Chapter II

THE PHYSIOLOGY OF FARM ANIMALS

1. Introduction

Physiology is defined as the study of function of the parts or organ

systems of the body; a study of function of living matter. It attempts to explain the
physical and chemical factors that are responsible for the origin, development and
progression of life. The study of physiology will provide knowledge on the structure
and function of the body and consequently, the care of the body.

Homeostasis is a key word in modern physiology. It is used to mean the

maintenance of static or constant conditions in the internal environment. For
example, the temperature of our body remains relatively constant at 37oC. Essentially
all the organs and tissues of the body perform functions that help maintain this
constant condition. The animal is kept alive by the coordination of the functions of
the different organ systems. For example, the lung or the respiratory system provides
oxygen required for the metabolic activities of the cells; the guts or digestive system
provides the nutrients; the heart or circulatory system circulates the blood that carries
oxygen, nutrients, hormones, and other metabolites required by the cells as well as
metabolic waste products produced by the cells for proper disposal by the excretory
organs. The nervous and endocrine systems perform the control, coordination and
integration of the functions of other organ systems in ways that make them work
together like parts of one machine to accomplish homeostasis.

2. The Nervous System

In general, the nervous system controls the rapid activities of the body such
as muscular contraction, secretions of some endocrine glands, heart rate, respiration
rate, gastro-intestinal motility, to mention a few. Your rapid reflex action to avoid
danger is due to nervous system activities.

Basic unit of the nervous system

The nervous system is composed of the Brain, the Spinal Cord and the
Nerves. Essentially all parts of the body are supplied with nerves. The nerve cells or
neurons specialize in impulse conduction or the relay of messages from effector
organs to the nervous system and vice versa. The human brain contains about 100
billion neurons or about the same number of stars in our galaxy. Neurons may be
classified according to the direction of impulse conduction as follows: (1) Afferent
(sensory) neurons – transmit nerve impulses from effector organ to the spinal cord or
brain; (2) Efferent (motor) neurons – transmit nerve impulses away from the brain or
spinal cord to or towards muscles or glands (effector organs); and (3) Interneurons –
conduct impulses from an afferent neuron within the central nervous system (CNS)

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which is made up of the brain and the spinal cord. The effector organ could either be
the skeletal muscle, cardiac muscle, smooth muscle or some other glands.

Structure of Neurons

All neurons consists of a cell body, one axon and one or more dendrites.
Axon and dendrites are threadlike extensions from the cell body and are often called
nerve fibers. The distal end of dendrites of sensory neurons are called receptors
because they receive the stimuli that initiate the conduction of impulses to the cell
body of the neuron. The axon is a single process that extends out from the cell body,
and may end up on a synapse or on any effector organ.

Neurons or nerve cells do not come in direct contact with one another;
instead, there is a small gap of about 200 A (Angstrom = 1/19 6 inch) between them.
This gap is called a synapse where nerve impulses are transmitted from one nerve cell
to another. Thus, synapses are located between the axon terminals of one neuron
(presynaptic or preganglionic cell) and the cell body or dendrites of another neuron
(postsynaptic or postganglionic cell). The transmission of nerve impulse across the
synapse involves the release from presynaptic neuron of a chemical mediator or
neurotransmitter (mostly acetylcholine) which crosses the synaptic cleft and brings
about a generation of signal or initiation of impulse in the postsynaptic neuron.

The Central Nervous System (CNS)

The brain and the spinal column jointly control the activities of the skeletal
or involuntary smooth muscles of the body in response to stimuli in their
environment. Information about the internal and external environment reaches the
CNS via a variety of sensory receptors. These receptors are transducers that convert
various forms of energy in the environment into action potential or nerve impulse in
the neurons.

The sensory receptor could be a part of a neuron or a specialized cell that

generates action potential in neurons. The receptor is often associated with non-
neural cells that surround it, forming a sense organ. The forms of energy converted
by the receptors include, for example, mechanical (touch-pressure), thermal (degrees
of warmth), electromagnetic (light), and chemical energy (odor, taste and 02 content
of blood). The receptors in each of the sense organs are adapted to respond to one
particular form of energy at a much lower threshold than other receptors respond to
this form of energy. The particular form of energy to which a receptor is most
sensitive or which is able to evoke an action potential is called adequate stimulus.

The Sensory Modalities

This consists of the various sense organs of the body. This includes the
senses of smell, vision, hearing, rotational and linear acceleration, taste, and
cutaneous senses with receptors in the skin to monitor touch-pressure cold, warmth

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and pain. There are, in addition, a large number of sensory receptors which relay
information that does not reach consciousness. Table 6 lists the principal sensory
modalities.

The rods and cones of the eyes for example, respond maximally to light of
different wavelengths, and there are different cones for each of the primary colors.
There are 4 different modalities of tastes – sweet, salt, sour and bitter – and each is
perceived by a more or less distinct type of taste bud in the tongue. Sounds of
different pitches are heard primarily because different groups of hair cells in the organ
of Corti are activated maximally by sound waves of different frequencies.

The sensation evoked by impulses generated by a specific receptor is

interpreted by a specific part of the brain which it ultimately activates.

The Somatic Nervous System

The striated or skeletal muscles of the body are innervated by the somatic
division of the nervous system. This part of the nervous system brings about quickly
adjustments of the muscles to changes in the environment. When we burn our finger,
receptors in the skin transform this stimulus into nerve impulses, which are carried by
different nerve fibers to the spinal cord and the higher nerve centers, which in tern
send nerve impulses by way of efferent fibers to the muscles of the hand which cause
the finger to be removed from the source of the heat. This is a form of reflex arc.
Adjustments of this type can be made with remarkable speed. Some nerve impulses
of this type travel at a rate of about 40 meters per second.

Autonomic Nervous System

The autonomic nervous system, like the somatic has afferent components,
central integrating stations, and effector pathways. The glands and visceral
musculature of the body receive efferent fibers from the autonomic nervous system.
The adjustments in the gland and visceral musculature are made by means of
chemical mediators, acetylocholine and epinephrine and norepinephrine released by
the terminal neurons of the autonomic fibers.

There are two major divisions of the autonomic system, the sympathetic and
parasympathetic. The cell bodies of the preganglionic neurons of the sympathetic
division are contained in the gray matter of the thoracic and lumbar regions of the
spinal cord, while the cell bodies of the preganglionic neurons of the
parasymphathetic system and situated within the midbrain, hindbrain and in the sacral
region of the spinal cord. The preganglionic neurons extend from the gray matter of
the CNS to one of the ganglia where they make synaptic junctions with post-
ganglionic neurons which extend from the ganglia to the effector organs.

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Table 6. Principal sensory modalities

===============================================================
Sensory Modality Receptor Sense Organ
___________________________________________________________________________________________________________

Conscious Sensations:

Vision Rods and cones Eye

Hearing Hair cells Ear (Organ of Corti)
Smell Olfactory neurons Olfactory mucous membrane
Taste Taste receptor cell Taste bud (tongue)
Rotational Acceleration Hair cells Ear (semicircular canals)
Linear acceleration Hair cells Ear (utricle and saccule)
Touch-pressure Nerve endings Various
Warmth Nerve endings Various
Cold Nerve endings Various
Pain Naked nerve endings Various
Joint position Nerve endings Various

Unconscious Sensations:

Muscle length Nerve endings Muscle spindle

Muscle tension Nerve endings Golgi tendon organ
Arterial blood pressure Nerve endings Stretch receptors in carotid
sinus and aortic arch
Central venous pressure Nerve endings Stretch receptors in walls of
Great vains, atria
Inflation of lung Nerve endings Stretch receptors in lung
Parenchyma
Temperature of blood in head Neurons in hypothalamus

Arterial PO2 Nerve endings (chemo- Carotid and aortic bodies

receptors)
pH of CSF Receptors on ventral surface
of medulla oblongata
Osmotic pressure Cells in OVLT and possibly
other circum-ventricular
organs in anterior hypothalamus
Arteriovenous blood
glucose difference Cells in hypothalamus (glucostats)
===============================================================

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 The preganglionic neurons of both the sympathetic and parasympathetic
division produce acetylcholine as do the postganglionic neurons of the
parasympathetic division produce mainly norepinephrine, and this compound is
generally considered as the adrenergic neurohumor, although small amounts of
epinephrine may also be produced by certain neurons. In general, the two types of
fibers produce antagonistic effects in the end organs they innervate, and many organs
receive fibers from both systems. The effects of stimulation of both cholinergic
(parasympathetic) and adrenergic (sympathetic) fibers on several organs are listed in
Table 7.

Table 7. Effects of sympathetic and parasympathetic stimulation on various effector

organs

=============================================================
Organ Sympathetic Stimulation Parasympathetic Stimulation
_______________________________________________________________________________________________________

Eye Dilatation of pupil Constriction of pupil

Salivary glands Vasoconstriction Vasodilatation
Lungs Relaxes muscles of Contracts muscles of bronchioles
bronchioles
Cardiovascular Accelerates heart, constricts Inhibits heart, dilates certain blood
arterioles vessels
Adrenal medulla Excitation
G.I. tract Inhibits motility, constricts Excites motility, relaxes sphincters
sphincters
Liver Glycogenolysis
Spleen Contracts capsule Relaxes capsule
Skin Sweat secretion
Erection of hairs
Spleen bladder Relaxes Contracts

=============================================================

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3. The Endocrine System

The endocrine glands secrete chemical mediators called hormones that

regulate growth and development, metabolism, reproduction, stress responses, body
and tissue fluid, and electrolyte balance. Thus, the endocrine system would enable
the animal to adjust to changes in its environment, and endocrinology deals largely
with this phase of environmental adjustments.

Endocrinology is defined as a branch of physiology dealing with the

coordination of various body tissues by chemical mediators (hormone) produced by
restricted areas of the body (endocrine gland) and transported through the circulatory
system to the organ or tissue on which they exert their effects.

A hormone is defined as a substance or chemical mediator produced by the

endocrine glands and carried by the blood to some distant part of the body where it
exerts its effect.

Hormones do not initiate reaction in a cell but only excite or inhibit the on-
going cell reaction. The cell must have all the materials or ingredients, the enzyme
systems, and the required environment to carry on the reaction de novo. Thus, the
hormone has only either an excitatory or inhibitory effect on existing cell reaction. It
is effective in very minute quantities (biocatalytic amount) but unlike enzyme which
has also a biocatalytic effect, hormones are destroyed in the process of participating
in the reaction, whereas, enzymes are not.

Some hormones exert their effect by participating in, or affecting

enzymatically controlled reaction in the animal. Some facilitate the passage of
important metabolites across cell membranes. Hormones are not secreted in regular
amounts; the amount of secretion depends on the need of the animal.

Hormones may be classified as simple protein, glycoprotein and steroids but

they all have common characteristics and functions:

(1) Hormones appear to regulate rather than initiate reactions;

(2) Hormones are effective in biocatalytic amounts;
(3) Hormones are not secreted in uniform rates;
(4) Hormones are inactivated rapidly either at the site where it exerts its effect or at
some other glands or organs; and
(5) Hormones are transported through the circulatory system or blood stream.

Not all hormones have specific target organs, like growth hormone or
somatotropin (STH); but for those with specific target organs, the cells in the target
organ contain receptors that specifically recognize the hormone. Hormone receptors
bind specific hormone and directly or indirectly trigger a metabolic effect.

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Hypophysis or Pituitary Gland

This is located at the base of the brain in a concavity of the sphenoid bone
called Sella turcica, which protects it from outside pressure. It has three lobes or
portion:

(1) Anterior Pituitary Lobe or Adenohypophysis; (2) Intermediate Lobe or Pars

Intermedia; and (3) Posterior Pituitary Lobe or Neurohypophysis.

The Adenohypophysis secretes the following hormones:

(1) Growth hormone or Somatotropic hormone (STH) - promotes growth of the long
bones before the epiphyseal – diaphyseal plate is fused together in adulthood.
Over secretion of STH results in Gigantism when this happens before adulthood
and Acromegaly when this happens after adulthood in human. Dwafism occurs
when there is a deficiency of STH during growth development.

(2) Adrenocorticotropic Hormone or ACTH – stimulates the adrenal cortex to

produce glucocorticoids such as cortisol, cortisone and corticosterone.

(3) Thyroid Stimulating Hormone or TSH – stimulates the thyroid gland to produce
thyroid hormones (T4 and T3).

(4) Prolactin or Luteotropic Hormone (LTH) – stimulates milk secretion in lactating

mammary gland.

(5) Follicle Stimulating Hormone (FSH) – stimulates the ovary to produce graafian
follicle; in the male, it maintains the integrity of the seminiferous tubules of the
testis.

(6) Luteinizing Hormone (LH) – stimulates ovulation in maturing graafian follicle

and the formation of corpus luteum in ovulated follicle as well as the production
of progesterone by the corpus luteum. In the male, it stimulates the cells of
Leydig or instertitial cells to produce testosterone, a male sex hormone.

The neurohypophysis secretes two hormones:

(1) Oxytocin – stimulates milk-ejection in lactating females; and

(2) Vasopression or Antidiuritic Hormone (ADH), which is important in conserving
body water by reducing urine formation.

Thyroid Gland

This gland is located at the neck area just below the Larynx. There are two
lobes of thyroid connected to each other by a bridge of tissue called Isthmus.

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 The thyroid gland maintains the level of metabolism in the tissues that is
optimal for their normal function. It secretes the hormone, thyroxine (T4) and
triiodothyronine (T3) which stimulates the oxygen consumption of most of the cells
in the body, helps regulate lipid and carbohydrate metabolism, and is necessary for
normal growth and maturation. Thyroxine increases the basal metabolic rate (BMR)
of an individual.

The thyroid gland is made up of follicles which are filled with colloids.
When the gland is inactive, the colloid is abundant, the follicles are large, and the
cells lining them are flat. When the gland is active, the follicles are small, and the
cells are cuboid or columnar.

The thyroid gland is not essential for life, but in its absence, there is poor
resistance to cold, mental and physical slowing, and, in children, mental retardation
and dwarfism (cretinism). Conversely excess thyroid secretion leads to body wasting,
nervousness, tachycardia, tremor, and excess heat production. The common disease
associated with oer activity of the thyroid gland is thyrotoxicosis such as Graves’
disease (exopthalmic goiter) caused by thyroid-stimualting immunoglobulins (TSI).
There is marked stimulation of the secretion of thyroid hormones, and the high
circulating T4 and T3 levels inhibit TSH secretion, so the circulating TSH is
depressed. The exophthalmos in Graves’ disease is due to the swelling of the tissue,
particularly the extraocular muscles, within the rigid bony walls of the orbits. This
pushes the eyeballs forward.

In the case of hypothyroidism such as simple goiter, there is lack of thyroxine

secretion due to a deficiency of iodine in the diet. Iodine is an important component
of thyroxine, thus iodine deficiency will concomitantly result in thyroxine deficiency.
The low level of thyroxine in circulation will stimulates TSH production by the
pituitary in an effort to increase thyroid activity. In the process there will be
hypertrophy and hyperplasia of the thyroid gland resulting in the production of goiter.

The secretion of T 4 or thyroxine is controlled by TSH of the pituitary.

Whenever T4 level is low, TSH production is increased and this will in turn,
stimulates increased production of T4. The high T4 in circulation will inhibit further
secretion of TSH and TSH level will decrease in circulation. This type of control is
known as negative feedback mechanism.

The Pancreas

This gland is located at the duodenal lope of the small intestine. It is both an
exocrine and an endocrine gland. It functions as exocrine gland when its acinar cells
secrete pancreatic juice containing digestive enzymes. The endocrine function is
limited to the cells of the islets of langerhans which are found throughout the
pancreas. The alpha cells of the islets of langerhans secrete glucagons which is
responsible for increasing blood sugar level; and the beta cells secrete insulin which
is responsible for lowering blood glucose level.

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 Insulin facilitates the transport of glucose from the blood into the cells of the
tissues, thus, increasing glucose utilization by the cells. It is anabolic, increasing the
storage of glucose, fatty acids, and amino acids. On the other hand, glucagons is
catabolic, mobilizing glucose, fatty acids, and amino acids from the stores into the
bloodstream. The two hormones are thus reciprocal in their overall action and are
reciprocally secreted in most circumstances. Insulin excess causes hypoglycemia,
which leads to convulsions and coma. Insulin deficiency, either absolute or relative,
causes diabetes mellitus, a complex and debilitating disease that if untreated is
eventually fatal. Glucagon deficiency can cause hypoglycemia, and glucagons excess
makes diabetes worse. A third hormone, somatostatin play a role in the regulation of
islet cell secretion. It produces hyperglycemia and other manifestations of diabetes
when there is hypersecretion or overproduction of somatostatin.

The Adrenal Gland

There are two endorsing organs in the adrenal gland, one surrounding the
other. The main secretions of the inner adrenal medulla are the epinephrine and
norepinephrine; the outer adrenal cortex secretes steroid hormones such as: (1)
aldosterone which regulate sodium metabolism by reabsorbing sodium from the
kidney tubules; (2) glucocorticoids (cortisol, cortisone and corticosterone) which
stimulate glycogenolysis and gluconeogenesis, thus, hyperglycemic.

The adrenal medulla is in effect a sympathetic ganglion in which the

postganglionic neurons have lost their axons and becomes secretory cells. The cells
secrete epinephrine and norepinephrine which are not essential for life, but they help
to prepare the individual to deal with emergencies.

The secretion of glycocorticoids is controlled primarily by ACTH from the

anterior pituitary. When there is low level of glucocorticoids in circulation. When
there is low level of glucocorticoids in circulation, ACTH secretion is increased
which in turn, stimulates increased production of glucocorticoids. The increased level
of glucocorticoids in circulation will in turn inhibit further secretion of ACTH.
Stressful stimuli will also stimulate the production of ACTH which is independent
from that elicited by the level of glucocorticoids in circulation.

The secretion of aldosterone is not under the control of ACTH but by

circulating factors such as blood pressure and/or the extra cellular fluid volume
(ECF). When the blood pressure or the ECF is low, this would stimulate the adrenal
cortex to produce aldosterone which in turn will act in the kidney tubules to
reabsorbed sodium as well as water which has a close affinity to sodium, thus,
increasing ECF volume and eventually blood pressure.

The adrenal cortex has three types of cell making up the three zones of the
cortex; the zona glomerulosa which secretes aldosterone, and the zona fasciculate and
zona reticularis which both secrete the glucocorticoids.

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Parathyroid Gland

In humans there are usually four parathyroid glands which are embedded in
the thyroid gland. However, the location of the individual parathyroids and their
number vary considerably. Parathyroid tissue is sometimes found in the
mediastinum. There are two distinct types of cells making up the parathyroid: (1)
The chief cells which have clear cytoplasm secrete the parathyroid hormone or PTH,
and (2) the less abundant and larger oxyphil cells which have oxyphil granules in
their cytoplasm, contain large numbers of mitochondria. The function of oxyphil cell
is unknown.

Parathyroid hormone or PTH mobilizes calcium from bone and increases

urinary phosphate excretion, thus in effect increases blood calcium level.
Hyperparathyroidism due to hyper-secretion of a functioning tumor in humans is
characterized by hypercalcemia, hypophosphatemia, hypercalciuria, and
hyperphosphaturia. There will be demineralization of the bones and the formation of
calcium-containing kidney stones. In young animals, demineralization of the bones
results in rickets but in adults, it is known as osteomalacia. In rickets or
osteomalacia, the amount of mineral accretion in bone per unit of bone matrix is
deficient. When there is a decrease in bone mass with preservation of the normal
ratio of mineral to matrix, the condition is known as osteoporosis. Likewise, these
bone diseases will also manifest in severe vitamin D deficiency. Thus, vitamin D is
closely associated with the function of the parathyroid gland. Vitamin D is
metabolized in the kidney tubules into 1, 25 dihydroxycholecalciferol which
increases the efficiency of calcium and phosphate absorption into the intestinal wall,
thus, making these minerals available for bone formation. On the other hand, when
there is vitamin D deficiency, limited amounts of calcium and phosphate are absorbed
from the intestines resulting in low blood calcium level, thus, stimulating the
parathyroid gland to secrete PTH, resulted in calcium mobilization from the bones.

In hypoparathyroidism, osteoclerosis may set in due to increased amount of

calcified bone. Symptoms manifested include hypocalcemia, hyperphosphatemia,
hypercaliurea and hypophosphaturia. One should understand that there is a constant
ratio of calcium and phosphorus being maintained in the blood circulation. When the
blood level of phosphorus is high, calcium level is low and vice versa.

Calcitonin or also known as Thyrocalcitonin is a hormone that lowers

calcium level in the blood, thus, has an opposite effect to that of parathormone which
increases calcium level in the blood. Thyrocalcitonin is secreted by the thyroid gland
upon stimulation by a secretion coming from the parathyroid gland in response to a
high calcium level in the blood perfusing the parathyroid gland. Thyrocalcitonin
lowers blood calcium level by preventing bone resorption through the activation of
the osteoblast cells which stimulate bone formation. Also, thyrocalcitonin increases
calcium excretion in the urine, thus, contributing to the lowering of blood calcium
level.

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 There are three types of cells associated with bone formation and bone
resorption: (1) osteoblast stimulates bone formation; and (2) osteoclast and (3)
osteocyte are both associated with bone resorption. When there is hypersecretion of
PTH, osteoclast and osteocyte cells predominate to cause bone resorption; on the
other hand, when there is hyposecretion of PTH or hypersecretion of calcitonin,
osteoblast cells predominate to cause bone formation.

4. The Cardiovascular System (CVS)

The CVS includes the heart, the blood and the blood vessels through which
the blood flows in circulation. The CVS has the following functions:

(1) To convey the nutrients absorbed from the digestive tract to the tissues;
(2) To carry O2 from the lungs to the tissues and CO2 from the tissues to the lungs;
(3) To remove the waste products of metabolism and take them to the excretory
organs for disposal;
(4) To transport hormones from one part of the body to another;
(5) To help in maintaining the water equilibrium of the body;
(6) To assist in keeping the normal temperature of the body;
(7) To regulate the hydrogen ion concentration in the body; and
(8) To assist in overcoming diseases by the antibodies contained in the blood.

The Heart

The heart is located in the middle mediastinal space. It is enclosed with a

pericardium or pericardial sac. The mammalian heart has 4 chambers; the upper 2
chambers are the atria, and the lower 2 chambers, are the ventricles (Fig. 5). There is
a complete septum separating the left and the right side of the heart. However, free
communication exists between the atrium (auricle) and the ventricle on the same side
of the heart. The atrio-ventricular valve or A-V valve prevents the backflow of
blood from the ventricle to the atrium during ventricular systole. The vale on the
right side is called tricuspid value and the one on the left side is known as bicuspid or
mitrral valve. A valve also stands at the aortic orifice (aortic valve) and at the
pulmonary orifice (pulmonary valve). These valves prevent the backflow of the
blood from these blood vessels (aorta and pulmonary artery) into the ventricles during
diastole.

The heart normally beats in an orderly sequence: contraction of the atria

(atrial systole) is followed by the contraction of the ventricles (ventricular systole),
and followed by diastole, in which all the 4 chambers are in isometric relaxation, the
AV valves open, thus, allowing the blood to fill up the ventricles. In fact, ¾ of
ventricular filling occurs during diastole and complete filling occurs during
atrial systole. At the start of ventricular systole, the AV valves are closed, and the
aortic and pulmonary valves are opened to allow the flow of ventricular blood into the
aorta and pulmonary artery, respectively. However, not all ventricular blood are

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Figure 5

24
 ejected at the end of the ventricular systole; about 50 ml of blood are left in each
ventricle in human heart as end systolic ventricular volume.

Contraction of the heart is spontaneous and is initiated by the depolarization of

the Sino-atrial node (SA node) (see Fig. 6). The depolarization spreads radially
through the atria resulting in atrial systole, and converges on the atrio-ventricular
node (AV node). From the AV node, the wave of depolarization passes through the
bundle of His, then through the Purkinje system to the ventricular muscle, causing
ventricular systole. The SA node is the cardiac pacemaker and its rate of discharge
determines the rats at which the heart beats. However, vagal stimulation results in
bradycardia or slowing of heart rate and stimulation of the sympathetic cardiac nerve
results in tachycardia or increased heart rate. Temperature also influences the rate of
discharge of SA node. Increased temperature results in tachycardia.

Heart Sound

Two sounds are normally heard through a stethoscope during each cardiac
cycle: a low, slightly “lub” sound (first sound), caused by the closure of the mitral
and tricuspid valves; and a shorter, high pitch “dub” sound (second sound), caused by
the closure of the aortic and pulmonary valves just after the end of ventricular systole.

The blood forced into the aorta during systole not only moves the blood in the
vessels forward but also sets up a pressure wave which travels down the arteries. The
pressure wave expands the arterial wall as it travels, and the expansion is palpable as
the pulse. Thus, the pulse is a wave of dilation of an artery originating from the aorta
as the blood flows into it from the heart. The rate of heart beat is usually measured by
determining the pulse rate. The average pulse rate per minute in different classes of
animals is as follows:
Elephant - - - - - - - - 30-45 pulse rate/minute
Horses - - - - - - - - 38 pulse rate/minute
Carabao and Cattle - 54 pulse rate/minute
Goat -------- 78 pulse rate/minute
Chicken - - - - - - - - 200-400 pulse rate/minute
Mouse - - - - - - - - 600 pulse rate/minute

Pulse rate may be taken by feeling the artery on the following animals:

Horse – external maxillary artery or about the middle of the lower jaw
Cattle and Carabao – similar location as in the hose but slightly on the outer surface;
coccygeal artery at the base of the underneath of the tail
Sheep, Goat, Dog and Cat – femoral artery
Pigs and others – auscultation method using stethoscope at the cardiac or chest region

25
Figure 6

26
The Blood Vessels

In general, the blood vessel that carries blood away from the heart is called
artery; and that which carries blood back to the heart is called vein. Also, the blood
running through the artery is oxygenated blood; and that which runs through the vein
is unoxygenated blood. The only exceptions to the principle are the pulmonary artery
which carries unoxygenated blood from the right ventricle to the lungs, and the
pulmonary veins which carry oxygenated blood from the lungs to the left atrium of
the heart. The aorta or aortic artery carries blood from the left ventricle to the
different systemic circulations, such as the head, neck, trunk, limbs, and the visceral
organs. The aorta gives off to smaller branches of arteries which in turn give rise to
several arterioles. An arterile gives rise to a bed of capillaries which eventually join
together to form a venule. A venule joins to a bigger vein which eventually end up on
the vena cava which returns unoxygenated blood from several systemic circulations
to the right atrium of the heart.

Blood Circulation

Venous blood coming from the different parts of the body is returned back to
the heart via the vena cava to the right atrium (Figure 7). From the right atrium it
goes to the right ventricule through the tricuspid valve. Then it passes through the
pulmonary valve and goes to the pulmonary artery which carries the blood to the
lungs (pulmonary circulation). In the lungs, exchange of gases takes place: carbon
dioxide is given off and oxygen is taken in by the circulating blood. The oxygenated
blood is returned back to the heart by the pulmonary veins which enter the heart at the
left atrium. From the left atrium, the blood goes to the left ventricles through the
mitral or biscuspid valve. Then it goes through he aortic valve to the aorta which
carries the blood to the different systemic circulations. In systemic circulation, the
oxygen is taken in by the tissues and carbon dioxide is given off by the tissues to the
circulating blood. These cellular exchange of gases take place at the different
capillary beds. Then all the venous blood from the systemic circulations are returned
back to the heart via the vena cava.

The systemic circulation includes the following special systems of blood

circulation:

(1) The coronary circulation – is a part of systemic circulation; it supplies blood to

the heart itself.
(2) Hepatic circulation – is a part of systemic circulation; it supplies arterial blood to
the liver.
(3) Cerebral circulation – supplies arterial blood to the brain.
(4) Renal circulation – supplies arterial blood to the kidney.
(5) Splanchnic circulation – supplies arterial blood to the digestive tract.

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Figure 7.

28
The Blood

Blood is a thick suspension of cellular elements in an aqueous solution of

electrolytes and some non-electrolytes. By centrifugation, the blood is separated into
the two categories of plasma and cells.

1). Plasma – the fluid portion of the blood containing a number of ions, inorganic
molecules, and organic molecules which are in transit to various parts of the body
or which aid in the transport of other substances. Blood plasma is composed of
the following important constituents: water, gases (oxygen, carbon dioxide,
nitrogen), proteins (albumin, globulin, fibrinogen), glucose, lipids (fats, lecithin,
cholesterol), non-protein nitrogen substances (amino acids, urea, uric acid,
creatine, creatinine ammonia, salts, etc., inorganic salts and minerals (chlorides,
bicarbonates, sulfates, phosphates of sodium, potassium, calcium, magnesium,
iron, and traces of manganese, cobalt, copper, zinc, etc.), enzymes, hormones,
vitamins, immune substances, etc. The normal plasma volume is about 3 to 5% of
the body weight.

2. Blood cells – made up the white blood cells or WBC (leukocytes), the red blood
cells or RBC (erythrocytes) and the platelets, which are all suspended in the
plasma. The number of each kind of cell present in the blood is determined by
means of a haemocytometer.

a) The white blood cells are of three types: Granulocytes, Lymphocytes and
Monocytes. Of these, the granulocytes or polymorphonuclear leukocytes
(PMN) are the most numerous. The granulocytes are subdivided into
neutrophils, eosinophils and basophils based on their affinity to either neutral,
acidic or basic dyes, respectively. They are formed from stem cells in the
bone marrow, mature rapidly and enter the circulation where they survive for
no more than 2 weeks. Their main function is phagocytic in nature. At least
in the neutrophils and eosinophils, the granules appear to be lysosomes and
function in the digestion of material (like bacterial) taken into the cells by
phagocytosis. Old granulocytes are normally destroyed in the spleen and
other portions of the reticulo-endothelial system.

Lymphocytes are mostly formed in the lymph nodes, spleen and

thymus and to some extent also in the bone marrow. They enter the blood
circulation for the most part via the lymphatics. They are believed to produce
antibodies and counteract toxins.

Monocytes are large none-nuclear leukocytes. They are also called the
transitional cells and have well developed motility. They are believed to come
from the reticulo-endothelial cells. Like neutrophilic leukocytes, they are
actively phagocytic and are capable of ingesting all sorts of foreign matter.

29
 b) The erythrocytes are biconcave disks manufactured in the bone marrow. In
mammals, they lose their nuclei before entering the circulation. These non-
nucleated cells are soft and contain hemoglobin. Hemoglobin is a complex
iron-containing conjugated protein with a molecular weight of about 68,000.
It is globular molecule made up of 4 subunits, and each unit contains a red
pigment, iron-containing derivative porphyrin called heme moiety conjugated
to a polypeptide, globin. The oxygen-carrying property of hemoglobin is due
to the iron content in the pigment.

Hemoglobin binds O 2 to form oxyhemoglobin, O2 attaching to the Fe++

in the heme. Since hemoglobin contains 4 Hb units, the hemoglobin molecule
actually reacts with 4 molecules of O2 to form Hb408.

Hb4 + 4O2 Hb4O8

This reaction is oxygenation (not an oxidation) and requires less than

0.01 second. Oxygen is afterwards readily given off to the tissues as the blood
goes to the systemic capillaries. In the muscles, oxygen is taken up by
myohemoglobin.

c) Platelets or thrombocytes – are small oval disk-like granulated bodies 2-4

microns in diameter. There are about 300,000/cu mm of circulating blood.
The megakaryocytes, giant cells in the bone marrow form platelets by
pinching off bits of cytoplasm, and extruding them into the circulation. When
blood vessel walls are injured, platelets collect at the site, sticking to the
vessel wall and liberating serotonin, which leads to local vasocontriction.
They also liberate thromboplastin which aids in blood clotting, and they play
a role in clot reactions.

Blood Coagulation

The essential process in coagulation is the conversion of the soluble plasma

protein, fibrinogen, into the insoluble protein, fibrin, a reaction that is catalyzed by
the enzyme thrombin. Thrombin is formed from its inactive circulating precursor,
prothrombin, in the presence of calcium ions by the action of activated
thromboplastin. Prothrombin is synthesized in the liver, and vitamin K is essential in
the hepatic synthesis of prothrombin. This is precisely why vitamin K is essential in
blood clotting mechanism.

Activated thromboplastin is made available at the site of the injury in the

presence of Ca++ by reaction involving platelets and some clotting factors. The
schematic mechanism in blood clotting is shown below:

Vitamin K
Liver Prothrombin
Factor VII and X

30
 Ca++
Prothrombin Thrombin
Activated
thromboplastin
(Platelets)

Thrombin
Fibrinogen Fibrin (clot)

Lymphatic System

The circulatory system and the lymphatic system are related to the body fluid
compartments. The animal body is made up of 60-70% water. This is distributed as
intracellular fluid (ICF) and extracellular fluid (ECF). The ICF is about 40-50% of
the body weight and the ECF is about 20% of the body weight.

In animals with closed vascular system, the ECF is divided into 2

components: The Interstitial fluid which consists of the cerebrospinal fluid, synovial
fluid and the lymph; and the blood plasma. The interstitial fluid is about 15% of the
body weight and the blood plasma is about 5% of the body weight.

The lymphatic system is composed of lymph node, lymph vessel and the
lymph.

The lymph nodes and its function. The lymph nodes are small bodies of
lymphoid tissues which are ovoid or bean shaped and located in strategic points of the
body through which the lymph passes on its way to the blood stream. It is generally
agreed that lymph nodes have at least 2 functions. One of these is the production of
lymphocytes of which the lymph nodes contain large numbers. Another function is
to stop foreign materials that come to them in the lymph. This filtration is said to be
accomplished mechanically and by the phagocytic activity of the reticulo-endothelial
cells. They become swollen or inflamed during severe bacterial infections.

The lymph vessels. The lymphatic drain from the lungs and from the rest of
the body tissues via a system of vessels that end in the venous system. The lymph
vessels begin in the tissues as blind lymph capillaries, similar in structure to blood
capillaries. By the convergence of lymph capillaries, smaller lymph vessels are
formed, and these in turn unite to form larger lymph vessels. Like the veins, the
lymph vessels contain valves which prevent the back flow of its content, but have
thinner walls than the veins. Ultimately, all the lymph vessels drain into either the
thoracic duct or the right lymphatic duct, which empty into the venous system
anterior to the heart. Lymph from the right side of the head and neck, the right
forelegs and the right side of the thorax drain to the right lymphatic duct, which
empty into the venous system anterior to the heart. Lymph from the right side of the

31
head and neck, the right forelegs and the right side of the thorax drain to the right
lymphatic duct; that from the rest of the body, to the thoracic duct.

Flow of lymph. The tissue fluid is in communication with the blood in the
capillaries, the intracellular fluid, and the lymph capillaries. The latter remove from
the tissue spaces materials that do not or cannot enter the blood capillaries. Water and
crystalloids can move either way. Particulate matter and large molecules such as
proteins and lipids cannot enter the blood capillaries but can penetrate the much more
permeable wall of the lymph capillaries. The flow of lymph in the lymph vessels is
sluggish and in one direction only, from the tissues towards the heart. The factors
concerned in lymph flow are: (1) the difference in pressure at the two ends of the
lymph system; (2) the massaging effect of muscular movements; and (3) the presence
of the lymph vessels of valves, which permit flow in only one direction, that is,
towards the heart.

Composition of Lymph

Tissue fluid and lymph proper, that is, the fluid in the lymph vessels are
different. Lymph, derived largely from the blood, is similar in composition to blood
plasma. The plasma of the blood passes through the thin wall of the blood capillaries,
enters the tissue spaces, and becomes tissue fluid or lymph. The cells of the tissues
themselves also contribute somewhat to the composition of the lymph, for there is
free interchange between intracellular fluid and the tissue fluid. In this way the cells
rid themselves of the waste products of metabolism and absorb foodstuffs.

The composition of lymph varies with the state of activity of the digestive
organs, lymph derives from the intestine during fat absorption has a milky appearance
because of the fat that it contains and is known as chyle.

Ordinarily, lymph is colorless, clear, watery liquid having a specific gravity

of about 1.015. It contains a few red cells normally, and lymphocytes are present.
The latter cells are more abundant in lymph that has passed through lymphoid tissue.
Whether or not monocytes are present is uncertain. Neutrophilic leukocytes are
ordinarily absent; however, they may be present in great number in infections.
Platelets are said to be absent; nevertheless, lymph will clot, though feebly. Lymph
contains water, glucose, gases, proteins, non-protein nitrogenous substances,
inorganic substances, hormones, coenzymes, vitamins, and immune substances. The
proteins are the same kind as in blood plasma but the amount is less. This is
especially true of lymph from the limbs, for the capillary walls in these regions are
less permeable to the blood proteins than in other regions.

Inter-relationship between the circulatory system and the lymphatic system

All body tissues are supplied with blood capillaries as well as lymph
capillaries. The blood capillaries absorb substances produced by the cells and other
nutrients, and metabolites present in the interstitial fluid which require the circulatory

32
 system for their distributions to other parts of the body. However, there are
substances which cannot readily enter the walls of the blood capillaries because of the
size of their molecules, such as protein molecules of certain hormones and enzymes.
These protein molecules can still join the circulatory system by way of the lymphatic
system. Since the lymph capillaries have more permeable walls than the blood
capillaries, all metabolites of big molecular size which cannot be absorbed by the
blood capillaries will be absorbed by the lymph capillaries. Eventually, the lymph
fluid will enter the circulatory system through the right lymphatic duct and the
thoracic duct.

5. The Respiratory System

The main function of respiration is to provide oxygen to the cells of the body
and to remove excess carbon dioxide from them. Different species achieve this in
different ways. Unicellular organisms get their O2 by diffusion from the fluid
surrounding them and eliminate CO2 in the same way: larger organisms cannot.
Some larger organisms that live in air (certain insects) do get enough O 2 by diffusion
alone, but they have a special system of air tubes (trachea or spiracles) that pipe air
directly to many regions of the body, so that the distance O 2 must diffuse to reach
tissue cells are short. Large animals, including man, make use of two systems:

(1) A blood circulatory system to carry to and from the tissue cells large
quantities of O2 and CO2, with the help of hemoglobin; and

(2) A respiratory system, a gas exchanger, to load the blood with O 2 and
remove excess CO2. In fish, blood flows through gill vessels and extracts O 2 from
water flowing around them. In man and other farm animals, the respiratory surfaces
are folded within the body to prevent drying of the delicate membranes; air saturated
with water vapor is drawn into intimate contact with the blood flowing through the
pulmonary capillaries, and gases are exchanged.

These two systems cooperate to supply the needs of the tissues. One system
supplies air; the other supplies blood. The ultimate purpose is the transfer of gases
between air and cells. The respiratory system is an air pump which draws fresh air
through the air tubes to small air sacs (alveoli) that have vary thin membranes. The
circulatory system is a blood pump which drives the whole output of the heart
through fine thin-walled blood tubes (capillaries) surrounding the alveoli.

The Respiratory Apparatus

The nasal cavity has two nasal tubes (some a third tube, the mouth, is also
used), and then becomes one, the trachea. The trachea is always kept open by the
presence of rings of cartilage in its wall. It subdivides into two main branches, the
right and left bronchi, which are similar in structure and function as trachea. Each of
the two bronchi divides into two more, and each of these into two more, and so on
until there have been 20-23 subdivisions in all. A sim0ple calculation shows that 20

33
subdivisions of this type produce about a million terminal tubes. At the end of each
are numerous blind pouches, the alveoli or alveolar sacs; here gas exchange occurs.
There are about 300 million of these in the two lungs of man; their diameter varies
from 75 to 300 microns.

The lungs may be regarded as two elastic membranous sac those interior (in
free communication with the outside air through the respiratory passages) is highly
modified and enlarged by the presence of numerous alveoli. The wall of the alveolus
is composed of a single layer of respiratory epithelium. Across this layer of cells and
the endothelium of the blood capillaries, gaseous exchange between the air in the
alveoli and the blood in the numerous adjacent capillaries takes place. The total area
of the alveolar walls in contact with the capillaries in both lungs is estimated to be 70
square meters in man, which is about 40 times the surface area of the body.

The thoracic cavity contains the lungs and the mediastinal organs. This
cavity is completely separated from the abdominal cavity by the diaphragm. The
pleura, a serous membrane, line the thoracic cavity, forming the lateral walls of
mediastinum and are reflected from there on the lungs, thus forming a pleural cavity.
The pleural cavity is merely a capillary space, occupied by a thin film of fluid, which
serves to moisten and lubricate the two pleural layers. The pressure in the pleural
cavity is negative. Therefore when the pleural cavity is opened, air rushes in and the
lungs will collapse.

The inspiratory muscles consist of the diaphragm and the external intercostals
muscles. The movement of the diaphragm accounts for 75% of the change in
intrathoracic volume during quiet inspiration. The diaphragm is attached around the
bottom of the thoracic cage and arches over the liver and moves downward like a
piston when it contracts. The distance of movement is about 1.5 to 7.0 cm.

The external intercostals muscles run obliquely downward and forward from
rib to rib. The ribs pivot as if hinged at the back, so that when the external
intercostals muscles contract, they elevate the lower ribs. This pushes the sternum
outward and increases the antero-posterior diameter of the chest.

The expiratory muscles consist of internal intercostals muscles and the

muscles of the anterior abdominal wall. The internal intercostals muscles pass
obliquely downward and posteriorly from rib to rib, and therefore, pull the rib cage
downward when they contract. The muscles of the anterior abdominal wall also aid
expiration by pulling the rib cage downward and inward; and by increasing the extra
abdominal pressure which pushes the diaphragm upward.

34
The Respiratory Center

There are at least three major parts of the respiratory center. They are:

1. Medullary center – capable of initiating and maintaining sequences of the

respiratory cycle. This contains the minimal number of neurons necessary for
the basic coordinated sequence of inspiration, expiration inspiration. This
center is often divided into an Inspiratory Center and an Expiratory Center,
because maximal sustained inspiration follows electrical stimulation of some
region; and maximal expiration follows stimulation of adjacent regions. At the
lateral sides of this region, there are special receptors which are believed to
respond to H+ concentration. A rise in H+ results in hyperventilation.

2. Pneumotaxic center – located in the upper pons above the medullary center.
Stimulation of this center accelerate respiration, especially expiration. It is
postulated that inspiration sets up impulses that ascend from the medullary
inspiratory center to the pneumotaxic center, where they generate impulses that
descend to the expiratory center and inhibit inspiration, a negative feedback
mechanism.

3. Apneustic center – located in the lower pons, between the pneumotaxic center and
the medullary center. The role of this center is revealed when both the
pneumotaxic center and the vagi are inactivated; prolonged apneusis then
results. (Apneusis is the cessation of respiration in the inspiratory position).

Regulation of Respiratory Center Activity

Respiration would increase whenever cells of the body need more O 2 or form
more CO2 and would decrease whenever they need less O 2 or from less CO2. There
are many sensory receptors, in many locations which can influence respiration rate;
appropriate electrical stimulation of almost any sensory nerve and of many parts of
the brain can affect respiration. However, some receptors appear to be highly
specialized for the task of respiratory regulation. These receptors are sensitive to
chemical changes in their environment – therefore called generally as
chemoreceptors. The well known of the chemoreceptors are:

1. Medullary chemoreceptors – believed to be located on the ventral surface of the

brain stem. It is believed to monitor the H+ concentration of the cerebrospinal
fluid or, possibly the brain interstitial fluid. An increase in H + concentration
stimulates respiration.

2. Carotid bodies – are small, pinkish nodules located just beyond the bifurcation of
the common carotid artery into the external and internal carotids. The carotid
bodies are completely different from the carotid sinuses in structure and in
function. The carotid sinuses contain mechanoreceptors that respond to changes

35
 in stretch or deformation of the carotid artery wall; the carotid bodies contain
chemoreceptors that respond to certain changes in their chemical environment.

3. Aortic bodies – contain chemoreceptors that function separately from aortic

pressoreceptors, which are in the wall of the ascending arch of the aorta. Most of
the aortic chemoreceptors lie between the arch of the aorta and the pulmonary
artery or on the dorsal aspect of the pulmonary artery.

The carotid and aortic chemoreceptors are sensitive to changes in PO2, PCO2
and H+ concentration in arterial blood. When the PCO 2 or H+ in arterial blood is
increased, or when arterial PO2 is decreased, the carotid and aortic chemoreceptors
are stimulated and the respiratory center activity increases.

Application of acetylcholine or nicotine to the chemoreceptor areas stimulates

respiration; whereas, application of cyanide or procaine reduces or abolishes
respiration.

The non-chemical influence of respiration can be shown by the fact that

breathing can be controlled. Irritation on the walls of the trachea or bronchi produces
coughing which begins with a deep inspiration followed by forced expiration against
a closed glottis. The glottis is then suddenly opened, thus, producing an explosive
outflow of air at velocities up to 600 miles per hour. Sneezing is a similar expiratory
effort with a continuously open glottis. This illustrates that non-chemical factors can
influence the activity of the respiratory center in response to some mechanical
stimuli.

The respiratory adjustments during vomiting, swallowing and gagging are

other examples of non-chemical control of respiratory. Inhibition of respiration and
closure of the glottis during these activities not only prevents the aspiration of food or
vomitus into the trachea but, in the case of vomiting, fixes the chest so that
contraction of the abdominal muscles increases the intra-abdominal pressure.

Mechanisms of Inspiration

Inspiration is an active process. Contraction of the diaphragm increases the

longitudinal diameter of the chest. Also, the contraction of the external intercostals
muscles elevates the ribs, resulting in an increased transverse diameter of the thorax.

At the start of inspiration, the intrapleural pressure is about –2.5 mmHg

(relative to atmospheric pressure). When the chest volume is increased, the
intrapleural pressure is further decreased to about –6 mmHg, and the lungs are pulled
into a more expanded position. The pressure in the airway becomes slightly negative,
and air flows into the lungs.

36
Mechanism of Expiration

Following an inspiration, the enlarged thorax may return to its resting

position by purely passive forces, that is, without muscular effort. At the end of
inspiration, the lung recoil pulls the chest back to the expiratory position where the
recoil pressures of the lungs and chest wall balance. The pressure in the airways
becomes slightly positive, and air flows out of the lungs.

Although in quite breathing expiration is passive, labored breathing is

accompanied by active expiration, that is, the return of the thorax to other resting
position being hastened. This is accompanied by the contraction of the expiratory
muscles (Internal Intercostal Muscles and the muscles of the anterior abdominal
wall). Very active expiration is seen also in coughing, talking, laughing, barking,
etc.).

Regulation of Respiration

The muscles of respiration posses no inherent rhythm. Spontaneous

respiration is completely dependent upon the discharge of the respiratory center in the
Medulla oblongate. If the efferent connection from the center to the respiratory
muscles are interrupted, or if the center is destroyed, breathing stops.

Lung Volume

The amount of air that moves into the lungs with each inspiration (or the
amount that moves out with each expiration) is called the tidal volume or TV. The air
inspired with a maximal inspiratory reserve volume (IRV). The volume expelled by
an active expiratory effort after passive expiration is the expiratory reserved volume
(ERB), and the air left in the lungs after a maximal expiratory effort is the residual
volume (RV). The space in the conducting zone of the airways occupied by gas that
does not exchange with blood in the pulmonary vessels is the respiratory dead space.
The vital capacity, the greatest amount of air that does not exchange with blood in the
pulmonary vessels is the respiratory dead space. The vital capacity, the greatest
amount of air that can be expired after a maximum inspiratory effort, is frequently
measured clinically as an index of pulmonary function. It gives useful information
about the strength of the respiratory muscles and other aspects of pulmonary function.
The fraction of the vital capacity expired in 1 second (timed vital capacity; also called
forced expired volume in 1 second, or FEV 1”) gives additional information; the vital
capacity may be normal but the timed vital capacity is greatly reduced in diseases
such as asthma, in which the resistance of the airways is increased owing to bronchial
constriction. The amount of air inspired per minute (pulmonary ventilation,
respiratory minute volume) is normally about 6 L (500 ml/breath x 12 breaths/min).

37
 Gas Exchanges

Oxygen continuously diffuses out of the gas in the alveoli (alveolar gas) into
the blood stream, and CO2 continuously diffuses into the alveoli from the blood. In
the steady state, inspired air mixes with the alveolar gas, replacing the O 2 that has
entered the blood and diluting the CO2 that has entered the alveoli. Part of this
mixture is expired. The O2 content of the alveolar gas then falls and its CO 2 content
rises until the next inspiration. Since the volume of gas in the alveoli is about 1.8-2.0
L at the end of expiration (functional residual capacity), each 350-ml to 500 ml
increment of inspired and expired air changes the PO2 and PCO2 very little. Indeed,
the composition of alveolar gas remains remarkably constant, not only at rest but in a
variety of condition as well.

Oxygen moves from the alveoli to the capillaries across the thin membrane
formed by the epithelial cells, the endothelial cells, and their fused basement
membrane. The PO2 of the alveolar air is 100 mmHg, whereas, that in the venous
blood in the pulmonary artery is 40 mmHg. There is no evidence that any process
other than passive diffusion is involved in the movement of O2 into the blood along
this pressure gradient. Diffusion into the blood must be very rapid, since the time
each milliliter of blood is in the capillaries is short. Nevertheless, O 2 diffusion is
adequate in health to raise the PO2 of the blood to 95 mmHg, a value just under
alveolar PO2. On the other hand, CO2 diffuses from the capillaries to alveolar air,
thus, reducing the CO2 content of the blood which returned to the heart at 40 mmHg.
In short, the arterial blood distributed to the different systemic circulation has a PO 2
of 95 mmHg and PCO2 of 40 mmHg. The tissues which would receive the arterial
blood has PO2 of 40 mmHg and PCO2 of 46 mmHg. Thus, at the tissue level O2
diffuses out of the blood capillaries to the tissues, whereas, CO2 diffuses from the
tissues to the blood capillaries. The resultant venous blood has PO 2 of 40 mmHg and
PCO2 of 46 mmHg when returned back to the right atrium of the heart.

6. The Excretory System – Kidney and Body Fluids

The kidneys play a prominent role in regulating (a) the concentration of

metabolic wastes; (b) the osmotic pressure; (c) the fluid volume; and (d) the ionic
composition of our internal environment. The kidneys are commonly described as
excretory organs’ but actually, they are primarily organs which regulate volume and
composition of the internal fluid environment. Their excretory function is incidental
to their regulatory function.

The kidney aids in keeping the composition of blood plasma constant by:
1. The excretion of urea and other nitrogenous waste products of metabolism;
2. The elimination of excess inorganic salts;
3. The elimination of excess water; and
4. The elimination of non-volatile, soluble foreign substances that may have gained
entrance to the blood.

38
Structure of the Kidney

The kidneys are paired, somewhat flattened bean-shaped organs which lie
retroperitoneally on either side of the vertebral column against the posterior
abdominal wall. Each kidney is supplied with blood by a single renal artery which
arises from the abdominal aorta. Each renal artery divides the pelvis. These arterial
branches pass between the calyxes and penetrate the parenchyma. Within the
parenchyma, these arteries are designated as interlobar because they course between
the lobes or pyramids.

At the junction of cortex and medulla, the interlobar arteries bend over the
bases of the pyramids to form a series of incomplete arches, the acriform arteries.
Interlobar arteries arise at right angles from the acriform arteries and run radially
toward the periphery in the cortical medullary rays. In their course through the cortex
they give rise to short lateral branches, the afferent arterioles, each of which supplies
a glomerulus. The venous system which drains blood out of the kidney runs parallel
with the arterial system and comes out of the kidney as renal vein.

Each kidney is composed of over a million units of nephrons. A nephron

consists of several parts (Figure 17):

(1) The glomerulus is formed by the invagination of a tuft of capillaries into the
dilated, blind end of the nephron called the Bowman’s capsule. The glomerulus
and the Bowman’s capsule is collectively called as the renal corpuscle or
malphighian.

(2) The proximal tubule is joined to Bowman’s capsule by a short connecting

segment. The convoluted portion of the proximal tubule (pars convuluta) enters a
cortical medulla ray to penetrate the deeper layers of the cortex and become the
loop of Henle. The proximal tubule is composed of a single layer of cuboidal or
trancated pyramid cells resting on a basement membrane. The cells are coarsely
granular; nuclei are large and basally located. The apical surfaces of the cells
which bulge into the tubular lumen are covered with numerous cytoplasmic
filaments which are revealed by electronmicroscopy as microvilli.

(3) The loop of Henle includes: the descending thick limb (pars recta) of the
proximal tubule; the descending and ascending thin limbs; and the ascending
thick limb, to be described later as a part of the pars recta of the distal tubule. The
thin segment of the loop of Henle arises abruptly from the descending thick
segment of the loop, the so-called pars recta of the proximal tubule. The diameter
of its lumen is less than that of either the proximal or the distal segment. The
cells are flattened and thin except in the nuclear region, which bulges into the
lumen. The descending limb of the lob of Henle bends upward forming the
ascending limb of the loop of Henle, which communicate with the pars recta of
the distal convoluted tubule.

39
(4) The distal tubule is shorter than the proximal tubule, and its convolutions are less
complex. The cells of the distal tubule are cuboidal in the region of the pars recta
and become more columnar in the cortical convolutions.

(5) Several distal tubules coalesce to form collecting ducts which pass through the
renal medulla to empty into the pelvis of the kidney at the apices of the medullary
pyramids. The renal pelvis drains into the ureter and the latter enters the urinary
bladder.

Urine Formation

In the kidneys, a fluid that resembles plasma is filtered through the

glomerular capillaries into the renal tubules (glomerular filtration). As this
glomerular filtrate passes down the tubules, its volume is reduced and its composition
altered by the processes of tubular reabsorption, active transport, diffusion and
osmosis to form urine.

Each day, about 160 to 180 liters of water are filtered through the glomeruli
of normal man. Each liter contains 300 mOsm/L of solute, consisting largely of
sodium, chloride and bicarbonate ions. As the filtrate flows along the proximal
convoluted tubules, sodium is actively extruded into the interstitium of the cortex.
Chloride follows sodium passively down an electrical gradient and water is
reabsorbed by osmosis. The ions and water deposited in the interstitium are rapidly
carried away by blood perfusing the cortical capillaries. Although the volume of the
tubular fluid is sharply reduced to perhaps 20% of that of the filtrate at the ends of the
thick descending limbs of Henle’s loops, the osmolar concentration remains
unchanged at 300 mOsm/L.

As the tubular fluid progresses down the thin descending limbs of Henle’s
loops, water diffuses out into the hypertonic interstitium of the medulla and papilla;
and sodium diffuses in. Volume decreases and osmotic pressure increases
progressively to the bends of the loops. In the ascending limbs of Henle’s loops,
sodium is extruded into the interstitium. Because the ascending limbs are
impermeable to water, the osmolar concentration of the tubular fluid is reduced. At
each level, a gradient of about 200 mOsm/L is established between tubular contents
and hypertonic interstitium. Indeed, it is this capacity of the tubular epithelium to
establish a modest osmolar gradient at each level which accounts for the much more
significant gradient of 300 to 1200 mOsm/L developed along the length of the loops.
This process is termed as countercurrent multiplication of concentration.

The fluid which enters the distal convoluted tubules is hypotonic to the
surrounding cortical interstitial fluids. Its volume is perhaps 15% of that of the
glomerular filtrate. The continued active extrusion of sodium and the passive osmotic
diffusion of water continue in the distal tubule, reducing volume to a few percent of
that of the filtrate as the fluid enters the collecting duct. This fluid becomes
progressively concentrated as it flows along the collecting duct and gives up water to

40
 the hypertonic medullary and papillary interstitium. The final urine entering the renal
pelvis, is essentially as concentrated as the interstitial tissue of the tips of the papillae.

The water which diffuses out of the descending limbs of Henle’s loops and
out of the collecting ducts, and the sodium, which is pumped out of the ascending
limbs of Henle’s loop, are removed by blood perfusing the vasa recta of the medulla
and papilla. These vessels serve as countercurrent exchangers to reduce excessive
loss of osmotically active solutes from the medulla and papilla.

In water diuresis, in which the titer of circulating ADH is low, the epithelium
of the distal tubules and collecting ducts is impermeable to water. The hypotonicity
of the tubular urine leaving the loops of Henle is maintained through out the
remainder of the nephron and is increased by the continued active extrusion of ions.
The final urine is dilute and its volume large.

Role of ADH in Water Conservation

Vasopressin or ADH conserves body water by the following actions:

(1) Since ADH is a vasoconstrictor, it reduces medullary and papillary blood flow in
the renal interstitium, thus increasing tissue hypertoni-city of the interstitium.
This would increase the concentration gradient of the solute between the
interstitium and the fluid in the lumen of the kidney tubules, thus, the reabsorption
of water by osmosis would increase.
(2) ADH may stimulate the “sodium pump” of the ascending loop of Henle, thus,
increasing both the rate of sodium transport from the tubule lumen to the
interstitium and the concentration gradient of sodium between the interstitium and
the fluid in the lumen of the tubules.
(3) ADH may dilate the “pres” of the collecting duct, thus, facilitates water
reabsorption.

7. The Reproductive System

There are two forms of reproduction: asexual and sexual reproduction. The
asexual reproduction does not require the sex organ to facilitate the perpetuation of
the species. Asexual reproduction is very common in unicellular organisms as well as
in some species of plants. Unicellular organisms may multiply by fission wherein the
individual organism may divide into two individual cells. In certain plants,
propagation may be achieved by using cuttings as planting materials or by budding,
marcoting or by propagating plant cell in culture media (Tissue Culture Technique).
All these forms of reproduction do not use sex organs in the process of propagation,
thus, they are asexual.

In the sexual reproduction, male and female sex organs are involved in the
process of propagation. The union of sex cells, ovum and sperm cell, are involved to
form a new individual. This is the usual form of reproduction in farm animals,

41
including man. Nature has made this form of reproduction very fulfilling to both
sexes such that the process of perpetuating the species becomes a very pleasant
experience.

The Female Reproductive System

The principal sex organs of the female are the ovaries. The ovaries have two
main functions: (1) the production of sex cell or ovum; and (2) the production of
female sex hormone – estrogen. The female reproductive system includes the pair of
ovaries and the accessory reproductive tract.

The ovaries are almond-shaped bodies attached by the broad ligament to the
dorsal wall in the sublumbar region of the body cavity. The outer layer (cortex) of the
ovary is made up of germinal epithelium with a very large number of primary
follicles each of which contains blood vessels, nerves, ganglion cells, stroma and
embryonic vestiges.

The accessory reproductive tract includes the infundibulum, oviducts, uterus

(horn and body), cervix, vagina and vulva (Figure 8). The infundibulum is a funnel-
shaped structure which picks up the egg when released by the ovary. The “picking-
up” of the egg is believed to evolve the active participation of the celia-like structures
at the rim of the infundibular funnel which attract the eggs into the infundibulum.

The oviduct is a tubular structure connecting the infundibulum to the horn of

the uterus. It serves as the passage way of the egg on its way to the uterus. It is the
site of fertilization and the beginning of embryonic development after fertilization.

The horn of the uterus is the organ which serves as the site of implantation for
the fertilized egg. This is where the fetus would develop during the stage of
pregnancy in gestating animals. The body of the uterus unites the two horns of the
uterus and connects them to the cervix.

The cervix is sometimes considered as the neck of the uterus. Its opening, os
uteri, closes when the animal gets pregnant to protect the uterine contents. It serves
as sperm receptacle in certain animals.

The vagina is the primary organ of copulation. It serves as the receptacle of

the sperm cells in many species. It also comprises a part of the birth canal of the
animal at parturition.

The vulva is a common passage way for the products of reproduction and for
urine. The vulva of mammals is comparable to the ventral portion of the cloaca of
birds. It is also homologous with the scrotum of the male, since both are derived
from the same embryological structure.

42
Figure 8

43
 The clitoris is a rudimentary organ located in the ventral commissure of the vulva
in mammals. It is homologous to the glans penis of the male.

The broad ligament suspends the female genital system from the dorsolateral
wall of the pelvic canal. Three specialized regions of the broad ligament are
recognized: (1) mesometrium is that portion of the broad ligament which suspends
the anterior portion of the vagina, the cervix and the uterus and comprises the major
portion of the broad ligament; (2) mesosalpinx is a lateral fold on the anterior portion
of the broad ligament which suspends the oviduct (Salpinx); and (3) mesovarium is a
specialized portion of the anterior edge of the broad ligament which suspend the
ovary proper. In some species a pocket-like structure called Bursa is formed by
fusion of two or more of the mesovarium, mesosalpinx and the infundibulum. It
encloses, more or less completely the ovary.

A blind pouch opening into the floor of the genital tract at the external
urethral opening of the junction between the vagina and the vulva is the sub-urethral
diverticulum. The neck of the bladder opens through the roof of the sub-urethral
diverticulum. This relationship of the neck of the bladder to the blind pouch and
hence to the external urethral opening would appear to be a safety feature to prevent
entrance of foreign objects into the urinary bladder proper.

The Male Reproductive System

The main sex organ of the male is the testis. In birds, the two testes are
located within the body cavity in the dorsolumbar region. In farm animals, the testes
are located outside the body cavity within the scrotum. The scrotum is the cutaneous
sac that serves as the external covering of the testes. It protects the testes from direct
mechanical injuries and provides an environment which is a few degrees (6-9 oF)
cooler than the body temperature which is required for normal spermatogenesis. The
thermoregulatory muscles of the testes are the cremaster muscle and the dartos
muscle.

During fetal development, the initial development of the testes starts inside
the body cavity. As the fetus grows, the testes start to descent to the scrotum through
the inguinal canal. In certain instances, when both testes failed to descent to the
scrotal sacs, the individual is said to be a bilateral cryptorchid, hence is sterile. If
only one of the testes failed to descend to the scrotal sac, the individual is said to be a
unilateral cryptorchid, but capable of fertilization. However, cryptorchid animals
should be culled and not allowed to mate because this condition is heritable.

The testes have two main functions: (1) production of sperm cells; and (2)
production of sex hormone – testosterone. Testosterone is the male sex hormone
responsible for the development of the secondary sex characters. Male characteristics
such as muscular development at the rear quarters and shoulders, aggressiveness and

44
libido when confronted with an in heat female animal of the same species are
governed or influenced by the hormone testosterone.

The seat of spermatozoa production in a testis is the seminiferous tubules

(Figure 9). The seminiferous tubules join together to form the rete testis and come out
of the testis as vasa efferentia. The vasa efferentia converged to form the head of
epididymis, then the body and tail of the epididymis. The convoluted epididymis
straightens up to form the vas deferens which then enters the inguinal canal and
enlarges to form the ampulla. The ampulla joins with the urethra of the penis. The
penis is the male organ of copulation and serves to introduce the spermatozoa into the
female reproductive tract.

There are three accessory glands which contribute to the bulk of the semen
ejaculate; these are the (1) seminal vesicles, (2) prostate gland, and (3) Cowper’s
glands or bulbo-urethral gland.

Semen consists of the sperm cells plus the secretions of the three accessory
glands. In vasectomized animals, the vas deferens are severed, thus, the ejaculate
consists only of the secretions of the accessory glands – making the vasectomized
male sterile but without losing libido. On the other hand, in castrated animals both
testes are removed, thus, rendering the male sterile with loss of libido.

The secretions of the accessory glands serve as the vehicle for the transport of
the spermatozoa from the vagina to the oviduct. It stimulates also spermatozoa
activity and served as the lubricating substance during copulation particularly the
secretions of the bulbo-urethral gland. The seat of spermatogenesis is the
seminiferous tubules. FSH stimulates the germinal epithelium lining the seminiferous
tubules to initiate spermatogenesis up to the secondary spermatocytes stage. On the
other hand, LH stimulates the interstitial cells or the cells of Leydig to secrete
testosterone; and testosterone is required for the final maturation of the spermatozoa.
Therefore, both FSH and LH are required for normal spermatogenesis. FSH has a
direct influence on spermatogenesis, whereas, the influence of LH on spermatozoa
production is through its stimulating effect on testosterone secretion, which is
required for the final stages of spermatogenesis. The level of testosterone in
circulation also serves as the negative feedback in controlling LH production by the
sertoli cells in the seminiferous inhibin, produced by the sertoli cells in the
seminiferous tubules, has a negative feedback effect on FSH secretion.

Testicular and epididymal sperm cells are non-motile. They become motile
only when they are suspended in a fluid and this occurs when they come in contact
with the secretion of the accessory glands.

The normal spermatozoa consists of a head, neck or mid-piece and a tail. The
head is covered by a protoplasmic cap (galea capitis) and the shape is flattened ovoid
in bull, ram, boar and rabbit and rounded in man. It varies with species like rat,
rooster and salamander.

45
Figure 9. The male reproductive system

46
 The mid-piece and the tail are composed of several strands or fibrils which
are covered by a sheath. At the tip of the tail, these fibrils flare out into a naked
brush.

The common abnormalities encountered in a semen sample are sperm cells

with protoplasmic droplets at the mid-piece; headless or tailless; giant and miniature
heads; bent, coiled and shoe-hooked tailed sperm cells. When sperm abnormalities
in a semen ejaculate are about 50% of the total sperm cell counts, the male is usually
sterile. Abnormal sperms are often seen in males suffering from fever, males used too
frequently for breeding, or too young males. In the same manner, if the number of
dead sperms in a semen ejaculate, as determined by a dead-alive staining technique, is
50% or more, the male has impaired fertility if not sterile.

The life-span of ejaculated spermatozoa in the female reproductive tract is

about 24 hrs (20-30 hrs) in most mammals and about 14 days in chickens. Of so
many million sperms per ejaculate, only a few thousand reach the oviduct and few
dozens actually reach the vicinity of the ovum. Several may penetrate the zona
pellucida, but only one sperm cell enters the ovum proper and accomplishes
fertilization.

Puberty and Estrous Cycle

The female reproductive tract starts to function when a female reaches the age
of puberty. Puberty indicates that the female has reached sexual maturity – capable
of producing offsprings. The age of puberty varies between breeds of animal and
among female animals of the same breed. The first manifestation to indicate that the
female animal has reached the age of puberty is when it starts to show signs of estrus.
In human, a girl does not show signs of heat, instead she shows or manifests signs of
menstruation to indicate that she has reached the age of puberty.

When the animal reaches puberty, the anterior pituitary gland starts to secrete
gonadotrophic hormones which could affect the ovaries. The first gonadotropin
secreted in significant amount is follicle stimulating hormone (FSH) with little
luteinizing hormone or LH. FSH causes the growth and development of the Graafian
follicle (GF) in the ovary. In turn, this developing follicle secretes a hormone known
as estrogen. This is the hormone which causes estrus in female. The presence of
estrogen stimulates the production of LH. At the peak of estrogen production LH
production increases which coincides with the production if inhibin hormone from the
ovary, which inhibits FSH production. Usually, LH is also produced with little FSH
because the cells in the anterior pituitary which secrete the former are the same cells
that secrete the former are the same cells that secrete the latter. These cells are the
basophils of the anterior pituitary.

LH is the hormone that causes ovulation of maturing follicles. It also initiates

the formation of corpus luteum (CL) by converting the cells of the stratum
granulosum into lutein cells. Eventually, upon the action of LH, what used to be the

47
GF will be filled up with lutein cells and becomes the corpus luteum or CL. So, the
GF with the action of LH becomes the CL.

The CL secretes a specific hormone known as progesterone. It has a strong

inhibitory effect on FSH production. As long as the CL is secreting progesterone,
estrus is inhibited. Progesterone prepares the endometrium of the uterus for
implantation of the fertilized egg. It also maintains normal pregnancy until birth.
Thus, CL also maintains normal pregnancy until birth. Thus, CL persists if there is
pregnancy; however, if there is no pregnancy, the CL will regress. It is now known
that the uterus secretes prostaglandin F2 alpha which could destroy the CL. In a
normal cycle or if there is no pregnancy, prostaglandin F2 alpha is secreted by the
uterus and causes the luteolysis or regression of the CL.

To summarize, in an estrous cycle, the sequence of events are as follows: the

anterior pituitary secretes FSH which causes the formation of GF in the ovary. In
turn, the GF secretes estrogen which stimulates LH production and at the same time
causes heat or estrus in the female animal. This is the best time to breed the animal.
Then, the surge of LH released by the anterior pituitary causes the ovulation of
maturing GF. At the same time, LH would initiate the formation of CL and causes the
production of progesterone by the CL. Progesterone would prepare the uterus for
implantation of the fertilized egg if fertilization had taken place. Also, progesterone
maintains pregnancy, but if the animal is not pregnant, the uterus then would produce
prostaglandin which causes the destruction of the CL, resulting in the cessation of
progesterone production. When this happens, FSH production would again continue
and a new cycle begins. The period from one estrus to the next estrus is known as the
estrous cycle. In many farm animals like carabao, cattle, pigs, and horses estrus
comes every 21 days if the female animal is cycling regularly, although it could vary
from 12 to 30 days.

Animals may be classified based on the occurrence of their estrus cycle as:
(1) Monoestrus, if the animal comes in heat only once a year, such as dogs; (2)
Seasonally polyestrus, if it comes in heat at certain seasons only, such as sheep; and
(3) Polyestrus, if it comes in heat all throughout the year, like cattle, swine and
carabao.

An estrous cycle may be divided into four portions:

(1) Proestrus which is characterized by follicular growth;

(2) Estrus which is under the influence of estrogen;
(3) Metestrus is characterized by the formation of CL; and
(4) Diestrus which is under the influence of progesterone secreted by the CL.

If you take rat vaginal smear and examined it under the microscope, you will
find nucleated epithelial cells during proestrus. During estrus, the vaginal smear will
show cornified cells; and during metestrus, you will find some leucocytes among

48
cornified cells. At diestrus, the leucocytes predominate among the nucleated
epithelial cells.

Animals may be classified also as spontaneous ovulators and induced

ovulators. Spontaneous ovulators are those that ovulate spontaneously during or at
around estrus, such as cattle, carabao, goat, sheep, hogs etc. Induced ovulators are
those animals that do not ovulate unless there is copulation, such as in rabbit.

Signs of Heat or Estrus

Estrus is perceived through physiological manifestations that the female may

show or exhibit. The manifestations of heat are as follows: (1) reddening and
swelling of the vulva; (2) mucus discharges from the vagina; (3) frequent urination;
(4) the animal becomes restless and lacks appetite; (5) mounting other animals in the
herd; and (6) the female stands still when mounted by the male.

The mucus discharge is watery at first and towards the end of estrus becomes
sticky. By taking a drop of the cervical mucus discharge and placing it on a glass
slide to dry, a ferning pattern will be observed under the microscope if the animal is
in heat. The vaginal lining of in heat animal is pinkish and it is pale when not in heat.
However, the best sign of heat is when the animal stands still when mounted by a
male animal of the same species.

When to Breed or Inseminate

In cattle and carabaos, ovulation takes place about 15-18 hours from the end
of estrus. The best time to inseminate a female animal is towards the end of its estrus.
But since duration of estrus is variable and no one can tell exactly when the animal
started its heat, it is then recommended that once the animal is observed to be in heat,
then inseminate it immediately. If the animal is observed to be in heat in the morning,
inseminate immediately and if it is still in heat in the afternoon, inseminate again.
Likewise, if it is observed to exhibit heat in the afternoon, inseminate immediately
and if it is still in heat the following morning, give the second insemination. This is
the thumb rule that should be followed. The duration of estrus in carabao lasts from 5
to 36 hours or an average of 18 hours (less than a day).

In swine, the duration of estrus is about 2-3 days. The best time to breed or
inseminate a sow is on the second and third day of estrus.

In mares (female horse that has already given birth), estrus duration is about
6-7 days. Mating or insemination is recommended on the fourth and fifth day of
estrus.

49
Fertilization and Pregnancy

During mating, the sperm cells are deposited in the vagina in most farm
animals. The sperm cells must reside in the female reproductive tract before
becoming capable of attaching to and penetrating the ovum. This process is known as
sperm capacitation and is believed to start in the uterus. However, the major site of
capacitation appears to be in the oviduct. Capacitation leads to acrosomal changes
needed for sperm penetration of the ovum; therefore, capacitation should function to
prevent pre-mature acrosome activation until the spermatozoa has reached the site of
fertilization and can be in contact with the ovum. Fusion and vesiculation of the
acrosome release hydrolytic enzymes, e. g., hyaluronidase and acrosin which are
required in penetration of the ovum. Thus, it is important that mating or insemination
should be made at the time that ovulation would likely take place, or as close as
possible to expected ovulation time. Regardless of the timing of ovulation, high
conception rates result if spermatozoa are present in the oviduct shortly before
ovulation. Insemination too early reduces conception rate due to the loss of sperm
viability and the number of sperm at the site of fertilization, while loss of ovum
viability can result from insemination after ovulation even though fertilization occurs.

Fertilization in mammals requires three critical events: (1) sperm migration

from the site of deposition to the oviduct and convergence of the spermatozoa
towards the ovum; (2) sperm attachment and penetration through the zona pellucida;
and (3) fusion of sperm and ovum plasma membranes.

Attachment of the sperm head to zona pellucida appears to be regulated by

receptors sites on the zona surface. Evidence that sperm receptor site are present on
the zona are supported by species specificity of sperm attachment to zona surface. A
sticky substance from the equatorial segment of the acrosome of hamster sperm cell
appears to be responsible for binding of the sperm head to the zona surface.

Penetration of the zona by the spermatozoa occurs within 5-15 minutes after
sperm attachment. The acrosome reaction may occur before or after attachment of
the sperm head to the glycoprotein receptors on the zona, but acrosomal activation is
essential for sperm penetration of the zona. The mammalian acrosome contains
enzymes such as hyaluronidase and acrosin which digest a pathway through the zona
pellucida through which the sperm cell can penetrate the ovum. Once the sperm has
traversed the zona pellucida, the head moves into the ovum and comes in contact with
the vitelline membrane covering the ovum. The activated ovum completes meiosis
and expells the first and/or second polar body. The remaining maternal haploid
chromosome are then enclosed by a pronuclear envelope forming the female
pronucleus. Subsequently, the surface of the sperm head is incorporated into the
plasma membrane of the ovum. The sperm plasma membrane becomes intermixed
with the ovum plasma membrane effecting fertilization. The ovum surface changes,
forming a fertilization membrane, to prevent fusion of additional spermatozoa. The
male and female chromosome aggregate to form the first cleavage division, resulting
the formation of a zygote and restoration of the diploid chromosome number. Thus,

50
the process of fertilization allows combining paternal and maternal hereditary
elements.

The cleavage of the fertilized egg continues to progress as the zygote travels
from the oviduct to the horn of the uterus where implantation takes place. Cleavage
divisions are always mitotic with each daughter cell (blastomeres) receiving the full
compliment of chromosomes. The blastomeres from 2- to 8-cell stage are capable of
giving rise to an intact embryo, e.g., complete cleavage of 2-cell stage can give rise to
twins and that of 4-cell stage can give rise to quadruplets. After the 8-cell stage,
however, the blastomeres appear to differentiate according to their position in the
morula. When the embryo has developed into the 8- to 16-cell stage, it is transported
to the uterus where it continues to proliferate.

Shedding off the zona pellucida is followed by a rapid phase of blastocyst

development and growth. This occurs on day 11 postestrus in the sheep and pig and
day 13 in the cow (Table 8). The blastocyst undergoes a logarithmic elongation
phase. For example, cow blastocyst transforms from a 3-mm spherical shape on
approximately day 13, to a 25-cm filamentous threadlike form, on day 17. Initial
placentation follows after the stage of blastocyst elongation.

In polytocous species such as pig, the embryos undergo uterine migration and
equidistant spacing prior to implantation. The pig blastocyst begins to attach to the
uterine surface on day 13 with attachment completed across the trophoblastic surface
between days 18 to 24. The trophoblastic surface is modified to form specialized
absorptive structures called areolae that allow nutrient uptake by the developing
conceptus.

In ruminants, like cattle and sheep, the placental attachment is through the
caruncles or cotyledons. These are finger-like villi or pappillae that allow nutrient
uptake by the developing conceptus. Initial placentation occurs at day 22 in cattle
and day 15 in sheep (Table 8).

In the mare, implantation or attachment does not occur until days 24-40. Early
attachment is through interdigestation between surface epithelium of the embryonic
vesicle and uterine lining. Specialized chorionic girdle cells form around the
spherical vesicle, detach around day 38 and invade the uterine endometrium to form
the endometrial cups that produce equine chorionic gonadotrophin.

51
Table 8. Time of events in early embryonic development
===============================================================
Species
________________________________________________
Parameter Cattle Horse Sheep Swine
________________________________________________________________________
Gamete Longevity (hours)
Sperm 30-48 72-120 30-48 34 -72
Ovum 20-24 6- 8 16-24 8- 10

Embryonic Development (days)*

2-cell 1 1 1 .6 - .8
4-cell 1.5 1.5 1.3 1
8-cell 3 3 1.5 2.5
Blastocyst 7-8 6 6-7 5-6
Hatching 9-11 8 7-8 6

Blastocyst transport to uterus

Hours 72-84 140-144+ 66-72 46-48
Cell stage 8-16 Blastocyst 8-16 4
Blastocyst Elongation (days) 13-21 NE++ 11-16 11-15
Initial Placentation (days) 22 37 15 13
Gestation period (days) 278-290 335-345 145-155 112-115
===============================================================
*Days after ovulation
+Unfertilized ova remain in the oviduct
++No elongation occurs to form filamentous blastocysts

52
Maternal Recognition of Pregnancy

Implantation allows the conceptus and uterine endometrium to achieve intimate

contact for nutrient exchange and endocrine communication. At appropriate time, the
conceptus must produce steroid hormones and/or proteins to signal its presence to the
maternal system. This signal is necessary for corpus luteum maintenance, production of
progesterone and continued endometrial development and secretory activity. If the
conceptus falls to signal its presence of exactly the correct time, the function of the
corpus luteum is terminated by the luteolytic action of prostaglandin secreted by the
uterus. This ensures that the female will return to estrus and mate at frequent intervals
until a successful pregnancy is established. Uterine prostaglandin is produced by
endometrium of cows, ewes, mares and sows and causes morphologic regression of the
corpus luteum and cessation of progesterone production. The luteolytic potential of
endometrial progesterone must be blocked if pregnancy is to be established since a
functional corpus luteum is essential for the initiation and maintenance of pregnancy in
all farm animals. Secretion of LH from the anterior pituitary is also essential for corpus
luteum maintenance and function during pregnancy.

Placental Development

A unique feature of early mammalian development is the provision of nutrients

from the maternal organism by way of the placenta. The placenta is an opposition or
fusion of the fetal membranes to the endometrium of the uterus to permit physiologic
exchange between fetus and mother. The placenta differs from other organs in many
respects. It originates as a result of various degrees of fetal-maternal interactions and is
connected to the embryo by a cord of blood vessels. The size and functions of the
placenta change continuously during the course of pregnancy, and the organ is eventually
expelled. For the fetus, the placenta combines in one organ many function activities that
are separate in the adult.

From the side of the embryo a fold grows up and over it, fusing at the top and
ultimately enclosing the embryo in a double-layered sac which is known as the amnion
(water bag). This is filled with a clear watery fluid in which the embryo is suspended.
Its purpose is to form a protective cushion against external shocks and pressure of the
adjacent body organs and to prevent adhesions between the surface of the embryo and the
surrounding membrane. At parturition, the amnion acts as a wedge to dilate the cervix.

The allantois is formed as an outpouching of the hind gut of the digestive tract
and functions as the urinary receptacle for the embryo, and also collects some solid
waste. The chorion, the outer membrane, completely surrounds the embryo, amnion and
allantoic cavity. It is rich in blood vessels and lies appositon to the uterine mucosa. The
allantois which fuses with the chorion becomes the fetal placenta - chorigallantois
placentation - characteristics of all farm animals. Nutrients from maternal circulation are
acquired by the fetus by simple transfusion. The concentration of nutrients is not
necessarily the same in both blood. Nutrients also pass from the dam to fetus in utero by

53
ingestion through mouth from the amniotic fluid, and by occasional passage of blood
cells from dam to fetus and vice versa.

There are two general types of placentation among farm animals. The sow and
mare have a diffuse placenta, which consists of a simple apposition of fetal and maternal
epithelia, although in the mare, the complex folding and branching of the two surfaces
give rise to the formation of microcotyledons between 75 and 110 days of gestation.

The sheep, goat, cows and carabao have cotyledonary placenta. The cotyledons
from the fetal placenta are attached to the caruncles of the maternal placenta through
which uterine blood flows. A caruncle with the attached cotyledon is called placentome.
At parturition, the chorionic villi of both types of placentation are merely withdrawn and
there is no extensive destruction of the uterine tissue.

In summary, the fertilized egg reaches the uterus through the oviduct as blastocyst
and remain their for a while bathed in uterine secretion. At the same time, changes
occurs in the uterus itself. the process of implantation follows; trophoblastic cells of the
blastocyst attached themselves between the epithelial cell of the uterus. Nutrition of the
embryo with the uterine wall forming a sort of placenta. The second stage in the
development of the zygote is the embryonic period which extends from the time of the
attachment of the egg to the wall of the uterus until the form of new individual is laid
down. Early in this period, the extra embryonic membrane starts to developed which are
the allantois, amnion and the chorion. The embryo is enclosed in two fluid filled sacs.
The last stage of development is the fetal period which extends from the end of
embryonic period to the time of birth.

Parturition and Lactation

Parturition is defined as the physiologic process by which the pregnant uterus

delivers the fetus and placenta from the maternal organism.

Most signs of approaching parturition relate to changes in the pelvic ligaments,

enlargement and edema of the vulva, and mammary activity. The signs are useful as a
guide but they are too variable for an accurate prediction of the date of parturition.
Obvious enlargement of the mammary gland occurs in all farm species. The teats
becomes swollen and secretions may escape through the teat meatus. Waxing occurs in
most mares between 6-24 hours before parturition and is replaced by drips of milk 12-24
hours later. Thus, presence of milk in the mammary gland is a strong indication of
approaching parturition.

Parturition is triggered by the fetus and is completed by complex interaction of

endocrine, neural and mechanical factors, but their precise roles and interrelationships are
not fully understood. Sequence of events leading to uterine contractions is triggered by
hormonal changes and myometrial stretch. An increase in estrogen, decrease in
progesterone and an increase in oxytocin plus myometrial stretch due to fetus
development can stimulate the production of prostaglandin from the maternal

54
myometrium. The elevated prostaglandin lowers cyclic AMP and the threshold of the
uterine muscle to respond to oxytocin in initiating uterine contraction (Figure 24).
Myometrial contraction of low amplitude and frequency occur during the major part of
gestation, and at the onset of parturition, these are replaced by the expulsive form
characteristic of delivery.

At the time of parturition, fetal cortisol production increases which stimulates the
placenta to convert progesterone to estrogen, thus, descreasing progesterone and
increasing estrogen levels both at the placenta and maternal circulation. Estrogen
stimulates prostaglandin synthesis from the placental membrane and myometrium.
Elevation of prostaglandin lowers the threshold to oxytocin, and oxytocin and/or
prostaglandin lead to myometrical contractility. Also, prostaglandin destroys the CL
thus, cutting off progesterone secretion from the CL. It also stimulates the ovary to
produce relaxin which softens the cervix and the pelvic bones (Figure 25).

Parturition may be divided into three stages: (1) the dilation of the cervix, (2)
expulsion of the fetus, and (3) expulsion of the placenta (Table 9). During the first stage,
uterine contractions are painful causing restlessness and signs of abdominal discomfort.
At the fetus progresses through the cervix, the allantochorion ruptures, releasing the
urine-like fluid that marks the end of the first stage of labor.

The distention of the cervix and vagina by the conceptus initiates the
neurohumoral reflex, which produces the expulsive force of abdominal muscular
contractions (straining) and the release of oxytocin, which in turn accentuates myometrial
contractions. The combined forces of intra-abdominal and intra-uterine pressure mark
the beginning of the second stage of labor. Straining consists of a few contractions
followed by a few minutes of rest. The fetus enclosed in the amnion is propelled through
the birth canal and appears at the vulva. As straining continues, the amnion ruptures.
The greatest effort is associated with the emergence of the head, forelegs and chest. All
farm species assume lateral recombency with limbs extended during delivery. The
umbilical cord breaks as the neonate or the dam moves.

Rhythmic uterine contractions originating at the apex of the uterine horn continue
after birth (third stage) and cause the inversion of the chorioallantois in ruminants. The
presence of the detached placenta within the birth canal then initiates further straining
and expulsion of the placenta. The expulsion of the placenta is rapid in the mare but is
slower is ruminants due to the cotyledonary type of placentation. The placentas of
adjacent piglets are usually fused and often expelled as one or more masses interspersed
with the birth of piglets. The largest mass of placenta, however, is usually expelled 3 to 4
hours after the delivery of the last piglet.

The onset of milk secretion follows parturition. In preparing the mammary gland
for lactation, hormonal interactions were needed even long before fertilization. It starts
with the development of the duct and the lobulo alveolar systems of the mammary gland
when the animal reached the age of puberty. The cyclic production of estrogen and
progesterone during the estrous cycle promotes the development of the mammary gland.

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Table 9. Stage of labor and related events in farm animals

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Stage of Mechanical
Labor Forces Period Related Events
______________________________________________________________________________

I Regular uterine Beginning of uterine Maternal restlessness, elevated

Dilation contractions contractions until cervix pulse and respiratory rates.
of Cervix is fully dilated and Changes in fetal position and
continuous with vagina posture.

II Strong uterine and From complete cervical Maternal recumbency and

Expulsion abdominal dilation to end of straining. Rupture of allanto-
of fetus* contractions delivery of fetus chorion and escape of fluid
from vulva. Appearance of
amnion (waterbag) at vulva.
Rupture of amnion and
delivery of fetus.

III Uterine Following delivery of

Maternal straining ceases.
Expulsion contractions fetus to expulsion of
Loosening of chorionic villi
of placenta decrease in placenta from maternal crypts.
amplitude Inversion of chorioallantois.
Straining and expulsion of
fetal membranes.
===============================================================

*In polytocous species (sow) and twin-bearing species (sheep and goat), cannot be
separated from the third stage.

Estrogen stimulates the development of the duct system; and progesterone stimulates the
development of the lobulo-alveolar system. With the onset of pregnancy, the sustained
production of progesterone by the CL and the placenta to maintain pregnancy until birth,
also maximize the development of the secretory units (alveoli) of the mammary gland.
Towards the end of pregnancy, there is a major surge in secretion of prolactin
immediately before parturition. Prolactin initiates milk secretion in the mammary gland.
Thus, when the fetus is born, the mammary gland is also ready to produce milk needed
for the survival of the newborn animal.

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