Homeostasis

G R O U P 2 – R E PO RTI N G I N I C H T H Y O L O G Y
Table of Contents
I. Endocrine System
A. Endocrine Gland
B. Hormones
C. Parts and Function of endocrine glands

II. Automatic Nervous System in controlling Various responses

A. Involuntary Physiological Function
B. Function of Gastrointestinal system
C. Gas Bladder

III. Temperature relationship

A. Ectothermic
B. Coping with temperature and Fluctuation
C. Heterothermic Fishes
D. Coping with temperature

IV. Water, Solute & pH Balance

A. Osmoregulation in Fishes
B. Control of osmoregulation and excretion
C. pH balance

V. Immune System
A. Immunity
B. Immunity System

VI. Stress
A. Biological Response
B. Physiological Responses
C. Primary Responses
Endocrine System
• Many endocrine functions are ultimately controlled by the hypothalamus of the brain regulating the many functions of the pituitary
which, in turn, helps regulate many other endocrine tissues in the body. The pituitary has two main functional regions.

• The endocrine system or internal

secretion system is a set of organs
and tissues of dissimilar origin,
structure and location, which
secretions called hormones, enter
the bloodstream directly without a
duct so they can reach distant
organs. Hormones are signaling
molecules that regulate many
physiological and behavioral
functions in functions including
growth, reproduction and water
and mineral balance, among
others.
Endocrine Gland of Fish
• The glands that secrete their • The pituitary gland or Hypophysis
products into the bloodstream and
body tissues along with the central • Thyroid Gland
nervous system to control and • Adrenal gland
regulate many kinds of body
functions are known as endocrine • Corpuscles of Stannius
gland. In fishes various endocrine
gland has been found associated • Ultimobranchial Glands
with different tasks and functions. • Urohypophysis
• Endocrine glands of fishes:
• Pancreatic islets
Different types of endocrine glands
are found in fishes; such as- • Pineal gland
Endocrine glands

• Of increasing complexities are found in cyclostomes, elasmobranchs and osteichthyes.

Elasmobranchs (sharks) possess well developed endocrine glands but these show some
interesting differences from those of higher chordates. However, osteichthyes (bony fishes)
have endocrine glands rather more similar to higher chordates.
• The difference between fish and mammal endocrine glands is probably due to the
development and modification of various body systems in these two classes, and also due to
exigencies of an aquatic mode of life.
• Mammalian endocrine glands are well advance and well-studied but fish endocrinology is
limited to the work on its influence on chromatophores, action of sex cells, function of
pituitary and thyroid and control on migration.
• Unlike nervous system, the endocrine system is basically related to comparatively slow
metabolism of carbohydrate and water by adrenal cortical tissue, nitrogen metabolism by
adrenal cortical tissue and thyroid glands and the maturation of sex cells and reproductive
behaviour by the pituitary gland and gonadal hormones.
The components of endocrine system can be classified
on the basis of their organization, which is as follows:
• Discrete Endocrine Glands:
• These include pituitary (hypophysis), thyroid and pineal (Fig. 19.1).
• Organs containing both endocrine and exocrine functions:
• In fishes, it is kidney, gonads (Fig. 19.1) and intestine. Kidney contains
heterotopic thyroid follicles, inter-renal, and corpuscles of Stannius.
• Scattered Cells with Endocrine Function:
• They are known as diffused neuro-endocrines. They are present in digestive tract
(Fig. 19.1). They are generally called as paracrines (e.g., somatostatin). There are
gastrointestinal peptides whose definite classification as hormone or paracrine
agent has not yet been established, these are designated as putative hormones.
Chemically, hormones can be divided into three
classes:

I. Steroid hormones (testosterone & estradiol)

II. Protein (peptide) hormones (e.g., insulin) and peptide hormones are
secreted by hypophysis, thyroid, internal tissue and pancreatic
tissue.
III. The amino acid analogues are norepinephrine and epinephrine,
collectively called catecholamines.
Pituitary Glands:

Origin of Pituitary Glands:

• The pituitary gland occupies the same central part in the endocrine signalling
system of fish that it has in mammals. This master endocrine gland originates
embryo-logically from the two sources. One as ventral down-growth of a neural
element from the diencephalon called the infundibulum to join with another, an
ectodermal up-growth (extending as Rathke’s pouch) from primitive buccal cavity.
Location of Pituitary Glands:
• The pituitary gland is located below the diencephalon (hypothalamus), behind the
optic chiasma and anterior to saccus vasculosus, and is attached to the
diencephalon by a stalk or infundibulum. The stalked pituitary is found in Barbus
stigma and Xiphophorus maculatus.
Anterior Pituitary

• The anterior pituitary, largely under the control of the hypothalamus,

manufactures and releases hormones that control many physiological functions
elsewhere in the body, including many other endocrine tissues. For example, the
anterior pituitary releases adrenocorticotropic hormone (ACTH), which influences
the production and release of cortisol from the interrenal tissue, and thyroid-
stimulating hormone (TSH), which stimulates the thyroid gland to release
thyroxin, gonadotropins (which stimulate the gonads), and growth hormone (GH)
which affects various tissues throughout the body (Takei & Loretz 2006).
• Fishes are the only jawed vertebrates known to possess a caudal neurosecretory
system. Located at the caudal end of the spinal cord, this region of neuroendocrine
cells, the urophysis, is most highly developed in the ray-finned fishes and
produces urotensins that help control smooth muscle contraction, osmoregulation,
and the release of pituitary hormones.
Thyroid Tissue

• The thyroid tissue of most fishes is scattered as small clusters of cells in

the connective tissue of the throat region. When stimulated by TSH from
the anterior pituitary, these cells produce thyroxin, which plays an
important role in growth, development, and metabolism in many fishes.
• Thyroxin is quite important in development, including the sometimes
extreme morphological and physiological changes associated with
metamorphosis – such as the transformation of flounder from larvae with
an eye on each side of the head to flatfish with both eyes on one side of
the head. It also initiates seaward migratory behavior and the
accompanying osmoregulatory adaptations of juvenile salmonids during
their seaward spawning migration.
Melatonin

• Produced by the pineal gland and the retina of the eye, is secreted
during the dark phase of daily light–dark cycles and helps regulate fish
responses to daily and annual cycles of daylight. This hormone
influences many physiological processes and behaviors through its role
in the maintenance of circadian activity cycles, daily changes in
temperature preference, and changes in growth and coloration
associated with changes in photoperiod and temperature
II. Automatic Nervous System in controlling Various
responses

• Involuntary physiological functions, such as control of internal organ function, are at least in part
controlled by the autonomic nervous system or ANS. Neural signals from the central nervous system
(brain and spinal cord) travel to ganglia of the ANS that are located either along the spinal cord or
near or within the target organs.
• The ANS often works together with the endocrine system to control involuntary physiological
functions such as heart rate, blood pressure, blood flow through the gills, and many functions of the
gastrointestinal system that are important to digestion and nutrition.
• The autonomic nerves in fishes, control the aperture of iris, blood pressure, blood flow through gills
for oxygenation and blood supply to various parts of the body automatically. It controls heart
performance, gastric motility and control the function of swim bladder. It also controls the colour
changes and release of catecholamines from the chromaffin tissue.
• The autonomic nervous system in strict sense can be defined as that portion of the peripheral nervous
system which conduct impulse to visceral organs, glands and blood vessels or in other words smooth
muscles, cardiac muscles and glandular epithelium. The effectors are present in the organs.
Types of Autonomic Nervous System:

It is sub-divided into two types. They are as follows:

• Sympathetic Nervous System,
• Parasympathetic Nervous System.
Sympathetic Nervous System
 There are two sympathetic chains extending from the
first spinal nerves to the end of the tail. On each
sympathetic chain there is ganglionic swellings, known
as sym­pathetic chain ganglia or paravertebral ganglia Organization of Sympathetic
(Fig. 13.2). In dipnoans the ganglionic swellings are Nervous system
not distinct on the sympathetic trunk. Actually the
sympathetic cells are scattered along the cord or
grouped near the junctions with the remi
communicanes.
 Generally there are two sympathetic ganglia on each
spinal segment. There are two separate sympathetic
cords in the anterior trunk region, but these trunks
fuse to form a single cord between the kidneys.
 The first two sympathetic ganglia give out rami commu­
nicanes, which are made up of postganglionic fibres,
most of the preganglionic fibres to these ganglia run
out at more posterior levels, and travel forwards in
sympathetic chain. Preganglionic fibers do not
terminate in the nearest ganglion but may cross over to
the opposite sympathetic cord to run to higher or lower
levels in the sympathetic chain.
 Postganglionic fibres in the grey rami run through spinal nerves to blood vessels and chromatophores and
small non-medullated nerves may innervate directly from the sympa­thetic chain to the arteries.

 The splanchnic nerve arises on the right side from the first two sympathetic ganglia and a commissure at
the same level to contribute fibres from the left sympathetic cord. This splanchnic nerve provides the entire
sympathetic innervation of the gut and its appendages.

 In the anterior trunk region the sympathetic ganglia give out many small nerves, which innervate. In the
posterior trunk region the sympathetic ganglia give out genital nerves which innervate the gonads.

 Vesicular nerve also arises from sympa­thetic ganglia which innervate the urinary bladder and mesonephric
ducts. In teleosts the presence of sympathetic cardiac nerves is not evident. The sympathetic fibres enter
the heart through vagus.
Parasympathetic Nervous System
 There is no evidence of spinal parasympathetic outflow in fish. Autonomic pathway in the Diagram of autonomic
III or oculomotor cranial nerve gives out preganglonic nerve heads of selachian fish Nervous system in the
fibres to the ciliary ganglia and from the ciliary ganglia head of teleost fish
postganglionic nerve fibres innervate, the eyeball.
 Oculomotor nerves are absent in forms with reduced eyes. In
Polydon ancillary ganglion has been observed, but in
Scohirhynchus there are ciliary ganglia on the ventral division
of the oculomotor nerves.
 In teleosts, parasympathetic autono­mic fibres are present only
in oculomotor III and X (vagus) (Fig. 13.5). In dipnoans, the
cranial autonomic fibres are present only in vagus (X) in
Protopterus and Lepidosiren while the Neoceratodus
oculomotor and vagus are given out from the cranial outflow.
 The ocular outflow has preganglionic fibres, which form
synaptic relations with postganglionic ciliary neurons,
proceed to the eyeball and neighbouring arteries. A large
number of sympathetic fibres join the vagi forming
vagosympathetic trunk, which carries both cranial and spinal
autonomic fibres to gills, heart, stomach and swim bladder.
 Adrenergic
 The adrenergic neurons in teleost fish contain both adrenaline and noradrenaline with predominance of
adrenaline. The adrenergic transmitters act as adrenoreceptor (adrenergic receptors) either the alpha or beta
type in effector organs.

Cholinergic
 Generally the pre- and postganglionic nerves secrete ACh. The receptors of the postganglionic neurons are of
the nicotinic type. ACh released from postganglionic cholinergic nerve endings acts as muscarinic receptor in
effector organs in teleosts, elasmobranch and dipnoans in all higher vertebrates.

Cholinergic tissue
 The chromaffin tissue is present in the sympathetic ganglia of elasmobranch and contains more of
noradrenaline than adrenaline.
 III. TEMPERATURE RELATIONSHIPS

 The view that many people have of fishes as “cold-blooded” is not accurate. Most fishes are about the same
temperature as the surrounding water, which may be cold or warm depending on the habitat. That
temperature can change, but usually any change is slow due to the thermal stability of water. Animals that
rely primarily on external heat sources are referred to as ectotherms, and include most fishes.

 Most fishes are ectothermic because they lack any mechanism for heat production and retention. In addition,
when blood flows through the gills it becomes the same temperature as the surrounding water due to the thin
gill membranes, before then flowing to the rest of the fish’s body. There are, however, interesting exceptions
of heat production or conservation in some fishes, a condition often referred to as either heterothermy or
regional endothermy.
1. Coping with Temperature Flactuations
 Fishes that experience changing environmental temperatures, such as those characteristic of diel or seasonal
changes, have several cellular and subcellular mechanisms for adapting to the new set of conditions.
 Physiological say adjustments are the result of switching on or off genes that are responsible for the
manufacture of particular proteins. For example, acute heat stress initiates the synthesis of stress proteins,
also known as heat shock proteins or HSPs, which maintain the structural integrity of proteins that
otherwise would become denatured at higher temperatures, thereby allowing them to function
biochemically.
 In some fishes alternative enzymes (termed isozymes) may be produced to catalyze the same reaction more
efficiently at different temperatures. Isozymes are regulated by switching on or off the different genes that
control their production.
 Polyploid species have extra sets of chromosomes, and may have a better capacity to cope with a wide range
of temperatures; perhaps the multiple copies of genes provide more opportunities for evolution to bring
about changes in alleles that may prove to be beneficial. For example, among cyprinids, Goldfish and
Common Carp are both polyploid and can tolerate a wide range of temperatures, and the polyploid Barbel
can acclimate better to different temperatures than can the diploid Tinfoil Barb.
 Some fishes exhibit allozymes, alternative forms of the same enzyme that are controlled by different alleles of
the same gene. Different populations of the species may exhibit higher or lower frequencies of the appropriate
alleles depending on their geographic location.
IV. Water, Solute & pH Balance
Osmoregulation
 Osmoregulation is the process of maintaining an internal balance of salt and water in a fish’s body. A fish is, after all, a
collection of fluids floating in a fluid environment, with only a thin skin to separate the two.
 One of the most important homeostatic functions of living organisms is proper regulation of the internal osmotic
environment. Deviation from the normal range can jeopardize proper physiological function through water loss or gain,
the changing of internal ionic concentrations, and shifts in ionic and osmotic gradients.

 Most fishes, like all other vertebrates, are osmoregulators – they regulate their internal osmotic
environment within a fairly narrow range that is suitable for proper cellular function, even if the external
osmotic environment fluctuates.
 Gills are an important osmoregulatory and excretory organ for fishes. Their large surface area, thin
membranes, and highly specialized cell types make them well suited for this role.
 Fishes that can tolerate only small changes in the solute concentration of their external environment are
referred to as stenohaline, whereas those with the ability to osmoregulate over a wide range of
environmental salinities are euryhaline.
1. Osmoregulation in fishes

Osmoregulation in Agnathans

 Hagfishes (Myxinidae, Myxiniforms), are osmoconformers, similar to many marine invertebrates. Their overall internal
osmotic concentration is about the same as that of sea water. Because they live in fairly stable osmotic conditions near the
bottom in relatively deep water, they do not Homeostasis have to contend with internal osmotic instability. Although the
overall internal osmotic concentration of hagfishes is the same as the ocean, there are differences in the concentrations of
some individual ions.

Osmoregulation in Elasmobranchs

 To prevent osmotic stress in the hyperosmotic marine environment, marine elasmobranchs convert their nitrogen wastes
into urea and retain high concentrations of it in their blood. This, in addition to trimethylamine oxide (TMAO), which helps
to stabilize proteins against the denaturing effect of urea, gives elasmobranch blood an osmotic concentration slightly higher
than that of sea water. Elasmobranch gills are not readily permeable to urea, and this is probably enhanced by the cells
transporting urea back into the blood and thereby reducing the gradient between the cell and the surrounding water
(Marshall & Grosell 2006). As a result of this urea retention, elasmobranchs are hyperosmotic to sea water and gain water by
diffusion across their gills.
Osmoregulation in Sarcopterygians
 The coelacanths (Coelacanthidae) are extant marine sarcopterygians that also maintain elevated levels of urea
and TMAO in their blood to offset the high ionic concentration of the external environment, as do the marine
elasmobranchs. The African and South American lungfishes (Dipnoi) are freshwater sarcopterygians that can
survive long periods of drought by estivating in mud burrows. During this estivation period they produce and
retain high levels of urea, perhaps as a way of storing their nitrogen wastes in a form that is less toxic than
ammonia and perhaps to help retain water. The phylogenetic distance between the sarcopterygians and the
elasmobranchs, and the fact that both groups use urea as a nitrogenous waste and osmolyte, indicates an
example of convergent evolution in the face of similar physiological challenges.

Osmoregulation in Freshwater Teleosts

 Freshwater teleosts are hyperosmotic to their
environment and therefore tend to gain water
and lose solutes by diffusion across the thin
membranes of the gills and pharynx. Solutes
also are lost in the urine. If left unchecked,
the fish’s cells would swell and burst from the
constant influx of water. To prevent this,
freshwater fishes excrete a large volume of
dilute urine and actively transport solutes
back into their blood. Some of these solutes
are recovered from urine as it is being formed
in the kidney tubules.
Osmoregulation in Marine Teleosts

 Teleosts that migrate between fresh and salt water, such as salmonids, must make appropriate adjustments in
the mitochondria-rich cells of the gill epithelium to physiologically adapt to the dramatic change in
osmoregulatory environment. For example, as Arctic Char migrate from the ocean into rivers, the membrane
proteins of the gill epithelial cells responsible for sodium–potassium exchange increase in abundance, and so
do the plasma sodium concentration and blood osmolarity (Bystriansky et al. 2007). This apparently is the
result of increased activity of the genetic and molecular mechanisms responsible for creating these proteins,
and also provides evidence that this sodium–potassium exchange plays an important role in sodium uptake in
freshwater fishes. Bystriansky et al. (2007) also note that there are apparently two forms of this sodium–
potassium exchange protein, one that excretes excess sodium from saltwater fishes and another that assists
with sodium uptake in freshwater fishes
2. Control of osmoregulation and excretion