Mammary gland

Mammary gland is a cutaneous gland. It is a compound tubule-alveolar type in advanced

mammals that originates from ectoderm. There is wide species variation in the appearance of the
mammary gland.

Functional organization and anatomical structure of mammary gland

a) External structure of mammary gland

The mammary gland of cattle, goats, mare and whales are located in inguinal region, of primates
and elephants in thoracic region and those of pigs, rodents, bitch and carnivores are located along
the ventral surface of both thoracic and abdominal region.

The mammary gland and teats together are called ‘udder’. A cow’s udder is having two halves
and each half has two teats. Each teat is joined with separate gland; called quarter and each
quarter are separated by connective tissues. This means cows has four glands or quarters having
four teats. In addition to four normal teats in cow, there may be supernumery teats having non-
secretory functions. Supernumery teats are also found in sheep, goat, swine and mare. Sheep and
goat has two glands and teats. In these species, each teat has streak canal. But in pigs and horses,
each teat has two streak canals and in rodents, carnivores, primates, each teat has numbers of
streak canals that ranges from 10-20.

In male, except stallion, rudimentary teats are usually found. The weight of lactating bovine
udder ranges from 14-32kg.

b) Internal structure of mammary gland

The two halves of the bovine udder are separated by median suspensory ligament. The posterior
extremity of this ligament is attached to prepubic tendon. The bovine teat is opened to exterior
through a canal called Galactophore. The bovine teat has a small cistern called ‘Teat cistern’ and
is continued to teat canal called ‘streak canal’ that opens to outide through Galactophore.
Radiating upward from the streak canal is structure called ‘Furstenberg’s rosette’, which is
composed of 7- folds of mucus membrane, which helps to retain milk in the teat. However, in
cattle, the primary structure responsible for retention of milk is ‘sphincter muscle’ surrounding
streak canal. Above each teat, there is ‘gland cistern’ to which teat cisterns are joined. Again,
numerous large ducts of the quarter are attached with gland cistern. These ducts branch
profusely, ultimately ending in the secretory units called alveoli or acini. The milk is formed in
the epithelial cells of alveoli. The alveoli are grouped together in units called lobules, each of
which is surrounded by distinct connective tissue septum. Lobules in turns are grouped into
larger units called lobes.
The alveoli are surrounded by myoepithelial cells that are involved in the milk ejection (milk
letdown) reflex. The nature of epithelial cells which surrounds udder and teat outside to inside
are:

• Skin epithelial of udder-stratified squamous

• Streak canal epithelium- Transitional epithelium

• Teat and gland cisterns-two layers epithelium-one layer of cuboidal and another
cylindrical epithelium.

Growth and development of mammary gland

a) Prepubertal development- At birth bovine udder has distinct teat and gland cistern but
original ducts are developed to less extent. The muscular and lymphatic systems are developed.
From birth to before puberty, the mammary gland undergoes little development, It increases in
size due to increase in connective tissues and fat and extension of duct system.

b) Growth during puberty and estrus cycle- At puberty and estrus cycle, growth of duct
during follicular phase and lobulo-alveolar development during luteal phase occurs.

c) Development during pregnancy- During pregnancy, mammary glands develop rapidly.

Glandular tissues spread by fifth month and lobules are formed.

The development of duct system and lobulo-alveolar development is completed at the time of
parturition in mammals.

Hormonal control of mammary gland

There are numbers of hormones which are responsible for the growth and development of
mammary gland (mammogenesis) either directly or indirectly which are as follows.

1) Pituitary Gonadotropins

Mainly mammary growth is under the control of estrogen and progesterone. FSH causes
development of follicles and these follicles are source of estrogen and LH causes ovulation and
forms CL which is the source of progesterone. Estrogen alone causes duct development and
estrogen and progesterone together cause lobulo-alveolar growth. However, there is some
species variation;

• In bitch estrogen alone cannot cause of growth of duct system.

• In goat and sheep estrogen alone can cause lobulo-alveolar development.

2) Role of anterior pituitary hormones

It has been known that anterior pituitary hormones are essential for optimum mammary growth.
They are- GH, ACTH, TSH, Prolactin.

3) Role of adrenal gland hormones

The development of mammary glands by estrogen and progesterone are stimulated further by the
presence of adrenocorticotophic hormones. The adrenal steroids play an important role in the
regulation of mammary gland development.

4) Role of placenta

The mammary gland shows remarkable developmental changes during pregnancy and placenta
partly plays the functional role in hypophysis and ovary enhancing the development of mammary
gland.

Lactogenesis

The process by which the alveolar cells of the mammary gland are tuned with the ability to
secrete milk constituents is known as lactogenesis. There different stages involved in it and they
are;

In the first stage, there occurs an enzymatic and cytologic change of alveolar epithelial cells and
a limited secretion of milk before parturition occurs. This is followed by second stage in which
all milk components are secreted just immediately before parturition and the process extends for
several days period. The onset of milk secretion after parturition occurs to supply nutrients to
new born which is a third stage. This onset of milk secretion is under the control of Prolactin,
Glucocorticoids and sudden fall in Progesterone level at parturition. Placental lactogen is slightly
responsible for this.

Galactopoiesis

It is the process of maintenance of milk secretion and it depends on alveolar cell number, their
synthetic ability and let down of milk. Periodic removal of milk by hand or suckling by calf is
important otherwise normal hormones production responsible for Galactopoiesis, ceases. There
are numbers of hormones responsible for this which are-Oxytocin, ACTH, GH, Glucocorticoids,
TSH, Prolactin, Insulin etc.

Let down of milk

Milk let down (ejection) is a natural process used by the cow to help remove milk from the
udder. Milk is initially secreted into small sacs within the mammary gland called alveoli, from
which it must be ejected for consumption or harvesting. Mammary alveoli are surrounded by
smooth muscle (myoepithelial) cells which are a prominent target cell for oxytocin. Oxytocin
stimulates contraction of myoepithelial cells, causing milk to be ejected into the ducts and
cisterns above the teat. Oxytocin reaches the udder where it causes the myoepithelial cells
around the alveoli to contract. Upon contraction of the myoepithelial cells, the alveoli collapse
and milk is squeezed out into the small ducts. These ducts shorten and widen, and the milk
rushes through into the gland cistern. Only alveolar and ductal milk is expelled by the action of
oxytocin on the myoepithelium. There is no contraction of large ducts or cisterns.

The pressure of milk being forced into the ducts/cistern and down towards the teat causes the teat
to swell with milk and become ‘plump’. It takes 60 - 90 seconds for teats to become plump after
let down has been initiated. Cows with well-filled udders require a shorter period of stimulation
to elicit milk let down response than cows with less-filled udders. The action of oxytocin is
essential for emptying of the udder during milking. The action of oxytocin only lasts for 6 to 8
minutes, because its concentration in the blood decreases rapidly.

Mechanical stimulation of peripheral neural receptors in skin of teat (during milking/suckling)→

neural stimulus. ↓

Stimulus travels from teat to spinal cord and to the supraoptic and paraventricular nuclei of
hypothalamus. ↓

Synthesize oxytocin hormone →release through posterior pituitary.

Oxytocin-released to b/d-transported to target t/s (myoepithelial cells of mammary gland).

Contraction of gland and ejection or let down of milk.

Factors affecting the let down of milk

1. Inadequate udder preparation

2. Delayed attachment of teat-cups for minute after preparing the udder or failure of milking
equipment to operate properly.

3. External stimuli: The milk let-down reflex of cows is very easily retarded or prevented by
external stimuli. It is often said that a cow "holds up" her milk. This is not possible since
the cow has no voluntary control over the let-down process, and she cannot thus put the
process into reverse.

 Milking routine: feeding, barn noises, irregular time of milking, method of

milking and the sight of the calf.

 Emotionally disturbing stimuli: dog barking, outer loud and unusual noises
  Stressors: death of the calves, loud and various unfamiliar noises, visit of
unfamiliar persons in the milking parlor or at the site of milking, change of
milker and beating or rough handling at the time of milking especially in
first calvers. The basal concentrations of cortisol and β-endorphin were
higher in the cows milked in unfamiliar surroundings compared to when
milked in familiar.

Such stimuli cause the brain to release adrenalin. This hormone works against oxytocin by:

• Blocking oxytocin release from the brain.

• Constricting blood vessels and preventing oxytocin from reaching the udder.

• Directly counteracting the effect of oxytocin on the contraction of myo-epithelial cells.

If adrenalin release occurs before the milk-ejection stimulus, the ejection will be virtually
completely blocked. When adrenalin is released after milk ejection has commenced, it will
result in large amounts of milk being retained in the udder with associated negative effects on
milk yield.

4. Udder Edema: This presents itself as a failure of milk let down associated with excess
fluid in the mammary tissues. It is characterized by a clinically normal animal with no fever
or loss of appetite. The distinguishing features are a firmness of all the glands, discomfort on
high pressure but no actual pain. The edema or fluid can be both in the skin and deep in the
udder tissue.

5. Mastitis: It may causes the painful let down of milk.

6. Disturbance of milk removal has been observed in primiparous cows immediately after
parturition and during estrus.

Composition of Milk
Milk is considered to be almost a perfect food for consumption. From the chemist point of view,
milk can be defined as a complete food consisting of fat, proteins, lactose and minerals and
water, as major constituents, and vitamins, enzymes, pigments, gases and several other organic
and inorganic substances.
Role of Constituents of Milk
As already mentioned, milk constituents are divided under two groups: major and minor. Their
role in the milk is described as follows:

A. Major constituents
The following are the major nutrient constituents in the milk that playa significant role in the
diet.

1. Water

The water in the milk is the same as any other water. Its function is to hold the solids of the milk
partly in solution and partly in suspension.

2. Milk fat

The fat exists in milk as an emulsion. When a milk droplet is examined under a microscope,
butterfat globules are visible in the form of small spherical myriads. The size of these globules
varies from 0.1 to 2 microns, the average being 3 microns. The butterfat globule is surrounded by
a thin membrane. This membrane is important in preventing the butterfat from becoming free oil
when it is heated. The membrane may also serve as some protection against the action of
enzymes, notably lipase, in causing butterfat to become rancid. The size and arrangement of the
globules affect the creaming ability of milk, the viscosity of cream, the case of whipping cream
and churning of cream to butter. Fat is a mixture of the glycerides of certain fatty acids. Milk fat
contains 19 or more fatty acid. These fatty acids may be classified as soluble and the insoluble,
volatile and the non-volatile, saturated and the unsaturated. Milk fat is the lightest constituent of
milk. It is because of this fact that the cream, which contains most of the milk fat in milk, rises to
the top to form the cream layer. Milk fat is very rich source of energy and also a good source of
vitamin A and D. Milk fat is one of the principal constituents of almost all dairy products, such
as feed, cheese, ice-cream, etc.

3. Proteins

The three principal classes of proteins of milk are casein, lactalbumin and lactoglobulin. The
caseins account for 80 per cent of total protein content, the lactalbumin and lactoglobulin (whey
proteins) almost 20 per cent.

a. Casein

Casein is a phosphor-protein which is found in no other product than milk. It exists in milk in
combination with calcium, hence is often spoken as calcium caseinate complex. It is present in
milk in a finely divided suspension. Such suspension is known as "Colloidal" dispersion. The
casein is precipitated by acids at its iso-electric point of pH 4.6. It is precipitated out of milk,
when the milk source or when acid or rennet is added to it. Because casein is a protein, it
supplies the body with muscle-building material. It is also used as a source of protein in many of
the imitation (artificial) milk products in the form of sodium caseinate.

b. Whey proteins .

The proteins lactalbumin and lactoglobulin are present in small amounts, dispersed in the water
of milk. The so called whey protein that remains in solution or suspension after the precipitation
of the casein by acid or rennet may be salted out by various combinations of inorganic salts.

Usually the first milk secreted after calving (colostrum) contains sixteen times more whey
proteins than the normal milk. This amount decreases rapidly to normal during the first few
milkings.

4. Lactose

Lactose, a carbohydrate found only in milk, it is synthesized in mammary gland. It is it

disaccharides composed of glucose and galactose, which are liberated when lactose is
hydrolyzed either by the enzyme lactase of the digestive tract or by lactase of bacterial origin.

Lactose serves the body as a source of energy, but is not as rich as butterfat in this respect lactose
is easily changed by bacterial action to lactic acid, a change which causes milk to taste sour. In
some dairy products, such as soft cheese butter- milk and butter, the characteristic flavour is
partly due to the acid fermentation. Since lactose is not very soluble, it some- times separates or
"crystalizes" in such dairy products as sweetened condensed milk and ice-cream, producing
"sandy texture" condition. Lactose has those nutritive properties which are not possessed by any
other sugar.

5. Mineral constituents

The mineral constituents or salts of milk constitute 0.6 to 0.8% of the total weight. The
potassium and chlorides of milk are in solution, whereas the calcium, phosphorus and
magnesium are partly in solution and partly in colloidal suspension. Because the metallic
elements are in excess as compared to the non-metallic elements, the minera1 content is alkaline
in character. Minerals of milk are very essential as a food for the young, as they help to build
bone and promote the proper development of the teeth. Although the average mineral content in
milk is 0.7%, the presence of this constituent answers one of the principal reasons why milk is
such an important food, especially for children and expectant mothers. Milk contains minerals
mainly as inorganic salts, although some of them are in organic combination. Numerous
elements in trace quantities are also found.

B. Minor constituents
The important minor elements of milk are pigments, cholesterol, phospholipids, enzymes, etc.
They are discussed here in detail.

1. Pigments

Milk contains two fat soluble pigments, "carotene" and "xanthophyll" and one water soluble
pigment, "lactoflavin" (lactochrome). Carotene is mainly responsible for the yellow colour of
cow milk. The amount of the water soluble pigment (lactoflavin) in milk varies with the breed
and the individual cow or buffalo.

2. Cholesterol

The amount of cholesterol in milk varies directly with the butter fat content of the milk.
Cholesterol is a yellow-white fatty chemical compound found in such foods as eggs, meat and
dairy products.

3. Phospholipids

The principal phospholids found in milk is called "lecithin". Lecithin is a fat-like substance
containing nitrogen and phosphorus. The lecithin is closely associated with milk fat, and when
milk is separated, some of the lecithin remains in the butter, though most of it passes into butter
milk.

4. Enzymes

Enzymes are complex organic compounds secreted by living plant and animal cells which
initiate chemical reactions but emerge unchanged. Enzymes are specific in their action. Lipase,
for example, acts only upon fat and proteins act only upon protein and its derivatives. They are
relatively unstable. High temperatures, unfavourable pH, light and certain chemicals tend to
inactivate them. The following are the principal enzymes in milk.

a. Catalase

This enzyme splits hydrogen peroxide into water. Temperature of 65° C to 67° C for 30 minutes
destroys this enzyme.

b. Amylase

Amylase, a starch splitting enzyme, is considered to be the least variable in quantity of all the
enzymes in normal milk.

c. Phosphatase

It is always present in milk. Its concentration is fairly constant. It has the ability to free phenel
from alcoholic esters of phosphoric acid.

d. Lipase

It is an enzyme in milk which breaks down milk fat, thus causing a strong or "rancid" flavour.

e. Galactase

Galactase slowly reduces milk protein to simple compounds; hence it plays a part in the
"ripening" of some cheeses.

f. Lactase

Lactase hydrolyses lactose into simpler sugar like glucose and galactose.

5. Gases

Milk contains some gases such as carbon-dioxide, oxygen and nitrogen. Carbon-dioxide enters
the milk in the cow's buffalo's udder, while oxygen, nitrogen and other gases get into the milk
during milking. Gases also may be formed by the growth of certain bacteria. The gases in milk
have no special significance, they slowly pass from the milk as it stands and almost entirely
escape during pasteurisation.

6. Nitrogenous substances

Other nitrogen containing compound besides the proteins, are found in milk in very small
quantities. These are fibrin, mucoid protein and an alcohol soluble protein. Other nitrogen
containing substances which are not proteins but are found on milk in very small amounts are
uric acid, urea, creation, creatinine and traces of some amino acids.