.

UNIT 12 ENDOCRINE SYSTEM

Structure
12.1 Introduction
Objectives
12.2 Endocrine Glands in Mammals
Pituitary
Thyroid
Parathyroid
Pancrcas
Adrenals
Gastrointestinal Hormones
Gonadal Hormones
12.3 Agnatha
12.4 Fishes
Elasmobranchs
Bonyfishes
12.5 Amphibians
12.6 Reptiles
12.7 Birds
12.8 Summary
12.9 Terminal Questions
12.10 Answers

12.1 INTRODUCTION
In the previous unit you have learnt about the nervous system of vertebrates. The nervous
and endocrine systems function to integrate the activities of the organisms. This is done
by their ability to synthesise and release chemical substances in the body. Certain nerve
cells, also known as neurosecretory cells produce substances, the neurohumours that act
locally. The secretions of the endocrine glands, the hormones, have a wide range of
action. In could be said that while regulation by nervous system is localised and rapid,
regulation by endocrine system is more general and relatively slow. The regulatory
functions of the hormones include homeostasis or the maintenance of the constancy of
the internal environment, general metabolism, morphogenesis, mental abilities, initiation
and maintenance of reproductive processes and behaviour associated with sex, that
includes courtship, nest building, parental care and migration.

In this unit, we shall first study the endocrine glands of the mammals, the hormones they
secrete and the functions they perform. This would be followed by a study of the
endocrine glands in different classes of vertebrates.

Objectives
Afler reading this unit you should be able to:

• describe the location, structure and functions of the endocrine glands of humans,
• list the secretions of the different endocrine glands in mammals,
• describe the endocrine structures of sub-mammalian species,
• discuss the variation in structure and function of endocrine glands of mammals
and other vertebrates.

12.2 ENDOCRINE GLANDS IN MAMMALS

A detailed description of endocrine glands of animals groups is provided in unit 10 of
Block 2, Physiology course (LSE-05) which you might have completed by now. In the
same unit the role of the hormones in energy metabolism is also discussed. In unit 15 of
Cell Biology course (LSE-01) you studied about the cellular mechanisms of hormone
action. It is advisable that you may go through these units before you begin a study of
this unit. In the present unit we shall describe the structure and functions of major
endocrine organs of vertebrates. We shall begin with the description of the endocrine
structures as found in mammals.

117
Functional Anatomy of
Chordates - II 12.2.1 Pituitary
The pituitary gland is a small unpaired structure found attached to the ventral side of the
brain. More specifically, themrgan, also known as hypophysis cerebri is found attached
to the hypothalamus on the floor of6tencephalon by a narrow stalk and is located in a
depression, sella turcica in the sphenoidal region of the skull (Fig.12.1).
Th ird ventricle Tubero-hypophyseal tract
Subarachnoid space
Mammillary body
Pia mater
Optic chiasma Cerebral peduncle

Durwmater Subarachnoid space

Supraoptical-hypophyseal Arachnoid membrane
tract Diaphragma sellae
Periosteum
P£lrsnervosa
Pars tuberalis
Pars intermedia
Dura mater Pons
Pars distal is Subdural space
Residual lumen Sphenoid bone "
Fig. 12. I: Diagrammatic representation or» Section through the hypophysis cerebri of human being
showing its relationship to adjacent structures.

The pituitary consists of two major parts, a larger anterior part, the adenohypophysis
(pars buccalis) and a smaller posterior part, the neurohypophysis (pars nervosa). Both
the structures are ectodermal derivatives. The hypophysis originates from two sources,
one as an outgrowth from the floor of the diencephalon - the infundibulum and the other
as an evagination of the roof of the stomodaeum - the Rathke's pouch. The distal part of
Rathke's pouch becomes separated form it and comes into close contact with the
infundibular outgrowth to develop into adenohypophysis (Fig.12.2). The infundibular
outgrowth is connected with the brain by the infundibular stalk and forms the
neurohypophysis.

Fig. 12.2: Stages in embryonic developiiieni of pituitary.

The adenohypophysis itself shows further divisions. The part of the adenohypophysis that
is in immediate contact with neurohypophysis is pars intermedia and the rest of it is
118
 greatly thickened to form pars distalis. The highly vascular adenohypophysis is formed Endocrine System
of cells arranged in a chord-like trabaculae separated by thin walled siniisoids. There are
two types of cells, the secretory, granular chromophil cells and the non-secretory, non-
granular chromophobe cells. The neurohypophysis is differentiated into pars nervosa,
the infundibular stem and the median eminence (Fig. 12.3). Pars nervosa, though
derived from the brain does not contain any nerve cells, but consists of neuroglial cells
and piginented cells called pituicytes (Fig. 12.3). The infundibular stem connects the pars
nervosa with the brain. The median eminence containing a rich supply of blood
capillaries is the swollen base of the infundibulum. "Fhe pars intermedia and the pars
distalis together form anterior pituitary, and the pars nervosa, infundibular stem and
median eminence collectively form the postefior pituitary. The anterior lobe has two
sources of blood supply. One source is the internal carotid artery and the other source is
the hypophysial portal vein. The posterior lobe is supplied with inferior hypophysial
artery.

F:g. I 2.3: rh‹ ii»i«-y gland — blood circulation.

The secretion of pituitary hormones is regulated by the neurosecretion from

hypothalamus, which lies on the floor of diencephalon. The neurosecretory materials
from the neurosecretory cells of hypothalamus are transported by the axons and enter into
the median em inence. The portal vein from the median eminence transports the materials
to the anterior pituitary where they regulate the secretion of specific hormones such as
adrenocorticotropic hormone, somatotropic hormone, follicle stimu ating hormone and
lutinizing hormone. Certain other neurosecretory fibres travel through the infundibular
tract and enter into the posterior lobe and release into it the neuropeptides oxytocin and
vasopressin. Table 12. l provides the list of hormones secreted from pituitary gland, the
site of synthesis and their possible functions.

Table 12.1: Hormones of the pituitary gland.

Region Hormone Site of Functions
synthesis
Pars Soinatolropin (STI-1) Acidophil Promotes protein synthesis and deposition of
Distal is Growth hormone cells glycogen in muscles and liver. Mobilises fats lrom
tissues. Promotes growth o1’ bones and muscles.

Adrenocoriicotropin Basophil cells Stimulatss secretion ot’adrenocortico steroids such

(ACTH) as glucocorticosteroids and mineralocorticosteroids
by adrenal cortex.

Thyiotropin (TSIJ) Basophil cells Stimulates thyroid gland to synthesise and release
thyroid hormone, the thyroxine.

119
Functional Anatomy of
Chordates - II Cionadotropins (CiH) Basophil cells Promotes gametogenesis and secondary sexual
(a) Follicle characters. Stimulates growth of ovarian follicles
stimulating and ovulation in females; stimulates testicular
hormone (FSH) development in males.

Basophil cells Along with FSH stimulates ovarian follicles and

(b) Lutinizing estrogen and progesterone production; helps in the
hormone (LH) formation of corpus luteum; in males stimulates
interstitial cells of testis and secretion of and rogens.

Acidophil Stimulates development of mammary glands and

(c) Prolactin or cells milk secretion. Essential for the maintenance of
luteotropic corpus luteum.
hormone (LTH)
Pars Stimulates production of melanin pigment in fishes
iiitermedia Melanophore and amphibians; in mammals tyrosine activation by
stimulating hormone the hormone induces melanin production.
(MSH) Melanotropin
Neurohy- Hypothalamic Aids in the resorption of water in kidney tubules.
pophysis Vasopressin or ai1ti- neurons A lso acts as a vasoconstrictor.
diuretic hgmone
(ADH)
Hypothalamic Causes contraction of sinootll muscles oùmammary
LX}’(OCif\ Neurons glands, IJterine wall.

Before we begin describing other endocrine glands, you may attempt the following SAQ.

SAQ 1
I. Fill in the blanks by using suitable words.
(a)............................................................................is found attached to hypothalamus
on the floor of diencephalon by a narrow stalk.
b) Tire two major parts of pituitary are...........................and
c) The two divisions of adenohypophysis are...........................................and

d) The three regions of posterior pituitary are .. ,

.............................. and ............... ..................
e) The two hormones stored in neurohypophysis are...........................................and

Il. List the hormones secreted by anterior pituitary and their functions.

................... ... ....................... ......................................................

........................... ......... .............. .................................................
......... ... . .... ............................... .. ..............................................
........................... ....... ................... ............................................ .
......................................... ..... .. ....................... .. .. ...................

12.2.2 Thyroid
Thyroid gland is generally located in the region of the neck. It is a two lobed structure,
the two lobes being connected by an isthmus on the ventral side of the larynx. Thyroid is
derived from endoderm and it arises as an unpaired median evagination of the floor of
the pharynx at the level of the first pair of pharyngeal pouches. Subsequently it detaches
itself from the pharynx and becomes a bilobed structure. The gland is formed of a number
of spherical follicles, each follicle having a central lumen which is surrounded by
secretory epithelial cells (Fig,12.4). The secretions of the cells are kept stored in the
lumen until they are released. In between the adjacent follicles, that is, in the
interfollicular spaces, there is a rich supply of blood vessels. The hormone thyroxine
secreted by the thyroid gland is a derivative of the amino acid tyrosine. The follicular
epithelial cells of the thyroid gland absorb the diffusible iodide ions (derived from the
food) from the blood vessels and convert them into iodine molecules. The free iodine is
then incorporated into tyrosine residues of thyroglobulin molecules leading to the
synthesis of triiodothyronine (T,) and thyroxine (+4) The T, and T4molecules are then
120 enzymatically released from the thyroglobulin molecule.
' Endocrine System

£''ig. 12.4: Thyroid glaad, a portion enlarged. a) •taia•d aectioa, b) di«grazezeatlc rcprñgcatatioa.

Thyroid secretions play a significant role in the maintenance of the normal meiatiolism of
the organism. It accelerates the mte of oxidative metabolism in the cells by uncoijpling
oxidation from phosphorylation. Thyroid also regulates the growth and morphogenesis
Metamorphosis is controlled by thyroxine in frogs. Thyroid facilitates carbohydfatñ
utilisation by promoting glucose release form glycogen molecules. Thyroidectomy in
young tadpoles prevents metamorphosis; similarly precocious metamorphosis is
witnessed on administration of thyroxine to young tadpoles. Thyroid is also known to
regulate water and salt content of the body in conjunction with the anti-diuretic hormone
of the pituitary gland. Thyroid also maintains normal activity of gonads and the
excitability of nerves and muscles.

12.2J Parathyroids
Parathyroid glands are found in close proximity to thymids and are found generally in
two pairs in mammals including man. They are endoderm derivatives from the third and
fourth pharyngeal pouches of the embryo and lose their connection with the walls of
the clefts in adults. The glandular epithelium is made of columnar cells and they release
their secretions in the numerous sinusoids or blood spaces between finn columns and
from there, they are carried to target organs. There are two types of cells; large and
non- granular chromophobe cells and the granular eosinophil cells (Fig. 12.5).
Parathyroid glands secrete two types of hormones — the parathormone and caleltonin.
The two hormones are involved in the regulation of calcium metabolism of the body
and are antagonistic in their functioning.

PBt¥tbyroid gland

'}g. t2 '\: Section of paratbyroid gland Ourronadtd by thyroid follicles.

121
Functional Anatomy of
Chordatcs - II
Whereas parathormone regulates the uptake of calcium and its deposition in bones,
calcitonin mobilises calcium from bones and promotes restoration of calcium levels of the
blood.

12.2.4 Pancreas
Pancreas has b6th exocrine and endocrine elements in it. The acinar tissue produces
pancreatic juice that is released into intestine for performing digestive functions. The
endocrine functions are performed by the Inlet tissue, the hormones of which are
released into blood stream. The tissue, more specifically known as iibtc of
Leagerhsns (Fig.12.6) has Evo types of cells, the alpha and beta cells, each of which
produces a specific hormone. Pancreas arises as an evagination of the gut and is thus
endodermal in origin.

y
Section tbroagh rat pancreas ebowlng both ezocriae (pancreatic accini) ae well ae eadocr oc
dssae (islcta of Langerbaoe).

The alpha cells produce a hyperglycemic hormone glucagon and the beta cells produce a
hypoglycemic hormone insulin. The two hormones in conjunction regulate the glucose
metabolism of the organisms. Insulin also promotes fatty acid synthesis in liver.

12.2.5 Adrenals
Adrenal glands in general lie in close association with the anterior end of kidneys. The
glands show two distinct regions, the outer cortex and the inner medulla. In primitive
vertebrates, like agnathans and fishes, the two regions are spatially separated, but in
higher forms the cells are interspersed. In mammals, the medulla is encapsulated in the
cortex (12.27 a). Nevertheless, the two regions of the gland differ in their origin, structure
and function.

Adrenal corten
In mammals cells of adrenal cortex are budded off from the mesodermal inter-renal
tissues. The cells are arranged in masses known as the inter-renal bodies in non-
mammalian vertebrates. In mammals they surround the medulla to form cortex. The
cortex shows three regions, the outer glomenilosa, the middle fasciculata and the inner
reticulosa (Figs12.7 b). The cortical tissue designated as steroidogenic tissue secretes
steroidal hormones, the glucocorticoids and mineralocorticoids. Glucocorticoids, such as
cortisol play a significant role in carbohydrate and protein metabolism. They promote
gluconeogenesis. Mineralocorticoids, on the other hand regulate water and electrolyte
balance of the body. Desoxycorticosterone and aldosterone are the. two
mineralocorticoids. Essentially, they cause the retention of sodium and increased
elimination of potassium and phosphates. They also cause a resorption of sodium in
kidney tubules and a reduction of resorption of potassium.

Adrenal medulla
Adrenal medulla arises from the neural crest cells of the neurectoderm. A group of cells
budded off frotn the neural crest cells, the phaecochromoblasts, become glandular and
form the cells of adrenal medulla (Fig.12.7). The medullary cells have a characteristic
histochemical reaction, staining dark brown with‘chromates and hence knowii as
chromafFm cells and the tissues composed of them is known as chromaffin tissue. The
secretions of these cells, known as catecholamines, are adrenaline and noradrenaline (also
known as epinephrine and norepinephrine respectively). Both the hormones are derived
122
 Endocrine System

Cortex Zona fasciculata

Medulla

Fig. 12.7: A section through part of adrenal glaad in a mammal, with division into cortex aad medullary
regionsi a) stained section, b) diagraminatic representation.

The effects of both the hormones are more or less the same. They cause a rise in
blood pressure by vasoconstriction. Adrenaline constricts vessels of the visceral area and
the skin; in small doses it dilates the vessels of muscles and in large doses constricts
them. lt has no effect on pulmonary and cerebral circulation and it dilates the coronary
arterioles. Noradrenaline has a constriction action on muscles of blood vessels. In man
the peripheral resistance is increased by noradrenaline and lowered by adrenaline. Both
the hormones stimulate the constriction of spleen thus causing an increase in the
circulating blood volume and the concentration of erythrocytes in circulation. Both the
hormones also increase the rate of heart beat. The two hormones cause an increase in
blood sugar levels, by effecting glycolysis in liver. Adrenaline also promotes indirectly
carbohydrate metabolism by stimulating the release of ACTH from pituitary. The
ACTH in turn releases glucocorticoids which regulate carbohydrate metabolism. The
secretion of adrenaline is regulated by sympathetic nervous system.

12.2.6 Gastrointestinal Hormones

There are a f'ew gastrointestinal hormones, polypeptides in nature, that are secreted by the
mucosa of gut and are concerned with the regulation of gastrointestinal activity in mammals.
Some of these polypeptides are gastrin, secretin and cholecystokinin. Many other
polypeptide hormones are being added to the list but their functions are not clearly defined.

Gastrin secreted by gastric mucosa regulates the secretion of HCI in the stomach. The
hormone is stimulated to be produced by the distension of the stomach as well as by some of
the food substances and is released into blood. Gastrin in turn acts on the parietal cells to
release HCI into the stomach. The HCI secreted serves as a feedback to inhibit further gastrin
production. Thus gastrin serves to regulate the volume of HCI production in the stomach. The
proper concentration of acid determines the peptic activity within the stomach.

Both secretin and cholecystokinin are secreted by duodenal mucosa. Secretin is released
when the stomach pH is low or in the presence of digested fat or bile. The hormone
initiates the release of large quantities of enzyme rich pancreatic juice. Cholecystokinin is
secteted under the ‘stimulation of partially digested proteins. The two hormones inhibit
stomach motility and secretion. Once the chyme leaves the stomach and arrives at
intestine, the activity of the stomach slows down and hormones production is inhibited.
123
Functional Anatomy of
C'hordatc0 - II 12.2.7 GonadBl Hormones
Gonads, the testis and ovary, perform dual functions. One, they are the sites of
gametogenesis, the sites at which the sperm and ova are produced respectively; and
two, they are the centers of intense endocrine activity — namely, they are the sites at
which the hormones which regulate the reproductive activity of the organism are
synthesised and secreted.

Testis
Testis is composed of seminiferous tubules in which sperm are produced and interstitial
cells or Leydig cells(Fig.12.8) which secretes the male hormones. Testosterone and
androsterone, the androgens, derived from testis play a significant role in male reproduction
and of the two; testosterone is seven to ten times more potent than androsterone. Androgens
promote growth stimulate the synthesis of cell proteins. They stimulate the adrenal
mineralocorticoids and cause the retention of sodium, potassium, phosphorus and sulphur
that are utilised building up of tissues. The'uynthesis and release of and organs is
under the control of lutinising hormone, otherwise known as interstitial cell stimulating
hormone (ICSH), a gonadotropic hormone. Reciprocally, the androgens regulate the
secretion of the gonadotropins, either by stimulating their synthesis or as in most cases,
inhibiting their secretion. Androgens in low titers are essential for spermatogenesis and
high titres suppress the gonadotropin release.

Pig. 12.6: Section a at tcstis. IC,1ntcrstitiaI cells of Leydig, 3-F, T Etic Tubule.

Ovary
Ovary consists of a fibrous tissue, the stroma and the Graafian follicles in which ova
develop. Besides the production of ova, ovary is also an endocrine centre. The two steroid
hormones produced by ovary are estrogens and progesterone. Ovary also secretes a
proteinaceous hormone, relaxin.

Fig. t2.9: Section ofrxt ovary to showing Graaflan follicle with granulosc cells (G) and fluid filled antrum
(A). thccql cells {T}. -
I24
Estrogens are a group of related chemical compounds; the important ones are esrradiol, estriol Endocrine System
and estrone. Estradiol is more potent than the other two. Estrogens regulate the development and
maintenance of the morphological and functional state of the reproductive system. They also
regulate the development of accessory genital organs and secondary sexual characters, such as the
development of uterine tubes, vagina and the mammary glands. Ovulation and the prepamtion of
the endometrium for the implantation of the ovum are other functions controlled by estrogens.
Besides, estrogens also influence nitrogen and electrolyte metabolism, water balance and
formation of bones. Estrogens also promote the retention of water and sodium chloride, decrease
phosphate levels in blood and reduce the elimination of calcium and phosphorus.

Progesterone‘is secreted by corpus luteum under the influence of luteotropic hormone

from pituitary. It is the ruptured follicle of the ovary thct tmnsforms into an yellowish
coloured corpus luteum and assumes an endocrine function. Progesterone is also
secreted by placenta, later during pregnancy. In pregnant women it can be detected in
urine. Like estrogens, the progesterone also performs a variety of functions. It prepares
the uterine wall for the implantation of fertilised ovum and maintains gestation.
Progesterone supplements the estrogen activity in the development of mammary gland.

Relaxin is a polypeptide hormone secreted under the influence of estradiol and

progesterone. The hormone relaxes the pubic symphysis at the time of parturition.

Thus far we made a general survey of some of the important endocrine secretions. Now
let us begin the study of hormones in specific groups of vertebrates. Before we do
so, please attempt the following SAQ.

SAQ 2
State whether the following statements are true or false.
a) Thyroid is derived form ectoderm and arises as an unpaired median
evagination on the floor of the pharynx.

b) Thyroxine is a derivative of the amino acid tyrosine.

c) Thyroidectomy of young tadpoles promotes metamorphosis.

d) In mammals parathyroid glands are paired sUuctures.

o)

g)

h) Cortisol regulates carbohydrate and protein metabolism.

i) Th: :
gon of HCI in the stomach is carried out by
a! he”se eetr:

j) Secretin and cholecystokinin are secreted by duodenal mucosa.

k) Synthesis and release of androgens are under the control of follicle

stimulating hormone produced from anterior piNitary.

i) yhno gesterone and protein are the steroid hormones produced

12.3 AGNATHA
In agnathans pituitary and adrenals are present and function as endocrine organs.
In lampreys (Fig. 12. 10) pituitary is located and pressed against the underside of
 hypothalamus. A h pothalamo-hypophysial portal system making a neurovascular link
between hypothalamus and hypophysis does not appear to be present. Essentially the
pituitary functions independently in lampreys. Pituitary lacks a median eminence but a
posterior thickening of the infundibular floor forms a neural lobe, the pars nervosa —a
structure comparable with the neural lobe of higher vertebrates. The neural lobe of the
pituitary, the neurohypophysis (Fig.12.10) contains a hormone that regulates the water
content of the body. The hormone, arginine vasotocin is produced in the neurosecretory
cells of the preoptic nucleus of the brain, carried down the axons and stored in the neural
lobe. Thus the lamprey pituitary may show many of the main features found in higher
vertebrates, although it may not secrete full complement of hormones. The control system
also appears to be a simple one. The pituitary is not regulated by the hypothalamic
regulating factors but is controlled by feedback from other tissues.

Neurol lobe

Rostriil pars distalis Pors intermedia

Proximal pars distalis

Fig. 12.10: Pituitary of lamprey.

Adenohypophysis, the glandular portion of the pituitary, shows three distinct regions.
One, the rostral and two, the proximal regions of the par distalis are separated from the
third region, the posterior region, the pars intermdeia by a connective tissue. Pars
intermedia is shown to secrete as in other vertebrates, the melanocyte stimulating
hormone (MSH) and the secretion appears to be under the control of the pineal eye by
way of hypothalamlus. Removal of hypophysis results in the non-development or gametes
and secondary sexual characters, suggesting that the pituitary is the source of
gonadotropic hormone. The basophil cells of the proximal pars distalis secrete the
gonadotropic hormone. The acidophil célls of the proximal pars dilstalis produce a
corticotropic hormone. The basophil cells located at the rostral parts of pars distalis tend
to proliferate at the time of metamorphosis, and were thought to be thyrotropes; but there
is no evidence of thyrotropin presence and metamorphosis does not appear to be under
the control of thyrqid gland.

Lampreys do not pessess compact adrenal glands, but cells which represent cortical and
medullary parts in higher vertebrates are present. The interrenal cortical cells are present
as a group in the region of pronephros just beneath the cardinal veins. Nevertheless
there is no evidence of their secreting any steroid hormones nor such hormones have
been detected in blood. The suprarenal medullary cells, in the form of chromaffin cells
are found in the walls of heart and large blood vessels. These cells answer positively
for the presence of catecholamines. Adrenaline and noradrenaline have been shown to be
present
in the tissues of time heart. ‘

12.4 FISHES
12.4.1 ElasmDbranchs
Pituitary
In elasmobranchs, usual parts of the pituitary, the pars distalis (anterior), the pars
intermedia and the heurohypophysis are present. In addition, the pituitary also has a
ventral lobe that is partly separate (Fig. 12.11). The ventral lobe is believed to be part
of anterior pituitary, containing a gonadotropin and thyrotropin. Removal of the ventral
lobe causes regression of the gonads in dogfish. Adenohypophysis is known to contain
126 lactotropic hormone and adrenocorticotropic hormone. The neurointermedia receives a
number of fibres from neurohypophysial tract. This region contains melanophore-
stimulating hormone (MSH). Arginine vasotocin that regulates water metabolism is also Endocrine System
known to be present. Besides, three neural octapeptides of unknown function are also
present.

Pre-optic nucleus
Saccus vasculosus

Median eminence
anterior postérior

T4euno
intermedia

O
O OO
Rostral pars distalis
@ Ventral lobe
Proximal pars distalis *

Fig. 12. I I: .4elachiati }situitary. SuiId arruws show reins, broken arrows — arterin and tbin arrows
portal veins.

Regulation of the pituitary is carried out by hypothalamus by way of a portal system that
passes through the median eminence (Fig.12.13). But the portal system does not reach the
ventral lobe of the pituitary. lt is believed that the regulation of ventral lobe is achieved
by way of the systemic blood stream by a relay in the pars distalis. In elasmobranchs,
unlike teleosts, there is no direct innervation of pars distalis.

Thyroid and Parathyroids

Thyroid in elasmobranchs arises as a downgrowth from the fioor of the pharynx. It
remains attached to the fioor of the pharynx by a narrow stalk containing a.small ciliated
pit. The functions of thyroid in elasmobranchs are not clearly known, but the gland shows
cyclical changes related to migration and reproduction. In dogfishes, removal of thyroid
results in non-deposition of yolk in eggs.

Hi fishes there are no parathyroids. Follicular structures known as ultimobranchial glands

are known to produce calcitonin, a hormone that regulates calcium metabolism.

Adrenals
In elasmobranchs, unlike in mammals, the cortical and medullary parts of the adrenal
glands are found widely separated. The part corresponding to medullary tissue is known
as suprarenal bodies. They occur as segmental series of glands and project into the
dorsal wall of the posterior cardinal sinus. The more anterior ones are fused to form an
elongated structure on either side of oesophagus. These structures are shown to be rich in
noradrenaline. The segmental series of suprarenal bodies continue along the whole length
of the abdomen, and more posteriorly they are embedded in the kidney tissue (Fig.12.12).
The posterior renal bodies, larger in males as compared to females stain positively for the
presence of adrenaline.

Interrenal bodies located in the kidney region represent the region corresponding to
adrenal cortex. In .some species they are unpaired and located medially, and in others
they are paired. The interrenal bodies are not associated with the suprarenal bodies in
anyway and it appears that the functioning of the two bodies is not interdependent in
elasmobranchs. The gland is stimulated by stress or by mammalian ACTH. Removal of
the interrenal bodies results in the death of the organism, but extracts of the tissue can
prolong the life. Secretions of interrenal bodies have been shown to regulate
carbohydrate metabolism of the animal and activity of the gonads; apparently they have
no role in electrolyte balance. The steroid secreted by the interrenal bodies has been
shown to be 1-It-liydroxycorticosterone and there is no evidence of aldosterone being
secreted.

127
Functional Anytoł»y cf
Chordntcs - II

big. 12.ł2: Saprarcnał »nd intcrrenal budics in hindcr n›csonephcric regian of dogfish.

Pancreas
Pancreas in elasmobranchs does not occur as islets, as it occurs in mammals but as outer
layer of some or the ducts. The pancreas of elasmobranchs secretes both insulin and
glucagon.

As in mammals, in elasmobranclls also, the gonąds have endocrine tissue that produces
steroid hormones. The interstitial cells ofthe testis secrete testosterone, androstenedione
and progesterone. Similarly ovarian follicles are shown to produce progesterone.

12.4.2 Bonyfisbes
Pituitary
Pituitary of bonyflshes, like other vertebrates shows two regions, the adenohypophysis
and the neurohypephysis. The adenohypophysis surrounds the neurohypophysis and
interdigitates with it. lt shows three divisions, the pars rostral distalis and the pars
proximal distalis and pars intermedia, the last region surrounding neurohypophysis
(Fig.12. 13). Rostral pars distalis contains acidophil lactotropes, qorticotropes and the
basophilic thyrotropes. The internal carotid artery that supplies arterial blood to the
pituitary forms the primary capillary plexus in the anterior neurohypophysis. It then
continues into the pars distalis region as a system of sinusoids among the secretory cells,
forming the secondary capillary plexus. Such an arrangement of blood supply in bony
fishes is in contrabt to the one found in other vertebrates in wlJich long extemal portal
vessels are (ound instead of a portal system of short capi llaries.

Ant

Neurohypophysis
Sacculus vasculosus 3“ ventricle
OTttCOtr€i}jgg

ars intermedia
Lactotrope follicles,
’Somatotropes
Gonadotropes niyrotropes
i
I28 ^*'^ ' Rostial pars distalis
Fig. 12.ł3: Pituitary of ecł, mid sagittał scction .
Two types of nerve fibres enter into pituitary from hypothalamus. Type A fibres. Type A .@docrine Syitem
peptidergic fibres end mainly around the capillaries or the cells of the neurohypophysis.
Type B cells that are mainly aminergic pasx on to innervate the cells of pars distalis.
Fishes possibly produce only one gonadotropin, the lutinizing hormone (LH).
Interestingly, injection of a fish gonadotropin into mammals produces some effects of
both FSH and LI-I; but only mammalian LH and not FSH induces gametogenesis and
steroid secretion in fishes.

Lactotropes, cells that secrete prolactin, are not regulated by hypothalamus. Thix is
evidenced by the fact that the lactotropes of the pituitary transplanted ectopically (away
from the bra:'n) are activated when transferred from dilute seawater to fresh water.
Prolactin of fishes ix concerned with osmoregulation. It reduces the exchange of sodium
across the surface of the body and gills. In some fishes, prolactin facilitates water
permeablity of the gills and enhances renal water excretion. The activity of lactotropes
increases and the prolactin titres are high in blood when migrant fishes move from
seawater to fresh water.

The pars intermedia produces melanophore-stimulating hormone that regulates the colour
changes. Pars intermedia has two types of cells and besides controlling colour changes,
the lobe is also known to regulate other activities such as water and electrolyte balance,
calcium levels of blood and probably reproduction as well.

Two octapeptides have been identified from the neurohypophysis of the bonyfishes. One
is the arginine vasotocin and the other is the isotocin or ichthyotocin, a homologue of
oxytocin. Both these peptides are secreted from the neurosecretory A fibres of pituitary.
But the functions of these hormones in bonyfishes are not known.

”thyroid
Thyroid gland is not a compact body but appears as scacered masses of tissue along the
ventral aorta. The thyroid secretes both triiodothyronine and thyroxine, hormones, which
are identical to those, found in mammals. The functions of thyroid in bonyfishes remain
uncertain. Thyroid tissues may occur in several parts of the body, kidneys, heart, eye and
other parts as well, particularly in those fishes that are deprived of iodine. The cyclic
activity of thyroid corresponds with the maturation of gonads implying a role for thyroid
in thereproductive process. However, thyioid does not appear to have any role in the
respiratory metabolism of bonyfishes, nor in growth or osmoregulation.

The suprarenal and interrenal tissues of adrenal gland of bonyfishes are found in masses
around the thickened walls of the posterior cardinal veins. Relaüvely littie is known about
the functions of the adrenal tissues of bonyfishes. Removal of the adrenal glands results
in the loss of sodium in freshwater and gain of sodium in seawater. In both cases, the
organisms fail to survive. A lthough adrenal tissues produce a number of steroids, there is
no evidence of aldosterone production. Dorsal to kidneys, a group of cells called
corpuscles of Stannius which do not produce any steroids, but which may be involved in
the regulation of calcium levels is present.

Ultimobranchial glands
Bonyfishes lack parathyroid glands. Calcitonin, which in higher vertebrates is produced
by parathyroid glands, is produced by a mass of cells developed from the floor of the last
branchial pouch, the ultimobranchial glands. The calcitonin of bonyfishes differs
markedly from that of mammals in the amino acid composition and is 25 times more
potent than the latter. Although the definite function of the calcitonin of fishes is not
clear, it is believed that it facilitates calcium transport.

Pancreas
Endocrine pancreas known as Brockerman bodies lie isolated from the exocrine tissue
and may occur scattered in the abdomen, in spleen and even in ovaries. Both insulin and
glucagon are produced by the specific types of cells.

129
Functional Anatomy of
Chordfltes - II UrO}3hysis

Urophysis (Fig.12.14) refers to masses of secretion produced by neurosecretory cells at

the hind end of the;spinal cord. These cells produce masses of neurosecretion. The
function of the cells appears to be regulation of the salt content of the body. At least two
hormones have been isolated from urophysis, urotensin I and urotensin II. These
hormones appear to regulate several aspects of the physiology of the organism including
osmoregulation, cardiovascular functions and reproduction.

Ependymal layer

Conducting&xons

Terminal storage bulbs

Fig. 12.IJ: \lrophysis in eel.

SAQ 3
Choose the correct word(s) from the alternatives provided.
a) 1n agnathans a hypothalamo-hypophyseal portal system linking hypothalamus and
anterior pituitrary is present /absent.
b) In lampreys/’ pituitary is regulated by hypothalamic regulating factors/feed back
from other tissues.
cj The pituitary of lampreys produces/does not produce a gonadotropic hormone.
d) Lampreys and mammals.have/do not have a similar type of compact adrenal gland.
e) The dorsal/ventral lobe of pituitary in elasmobranchs secretes a gonadotropin and
a thyrotropin,
Adrenal cortex/medulla in elasmobranchs is known as suprarenal bodies.
g) The steroid› hormone secreted by the interrenal bodies of elasmobranchs
is aldosterone/1-alpha-hydroxycorticosterone.
h) Melanophdre stimulating hormone of the bony fishes is secreted from the pars
distalis/pars intermedia region of the pituitary.
i) Thyroid gland in bony fishes occurs as a compact body in the neck
region/ scattered masses of tissue along the ventral aorta.
j) Removal of adrenal glands in bony fishes results in loss/gain of sodium in
freshwater fishes.

12.5 AMPHIBIANS
Pituitary
In amphibians the three main parts of the pituitary, pars distalis, pars intermedia and
pars nervosa are clearly distinguishable. Pars distalis lacks any divisions as witnessed in
fishes and resembles that of other tetrapods. "the releasing factors of the hypothalamus
pass through the median eminence via the portal system and control the secretions of
pars distalis. Pars distalis is known to produce two growth hormones (1) lactotropin
(LTH), a hormone that controls larval growth and (2) somatotropin (STH), that controls
adult growth. Metamorphosis is regulated by thyroxine whose secretion is regulated
by thyrotropin (TSF1) secreted by pituitary. The secretion of thyrotropin itself is regulated
by a releasing facter of hypothalamus, the TRF. ACTH is also secrete@by pars distalis
and its secretion is regulated by hypothalamic ACTH-Rf. Pars distalis also produces
130 two gonadotropins, FSH and LH.
"the intermediate lobe of pituitary produces a melanophore-stimulating hormone(MSH) Endocrine System
under inhibitory control from the hypothalamus. In frogs and toads, the neural lobe of
pituitary produces arginine vasotocin and mesotocin, which enhance water uptake by the
skin. Both in urodeles and anurans the hormones are known to reduce the glomerular
filtration. Such functions are more pronounced in landforms than the aquatic ones
suggesting that the evolution of these hormones have facilitated land life.

Thyroids and Parathyroids

Amphibians possess paired thyroids. These are located in close association with the
external jugular vein, one on each side. Thyroxine controls growth and metamorphosis in
amphibians. Thyroidectomy in young tadpoles keeps them in larval state throughout their
lives, but when fed on thyroid metamorphosis resumes and adult form emerges. In
amphibian adults there is no evidence that thyroids stimulate oxygen consumption.

Among vertebrates, parathyroids for the first time appear in amphibians as regulators of
mineral metabolism by secreting parathormone. The parathyroids are derived from
pharyngeal pouches. Parathyroidectomy in frogs results in reduction of the calcium levels
of blood and increase in phosphorous levels. lt is believed that parathyroids might have
evolved by the modification of the chloride secreting cells, the ionocytes of the gills of
fishes. Ultimobranchial bodies that develop from the floor of the last phary'ngeal pouch
secrete calcitonin, a hormone that has antagoni.stic functions to that of parathormone. The
hormone functions by mobilising calcium from the endolymphatic sacs, particularly at the
time of metamorphosis.

Adrenals
In amphibians, the ectodermal, adrenaline secreting chromaffin tissue of the suprarenals
is closely associated with steroid secreting mesodermal interrenal tissues. In fact
amphibians are the first vertebrates in which such an association is found. Whereas in
anurans the adrenals appear as compact orange masses on kidneys, in urodeles they form
an extended series. The action of amphibian adrenaline is similar to those of mammals,
namely, that it accelerates heart beat, enhances blood pressure and dilates lungs. It
also causes hyperglycemia and dilates pupils of eyes and causes the contraction of
melanophores. Adrenal cortex appears to have a role in carbohydrate metabolism as well
as in maintaining the water balance of the body. The details of hormones secreted are not
known.

Pancreas
Pancreas of amphibians contains both exocrine and endocrine tissues. The islets of
Langerhan's, which constitute the endocrine tissue, contain both o and § cells which
secrete glucagon and insulin respectively. The action of insulin in lowering blood sugar
levels and that of glucagon in enhancing it have been well demonstrated in amphibians.

12.6 REPTILES
Pituitary
As in amphibians, in reptiles also the pituitary contains the usual three parts. There is
also the portal system that carries blood from capillaries in median eminence to pars
distalis. Also as in amphibians the pars distalis contains five different types of
chromophilic cells that secrete different tropic hormones. Lactotropes that secrete
prolactin, somatotropes that secrete growth hormone, basophilic thyrotropes that secrete
thyrotropin, corticotropes that secrete corticotropin and gonadotropes that secrete
gonadotrpins are the five types of cells.

The pars intermedia that is distinctly large in those reptiles which change colour, secretes
melanophore stimulating hormone (MSH). In burrowing forms, the organ is very small.
The colour changing mechanism is very pronounced in certain lizards such as chameleon
and Anolis (also known as American chameleons). The colour may change with the
environment and thus serves the lizards to conceal themselves. Colour changes also occur
during courtship or when the organism is under threat. Changes in temperature or
environment are other factors that bring about varied colour patterns. The physiology of
colour changing mechanism varies with lizard. In Aiiolis the melanophores are not
innervated. Darkening effect is caused by MSH from pituitary and paling effect is caused
13 I
Functional Anatomy or
Chordotes - l t by adrenaline. In chameleons, however, the colour changing mechanism is under the
control of sympathetic nervous system. Colour changes are largely controlled through
eyes, and probably by some local control mechanism as well, as the covered areas of the
skin tend to become pale. Reptiles, like other tetrapods secrete the two gonadotropins,
FSH and LH. The neurohypophysis contains arginine vasotocin, the main anti-diuretic
hormone and mesotocin. ft appears that oxytocin-like hormone is present in
neurohypophysis, as the injection of the extracts of the organ to the viviparous lizard
Lacerla vivipara results in premature oviposition.

Thyroids and Parnthyroids

Reptiles have well-developed thyroid glands but their metabolic function is not clearly
understood. Injection of thyroxine (Tt) causes marked enhancement of oxygen
consumption at 3O°but has no effect at 20°. apparently the higher temperature is the
most
preferred temperature for the activity of thyroxine. Thyroid have a role in moulting, "
reproduction and egg-laying of lizards and snakes.

Parathyroids of reptiles are structurally similar to those of mammals. Parathyroidectomy

in lizards causes the blood calcium levels to be reduced to half its normal level and
phosphate levels to rise. In the absence of parathyroid gland, the animal becomes l
hyperexcitable and› has tetanic convulsions after exercise. Injections of the extracts of
parathyroid gland restore normally in the animal. Essentially there is a definite
mechanism for the regulation of calcium levels in the blood but the details are not as
clearly understood as in the case of mammals.

Adrenal glands
The adrenal glands are located very close to the gonads except in the case of turtles. 'Fhe
gland is a mixture of steroid secreting and catecholamine secreting tissues and the two
areas are not very clearly defined. As in mammals, the steroid secretion is under the
control of pituitary and hypophysectomy invariably causes atrophy of the steroid
secreting tissues.

Thymus
Reptiles have wel1•developed thymus glands. Thymus glands, as you are aware, are
closely associated with immune functions of the body. In reptiles also, the introduction of
any foreign material into its body results in the proliferation of cells as well as
antibody production. Challenging of the animal by a second injection of the foreign
material heightens the immunological response suggesting that there is immunological
memory as well. Such responses have been shown to be partly dependent on thymus.
Thymectomy or the removal of the thymus gland from young reptiles reduces the
capacity for adaptive response.

12.7 BIRDS

In birds the pars dlstalis of the pituitary lies away from the brain. It shows two regions,
the cranial and caudal regions. The median eminence is a separate structure and is also
divided into two parts. The anterior part receives the fibres of the hypothalamo-
hypophyseal tract!and these fibres reach the pars nervosa to release the hormones arginine
vasotocin and mesotocin, a derivative of oxytocin. These hormones are involved in the
regulation of water balance, heart rate and reproduction. Mesotocin is also involved in
enhancing fatty acids and sugar in the blood. The distal lobe of the pituitary has been
shown to contain the following hormones: Growth hormone or somatotropin, prolactin,
thyrotropin, follicle stimulating hormone, lutinizing hormone, adrenocorticotropic
hormone and melanotropin. Pars intermedia in other vertebrates produces the last two
hormones. Whereas the function of melanotropin is not clear, the other hormones have
the same actions as in other vertebrates. Prolactin influences the production of brood
pouches and broodiness, and in pigeons stimulates the production of milk in both sexes.

Adrenals

132 The cortical and medullary cells of adrenals may be found separately or intermingled
with one another. Corticosterone is the main cortical hormone in birds and aldosterone
may be secreted in small amounts. Marine birds have large adrenal glsnds.
Thyroids and Parathyroids Endocrine System

The paired thyroid glands produce Tt and Tt hormones. The turnover rate of both the
hormones is much higher than in mammals. This could be attributed to higher body
temperature and metabolism. Thyroids also control growth and moulting and
possibly have a role in migration and reproduction. Birds have large parathyroids, which
develop from fifth and sixth branchial pouches and generally remain separated from
thyroid. The parathormone of birds is a potent hormone and probably regulates the
mobilisation of calcium from bones to the eggshells at the time of egg laying.

Pancreas
Bird pancreas produces both insulin and glucagon as in mammals; and ten times less
insulin is produced than glucagon, which mobilises sugar from the liver and raises its
level in blood. Glucagon also promotes a strong lipolytic action. Essentially hormonal
activities in birds commensurate with their high metabolic demands.

SAQ 4
Match the endocrine gland with the hormone it secretes.
A B
a) Pars distalis of pituitary I) Triiodothyronine
b) Parathyroid 2) nor-epinephrine
c) Pancreas 3) anti-diuretic hormone
d) Adrenal medulla 4) calcitonin
e) Neurohypophysis 5) glucagon
f) Thyroid gland 6) somatotropin

12.8 SUMMARY
After a study of this unit you have
learnt:

• About the endocrine glands of different groups of vertebrates. We began the study mesot
with the description of the endocrine glands of mammals. In mammals the unpaired ocin,
pituitary gland consists of two major parts, the adenohypophysis and the the
neurohypophysis. The adenohypophysis shows further divisions, the pars intermedia pars
and the pars distalis. 'Fhe neurohypophysis is differentiated into pars nervosa, the distali
infundibular stem and the median eminence. Pars distalis secretes a number of tropic s
hormones that regulate the secretion of various endocrine glands in the body. Pars secret
es a
. intermedia secretes melanotropin and the neurohypophysis releases two numb
neuropeptides, oxytocin and vasopressin. er of
That agnathans lack a hypothalamo-hypophyseal portal system and pituitary appears tropic
to be independent of hypothalamus. Pars intennedia secrete melanophore-stimulating horm
hormone and the pituitary appears to secrete a gonadotropin as well. In ones
elasmobranch fishes the anterior pituitary secretes thyrotropin and a gonadotropin. inclu
Lactotropic hormone and ACTH are the other tropic hormones secreted by the ding
pituitary.
Neuropeptide arginine vasotocin is secreted by the neurohypophysis. Pituitary of
bony fishes has adenohyphysis and neurohypophysis, and the adenohypophysis
shows three regions, pars rostral distalis, pars proximal distalis and pars intermedia.
Lactotropin, corticotropin and thyrotropin are the three tropic hormones secreted
from rostral pars distalis. Lutinising hormone is the only gonadotropin that is -
produced. Arginine vasotocin and isotocin are the two neuropeptides secreted from
the neurohypophysis and the function of these hormones is not well understood.
Amphibian pituitary shows three main parts of the pituitary, pars distalis, pars
intermedia and pars nervosa. Amphibian pituitary secretes all the tropic hormones
produced by a mammalian pituitary including the two gonadotropins. The secretion
of thyrotropin and ACTH are regulated by the respective regulating factors produced
by the respective releasing factors released from the hypothalamus. The neural lobe
of the pituitary produces vasotocin and mesotocin, the two hormones that regulate
water metabolism. The reptilian pituitary is more or less similar in structure and
functioning to that of amphibians. In birds the pituitary shows cranial and caudal
regions. While the pars nervosa releases neuropeptides such as vasotocin and
133
 1
£’'unctionpl Anatomy of
melanotropin and ACTH, two hormones, which are, produced by pars intermedia in
other vertebrates.
That thyroid gland, an endoderm derivative, synthesises and releases a tyrosine
derivative, thyroxine. Thyroxine regulates growth and morphogenesis as well as
accelerates the oxidative metabolism. Thyroid does not appear to be present in
Agnatha , and both in elasmobranchs and bony fishes although it is present, its
functions are not clearly known. In amphibians thyroid gland regulates
metamorphosis. In reptiles, it appears higher temperature favours thyroid activity.
And in birds thyroid produces both T, and T‹ hormones. The turnover rate of both the
hormones is higher in birds than in mammals.
That parathyroids, which are found in close proximity to the thyroids, produce
two hormones that function antagonistically, the parathormone and calcitonin. The
two hormones regulate calcium and phosphate metabolism. Parathyroids are absent
in agnathans. In fishes also there are no parthyroids, but structures known as
ultimobranchial glands derived from the fioor of the last branchial pouch secrete
calcitonin that ›regulates calcium metabolism. ln amphibians, parathyroids derived
from pharyngeal pouches appear to regulate calcium levels of blood. Parathyroids of
reptiles are structurally very similar to those of mammals and their removal causes
lowering of bleod calcium levels in them. Further parathyroidectomy causes
hyperexcitability and tetanic conditions; administering extracts of parthyroids can
restore normalcy. Parathormone of the birds besides regulating blood calcium levels
also mobilises calcium for deposition into eggshells.
That pancreas have both exocrine and endocrine tissue the exocrine tissue produces
digestive enzymes while the endocrine tissue or islets of Langerhans produce
glucagons (by n cells) and insulin (by § cells). Pancreas are absent in jawless fishes
pancreas of bony fishes and elasmobranchs secrete both glucagons and insulin
though no islets cells are present. Pancreas of amphibians reptiles and birds are
similar to mammals in action.
That adrenal glands are present in all groups of vertebrates. The two regions of which
the gland is composed of, the cortex and medulla, are spatially separated in Agnatha
and lower fishes; but in higher forms the cells of the two regions are found
interspersed. The cortex, a mesodermal derivative, secretes steroidal hormones —
glucocorticostaroids and minealocorticoids. Glucocorticoids are concerned with
regulation of carbohydrate metabolism and mineralocorticoids are concerned with
the regulation ef sodium and potassium metabolism. Adrenal medullary region
secretes two catecholamines, the adrenaline and noradrenaline. Adrenaline acts as a
vasodilator in small doses and as a vasoconstrictor in large doses. Both the hormones
cause an increase in heart rate, blood volume and concentration in erythrocytes.
Adrenaline alse regulates carbohydrate metabolism. In Agnatha although cortical
cells could be located in the region of pronephros. no steroid hormone has been
identified. Medullary tissue represented in the form of chromaffin cells are found
in the walls of heart and large blood vessels. Catecholamines have been shown to
be present in the tlssues of heart. In fishes interrenal bodies located in the kidney
region represents the cortical tissue. They may be either paired or unpaired. The
steroid secreted by the› cortical cells, identified as 1-alpha-liydroxycorticosterone
regulates carbohydrate metabolism but Steroids that regulate mineral metabolism
have not been identified. In amphibians the cortical and medullary tissues are found
to be closely associated as in other higher vertebrates. Although the details of the
hormones secreted by the amphibian adrenals acre not known, the glands are known
to regulate both carbohydrate and mineral metabolism. Adrenals in reptiles is a
mixture of
steroid secreting and catecholamine secretirig tissues. The steroid secreting tissue
appears to be under the control of a pituitary hormone. In birds the cortical and
medullary tissues may be found separately or together. Both corticosterone and
aldosterone have been identified.
That ovary andf testis are the two endocrine centres that secrete the hormones that
regulate the reproductive activity of the vertebrates. Leydig cells of testis secrete
male hormones, testosterone and androsterone. The synthesis and release of these
androgens are under the control of the gonadotropic hormone, the interstitial cell
stimulating hormone from pituitary. Similarly the ovary synthesises and releases two
steroids estrogens and progesterone, besides a peptide hormone relax in. Estrogens
maintain the functional state of the reproductive system, besides regulating the
development and functions of accessory reproductive structures and secondary
134 sexual organs. Progesterone secreted under the influence of luteotropic hormone
 from pituitary is released from the ruptured follicle of the ovary that transforms into Endocrine System
corpus luteum. Progesterone besides preparing the uterine wall for the implantation
of the ovum, also maintains gestation.

12.9 TERMINAL QUESTIONS

1. Describe the structure and functions of pituitary gland of mammals.
. .... . .. .. . .. . . . ..... ... .... ...................... .. . ..........
... . . .................................................................. .....................
..... ... .. ...... . . .. .... . ..... ................................. .......................
2. Give a comparative account of thyroids of vertebrates.
.... ............. .............. .... . . .. . ..... . .. .. ............................ ......
. .. .... .... . . .. ........ . .... ............ ......... .......................................
. . .. . . ...... .. ............ ....... ................................... . . .. .................
3. Briefly write the functions of the following hormones.
(a) Adrenocorticotropic hormone (b) parathormone (c) aldosterone
(d) Testosterone (e) progesterone.
... . . . .... . . . . .......................... ............................................
. ... .. ...... . . .. . ... ............. .........................................................
. . ... .... .. ..... . ... . ....................... . ........... . ...............
. . . . . . ....... . . . . ..... . .. . ... . . . ... ......... ..
.... ...... .. . .. .. . . . . . .. . . . .............. . ...... ... ..... . . ..
. . . .... ... .. .. ................ .. . . ...... .. . .. ..................................
4. Byriefl y describe the structure and functions off adrenal cortex and medul Ia.

........................................ ............................................................
............ ... ........................... .. ............... . .................. ... .........

12.10 ANSWERS
Self-assessment Questions
1. 1. (a) Hypoph ysis cerebri, (b) adenohypophysis and neurohypophysis, (c)
pars distal is and pars intermedia, (d) pars nervosa, infundibular stem and
median eminence, (e) oxytocin and vasopressin
II. Refer to Table 12.1.
2. a) F, b) T, c) F, d) T, e) F, f) F, g) F, h)T, i) F, j) F, k) F
3. a) absent
b) feed back from other tissues
c) produces
d) do not have
e) ventral
f) medulla
g) 1-alpha-hydroxycorticosterone
h) pars intermedia
i) scattered masses of tissue along ventral aorta
j) loss
4. a) 6, b) 4, c) 5, d) 2, e) 3, f) 1

Terminal Questions
1. Refer to section12.2.1
2. Refer to sections 12.2.2, 12.4, 12.5, I 2.6 & 12.7
3. Refer to section12.2.1, 12.2.3, 12.2.5 and 12.2.7
4. Refer to section 12.2.5.
135
Functional Anatomy of
Chordates - II
GLOSSARY
Acinar: Having rounded compartments.
Acuity: Sharpness or resolution in sensory perception especially vision and hearing.
Adrenaline: Epinephrine.
Adrenal gland: An endrocrine gland next to kidney; suprarenal gland.
Afferent artery: An artery that carries blood towards an organ.
Aldosterone: An adrenal corticosteroid that regulates sodium, potassium and water
balance by the kidneys.
A macrine cells: Special neurons of the retina that interact with the transmission of
impulses between photoreceptor cells and bipolar cells or between bipolar cells and
•panglion cells.
Ampulla: Any membranous enlarged vesicle such as the ones on semicircular ducts of
the inner ear.
Androgen: Any hermcine that possesses masculinising activities, such as testosterone or
any of the other male hormones.
Antonomic: Self controlling, independent of outside influence.
A rchinephric due.t: The collecting duct of the archinephros or primitive kidney or
pronephros.
Arcualium: An embryonic, cartilaginous analogue parts of the adult vertebra.
Areola: The area around the mammary nipple that can be distinguished, often by colour,
from the nipple.
Auditory: Pertaining to the perception of sound.
Autostyle: Jaw suspension in which the jaws articulate directly with the braincase.
Axon: The process of a neuron that transports the nerve impulse away from the cel I body.
Basilar membrane: A basal membrane of the inner ear of mammals that supports the
organ of Corti and subdivides the cochlear channel into two subdivision, the scale media
and scale tympani.
Bipedal: Walking or running by means of only two hind legs.
Bipolar cell: A type of neuron that has processes at two ends.
Braincase: The part of the skull containing the cranial cavities and housing the brain.
Blind spot: The nonsensory area of the retina through which the axons of ganglion cell
coalesce and turn inward to form the optic nerve.
Centrum: The body or base of a vertebra.
Cephalic: Pertaining to head.
Cerebellum: The part of vertebrate hindbrain that coordinates body posture and
equilibrium.
Cerebrum: A major part of vertebrate forebrain; the two cerebral hemispheres united by
corpus callosum form the largest part of vertebrate brain in mammals and integrate most
of the sensory impulses and motor responses.
Chondrocranium: The part of the skull formed by endochrodal bone or cartilage that
underlies and supports the brain; also includes the fused or associated nasal capsules.
Choroid: Relating to a coat or membranes.
Choroid plexus: Any of the several convoluted vascular structures that projects into
lateral, third, and fourth ventricles of the brain, and that produce cerebrospinal fluid.
Chromaffin cells: Any cells that stain with chrominum salts, especially cells of
suprarenal glands adrenal medulla, paraganglion and carotid bodies. Chromaffin cells are
derived from neural crest.
136
 Cocfilea: The elongated and often coiled portion or inner ear of‘crocodiles, birds and Endocrine System
mammals containing sound receptive cells.
Coelom: A body cavity that develops within mesoderm as it migrates between endoderm
and ectoderin.
Commissure: Bond of tissue connecting corresponding halves of brain or spinal cord.
Cone cells: Photoreceptor cells in the vertebrate retina that are differentially sensitive to
light of different wavelengths.
Cornea: Transparent layer of skin that covers the eye in terrestrial animals; with the
crystalline lens forms the complex lens of the eye.
Corpus callosum: A broad band of nerve fibres that links right and left cerebral
hemispheres.
Corpus luteum: A yellow glandular mass in the vertebrate ovary formed by the cells of
an ovarian follicle that has matured and discharged its ovum; secretes progesterone.
Cortex: The outer part of a structure or organ that overlies (surrounds the medulla).
Crystalline lens: The lens of the eye, so named to distinguish it from the cornea, which
in tetrapods also functions as lens.

! Cupula: A gelatinous cap over a group of hair cells.

Decussation: A crossing of nerve fibres that carries impulses to or from one side of the
body from or to the opposite side of the brain.
Deferent duct: The part or the mesonephric tubule that carries sperm away from
epididynius; vas deferens.
Dendrite: Any fibre of the nerve cell that carries an impulse towards the cell body.
Dentition: A set of teeth.
Echolocation: The location of objects by detecting echoes reflected from them.
Efferent artery: An artery that carries blood away from a structure.
Efferent duct: A mesonephric duct that carries sperm away from the testis.
Endocrine: Secreting internally; applied to a ductless gland that secretes into blood or
lymph a substance that has a specific effect on another target organ or tissue.
Endoskeleton: The supportive or protective framework within the body that lies beneath
the integument.
Epididymis: The mostly convoluted tubule that drains the efferent ducts of the testis into
the urethra; the epididymis is derived from the mesonephric or Wolffian duct and efferent
ducts.
Epinephrine: Hormone secreted by adrenal medulla of mammals that mobilises glucose
in the tissues, among other functions; derived from the methylation of norepinephrine.
Estrogens: One of a group of female steroid sex hormones of which estradiol is the most
common, synthesised by placenta and ovary. Responsible for development of female
secondary sex characteristics and for maturation and functioning of female reproductive
organs.
Fenestra: An opening within the bony braincase.
Fossa: A pit or cavity.
Fovea: A small depression; fovea centralis.
Fovea centralis: Area of the vertebrate retina containing only cone cells, where the
most acute vision is achieved at high light intensities.
Free nerve endings: The terminal or sensory end of an afferent neuron that is not
ensheathed.
Funiculus (pleural funiculi): Any marginal part of the spinal cord that contains nerve
fibres that carry impulses up and down the cord. It is the white or myelinated column of
tetrapods.

137
Functional Anatomy of
Ganglion: A knott like mass of cell bodies of neurons located outside the nervous systein
in vertebrates; in invertebrates includes the swellings of the central nervous system.
Ganglion cell: A cell type that forms the outermost layer of retina and the optic nerve.
Genitalia: Sexual organs, especially copulatory organs, genitals.
Glans penis: Head or tip of penis.
Hemipenis (plural, hemipenes): One of the paired, grooved intromittent organs of
reptiles.
Hermaphrodite: An individual that has both male and female organs; monoecious;
bisexual.
Heterodont: Dentition in which the teeth are different in general appearance throughout
the mouth.
Homodont: Dentition in which the teeth are similar in general appearance throughout the
mouth.
Hormone: Substance produced in cells in one part of the body that diffuses or is
transported by blood stream to cells in other part of the body, where it regulates and
coordinates their activities.
Hypothalamus: A region of the forebrain, the floor of the diencephalon, that contains

” centres controlling viseral activities, water balance, temperature, sleep etc.

Hypothalamus Releasing Hormone: A hormone secreted in the anterior pituitary by
hypothalamic neurons that terminate there and that stimulates the release of other
hormones.
Incus: The middle ear bone of mammals derived phylogenetically from the quadrate.
1nterstitial tissue: Tissues that lies between other structures or tissues.
Intervertebral body: A pad of cartilage or fibrous connective tissue between articular
ends of successive vertebral centra.
Intervertebral disc: A pad of fibrocartilage in the adult mammal that has a gel like core
derived from the notochord and is located between articular ends of successive vertebral
centra.
Jaws: Skeletal elements of bone or cartilage that reinforce the lower borders of the
mouth.
Labia majora, Latiia majus (singular): Two lateral folds that surround the vulva.
La bia minora, Labium minus (singula r): Two small folds on the medial surface of the
labia majora that surround the vulva.
Lacrima I (Lachrymal): Of or pertaining to tears, tear glands, or ducts.
Lateral line: The line or lines along the body of fishes and other aquatic vertebrates that
contain mechanoredeptors that detect movement.
Lateral line organs: Any complex of mechanoreceptors in the lateral line; neuromest,
neuromost organ.
Leydig cells: Interstitial cells of the testis that produce testosterone.
Macroglia: The astrocyte component of neuroglia.
Malleus: One of tha three middle ear bones in mammals that is phylogenetically derived
from the articular bone of the lower jaw; hammer.
Meatus: Canal or channel.
Mechanoreceptors: A sensor that detects mechanical changes such as motion, pressure,
tension and compression.
Medulla: Central region of an organ or structure.
Melanin: The dark brown pigment produced by the pigment cells of many vertebrates.
Meninges: Sheets of tissue enclosing the central nervous system. In mammals these are
138 the dura mater, arachnoid and pia mater.
Microglia: Small neuroglial cells.
Myelin: The fatty substance that forms a sheath around axons of nerve cells in the white
matter of the central system and in most peripheral nerves.
Nephrotome: A narrowed mass of intermediate mesoderm (mesomere or nephromere)
that develops into kidney structures.
Nerve: A bundle of neuron axons connecting the central nervous system with peripheral
receptors and effectors.
Neural crest: Embryonic cells unique to vertebrates, associated with the neurectoderm
but subsequently widely migrating to participate in the formation of many tissues and
structures which are characteristic of the subphylum.
Neurocranium: That part of the brain case that contains cavities for the brain and
associated sensory capsules (nasal, optic, otic).
Neuroglia: Connecting and supporting cells in the central nervous system.
Neurosecretion: Production of hormone like substances by nerve cells.
Nidamental gland: Any of various maternal glands of the oviduct that secrete material
that covers an egg or cluster of eggs; shell gland.
Norepinephrine: A neurosecretion produced by axon terminals of post-ganglionic motor
neurons of sympathetic nervous system that mobalises glucose in tissues.
one:fication: "rhe process of calcium deposition in tissue.
Osteon: A highly ordered arrangement of bone cells into concentric rings with bone
matrix surrounding a central canal through which blood vessels and nerves run, the
Haversain system.
Oviduct: The tube that conveys egg celts to the exterior or to the uterus.
Parietal eye: A median eye with an aperture that is surrounded by the parietal bone. The
parietal eye forms the pineal gland.
Pars recta: The straight part of some structures, such as kidney tubules.
Pecten: A comb like process of the retina of reptile and birds.
Perichondrin: The sheet of fibrous connective tissue around cartilage.
Peritoneal canal: A canal off the abdominal coelom.
Peritoneal funnel: A funnel like opening from the coelom.
Peritoneum: A membrane that covers the abdominal viscera and inner abdominal wall.
Portal system: Portion of the venous system specialised for the transportation of
substances from the site of production to site of action. A portal system begins and ends
in capillary beds.
Ptaguim: A stretched fold of skin that forms an airfoil or flight control surface.
Quadrapedal: Walking or running by means of four legs.
Rete testis: The network of anastomosing tubules that carries sperm from testis.
Receptor: Sensory nerve ending in skin or sensory organ.
Rods: Photoreceptor cells in vertebrate retina specialised to fiinction effectively under
conditions of dim fight.
Sacculus: Small pouch; smaller of the two sacs in inner ear.
Sclera: Tough outer coat of eye.
Stapes: One of the three middle ear bones in mammals that is phylogenotically derived
from the columella (hyomandibula).
Suprarenal gland: Adrenal gland.
T : Triidothyronine.
T : Tetraiodothyronine.
139
Functional Anatomy of
Chordates - II Tapetum lucidum: (1) A reflective choroid in the eyes of nocturnal animals (2) A'
nervous pathway in the roof or the lateral ventricles of cerebrum.
Tympanum: Tire eardrum or tympanic membrane.
Utriculus: Larger of the two sacs in the inner ear.
Vertebra: One of the several bones or cartilage blocks firmly joined into a backbone
that defines the major body axis of vertebrates.
Zygapophysis: The projection of a neural arch that articulates with the adjacent neural
arch.

FURTHER READING
i. The Life of Vertebrates J.Z. Young (Third Edition) ELBS, Oxford University Press. •
2. The Vertebrate Body A.S. Romer and T.S. Parson (Sixth Edition) CBS
College Publishing.
3. Chordata Anatomy and Evoultion D. Jacob, A Sharma and K. Nandchahal (1994).
Ramesh Book Depot, Jaipur.

140