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Hormone, organic substance secreted by plants and animals that functions in the regulation of
physiological activities and in maintaining homeostasis. Hormones carry out their functions by
evoking responses from specific organs or tissues that are adapted to react to minute quantities of
them. The classical view of hormones is that they are transmitted to their targets in the
bloodstream after discharge from the glands that secrete them. This mode of discharge (directly
into the bloodstream) is called endocrine secretion. The meaning of the term hormone has been
extended beyond the original definition of a blood-borne secretion, however, to include similar
regulatory substances that are distributed by diffusion across cell membranes instead of by a
blood system.

General Features

Relationships between endocrine and neural regulation

Hormonal regulation is closely related to that exerted by the nervous system, and the two
processes have generally been distinguished by the rate at which each causes effects, the duration
of these effects, and their extent; i.e., the effects of endocrine regulation may be slow to develop
but prolonged in influence and widely distributed through the body, whereas nervous regulation
is typically concerned with quick responses that are of brief duration and localized in their
effects. Advances in knowledge, however, have modified these distinctions.
The Hormones Of Vertebrates

Hormones of the pituitary gland

The pituitary gland, or hypophysis, which dominates the vertebrate endocrine system, is formed
of two distinct components. One is the neurohypophysis, which forms as a downgrowth of the
floor of the brain. The other is the adenohypophysis and usually includes two glandular portions.
Functional analysis of these hormones also is difficult, for the targets of certain hormones of the
adenohypophysis, called tropic, or trophic, hormones, are other endocrine glands. The action of
such tropic hormones can be understood only in the light of the mode of function of the
endocrine glands they regulate.

Elements in a generalized mammalian pituitary gland.


Growth hormone (somatotropin)

The physical and chemical properties of growth hormone, which differ from species to species,
are associated with marked differences in biological activity.

Growth is such a complex process that definition of the growth hormone’s mode of action is
difficult. One of its known effects is an increase in the rate of protein synthesis, which is to be
expected, since growth involves the deposition of new protein material. In addition, growth
hormone affects the metabolism of certain ions (including sodium, potassium, and calcium),
promotes the release of fats from fat stores, and influences carbohydrate metabolism in ways that
tend to cause an increase in the level of glucose in the bloodstream.

Excess secretion of growth hormone does, however, have damaging effects in humans, for it
produces overgrowth of the skeleton. If this occurs in youth, before the closure of
the epiphyses (ends) of the long bones, it results in gigantism. If it occurs afterward, it
causes acromegaly, in which the disturbance is more serious, with enlargement of the bones and
soft tissues and consequent distortion of the skull.


Prolactin is a protein hormone that in female mammals initiates and maintains the secretion of
milk, the mammary glands having been previously prepared for this function by the action of
other hormones

Adrenocorticotropic hormone

Adrenocorticotropic hormone (ACTH; also called corticotropin) is present in all vertebrates. It

regulates the activity of part of the outer region (cortex) of the adrenal glands. In mammals its
action on the adrenal cortex is limited to areas in which important steroid
hormones (e.g., glucocorticoids, such as cortisol and corticosterone) are formed.

ACTH does not affect the synthesis of the mineralocorticoid hormone aldosterone, which takes
place chiefly in the outer cortical region. The relationship between ACTH and the adrenal cortex
is an example of the negative feedback characteristic of endocrine systems; i.e., a decrease in the
level of glucocorticoids circulating in the bloodstream evokes an increase in the secretion of
ACTH, which, by stimulating the secretory activity of its target gland (the adrenal cortex), tends
to restore to normal the level of glucocorticoids in the bloodstream.
Thyrotropin (thyroid-stimulating hormone)

Thyrotropin (also called thyroid-stimulating hormone, or TSH) regulates the thyroid gland
through a feedback relationship similar to that for ACTH; thyrotropin increases the secretion of
the hormones from the thyroid gland and, if its action is prolonged, evokes increases in cell
number (hyperplasia) and in size of the gland. One consequence of an overactive thyroid in
humans is a bulging of the eyes (exophthalmos).

Follicle-stimulating hormone

Follicle-stimulating hormone (FSH) is a type of gonadotropin; it is concerned with the regulation

of the activity of the gonads, or sex organs, which are endocrine glands as well as the sources
of eggs and sperm. FSH stimulates development of the graafian follicle, a small vesicle
containing an egg, in the ovary of the female mammal. In the male it promotes the development
of the tubules of the testes and the differentiation of sperm.

Luteinizing hormone (interstitial-cell-stimulating hormone)

Luteinizing hormone (LH; also called interstitial-cell-stimulating hormone, or ICSH) is another

gonadotropin. In the female mammal it promotes the transformation. In the male LH promotes
the development of the interstitial tissue of the testes and hence promotes secretion of the male
sex hormone, testosterone. It may be associated with FSH in this function.

An unexpected property of mammalian FSH and LH is that both have a thyrotropic action (i.e.,
stimulate secretion of thyroid hormones) in lower vertebrates. This so-called heterothyrotropic
effect has led to the supposition that FSH, LH, and thyrotropin may have evolved by
modification of a common ancestral glycoprotein molecule, resulting in an overlap of properties.

Melanocyte-stimulating hormone (intermedin)

Melanocyte-stimulating hormone (MSH) regulates colour changes in animals by promoting the

concentration of pigment granules in pigment-containing cells(melanocytes and chromatophores)
in the skin.
The pituitary gland secretes multiple hormones, including melanocyte-stimulating hormone
(MSH, or intermedin), adrenocorticotropic hormone (ACTH), and thyrotropin (thyroid-
stimulating hormone, or TSH).

Hormones of the thyroid gland


The two thyroid hormones, thyroxine and triiodothyronine, are formed by the addition
of iodine to thyroglobulin. Thyroglobulin is stored within the gland in follicles as the main
component of a substance called the thyroid colloid. This arrangement, which provides a reserve
of thyroid hormones, perhaps reflects the frequent scarcity of environmental iodine, particularly
on land and in fresh water. Iodine is most abundant in the sea.

Synthesis of the thyroid hormones is regulated by the level of circulating hormones (i.e., a
negative feedback mechanism) operating, as indicated earlier, partly by direct action on the
thyrotropin-secreting cells of the pituitary gland and partly by indirect action on the
hypothalamus and its thyrotropin-releasing hormone

One established effect of the thyroid hormones in mammals is an increase in metabolic rate and
in oxygen consumption, but the effects of the hormones undoubtedly are more wide-ranging than
this. On the one hand, impairment of the thyroid function in mammals results in disturbances in
the processes of growth and maturation. Both growth and maturation disturbances occur in the
cretinous dwarfism resulting from thyroid deficiency in newborn infants.


The discovery of calcitonin (thyrocalcitonin) in 1961 demonstrated the importance of

comparative studies in endocrinology. It originally had been thought that this hormone, which is
present in preparations made from mammalian thyroid glands, was secreted by the parathyroid
glands, which in some species are combined with the thyroid gland. Later, the hormone was
concluded to be a secretion of the thyroid gland itself. Calcitonin is secreted by the called light,
C, or parafollicular cells, which are found in the thyroid gland of mammals.

The C cells, or parafollicular cells, of the thyroid gland (indicated by the arrow marked “P”)
produce a hormone called calcitonin, which regulates serum calcium
Calcitonin lowers the level of calcium in the blood (hypocalcemic action) when it rises above the
normal level. Its secretion probably is regulated by a negative feedback relationship between the
gland and the blood plasma. The hormone affects bone, which is an active tissue. It undergoes
not only growth but also remodeling as it adapts to the changing patterns of stress to which it is
subjected; its calcium exchanges continuously with that of the plasma. The effect of calcitonin is
to decrease the mobilization (resorption) of calcium from the skeleton into the blood plasma. In
this respect, it is opposite in direction to the effect of parathormone of the parathyroid glands.

Parathormone of the parathyroid gland

The parathyroid glands secrete a hormone called parathormone (PTH), which regulates calcium
metabolism in conjunction with calcitonin; its evolution in terrestrial vertebrates may have been
an adaptation to the increased demand for continuous skeletal adjustments imposed by the
evolution of terrestrial locomotion. Skeletal adjustments must be made without disturbing the
delicate calcium balance of the rest of the body, for calcium is involved in maintaining the
transport of substances through cell membranes; hence, it has an important role
in muscle contractility, excitability of motor end plates in the nervous system, and coagulation of

Removal of the parathyroid glands in mammals causes a fall in the level of calcium in the blood
plasma, which, if sufficiently severe, is accompanied by convulsions and other symptoms
resulting from increased excitability of the motor nerves.

In general, therefore, the action of parathormone is opposite in direction to that of calcitonin.

Parathormone keeps the level of blood calcium up to its normal value; on the other hand,
calcitonin ensures, through its hypocalcemic action, that the level does not rise far above this
critical point. The combined actions of the two hormones serve to illustrate the importance of
endocrine regulation in homeostasis. Vitamin D is a third factor in calcium regulation; its
absence in young children results in skeletal malformations (rickets). Parathormone is unable to
regulate the absorption and mobilization of calcium in the absence of vitamin D, which is also
associated with the hormone in promoting mobilization of magnesium from bone and perhaps in
the movement of phosphate within the kidney tubule.

Hormones of the pancreas


The vertebrate pancreas contains, in addition to the zymogen cells that secrete digestive
enzymes, groups of endocrine cells called the islets of Langerhans. Certain of these cells (the B,
or beta, cells) secrete the hormone insulin, inadequate production of which is responsible for the
condition called diabetes
Injection of insulin lowers blood sugar (glucose) levels, but this so-called hypoglycemic effect is
only one expression of the wide-ranging influence of insulin on storage and mobilization of
energy, in which the target tissues of primary importance are muscle, adipose (fat) tissue,
and liver. The actions of insulin on these tissues are varied. First, it promotes the use of the sugar
glucose as an energy source; at the same time, it encourages the storage of excess carbohydrate
as glycogen, the storage carbohydrate of animals. Second, insulin reduces the use of fat as an
energy source and promotes its storage. Third, it reduces the use of protein as an energy source
and promotes the formation of proteins from amino acids.


It strongly opposes the action of insulin, primarily through a hyperglycemic (blood glucose-
raising) effect that results from its promotion of the breakdown of glycogen (glycogenolysis) in
the liver, a process that results in the formation of glucose.

Hormones of the adrenal glands

The adrenal gland of mammals is composed of an outer region, the cortex, which consists of
adrenocortical tissue that secretes steroid hormones (steroids are fat-soluble organic compounds),
and an inner region, the medulla, which is composed of chromaffin tissue, so called because its
cells contain granules that can be characteristically coloured by certain reagents.

Chromaffin tissue secretes two hormones, epinephrine and norepinephrine, which are members
of a class of compounds called catecholamines.

Human adrenal gland.

Epinephrine and norepinephrine evoke diverse and widespread responses but differ from each
other in certain of their effects. Both influence the heart and blood vessels in ways which,
although opposed to each other in a few respects, generally result in an increase in blood
pressure and in output of blood from the heart. Both hormones also have metabolic actions.
Epinephrine, for example, like glucagon, stimulates glycogenolysis (breakdown of glycogen to
glucose) in the liver, which results in the raising of the level of blood sugar; in addition, it
increases oxygen consumption and the output of blood from the heart, probably contributing
thereby to the regulation of body temperature in mammals. Epinephrine has effects on the
nervous system, which are recognizable subjectively in humans by feelings of anxiety and of
increased mental alertness.

Effects of adrenaline and noradrenaline in humans

Source: G.H. Bell, J.N. Davidson, and H. Scarborough, Textbook of Physiology and Biochemistry, 7th ed., 1968, used
by permission of Williams and Wilkins.

adrenaline noradrenaline

heart rate increase decrease

cardiac output increase variable

total peripheral resistance decrease increase

blood pressure rise greater rise

respiration stimulates stimulates

skin vessels constriction less constriction

muscle vessels dilation constriction

bronchus dilation less dilation

eosinophil count increase no effect

metabolism increase slight increase

oxygen consumption increase no effect

blood sugar increase slight increase

central nervous system anxiety no effect

uterus in late pregnancy inhibits stimulates

kidney vasoconstriction vasoconstriction

Adrenocortical tissue of the cortex

Many steroids have been isolated from the adrenal cortex, but in most vertebrate groups only
three of them are active as hormones—cortisol (hydrocortisone; compound F), corticosterone
(compound B), and aldosterone.

In contrast to the chromaffin tissue of the adrenal medulla, the adrenocortical tissue is essential
for life. Two primary functions of the corticosteroids are distinguishable in mammals. One,
which contributes to the regulation of carbohydrate metabolism, is an action of cortisol and
corticosterone, which are therefore called glucocorticoids. These hormones promote
gluconeogenesis in the liver and are thus important in maintaining normal blood sugar levels,
particularly during glucose shortage; lack of them results in low levels of blood sugar and an
increase in the sensitivity of the liver to insulin (whose effect there is to decrease
gluconeogenesis). In addition, lack of the glucocorticoids is associated with a decrease in the
entry of amino acids into muscles and an increase in their uptake by the liver, where enzymes
required to convert amino acids to glucose must be synthesized.

In contrast to glucocorticoid action is the so-called mineralocorticoid action of aldosterone,

which is manifested in mammals in the regulation of sodium metabolism. In the absence of
aldosterone, sodium is lost from the body by excretion in urine; secondary consequences include
a decrease in blood volume and in the filtration rate of substances through kidney structures
called glomeruli. Cortisol and corticosterone also play a minor part in mineral regulation, so that
slight overlap in function occurs between the two corticoid types.

Other actions of the corticoids are apparent in patients suffering from Addison disease, which is
caused by a general deficiency in corticoid production. A deficiency of corticoids causes
disturbances in urinary output and fat metabolism, diminished resistance to stress, muscular
weakness, and nervous disturbances manifested by depression and a general lack of mental

Hormones of the reproductive system

Female hormones


Two types of gonadal hormones, estrogens and progestins, are secreted in the female mammal.
Estrogens are substances that evoke the cyclical onset of heat, or estrus, during which the animal
is sexually active and receptive to the male. Estrus in this sense is not found in human females,
but estrogens contribute to the events of the menstrual cycle, bringing about cyclical changes in
the reproductive system that are comparable with those accompanying estrus in other mammals.

Hormones are secreted from the mammalian ovary by the ovarian follicle.

Progestins, of which the most important is progesterone, are concerned with the maintenance
of pregnancy.

Some progesterone is probably formed in the ovarian follicle, but the main site of production is
the corpus luteum, which is formed by a transformation of the follicle after ovulation.

The transformation of the follicle into the corpus luteum is an important turning point in the
diphasic menstrual cycle of women and in the ovarian cycles of other mammals, from which the
human cycle evolved. Progesterone prepares the uterus for the implantation of fertilized eggs,
and it is also needed for the maintenance of pregnancy once implantation has taken place

Male hormones

The sex hormones of the male follow a much simpler pattern than do those of the female. The
latter organs, the testes, secrete steriods called androgens, which are responsible for the
maintenance of male characteristics and behaviour.

Under the influence of LH (see pituitary gland), the interstitial tissue secretes the steroid
hormone testosterone, which is the most important vertebrate androgen.

In addition to promoting male characteristics, male behaviour, and the maintenance of the
spermatic tubules, testosterone, in the presence of normal amounts of growth hormone, also
promotes growth of the bony skeleton. The reason for rapid growth at puberty is that the
secretion of androgen markedly increases. The hormone brings about the closure of
the epiphyses (ends) of the long bones, which completes the process of growth (estrogens have a
similar action in the female).