic pressure in
pseudocoelom and the pumping action of pharynx push food
through alimentary canal.

Annelida:
 Body metameric, bilaterally symmetrical and worm like. The
anterior region of digestive tract is modified into a proboscis
that special protractor muscles and coelomic pressure can evert
through the mouth. Retractor muscles bring the proboscis back
into the Peri-stomium. In some, when proboscis is everted,
paired jaws are opened and may be used for seizing prey.
Predatory Polycheates may not leave their Burrow or coral
crevice. When prey approaches a burrow entrance, the worm
quickly extends its anterior portion, everts the proboscis and
pulls the prey back into the burrow. Some Polycheates have
poison glands at the base of jaw. Other Polycheates are
herbivores and scavengers and use jaws for tearing food.
Deposit feeding polycheats extract organic matter from the
marine sediments they ingest.
 The digestive tract consist of pharynx that when everted forms
the proboscis and a storage sac called crop, a grinding gizzard and
long straight intestine. These are similar to the digestive organs of
earthworm. Organic matter is digested extracellularly and
inorganic particles are passed through the intestine and released
as castings.
 Many sedentary and tube dwelling Polycheates are filter feeders.
They usually lack a proboscis but possess other specialised feeding
structures. Some tube dwellers called fan worms possess radioles
that form a funnel shaped fan. Cilia on the radioles circulate
water through the fan ,trapping food particle. Trapped particles
are carried along a food groove at axis of radiole. During
transport, a sorting mechanism rejects the largest particle and
transport the finest particle to the mouth.
 Oligochaetes are scavengers and feed primarily on fallen and
decaying vegetation which they drag into their burrows at night.
The digestive tract of oligochaetes is tubular and straight. The
mouth leads to muscular pharynx. In the earthworm,
pharyngeal muscles attach to the body wall. The pharynx act as
a pump for ingesting food. The mouth pushes against the food
and pharynx pumps the food into esophagus which is a narrow
and tubular and frequently expands to form a stomach, crop
and gizzard; the latter two are common in terrestrial species. A
crop is a thin walled storage structure and gizzard is a muscular
cuticle lined grinding structure.
 Many Leeches feed on the body fluids or the entire bodies of
other invertebrates. Some feed on the blood of vertebrates
including human blood. Leeches are sometimes called parasites.
The mouth of leech opens in the middle of anterior sucker. In
some leeches, the anterior digestive tract is modified into
protrusible proboscis, lined inside and outside by a cuticle. In
others the mouth is armed with three chitinous jaws. While
feeding a Leech attaches to its prey by anterior sucker and
 either extend its proboscis into the prey or use its jaws to slice
through the host tissues . Salivary glands secrete an
anticoagulant called hirudin that prevents blood from clotting.
Behind the mouth is a muscular pharynx that pumps the body
fluids of the prey into the Leech. The oesophagus follows the
pharynx and leads to a large stomach with lateral cecae. Most
leeches ingest large quantities of the blood or other body fluids.
Echinoderms:

The water vascular system is a series of water filled canals and

their extensions are called tube feet. It includes a ring canal
that surrounds the mouth. The ring canal really opens to the
outside or to the body cavity through a stone canal and sieve
like plate called madreporite which may serve as inlet to
replace water lost from the water vascular system and may
help equalize pressure differences between water vascular
system and outside. Five radial canals branch from ring canal.
Radial canals are associated with arms of star shaped
echinoderms.

 Tube feet are extensions of canal system and usually emerge

through openings in skeletal ossicles. When an ampulla
contract it forced water into the tube feet which then
extends. Valves present prevent the backflow of the water
from the tube feet into the lateral canal. A tube feet often
has a suction cup at distal end. When the food extends and
contracts solids substrate, muscles of the suction cup
contract and create a vacuum . The function of water
vascular system was probably feeding not locomotion.
 The stomach is divided into two regions, larger oral
stomachs sometimes called cardiac stomach release receive
ingested food. it joins smaller aboral stomach sometimes
called pyloric stomach. The aboral stomach give rise to ducts
that connect secretary and absorptive structural pyloric
cecae. Two pyloric cecae extend into each arm. A short
intestine lead into rectal caeca and to nearly nonfunctional
anus which opens on the aboral surface of central disc.
  Some Seastar ingest whole prey which are digested
extracellularly within stomach. Undigested material is
expelled through the mouth.

Arthropods:
 The grasshopper is a representative insect with a complete
digestive tract and extracellular digestion. During feeding,
the mandibles and maxillae first break up (masticate) the
food, which is then taken into mouth and passed to the crop
via the esophagus. During mastication, the salivary glands
add saliva to the food to lubricate it for passage through the
digestive tract.
 Saliva also contains the enzyme amylase, which begins the
enzymatic digestion of carbohydrates. This digestion
continues during food storage in the crop. The midgut
secretes other enzymes (carbohydrases, lipases, proteases)
that enter the crop. Food passes slowly from the crop to the
stomach, where it is mechanically reduced and the nutrient
particles sorted.
  Large particles are returned to the crop for further
processing; the small particles enter the gastric cecae, where
extracellular digestion is completed. Most nutrient
absorption then occurs in the intestine. Undigested food is
moved along the intestine to the rectum, where water and
ions are absorbed. The solid fecal pellets that form then pass
out of the animal via the anus.
 During this entire feeding process the nervous system, the

endocrine system, and the presence of food exert

considerable control over enzyme production at various
points in the digestive tract.
Key points:
1. Protozoa:
 Simplest unicellular
 Intracellular digestion
 Simple diffusion
 Autotrophic, saprozoic or heterotrophic
 Ciliary action pushes food to buccal cavity, cytosome and
cytopharynx
 Food vacuole formation
 Removal of excess water
 Lysosomes attatch and provide digestive enzymes
 pH is alkaline
 digestion and absorption of food
 residual vacuole removal by exocytosis.
2. Porifera:
 Pore bearing organisms
 Ex: Sponges
 Intracellular digestion
 Simple digestion
 Filter feeding
 Ostia : water enters
 Osculum : water leaves
 Spongocoel : Cavity
 Pinacocytes : regulate water circulation
 Choanocytes : flagellated collar cells , receive food
 Amebocytes : food digestion, reproduction
3. Cnideria:
 Ex : Hydra
 Bilateral symmetry
 Diploblastic
 Tissue level of organization
 Both intracellular and Extracellular digestion
 Intracellular : inside cells
 Extracellular : In gastrovascular cavity
 Sac like digestive system
 Cnidocytes : defense, feeding, attachment.
 Nematocysts: prey paralysis, food ingestion
 Peristalsis: removal of wastes
4. Platyhelminthes:
 Flatworms e.g. Planaria
 Dorsoventrally flattened
 Triploblastic, acoelomate, bilateral symmetry
 Sac like (incomplete) digestive system
 Both intracellular and Extracellular digestion
 Intracellular : inside cells
 Extracellular : in gastrovascular cavity
 Carnivores, scavengers or herbivores
 Sensory cells(chemoreceptors) : food detection
5. Mollusca:
 Ex: snails, mussels.
 Bilateral symmetry
 Herbivores, scavengers or predators.
 Tube like(complete) digestive system
 Bivalve molluscs: Both intracellular and Extracellular digestion
 Proboscis containing radula
 waste remove as Pseudofeces
 Stomach parts: crystalline style, gastric shield, diverticulae
 Digestion cycle: Feeding, Extracellular dig, intracellular dig.
6. Aschelmenthis:
 round worms eg.nematodes
 tube like digestive system
 pseudocoelomates
 pseudocoelom: cavity for digestion and locomotion
 Development of anus
  Rotifers: Mastax contain trophi.
 Salivary and digestive glands
 Stomach: Extracellular digestion
7. Annelida
 Body metameric, bilaterally symmetrical
 Proboscis, peri-stomium
 Polycheates: herbivores, scavengers ,filter feeding
 Poison glands
 Proboscis, pharynx, crop, gizzard, intestine
 Oligochaetes: scavengers
 Leeches: parasites, proboscis, Salivary glands, hirudin
8. Echinoderms:
 Ex . starfish, seastar
 water vascular system
 tube feet
 central ring canal
 five radial canals
 sieve like plate ; madreporite
 Stomach: cardaic , pyloric
9. Arthropods:
 Ex: insects
 Complete digestive system
 Mouth , crop, oesophagus, stomach, intestine, rectum
 Saliva: lubrication and digestive enzymes
 Amylase, lipase, protease
 Stomach: Extracellular digestion
  Intestine: absorption
 Rectum: removal of solid fecal pellets.

Digestive Structures In Vertebrates:

Embryonic Digestive Tract:

Archenteron:
The embryonic archenteron becomes the lining of the adult digestive
tract and of all its derivatives. Splanchnic mesoderm adds layers of connective
tissue and smooth muscles around the archenteron. Ectodermal invagination of
the head forms the stomodaeum leading into oral cavity, and a similar mid-
ventral ectodermal invagination forms proctodaeum, which leads into the
hindgut. The stomodaeum becomes the adult buccal cavity and gives rise to
teeth enamel, epithelial covering of tongue, glands, e.g., mucous, poison and
salivary, etc., and Rathke’s pouch of anterior pituitary gland. The proctodaeum
forms either a small terminal part of the cloaca in lower vertebrates and rectum
in mammals.

Figure 1: Alimentary canal and its chief derivatives in vertebrates

The alimentary canal in embryos from stomach to cloaca is attached to the

dorsal body wall by a double fold of peritoneum, called the dorsal mesentery,
and to ventral body wall by a ventral mesentery. In adults, dorsal mesentery
persists but the ventral mesentery disappears leaving only in the region of liver
and urinary bladder.

Digestive Tract of Adult:

The digestive tract differentiates for different functions into the
following regions- mouth, buccal cavity, pharynx, oesophagus, stomach, small
intestine, large intestine and cloaca. Following outgrowths arise from the
digestive tract- oral glands, Rathke’s pouch, thyroid gland, gill-clefts, tympanic
cavity, thymus and other glands of gill-clefts, trachea, lungs, swim bladder,
liver, pancreas, yolk sac, and urinary bladder.

Histology:
The wall of the alimentary canal is made of four concentric layers. They are:
 (i) An outermost visceral peritoneum or serous coat is made of
mesothelial cells and thin layer of connective tissue. It is lacking in the
oesophagus,

(ii) Below this is a muscular layer formed of smooth muscle fibres

arranged in outer longitudinal and inner circular muscle fibres.
Between the two layers of muscles is a network of nerve cells and
nerve fibres of the autonomic nervous system, known as myenteric
plexus or plexus of Auerbach.

(iii) Beneath the muscle layer is a submucosa made of connective tissue

having elastic fibres, fat, blood and lymph vessels, nerve cells and
fibres glands,

(iv) The innermost layer is a mucosa composed of three regions:

(a) Outer-most narrow muscularis mucosa of outer longitudinal and
inner circular smooth muscle fibres.

(b) Middle thin layer of lamina propria of connective tissue, blood

vessels, nerves and nodules of lymphatic tissue, and

(c) A basement membrane supporting a layer of columnar epithelial

cells which are often glandular and ciliated.

 Mouth:
Mouth is the opening leading into buccal cavity. In lampreys
(cyclostomes) it is a circular opening at the base of buccal funnel and remains
permanently open due to lack of jaws, etc. In gnathostomes it is terminal. Mouth
is bounded by lips which are immovable and formed of cornified skin in fishes,
amphibians and reptiles. In mammals these are fleshy and muscular.
  Buccal Cavity:

The space between the lips and the jaws is a vestibule. It may be
bounded on the outside by cheeks and on the inside by the gums. Mucous
glands of cheeks open into the vestibule. The mouth opens into a buccal
cavity, which is a space between the mouth and the pharynx. The exact
point where the stomodaeal ectoderm and pharyngeal endoderm merge is
variable and not easy to discern.

In elasmobranchs and most bony fishes the nasal cavities do not open into the
buccal cavity. In Chondrichthyes and tetrapoda (amphibians and most reptiles)
the nasal cavities open into the buccal cavity by choanae or internal nares,
which are primitively placed anteriorly, but in crocodiles, birds and mammals
they become posterior in the pharynx due to the formation of a secondary
palate, which effectively separates the respiratory nasal passage from the mouth
cavity or food passage.

In birds, this palate is cleft due to which nasal and buccal cavities communicate
with each other. In mammals, secondary palate is continued posteriorly as a
membranous soft palate. In human beings soft palate hangs into the laryngeal
pharynx in the form of fleshy process, called uvula.

Figure 2: Relation of nasal passages to buccal cavity in vertebrates

 Tongue:
 A true tongue, which has voluntary muscle and is movable, is found
in amphibian animals. As summarized by Youson (1981), a tongue-like
piston is found in the oral opening of the adult lamprey but this organ is
not homologous to the tongues of gnathostomes. When the lamprey grips
its prey with its oral disc, the teeth on the piston abrade and cut the host
tissue. The functional role of the piston is very similar to that of the
tongue of some tetrapods. The precursors to tongue muscle in amniotes
have been shown to be derived from the occipital somites. The muscle
precursors lose the features of epithelial cells and emigrate as a
condensed stream of cells, known as the hypoglossal cord.

Birds live in the air, on land, and on and around fresh water and sea water.
However, keratinization of the lingual epithelium is a common feature, in
particular on the ventral side of the tongue, where the so-called ‘lingual nail’ is
prominent in all species examined. In the ancestors of birds, the lingual
epithelium might have become adapted to dry conditions. In birds, the lingual
papillae also play an important role in feeding, and birds that eat hard foods
have structures similar to teeth in their upper and lower beaks.

Figure 3: Types of tongues in some vertebrates

In mammals the mucous membrane below the tongue forms a median fold,
called frenulum which joins the tongue to the floor of the mouth. In mammals
the upper surface of the tongue bears four kinds of papillae, (filiform,
fungiform, foliate and circumvallate), bearing taste buds except filiform
papillae.

Figure 4: Dorsal view of human tongue

 Teeth:
Vertebrates have two types of teeth attached to jaw bones- epidermal
teeth and true teeth. Epidermal teeth are best developed in cyclostomes. They
are hard, conical, horny structures derived from the stratum corneum. In
lampreys they are found on the walls of the buccal funnel and on the tongue.
Tadpole larvae of frogs and toads have serrated epidermal teeth in rows on the
lips. In mammals the adult duckbill platypus has epidermal teeth.

True Teeth:
Teeth are not found in baleen whales and anteaters in mammals, and agnathans,
sturgeons, some toads, sirenians, turtles and modern birds, etc. In lower
vertebrates (such as fish, amphibians and most reptiles) teeth may be replaced
continually an indefinite number of times, such teeth are called polyphyodont.
These teeth are homodont (similar type) and acrodont. (with the jaw bones).
In most mammals teeth are diphyodont, thecodont and heterodont. In some
mammals these are monophyodont having only one set of teeth, e.g., moles,
Indian squirrel. Teeth are similar in structure to the placoid scales of sharks,
formed of a central pulp cavity, around which is present a thick but soft layer,
the dentine, which is externally covered by a thin, extremely hard enamel. These
are supposed to have derived from bony scales of ostracoderms and placoderms.
For details readers may see the Dentition in Mammals.

Figure 5: Teeth in vertebrates

 Pharynx:
The part of the alimentary canal immediately behind the buccal cavity
is a pharynx, lined with endoderm. It is a common passage serving both for the
digestion and respiration. As a part of the digestive system it is used as a
passage for food from the buccal cavity to the oesophagus, its muscles initiate
swallowing.

In fishes, the pharynx is large and laterally perforated for gill-slits, while in
tetrapoda, it is short and bears openings of nostrils. In embryos, the wall of
pharynx gives off a number evaginations which develop into spiracles, gill-
clefts, air bladders, lungs, tonsils and a few endocrine glands (e.g., thymus,
thyroid and parathyroids).

 Oesophagus:
 The oesophagus is short in most fishes and amphibians because they
lack neck, but it is longer in amniotes due to presence of neck. The oesophagus
of reptiles is more elongate than that of fishes and amphibians. In granivorous
and carnivorous birds, a portion of the oesophagus is enlarged into a sac-like
pouch called crop which serves to store food which has been eaten quickly.

The crop is essentially lacking in digestive glands although in pigeons the crop
has 2 crop glands in both sexes, they are really not glands but cell-forming
structures, the cells form ‘pigeon milk’ which is fed to the young. In mammals,
the oesophagus is long, lacks glands and varies in relation to the length of neck.
It passes through the diaphragm, the portion below the diaphragm is covered
with visceral peritoneum which is lacking from the upper part. Oesophagus has
mucous glands. Its lining forms longitudinal folds, or finger-like fleshy papillae
(elasmobranchs) or horny papillae in marine turtles.

Histologically, the oesophagus differs from the rest of the alimentary canal
in three facts:
(i) It has no visceral peritoneum because it lies outside the coelom, its outermost
covering layer is a thin tunica adventitia.

(ii) The muscle fibres in its anterior part are striped, middle part has both striped
and unstriped muscles, and the posterior part has only unstriped muscles. But
there are exceptions in ruminating mammals, all the muscles are striped or
voluntary.

(iii) The mucous membrane lining is made of stratified squamous epithelial

cells and not of columnar cells.

 Stomach:
There is practically no stomach in cyclostomes, chimeras, lung fishes
and some primitive teleost fishes, since it has no gastric glands, but in most
fishes and tetrapoda it is dilated for storage and maceration of solid food, and
digestion of food because it contains gastric glands.

The first part of the stomach, next to the oesophagus, is the cardiac region and
the lower end near the intestine is the pyloric region, which has a pylorus or
pyloric valve in which the mucous membrane lining is surrounded by a thick
sphincter muscle which regulates the opening and closing of the pyloric
stomach into the intestine.

Stomach is straight in cyclostomes, gar, Belone, etc., and spindle-shaped in

Proteus, Necturus, some lizards and snakes. In turtles and tortoises, it is a wide
curved tube, and in elasmobranchs the stomach is J-shaped. In crocodiles and
birds the stomach has two parts, a proventriculus with gastric glands, and a
highly muscular gizzard, which represents the pyloric region and has a hard,
cornified lining for grinding food. In mammals the stomach lies transversely
and may be a simple sac or divided into 3 regions, namely cardiac, fundic and
pyloric and each region has its gastric glands. In many ruminants the stomach
has four chambers- a rumen, reticulum, omasum and an abomasum. It is
claimed that the first three chambers are modifications of the oesophagus, and
abomasum is the true stomach representing the cardiac, fundic, and pyloric parts
of the stomach.

It has been shown embryologically that all four chambers are modified regions
of the stomach. In camels, there is no omasum, the rumen and reticulum have
pouch-like water cells which were once believed to store water, but they are
probably digestive.
 Figure 6: Digestive track of vertebrates

Histologically, the stomach has the typical parts of the alimentary canal, but it
has two peculiarities, the muscularis mucosa is made of an outer longitudinal
layer and an inner circular layer of muscles. The epithelium lining is thick with
several types of glandular cells forming gastric glands of three types called
cardiac, fundic, and pyloric gastric glands.The cardiac and pyloric glands
secrete only mucus from their surface cells. Fundic glands (or cardiac glands in
some) have three kinds of cells, mucous neck cells produce mucus, oxyntic cells
produce hydrochloric acid, they may be present in the cardiac region also,
zymogen cells or peptic cells secrete pepsin.

In most animals the zymogen cells also secrete two proenzymes called
propepsin and prorennin which are converted by hydrochloric acid into pepsin
and rennin respectively. The secretions of all stomach cells form a mixture
called gastric juice.

 Small Intestine:
Small intestine is long, narrow and coiled tube after the pylorus. It is
the most important part of the digestive tract because the digestion and
absorption of food take place in it. In cyclostomes, the intestine is a short
straight tube with a spirally arranged longitudinal flap extending into it.

In elasmobranchs, it is divided into small and large portions, and the small
portion has a spiral valve which greatly increases the absorptive surface. A
spiral valve is also present in the small intestine of a few more primitive bony
fishes, but is lacking in higher forms in which the intestine is long and coiled.

In caecilians, it is little coiled and not differentiated into a small and large tract.
In frogs and toads it is relatively long and coiled. In reptiles, it is more coiled
than in amphibians. For the first time in vertebrates a caecum or blind
diverticulum arises at the junction of small and large intestines.However, this is
not permanent in all reptiles. In birds, the small intestine is coiled or looped and
one or two colic caeca are also present at the junction of small and large
intestines. In most mammals also the small intestine is proportionately long and
coiled. Its length, however, is correlated with food habits. In herbivores it is
relatively more longer in comparison to insectivores and carnivores.

There is a blind pocket or caecum at the junction of the colon and small
intestine which is generally small in carnivorous species and quite long in many
herbivores. The first part of the small intestine is the duodenum, which is short
beginning from pylorus and terminates beyond the entrance of pancreatic and
hepatic ducts.It has many folded villi and contains branching Brunner’s glands
in the submucosa which secrete mucus, some alkaline watery-fluid, and a little
enzyme. Duodenum also produces two hormones called secretin and
cholecystokinin which stimulate the pancreas and gall bladder to liberate their
juices. Ducts from the gall bladder and pancreas open into the duodenum.

Behind the duodenum is an ileum, which only in mammals is differentiated into

an anterior smaller jejunum and posterior longer ileum. A large number of small
digestive glands are present in the small intestine. They are tubular glands or
crypts of Lieberkuhn found through the entire length, they secrete mucus and a
succus entericus which has several enzymes.The lining of the small intestine is
folded to form small villi, which increase the surface area for secretion and
absorption. The villi are covered densely by minute finger-like projections,
called microvilli which assist in absorption into the villi. In mammals nodules
of lymphoid tissue called Peyer’s patches are found on the ileum.

Figure 7: Vertebrate shwing different glandular regions

 Large Intestine:
Large intestine has a larger diameter than the small intestine. It is
generally short in fishes, amphibians, reptiles, and birds, but in mammals it is
long. In lower forms the large intestine forms a rectum, but in tetrapoda it has a
colon and terminal rectum. In most fishes and amphibians, the terminal part of
the rectum leads into a cloaca formed by the proctodaeum.

The rectum, excretory ducts, and genital ducts open into the cloaca, and it opens
to the exterior by a cloacal aperture. But in many bony fishes and all mammals
(except monotremes) the rectum and urinogenital ducts have separate openings
to the exterior; the opening of the former is an anus.Rectum of mammals is not
homologous with the rectum of vertebrates since in mammals it is derived by
partitioning of embryonic cloaca. In most vertebrate embryos there is a postanal
gut as an extension of the intestine into the tail, but it disappears later.
In elasmobranchs the large intestine bears a pair of rectal glands which secrete
mucus and sodium chloride. In amniotes there is an ileocolic valve at the
junction of small and large intestines, which is absent in fishes. It prevents
bacteria to enter into ileum from colon.

In amniotes from this junction arises an ileocolic caecum which is two in birds.
It contains cellulose digesting bacteria. It is very long in herbivorous mammals
(rabbit, horse, cow, etc.). In primates caecum is small having a vestigial
vermiform appendix.

 Digestive Glands:
Liver:
The liver arises as a single or double outgrowth from the ventral wall of the
embryonic archenteron. This outgrowth forms a hollow hepatic diverticulum,
which soon differentiates into an anterior part, which proliferates to become the
liver and its bile ducts, and a posterior part, which gives rise to the gall bladder
and cystic duct. The bile ducts join to form a hepatic duct which unites with the
cystic duct to form a common bile duct or ductus choledochus. The region of
the archenteron from which the liver arises becomes the duodenum.The liver is
the largest lobed gland in the body, suspended by a double layer of peritoneum
from the transverse septum or its representative.

A liver is present in all vertebrates. In cyclostomes, it is small, single lobed

(lampreys) and two lobed in hagfishes. It is bilobed in elasmobranchs, two or
three lobed in bony fishes, amphibians, reptiles and birds and many lobed in
mammals. Liver is long, narrow and cylindrical in fishes, urodeles and snakes.

A gall bladder is for storage of bile secreted by the hepatic cells, lies in the liver
and drains into the duodenum through common bile duct formed by the union of
cystic duct and hepatic duct. A gall bladder is not indispensable and is lacking
in many birds and mammals. It is short, broad and flattened in birds and
mammals. A gall bladder and bile duct are present in larval cyclostomes but
they are absent in the adult. Fishes, amphibians, and reptiles generally have a
gall bladder, but it is lacking in many birds. Most mammals have a gall bladder,
but it is absent in Cetacea and Ungulata.

Figure 8: Vertebrate liver

The liver secretes a watery, alkaline bile but have no enzymes. It neutralises the
acidity of food entering the duodenum. It aids in the digestion of fats.

Pancreas:
Pancreas is formed from the endoderm of the embryonic archenteron.
A single dorsal diverticulum from embryonic duodenum and one or two ventral
outgrowths from the liver form pancreatic diverticula. The proximal parts of
diverticula form pancreatic ducts, but these ducts undergo reduction or fusion so
that only one or two pancreatic ducts remain in the adult, they open into the
duodenum either separately or after uniting with the common bile duct.

The distal parts of diverticula undergo budding to form the main mass of
pancreatic cells to which mesodermal derivatives are added. Thus, a single
gland is made which has several lobes forming either a diffuse or a compact
pancreas.The pancreas is both an exocrine and endocrine gland, bound together
by delicate strands of connective tissue. The exocrine part secretes digestive
enzymes which are poured into the duodenum through pancreatic ducts.
Whereas the endocrine part secretes hormones such as insulin and glucagon.

Figure 9: Evolution of pancreatic cells

A pancreas is present in all vertebrates. In lampreys, some bony fishes,

lungfishes and lower tetrapods, it is a diffuse organ embedded in the liver,
mesenteries and intestinal wall. Hagfishes have a small pancreas.
Elasmobranchs have a well defined bilobed pancreas. In higher tetrapoda it is
generally a compact gland. One or two pancreatic ducts open into the
duodenum.
INVERTEBRATE EXCRETORY SYSTEMS

Excretory system :

‘’The excretory system is a passive biological system that removes

excess, unnecessary materials from the body fluids of an organism, so
as to help maintain internal chemical homeostasis and prevent damage
to the body. ‘’

 The dual function of excretory systems is the elimination of the

waste productss of metabolism and to drain the body of used up
and broken down components

 Aquatic invertebrates
Aquatic invertebrates occur in a wide range of media, from freshwater
to markedly hypersaline water (e.g., salt lakes).

Generally marine invertebrates have about the same osmotic

concentration as seawater (i.e., they are osmo conformers). This
avoids any need to osmoregulate. Most water and ions are gained
across the integument, via gills, by drinking, and in food. Ions and
wastes are mostly lost by diffusion via the integument, gills, or urine.

Freshwater invertebrates are strong osmoregulators because it is

impossible to be isosmotic with dilute media. Any water gain is
usually eliminated as urine.
  Terrestrial invertebrates :
The most successful terrestrial invertebrates are the arthropods,
particularly the insects, spiders, scorpions, ticks, mites, centipedes,
and millipedes.

Overall, the water and ion balance of terrestrial invertebrates is quite

different from that of aquatic animals because terrestrial invertebrates
face limited water supplies and water loss by evaporation from their
integument.

 Invertebrate excretory mechanisms

 Contractile vacuoles :

 A contractile vacuole is an organelle, or sub-cellular structure,

that is involved in osmoregulation and waste removal.
 Previously it was known as pulsating vacuole.
 Contractile vacuoles should not be confused with vacuoles
which store food or water.
 Contractile vacuole, regulatory organelle, usually spherical,
found in freshwater protozoa and lower metazoans, such as
sponges and hydras, that collects excess fluid from the
protoplasm and periodically empties it into the surrounding
medium.
 Mechanism of excretion :
 A Contractile vacuole is found predominantly in protists and in
unicellular algae.
 In freshwater environments, the concentration of solutes inside
the cell is higher than outside the cell.
 Under these conditions, water flows from the environment into
the cell by osmosis. Thus, the CV acts as a protective
mechanism against cellular expansion (and possibly explosion)
from too much water; it expels excess water from the cell by
contracting.
 In some freshwater species, water. Contractile vacuoles are
energy-requiring devices that expel excess water from
individual cells exposed to hypoosmotic environment
The contractile vacuole helps prevent excessive water influx that
could harm and cause rupture (lysis) to the cell. The contractile
vacuole contracts to expel water out of the cell

Systole:
The period of water expulsion of contractile vacuole is referred to as
systole

Diastole:
The period at which water flows into contractile vacoule is called
diastole
PROTONEPHRIDIA

 A protonephridium (proto = "first") is a network of dead-

end tubules lacking internal openings
 These are found in the
phyla Platyhelminthes, Nemertea, Rotifera and Chordata’’

 Protonephridia likely first arose as a way to cope with a

hypotonic environment by removing excess water from the
organism (osmoregulation). Their use as excretory structures
likely secondarily

 Although a few groups of invertebrates possess no known

excretory structures, most have nephridia that serve for
excretion, osmoregulation, or both.
 Probably the earliest type of nephridium to appear in the
evolution of animals was the protonephridium
 Among the simplest of the protonephridia are flame-cell
systems, such as those in rotifers, some annelids, larval
molluscs, and some flatworms

 The protonephridial excretory system

The protonephridial excretory system is composed of a network
of excretory canals that open to the outside of the body through
excretory pores. Bulblike flame cells are located along the
 excretory canals. Fluid filters into the flame cells from the
surrounding interstitial fluid each of which have one or more
cilia
The beating cilia propel the fluid through the excretory canals
and out of the body through the excretory pores.

 Functions
 Flame-cell systems function primarily in eliminating excess
water. Nitrogenous waste simply diffuses across the body
surface into the surrounding water.
 The terminal cells are located at the blind end of the
protonephridium. Each cell has one or more cilia and their
beating inside the protonephridial tube creates an outward
going current and hence a partial pressurization in the blind
of the tube
  this pressurization drives waste fluids from the inside of the
animal, and they are pulled through small perforations in the
terminal cells and into the protonephridium
 The perforations in the terminal cell are large enough for
small molecules to pass, but larger proteins are retained
within the animal. From the bottom of the protonephridium
the solutes are led through the tube and exits the animal from
a small opening formed by the nephridiopore
 . Selective reabsorption of useful molecules by the canal cells
occurs as the solutes pass down the tubule

Metanephridia

 A more advanced type of excretory structure among

invertebrates is the metanephridium
 metanephridium (meta = "after") is a type of excretory gland
found in many types of invertebrates such as annelids
, arthropods and mollusca
 (In mollusca, it is known as the Bojanus organ.)
 Structure of metanephridium :

 A metanephridium consists of a ciliated funnel opening into the

bodycavity connected to a duct which may be variously
glandularized, folded or expanded which opens outside the body
 Protonephridia and metanephridia are two types of nephridia
that mainly occur in invertebrates.

 Generally, they are functionally analogous to the vertebrate

kidney and are responsible for the removal of metabolic waste
from the body.

 Functions :
 These ciliated tubules pump water carrying surplus ions
metabolic wastes and toxins from food and useless hormones
out of the organism by directing them down funnel-shaped
bodies called nephrostomes .
 This waste is passed out of the organism at the nephridiopore .
The primary urine produced by filtration of blood (or a similarly
functioning fluid) is modified into secondary urine
through selective reabsorption by the cells lining the
metanephridium.

 Metanephridia in earth worm :

 earthworm’s body is divided into segments and that each

segment has a pair of metanephridia.
 Each metanephridium begins with a ciliated funnel, the
nephrostome, that opens from the body cavity of a segment into
a coiled tubule
 . As beating cilia move the fluid through the tubule, a network
of capillaries surrounding the tubule reabsorbs and carries away
ions.
 Each tubule leads to an enlarged bladder that empties to the
outside of the body through an opening called the
nephridiopore.The excretory system of molluscs includes
protonephridia in larval stages and metanephridia in adults
  Difference between protonephridia and metanephrdia

Main points :

 The excretory system is a passive biological system that removes

excess, unnecessary materials from the body fluids of an
organism, so as to help maintain internal chemical homeostasis.
 Marine invertebrates have about the same osmotic
concentration as seawater (i.e., they are osmo conformers). This
avoids any need to osmoregulate.
 Freshwater invertebrates are strong osmoregulators because it
is impossible to be isosmotic with dilute media.

 A contractile vacuole is an organelle, or sub-cellular

structure, that is involved in osmoregulation and waste removal.
 Contractile vacuoles are energy-requiring devices that expel
excess water from individual cells exposed to hypoosmotic
environment and prevent them from lysis

 A protonephridium (proto = “first”) is a network of dead-end

tubules lacking internal openings

 simplest of the protonephridia are flame-cell systems, such as

those in rotifers, some annelids, larval molluscs, and some
flatworms

 The beating cilia propel the fluid through the excretory canals
and out of the body through the excretory pores.
 A metanephridium consists of a ciliated funnel opening into the
bodycavity.
 These ciliated tubules pump water carrying metabolic wastes
and toxins from food and useless hormones out of the organism
 by directing them down funnel-shaped bodies called
nephrostomes .
 This waste is passed out of the organism at the nephridiopore.

Crustaceans:

Crabs, lobsters, shrimps, and wood lice are among the best-known
crustaceans. Crustaceans are generally aquatic and differ from other
arthropods in having two pairs of appendages (antennules and
antennae) in front of the mouth and paired appendages near the mouth
that function as jaws.

Gills
In some crustaceans nitrogenous wastes are removed by simple
diffusion across the gills.Most of them release ammonia,although
some of them also release urea or uric acid.Thus, excretory organs of
freshwater species may be more involved with the reabsorption of
ions and elimination of water than with the discharge of nitrogeneous
wastes.
Excretory organs are called antennal glands or maxillary glands,
depending on whether they open at the base of the antennae or at the
maxillae.
Antennal gland (green gland)
Excretory organs in some of the crustaceans are antenna glands or
green glands.Named due to their location near antenna or green
color.It is a sac opening at the base of antenna in crustaceans.
Either of a pair of ducts (coelomoducts) found in the third segment of
a crustacean and opening to the exterior at the base of the second
antenna. They function as osmoregulatory organs.
For example; in the antennal gland of the freshwater crayfish
(Astacus), fluid is filtered from the blood into an end sac and passes
through a tubular labyrinth, where ions are reabsorbed, to produce a
hypotonic urine that passes via a renal tubule to the bladder.
Maxillary glands:
In some crustaceans excretory organs near maxillary segments are
maxillary glands. In these glands fluid collects within the tubules
from the surrounding blood of hemoceol and this urine is modified by
reabsorption and secretion as it moves through the excretory system
and rectum.
Malphigian tubules:
Malpighian tubule, in insects, any of the excretory organs that lie in
the abdominal body cavity and empty into the junction between
midgut and hindgut.
In species having few malpighian tubules, they are long and coiled; in
species with numerous (up to 150) tubules, they are short.
Functions:
 The tubule cells actively transport
initial urine constituents (potassium ions, water, urate ions,
sugar, amino acids) into the tubule.
 In some species urine is acidified in the distal end of the tubule
and an aqueous suspension of uric acid crystals is conducted
into the rectum, where water and nutrients are reabsorbed.
 In other species the urine is acidified in the rectum.
 Certain tubule cells may have special functions, as in the
secretion of the sticky substance that surrounds eggs of certain
leaf beetles or in the secretion of silk by certain immature
beetles.
Coaxal glands:
The coxal gland is a gland found in some arthropods, for collecting
and excreting urine. They are found in all arachnids (with the
exception of some Acari), and in other chelicerates, such as horseshoe
crabs. The coxal gland is thought to be homologous with the antennal
gland of crustaceans.
In certain arthropods, one of a pair of excretory organs consisting of
an end sac where initial urine is collected, a tubule where secretion
and reabsorption may take place, and an excretory pore at the base
(coxa) of one of the legs. Variations among the species include
highly convoluted tubule sections, doubling back of straight tubule
sections, and expansion of the terminal end into a bladder.

Vertebrate Excretory System:

Excretory systems regulate the chemical composition of body

fluids by removing metabolic wastes and retaining the proper
amounts of water, salts, and nutrients. Components of this system
in vertebrates include the kidneys

 Liver
 Lungs and
 Skin

Problems

Vertebrate face the same problems as invertebrates in controlling

 Water
 Ion blance
 Generally Water losses are balanced precisely by water gains.
 Vertebrates gains water by absorption from liquid and solid food
in the small and large intestines and by metabolic reaction that
yield water as an end product.
 They lose water by
 Evaporation from respiratory surfaces
  Evaporation from integument
 Sweating
 Panting
 Elimination in feces
 Excretion by the urinary system

Solute losses also must be balanced by solute gains.

Vertebrates take in solute by

 Absorption of minerals
 Through integument and gills
 From secretions of various gland
 By metabolism.

They lose solute in Sweat , feces,urine and gill secretions as

Metabolic waste.The major metabolic waste that must be eliminated
as

 Ammonia
 Urea
 Uric acid

Vertebrates live in saltwater , fresh Water , and on land.

How vertebrates achieve osmoregulation?

A variety of mechanisms has evolved in vertebrates to cope with their

osmoregulatory problems.
 Filtration in which blood passes through a filter that retains
blood cells , proteins , and other large solutes but lets small
molecule , ions, and urea passes through .
 Reabsorption in which selective ions and molecules are taken
back into the bloodstream from the filterate.
 Secretion where by select ions and end products of metabolism
that are in the blood are added to the filrerate for removal from
the body.
Points:
 Excretory system regulate the chemical composition of body.
 Vertebrates face the same problem as invertebrates.
 Water
 Ion balance
 Vertebrates achieve osmoregulation through
 Filtration
 Secretion
KIDNEY
 Location
 Anatomy
 Structure
 Function
 Diseases
 Causes
 Dialysis
 Maintaining kidney health

The kidneys are a pair of bean-shaped organs present in all

vertebrates. They remove waste products from the body, maintain
balanced electrolyte levels, and regulate blood pressure.

The kidneys are some of the most important organs in the body. The
ancient Egyptians left only the brain and kidneys in position before
embalming a body, inferring that they held a higher value than other
organs.

In this article, we look at the structure and function of the kidneys, the
diseases that affect them, and how to keep them healthy.

Location

The positioning Trusted Source of the kidneys is just below the rib
cage, with one on each side of the spine. The right kidney is generally
slightly lower than the left kidney to make space for the liver.
Each kidney is approximately 3 centimeters (cm) thick, 6 cm wide,
and 12 cm long. In males, the average weightTrusted Source of the
kidneys is roughly 129 grams (g) for the right one and 137 g for the
left. In females, the average weight Trusted Source of these organs is
108 g for the right kidney and 116 g for the left kidney.

Anatomy

Structure

The kidneys are two bean-shaped organs that are roughly the size of a
fist. A tough, fibrous renal capsule surrounds each kidney and
provides support for the soft tissue inside. Beyond that, two layers of
fat serve as further protection. The adrenal glands lie on top of the
kidneys.

Inside the kidneys are a number of pyramid-shaped lobes. Each

consists of an outer renal cortex and an inner renal medulla. Nephrons
flow between these sections. Each nephron includes a filter, called the
glomerulus, and a tubule. The glomerulus filters blood, which enters
the kidneys through the renal arteries and leaves through the renal
veins. The kidneys are relatively small organs, but they receive 20–
25%Trusted Source of the heart’s output.

The tubule returns necessary substances to the blood and removes

waste that then becomes urine. The kidneys excrete urine through the
ureter, a tube that leads to the bladder.

What does a kidney look like?

Function

The main role of the kidneys is maintaining homeostasis. They

manage fluid levels, electrolyte balance, and other factors that keep
the internal environment of the body consistent and comfortable.

These organs carry out a wide range of bodily functions.

Waste excretion

The kidneys removeTrusted Source various waste products and get rid
of them in the urine. Some major compounds that the kidneys remove
are:

 urea, which results from the breakdown of proteins

 uric acid from the breakdown of nucleic acids
 drugs and their metabolites

Reabsorption of nutrients

The kidneys reabsorbTrusted Source nutrients from the blood using

tubules and transport them to where they will best support health.
They also reabsorb other products to help maintain homeostasis.
Reabsorbed products include:

 glucose
 amino acids
 bicarbonate
 water
 phosphate
 chloride, sodium, magnesium, and potassium ions

Maintaining pH

In humans, the range of acceptable pH levels is 7.35–7.45. At levels

below or above this range, the body enters a state of acidemia or
alkalemia, respectively. In these states, proteins and enzymes break
down and can no longer function. In extreme cases, this can be fatal.

The kidneys and lungs help keep the body’s pH stable. The lungs
achieve this by moderating the concentration of carbon dioxide in the
blood. The kidneys manage the pH by reabsorbing and producing
bicarbonate from urine, which helps neutralize acids.

The kidneys can retainTrusted Source bicarbonate if the pH is

tolerable and release it if acid levels rise. They can produce new
bicarbonate by excretingTrusted Source acid.

Osmolality regulation

Osmolality is a measure of the body’s electrolyte-water balance,

which is the ratio between fluids and minerals in the
body. Dehydration is a primary cause of electrolyte imbalance.
If osmolality rises in the blood plasma, the hypothalamus in the brain
responds by passing a message to the pituitary gland. This gland
releases antidiuretic hormone (ADH). In response to ADH, the kidney
makes several changes, including:

 increasing urine concentration

 increasing water reabsorption
 reopening portions of the collecting duct that water cannot
normally enter, allowing water back into the body
 retaining urea in the medulla of the kidney rather than excreting
it, as this compound draws in water

Regulating blood pressure

The kidneys regulate blood pressure when necessary, but they are
responsible for slower adjustments.

They adjust long-term pressure in the arteries by causing changes in

the fluid outside of cells. The medical term for this fluid is
extracellular fluid. These fluid changes occur after the release of a
vasoconstrictor called angiotensin II. Vasoconstrictors are hormones
that cause blood vessels to narrow.

These hormones play a role in increasing the kidneys’ absorption of

sodium chloride, or salt. This absorption effectively increases the size
of the extracellular fluid compartment and raises blood pressure.
Anything that alters blood pressure, including excessive alcohol
consumption, smoking, and obesity, can damage the kidneysTrusted
Source over time.

Secretion of active compounds

The kidneys release several important compounds, including:

  Erythropoietin: This controls erythropoiesis, which is the
production of red blood cells. The liver also produces
erythropoietin, but the kidneys are its main producers in adults.
 Renin: This enzyme helps manage the expansion of arteries and
the volumes of blood plasma, lymph, and interstitial fluid.
Lymph is a fluid that contains white blood cells, which support
immune activity, and interstitial fluid is the main component of
extracellular fluid.
 Calcitriol: This is the hormonally active metabolite of vitamin
D. It increases both the amount of calcium that the intestines can
absorb and the reabsorption of phosphate in the kidney.

Diseases

A range of diseases can affect the kidneys. Environmental or medical

factors may lead to kidney disease, and they can cause functional and
structural problems from birth in some people.

Diabetic nephropathy

In people with diabetic nephropathy, damage occurs to the capillaries

of the kidney as a result of long-term diabetes. The symptoms may
not become apparent until years after the damage starts to develop.
They can include:

 fluid buildup
 sleep difficulty
 poor appetite
 upset stomach
 weakness
 difficulty concentrating
Kidney stones

Stones can form as a solid buildup of minerals in the kidneys.

They can cause intense pain and might affect kidney function if they
block the ureter.

Kidney infections

Kidney infections tend to result from bacteria in the bladder that

transfer to the kidneys.

The symptoms can include lower back pain, painful urination, and,
sometimes, fever. Changes in the urine may include the presence of
blood, cloudiness, and an unusual odor.

Kidney infections are more commonTrusted Source in females than in

males and more likely to affect those who are pregnant. The infection
often responds well to antibiotics.

Renal failure

In people with renal failure, the kidneys become unable to filter out
waste products from the blood effectively.

If an injury or another factor, such as the overuse of medication,

causes kidney failure, the condition may be reversible with treatment.

If the cause is a disease, however, kidney failure often does not have a
full cure.

Kidney hydronephrosis

Hydronephrosis means “water on the kidney.”

It usually occurs when an obstruction prevents urine from leaving the
kidney, causing intense pain.

In time, untreated hydronephrosis can put pressure on a person’s

kidneys and may result in kidney damage.

Interstitial nephritis

A reaction to medications or infection can cause inflammation of the

nephrons.The treatment usually involves addressing the cause
of inflammation or changing a course of medication.

Kidney tumor

These tumors can be benign or malignant. Benign cancers do not

spread or attack tissue, but malignant cancers can be aggressive.

The most common malignant kidney cancer is renal cell carcinoma.

Nephrotic syndrome

When damage to the kidney affects its function, this causes protein
levels in the urine to increase. This effect leads to a protein shortage
throughout the body, which draws water into the tissues. The
symptoms of nephrotic syndrome can includeTrusted Source:

 puffy eyes
 increased cholesterol levels
 weight gain
 fatigue
 loss of appetite
Lower back pain and changes in urination, especially on one
side, may be signs of kidney problems.

Causes

Some common causes of kidney damage may include:

 Analgesics: Using pain medication over a long period might

result in chronic analgesic nephritis. Examples include aspirin,
acetaminophen, and nonsteroidal anti-inflammatory drugs
(NSAIDs).
 IgA nephropathy: Also known as Berger disease, this occurs
when immunoglobin A (IgA) antibodies build up in the kidney.
IgA forms a vital part of the immune system, but a buildup can
be harmful. The disease progresses slowly, sometimes taking as
long as 20 yearsTrusted Source to develop. The symptoms
include abdominal pain, a rash, and arthritis. It can result in
kidney failure.
 Lithium: Doctors prescribe lithium to
treat schizophrenia and bipolar disorder. However,
lithium might cause nephropathyTrusted Source with long-term
use. Close medical supervision can help a person avoid the
negative effects of lithium.
 Chemotherapy agents: The most common type of kidney issue
in people with cancer is acute kidney injury. This might be due
to the intense vomiting and diarrhea that are common side
effects of chemotherapy.
 Alcohol: Alcohol alters the kidneys’ ability to filter the blood. It
also dehydrates the body, making it harder for the kidneys to
redress internal balances, and increases blood pressure, which
can also hinder the kidneys.
Dialysis

In the case of severe kidney damage, dialysis might be an option.

Doctors only use this treatment for end stage kidney failure involving
the loss of 85–90% of kidney function. Kidney dialysis aims to
complete some of the functions of a healthy kidney. These include:

 removing waste, excess salt, and water

 maintaining the correct levels of chemicals in the blood,
including sodium, bicarbonate, and potassium
 maintaining blood pressure

The two most common types of kidney dialysis are:

Hemodialysis

An artificial kidney, or hemodialyzerTrusted Source, removes waste,

additional fluids, and chemicals. The treating doctor makes an entry
point in the body by connecting an artery and a vein under the skin to
create a larger blood vessel.

Blood travels into the hemodialyzer, receives treatment, and then

returns to the body. This process usually takes place three or four
times a week.

Peritoneal dialysis

The doctor inserts a sterile cleansing solution into the abdominal

cavity around the intestine. This is the peritoneum, and a protective
membrane surrounds it.

In continuous peritoneal dialysis, the fluid drains through a catheter.

The individual discards these fluids four or five times a day. In
automated peritoneal dialysis, the fluid also drains through a catheter,
and the exchanges usually occur throughout the night while the
person sleeps.

Maintaining kidney health

The following are suggestionsTrusted Source for keeping the kidneys

healthy and helping avoid kidney disease:

 Eating a balanced diet: Many kidney problems result

from high blood pressure and diabetes. As a result, maintaining
a nutritious, well-balanced diet can prevent several common
causes of kidney disease. The National Heart, Lung, and Blood
Institute recommends the DASH dietTrusted Source for
maintaining healthy blood pressure.
 Getting enough exercise: Exercising for 30 minutes every day
can reduce the risk of high blood pressure and obesity, both of
which put pressure on kidney health.
 Drinking plenty of water: Fluid intake, especially of water, is
important. Many factors can affect how much water people
should drink, but about 4–6 glasses per day can help improve
and maintain kidney health.
 Taking supplements: Be careful when taking supplements, as
not all dietary supplements and vitamins are beneficial. Some
can harm the kidneys if a person takes too many.
 Limiting salt: Limit sodium intake to a maximum of 2,300
milligrams (mg)Trusted Source of sodium, and ideally 1,500
mgTrusted Source, each day.
 Moderating alcohol: Consuming more than one drink per day
for females and two per day for males can harm the kidneys and
impair renal function.
 Avoiding smoking: Tobacco smoke restricts blood vessels.
Without an adequate blood supply, the kidneys will not be able
to complete their normal work.
 Taking over-the-counter (OTC) medications: A drug is not
harmless simply because a person does not need a prescription
to get it. Overusing OTC drugs, such as ibuprofen and naproxen,
can damage the kidneys.
 Screening for health conditions: Anyone with high blood
pressure or diabetes should consider discussing regular kidney
screening with a doctor to help spot any possible health issues.
 Managing diabetes and heart disease: Following a doctor’s
recommendations for managing these conditions can help
protect the kidneys in the long term.
 Taking steps to get quality sleep and control stress: The
National Institute of Diabetes and Digestive and Kidney
Diseases (NIDDK) recommends getting 7–8 hours of
sleepTrusted Source each night and seeking out activities to
reduce stress.
Osmoregulation by Introductory article

Vertebrates in Aquatic Article Contents

. The Origin of Aquatic Vertebrates and Osmoregulation

Environments . Freshwater Vertebrates

. Seawater Vertebrates

David H Evans, University of Florida, Gainesville, Florida, USA . Summary

Because the salt concentration of the body fluids of aquatic vertebrates differs from that of
their freshwater or marine environment, they face net movements of water and salt across
their permeable membranes, most notably the gill. Various organs and transport processes
are involved in maintaining internal consistency in the face of these potential osmotic and
salt problems.

The Origin of Aquatic Vertebrates and

(which succeeded the amphibians as the truly terrestrial
Osmoregulation vertebrates) have returned to aquatic existence from their
terrestrial ancestry, and they may face similar problems of
The vertebrates arose from marine ancestors nearly 550 osmosis and diffusion. But these are reduced by the absence
million years ago, and many of these early forms of gills and the presence of scaled, feathered or furred outer
apparently entered fresh water soon after they evolved. body coverings. In sea water, however, salt loading may
This transition into an environment that contained far less take place because of salty food, whether plant or animal.
salt per volume than sea water was associated with a In summary, because of these water and salt gradients
decline in the salt (largely NaCl) concentration of the body and permeable gills or skin, freshwater vertebrates face a
fluids that was retained in subsequent vertebrates, with the net osmotic influx of water and net loss of salt by diffusion;
exception of the hagfishes. (These very primitive, marine marine forms face dehydration and a net influx of salt, the
parasitic fishes apparently descended from early ancestors opposite problems.
that never entered fresh water, and they have retained
blood salt concentrations that are essentially equivalent to
those in the ancestral marine invertebrates and their
surrounding sea water.) Despite this reduction in body Freshwater Vertebrates
fluid salt concentration, these early freshwater vertebrates
(fishes) were still hypertonic to their environment. Thus, Fishes
these fishes faced gradients of both water and salt that were
unknown to their marine invertebrate predecessors, In these descendants of early vertebrates, water influx
relatives of the modern starfish and sea urchins. Water across the gills must be balanced by water excretion by the
and salt move down their individual concentration kidney, but urinary (renal) salt loss must be minimized at
gradients, across permeable membranes of cells or organ- the same time in an attempt to minimize the diffusional loss
isms, by the process of osmosis and diffusion, respectively. of salt across the gills (Figure 1). In all vertebrates, urine is
The problem of osmotic and salt gradients is exacerbated formed by filtration of blood across the glomerulus in the
by the presence of the gill epithelium in many aquatic kidney, driven by blood pressure. Because of the need to rid
vertebrates. This large, thin tissue is modified for gas the body of excess fluid, glomerular filtration is high in
exchange; unfortunately, the very characteristics that freshwater fishes, and the kidney (renal) tubules reabsorb
facilitate gas exchange (large surface area, thin, highly little of the urine. Freshwater existence is associated with
supplied with blood) make it a site for passive movements the evolution of a distal tubule, specializing in the
of both water and salt. reabsorption of needed salt back into the blood. Such a
Most fish groups re-entered sea water and remain there terminal, diluting renal segment is also found in some
today, but one group remained in fresh water and emerged invertebrates that have evolved in fresh water (earth
on to land as the ancestor of the modern amphibians. As worms, and crustacea, for instance). The diffusional and
might be expected, water and salt balance problems exist renal loss of ions is balanced by a relatively small gain from
for the semiterrestrial amphibians because of the presence food, but largely by active transport of Na 1 and Cl 2
of gills in the larval stage of some species and a relatively inward, across the gill epithelium. The mechanisms of ionic
thin skin in the adult. Some reptiles, birds and mammals transport are still debated, but recent evidence suggests
that Na 1 is driven into the gills cells by a favourable

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net 1
 Osmoregulation by Vertebrates in Aquatic Environments

Water Salt Birds and mammals

Food In these very impermeable animals (cormorants and otters,
• salt
for instance), osmotic uptake of water is minimal, balanced
by increased urine flows if necessary, and any salt lost in the
Salt urine is compensated for by food. There is no evidence for
any skin uptake of salt from the environment.
Urine
• low salt
• large volumes of water

Figure 1 Osmoregulation in a freshwater fish. The net osmotic gain of

Seawater Vertebrates
water and diffusional loss of salt across the gills is balanced by excretion of
relatively dilute urine, active uptake of salt across the gill, and possibly some Fishes
ingestion of salt in the food. Blue arrows represent passive movements of
salt and water, and red arrows indicate active pathways of osmoregulation. The freshwater ancestors of modern marine fishes re-
See text for details. Outline drawing of a goldfish was redrawn from
entered the marine environment approximately 500 million
Greenwood HP, et al. (1966) Bulletin of the American Museum of Natural
History 131: 339–456. years ago and radiated into the huge array of marine bony
(teleosts) and cartilaginous (elasmobranchs: sharks and
relatives) fishes present in sea water today. Because of their
freshwater ancestry and attending reduction in the salt
electrochemical gradient produced by the active extrusion concentration of their body fluids, both groups face an
of H 1 by a transport protein termed H-ATPase which osmotic loss of water and diffusional gain of salts. But their
moves positive charges out of the fish. Cl 2 enters the cells solutions to these common problems are quite different.
from the fresh water in exchange for cellular HCO32 . These Teleosts balance the net loss of water across the gill
electrical and biochemical couples could, theoretically, be epithelium by obtaining water from the only source
involved in the net secretion of either H 1 or HCO32 . In available: the surrounding sea water (Figure 2). They ingest
terrestrial vertebrates, renal loss of acid or base equiva- sea water, and absorb the needed fluid across the intestine
lents, coupled with respiratory excretion of CO2, controls by transporting the salt across the gut lining, into the
acid–base balance, but in fishes the extrusion of either H 1 blood. This salt transport produces gradients, which
or HCO32 is the regulatory pathway. osmotically withdraw the water from the intestinal
contents at the same time. Marine teleosts limit their renal
loss of water by reducing glomerular filtration to near zero,
Amphibians and some species have evolved glomerular-free renal
tubules. Marine fishes can tolerate a very small urine flow,
In those species of amphibians that have tadpoles as larvae, because they can excrete unwanted nitrogen as ammonia
the osmotic adaptations to life in fresh water are basically across the gills. (Terrestrial vertebrates, including mam-
the same as those for fishes. In amphibians (adult frogs, mals, must excrete unwanted, and toxic, ammonia as urea
salamanders, etc.) that are gill-less, the skin has evolved
salt transport pathways similar to those in the gills of fishes
or tadpoles. Indeed, much early work on the biophysical
mechanisms of Na 1 transport involved study of the Salt
movement of Na 1 across frog skins. Water
Drink
• water
• salt

Reptiles Salt

Freshwater reptiles are relatively uncommon; turtles are Urine

the only group that has been studied in any detail. In • low water
• some salt
reptiles, net influx of water, which is minimized by the thick
skin, is balanced by variable urine flows. Urinary salt loss is Figure 2 Osmoregulation by a marine teleost fish. The net osmotic loss of
minimal (because of salt reabsorption by the distal renal water and diffusional gain of salt across the gills is balanced by ingestion of
tubule), as is diffusional loss across the skin. What salt is sea water, production of small volumes of urine that contains some salt, and
lost is balanced by either salt in the food or, in some species, active extrusion of salt across the gill. Blue arrows represent passive
movements of salt and water, and red arrows indicate active pathways of
active extraction of salt from water drawn into either the osmoregulation. See text for details. Outline drawing of a tuna was redrawn
mouth or anus, but the salt transport mechanisms involved from Bigelow HB and Schroeder WS (1953) Fisheries Bulletin, Fish and
are unknown. Wildlife Service 53: 1–577.

2 ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
 Osmoregulation by Vertebrates in Aquatic Environments

or uric acid via the kidney.) Marine teleosts increase their Water
Salt
renal loss of ions (to partially compensate for the net
diffusional gain) by increasing renal salt loss, but they are
not able to secrete a net amount of salt because they cannot Food
produce a urine that contains more salt per volume than • salt
the body fluids. (This requires the evolution of another Salt?
Rectal gland Urine
renal tubule, the loop of Henle, which did not appear in the • salt • some salt
vertebrates until the birds and mammals evolved from their • large volumes of water
reptilian predecessors.) Figure 3 Osmoregulation by a shark. The osmotic gain of water and
Net salt secretion in marine teleosts is managed by the diffusional gain of salt across the gills is balanced by production of large
gill tissue, which uses a suite of transport proteins that are volumes of urine that contains some salt, active secretion of salt via the
also found in our own kidney tubules. Na 1 enters the gill rectal gland, and possibly active extrusion of salt across the gill. Blue arrows
cell from the blood co-transported with K 1 and Cl 2 , represent passive movements of salt and water, and red arrows indicate
active pathways of osmoregulation. See text for details. Outline drawing of
driven by the favourable electrochemical gradient for a spiny dogfish shark was redrawn from Bigelow HB and Schroeder WS
Na 1 . Cl 2 exits the apical portion of the cell through a (1953) Fisheries Bulletin, Fish and Wildlife Service 53: 1–577.
channel that is very similar to the defective structure that
produces cystic fibrosis in humans. Na 1 is transported
back across the basolateral membrane into the blood by rectal gland, an extension of the terminal intestinal tract.
Na,K-activated ATPase and then diffuses out into the sea This gland is capable of secreting a concentrated NaCl
water via the relatively leaky paracellular pathways solution that contains more salt than either blood or sea
between adjacent gill cells, driven by a favourable water. The mechanisms of salt secretion are the same as
electrochemical gradient. The net result is secretion of described for the teleost gill tissue. Importantly, experi-
NaCl across the gill epithelium, balancing the net influx of mental removal of the gland is associated with a slight
salt. It is important to note that these net excretory NaCl increase in the plasma NaCl concentration, but the
pathways are not reversed freshwater uptake pathways; experimental animals (sharks) survive in sea water. This
they are quite different. Freshwater ionic uptake pathways experiment suggests that the gill also must be capable of net
for acid–base regulation may be present in marine fish gills, salt secretion.
even though they actually move NaCl into the marine fish.
Marine elasmobranch fishes have evolved a different
strategy for osmoregulation. To avoid the desiccating
effects of sea water, they retain urea (an ammonia Amphibians
detoxification compound) in the blood by reabsorption
in the kidney and the maintenance of a low gill permeability Amphibians are rarely associated with sea water. One
to this organic solute. Urea concentrations may approach exception is the crab-eating frog of southeast Asia, whose
500 mmol L 2 1, a concentration that is fatal in humans. tadpole osmoregulates like a marine teleost and whose
Elasmobranch enzymes are not damaged by such high urea adult osmoregulates like an elasmobranch, by storing urea.
concentration because another organic osmolyte, tri- How salt is secreted by these animals is not known.
methylamine oxide, counters the denaturing effects of the
urea when present at a ratio of 1:2 inside the cells. The
presence of these two organic solutes raises the total solute
concentration of elasmobranch blood slightly higher than Reptiles
that of sea water (these fishes are hypertonic), thus
avoiding the osmotic loss of water. The osmotic gradient Marine reptiles are common, and include turtles, snakes,
favours uptake of water across the elasmobranch gill, lizards, crocodiles, and even alligators that may have
thereby providing sufficient water for the relatively high entered the marine environment. In all cases, the desiccat-
urine flows seen in marine elasmobranchs. Despite being ing effects of sea water are minimized by their thick skin,
hypertonic to sea water, elasmobranch blood still contains but the salt in their food and ingested sea water must be
less Na+ and Cl 2 than sea water, so these ions diffuse excreted. The absence of a loop of Henle precludes net
inward across the gill, in the same way as they do across the renal salt secretion, so extrarenal pathways have evolved.
teleost gill epithelium. Thus, like teleosts, marine elasmo- In all cases, the secretory tissue is in the head region, with
branchs face a salt load, which must be excreted (Figure 3). orbital salt glands in turtles and lizards, sublingual glands
Like the teleost kidney, that of elasmobranchs is incapable in sea snakes, and supralingual glands in crocodiles and to
of producing a net secretion of salt, so extrarenal a much lesser extent, alligators. The specific mechanisms of
mechanisms have evolved. In this group, the gills probably net salt secretion by these salt glands is unknown, but
play a role in salt excretion, but this has been largely assumed to be similar to that described for the teleost gill
unstudied. Instead, research attention has focused on the tissue and shark rectal gland.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net 3
 Osmoregulation by Vertebrates in Aquatic Environments

Birds Summary
Marine birds have evolved the same strategies as the Because of the evolutionary history of aquatic vertebrates,
reptiles, including an orbital salt secretory gland (termed a the net movement of water and/or salt across their body
nasal gland because its secretory fluid drips from the nasal surfaces must be countered in order to maintain the volume
openings). The mechanisms of salt secretion have been and salt content of their internal tissues and body fluids. A
better studied in birds than reptiles, and the published variety of strategies has evolved to maintain this consis-
evidence suggests that the pathways are identical to those tency, involving organs as diverse as gills, intestine, kidney,
described for the teleost gill tissue and shark rectal gland. and specialized salt secretory glands. The underlying
In addition to extrarenal salt secretion, bird kidneys are mechanisms that transport salt across these structures are
able to elaborate a slightly salty urine because of the origin conserved throughout vertebrate evolution.
of the loop of Henle.
Further Reading
Mammals Campbell NA and Reece JB (2002) Biology, 6th edn. San Francisco:
Benjamin Cummings.
Like the outer covering of marine reptiles and birds, the Evans DH (1998) The Physiology of Fishes, 2nd edn. Boca Raton: CRC
relatively impermeable skin of mammals avoids many of Press.
the osmoregulatory problems of life in sea water. The salt Goldstein L (1999) Comparative physiology of osmoregulation: The
legacies of August Krogh and Homer Smith. Journal of Experimental
loading produced by ingestion of sea water and inverte-
Zoology 283: 619–733.
brate food is offset by the presence of a loop of Henle, NcNab BK (2002) The Physiological Ecology of Vertebrates. A View from
which allows the production of urine that is 2–3 times as Energetics. Ithaca: Comstock Publishing Associates.
salty as the blood. No extrarenal salt secretory mechanisms Randall D, Burggren W and French K (2002) Animal Physiology.
have been found, or are necessary. Mechanisms and Adaptations, 5th edn. New York: WH Freeman.

4 ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net