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 cows udder is having two halves and each half has two teats. ach teat is !oined with separate gland" called #uarter and each #uarter are separated by connective tissues. This means cows has four glands or #uarters having four teats. In addition to four normal teats in cow, there may be supernumery teats having nonsecretory functions. $upernumery teats are also found in sheep, goat, swine and mare. $heep and goat has two glands and teats. In these species, each teat has strea% canal. &ut in pigs and horses, each teat has two strea% canals and in rodents, carnivores, primates, each teat has numbers of strea% canals that ranges from '(-)(. In male, e*cept stallion, rudimentary teats are usually found. The weight of lactating bovine udder ranges from '+-,)%g. b) Internal structure of mammary gland The two halves of the bovine udder are separated by median suspensory ligament. The posterior e*tremity of this ligament is attached to prepubic tendon. The bovine teat is opened to e*terior through a canal called -alactophore. The bovine teat has a small cistern called Teat cistern and is continued to teat canal called strea% canal that opens to outide through -alactophore. .adiating upward from the strea% canal is structure called /urstenbergs rosette, which is composed of 0- folds of mucus membrane, which helps to retain mil% in the teat. 1owever, in cattle, the primary structure responsible for retention of mil% is sphincter muscle surrounding strea% canal. Above each teat, there is gland cistern to which teat cisterns are !oined. Again, numerous large ducts of the #uarter are attached with gland cistern. These ducts branch profusely, ultimately ending in the secretory units called alveoli or acini. The mil% 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. 2obules in turns are grouped into larger units called lobes.

The alveoli are surrounded by myoepithelial cells that are involved in the mil% e!ection 3mil% letdown4 refle*. The nature of epithelial cells which surrounds udder and teat outside to inside are5 $%in epithelial of udder-stratified s#uamous $trea% 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 e*tent. The muscular and lymphatic systems are developed. /rom birth to before puberty, the mammary gland undergoes little development, It increases in si6e due to increase in connective tissues and fat and e*tension 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) evelopment during pregnancy- 7uring pregnancy, mammary glands develop rapidly. -landular 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. !ormonal control of mammary gland There are numbers of hormones which are responsible for the growth and development of mammary gland 3mammogenesis4 either directly or indirectly which are as follows. ") Pituitary Gonadotropins Mainly mammary growth is under the control of estrogen and progesterone. /$1 causes development of follicles and these follicles are source of estrogen and 21 causes ovulation and forms 82 which is the source of progesterone. strogen alone causes duct development and estrogen and progesterone together cause lobulo-alveolar growth. 1owever, 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.

#) $ole of anterior pituitary hormones

It has been %nown that anterior pituitary hormones are essential for optimum mammary growth. They are- -1, A8T1, T$1, 9rolactin. %) $ole 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. &) $ole of placenta The mammary gland shows remar%able developmental changes during pregnancy and placenta partly plays the functional role in hypophysis and ovary enhancing the development of mammary gland. 'actogenesis The process by which the alveolar cells of the mammary gland are tuned with the ability to secrete mil% constituents is %nown as lactogenesis. There different stages involved in it and they are" In the first stage, there occurs an en6ymatic and cytologic change of alveolar epithelial cells and a limited secretion of mil% before parturition occurs. This is followed by second stage in which all mil% components are secreted !ust immediately before parturition and the process e*tends for several days period. The onset of mil% secretion after parturition occurs to supply nutrients to new born which is a third stage. This onset of mil% secretion is under the control of 9rolactin, -lucocorticoids and sudden fall in 9rogesterone level at parturition. 9lacental lactogen is slightly responsible for this. Galactopoiesis It is the process of maintenance of mil% secretion and it depends on alveolar cell number, their synthetic ability and let down of mil%. 9eriodic removal of mil% by hand or suc%ling by calf is important otherwise normal hormones production responsible for -alactopoiesis, ceases. There are numbers of hormones responsible for this which are-:*ytocin, A8T1, -1, -lucocorticoids, T$1, 9rolactin, Insulin etc. 'et down of mil( Mil% let down 3e!ection4 is a natural process used by the cow to help remove mil% from the udder. Mil% is initially secreted into small sacs within the mammary gland called alveoli, from which it must be e!ected for consumption or harvesting. Mammary alveoli are surrounded by smooth muscle 3myoepithelial4 cells which are a prominent target cell for o*ytocin. :*ytocin stimulates contraction of myoepithelial cells, causing mil% to be e!ected into the ducts and cisterns above the teat. :*ytocin reaches the udder where it causes the myoepithelial cells

around the alveoli to contract. ;pon contraction of the myoepithelial cells, the alveoli collapse and mil% is s#uee6ed out into the small ducts. These ducts shorten and widen, and the mil% rushes through into the gland cistern. :nly alveolar and ductal mil% is e*pelled by the action of o*ytocin on the myoepithelium. There is no contraction of large ducts or cisterns. The pressure of mil% being forced into the ducts<cistern and down towards the teat causes the teat to swell with mil% and become plump. It ta%es =( - >( seconds for teats to become plump after let down has been initiated. 8ows with well-filled udders re#uire a shorter period of stimulation to elicit mil% let down response than cows with less-filled udders. The action of o*ytocin is essential for emptying of the udder during mil%ing. The action of o*ytocin only lasts for = to ? minutes, because its concentration in the blood decreases rapidly. Mechanical stimulation of peripheral neural receptors in s%in of teat 3during mil%ing<suc%ling4@ neural stimulus. A $timulus travels from teat to spinal cord and to the supraoptic and paraventricular nuclei of hypothalamus. A $ynthesi6e o*ytocin hormone @release through posterior pituitary. A :*ytocin-released to b<d-transported to target t<s 3myoepithelial cells of mammary gland4. A 8ontraction of gland and e!ection or let down of mil%. Factors affecting the let down of mil( '. Inade#uate udder preparation ). 7elayed attachment of teat-cups for minute after preparing the udder or failure of mil%ing e#uipment to operate properly. ,. *ternal stimuli5 The mil% let-down refle* of cows is very easily retarded or prevented by e*ternal stimuli. It is often said that a cow Bholds upB her mil%. 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. Mil%ing routine5 feeding, barn noises, irregular time of mil%ing, method of mil%ing and the sight of the calf. motionally disturbing stimuli5 dog bar%ing, outer loud and unusual noises

 $tressors5 death of the calves, loud and various unfamiliar noises, visit of unfamiliar persons in the mil%ing parlor or at the site of mil%ing, change of mil%er and beating or rough handling at the time of mil%ing especially in first calvers. The basal concentrations of cortisol and C-endorphin were higher in the cows mil%ed in unfamiliar surroundings compared to when mil%ed in familiar. $uch stimuli cause the brain to release adrenalin. This hormone wor%s against o*ytocin by5

&loc%ing o*ytocin release from the brain. 8onstricting blood vessels and preventing o*ytocin from reaching the udder. 7irectly counteracting the effect of o*ytocin on the contraction of myo-epithelial cells.

If adrenalin release occurs before the mil%-e!ection stimulus, the e!ection will be virtually completely bloc%ed. Dhen adrenalin is released after mil% e!ection has commenced, it will result in large amounts of mil% being retained in the udder with associated negative effects on mil% yield. +. ;dder dema5 This presents itself as a failure of mil% let down associated with e*cess fluid in the mammary tissues. It is characteri6ed 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 s%in and deep in the udder tissue. E. Mastitis5 It may causes the painful let down of mil%. =. 7isturbance of mil% removal has been observed in primiparous cows immediately after parturition and during estrus. )omposition of Mil( Mil% is considered to be almost a perfect food for consumption. /rom the chemist point of view, mil% can be defined as a complete food consisting of fat, proteins, lactose and minerals and water, as ma!or constituents, and vitamins, en6ymes, pigments, gases and several other organic and inorganic substances.

$ole of )onstituents of Mil( As already mentioned, mil% constituents are divided under two groups5 ma!or and minor. Their role in the mil% is described as follows5 *+ Ma,or constituents The following are the ma!or nutrient constituents in the mil% that playa significant role in the diet. "+ -ater The water in the mil% is the same as any other water. Its function is to hold the solids of the mil% partly in solution and partly in suspension. #+ Mil( fat The fat e*ists in mil% as an emulsion. Dhen a mil% droplet is e*amined under a microscope, butterfat globules are visible in the form of small spherical myriads. The si6e of these globules varies from (.' to ) microns, the average being , 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 en6ymes, notably lipase, in causing butterfat to become rancid. The si6e and arrangement of the globules affect the creaming ability of mil%, the viscosity of cream, the case of whipping cream and churning of cream to butter. /at is a mi*ture of the glycerides of certain fatty acids. Mil% fat

contains '> or more fatty acid. These fatty acids may be classified as soluble and the insoluble, volatile and the non-volatile, saturated and the unsaturated. Mil% fat is the lightest constituent of mil%. It is because of this fact that the cream, which contains most of the mil% fat in mil%, rises to the top to form the cream layer. Mil% fat is very rich source of energy and also a good source of vitamin A and 7. Mil% fat is one of the principal constituents of almost all dairy products, such as feed, cheese, ice-cream, etc. %+ Proteins The three principal classes of proteins of mil% are casein, lactalbumin and lactoglobulin. The caseins account for ?( per cent of total protein content, the lactalbumin and lactoglobulin 3whey proteins4 almost )( per cent. a+ )asein 8asein is a phosphor-protein which is found in no other product than mil%. It e*ists in mil% in combination with calcium, hence is often spo%en as calcium caseinate comple*. It is present in mil% in a finely divided suspension. $uch suspension is %nown as B8olloidalB dispersion. The casein is precipitated by acids at its iso-electric point of p1 +.=. It is precipitated out of mil%, when the mil% source or when acid or rennet is added to it. &ecause 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 3artificial4 mil% products in the form of sodium caseinate. b+ -hey proteins + The proteins lactalbumin and lactoglobulin are present in small amounts, dispersed in the water of mil%. 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. ;sually the first mil% secreted after calving 3colostrum4 contains si*teen times more whey proteins than the normal mil%. This amount decreases rapidly to normal during the first few mil%ings. &+ 'actose 2actose, a carbohydrate found only in mil%, it is synthesi6ed in mammary gland. It is it disaccharides composed of glucose and galactose, which are liberated when lactose is hydroly6ed either by the en6yme lactase of the digestive tract or by lactase of bacterial origin. 2actose 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 mil% to taste sour. In some dairy products, such as soft cheese butter- mil% and butter, the characteristic flavour is partly due to the acid fermentation. $ince lactose is not very soluble, it some- times separates or Bcrystali6esB in such dairy products as sweetened condensed mil% and ice-cream, producing

Bsandy te*tureB condition. 2actose has those nutritive properties which are not possessed by any other sugar. .+ Mineral constituents The mineral constituents or salts of mil% constitute (.= to (.?F of the total weight. The potassium and chlorides of mil% are in solution, whereas the calcium, phosphorus and magnesium are partly in solution and partly in colloidal suspension. &ecause the metallic elements are in e*cess as compared to the non-metallic elements, the minera' content is al%aline in character. Minerals of mil% 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 mil% is (.0F, the presence of this constituent answers one of the principal reasons why mil% is such an important food, especially for children and e*pectant mothers. Mil% contains minerals mainly as inorganic salts, although some of them are in organic combination. Gumerous elements in trace #uantities are also found. /+ Minor constituents The important minor elements of mil% are pigments, cholesterol, phospholipids, en6ymes, etc. They are discussed here in detail. "+ Pigments Mil% contains two fat soluble pigments, BcaroteneB and B*anthophyllB and one water soluble pigment, BlactoflavinB 3lactochrome4. 8arotene is mainly responsible for the yellow colour of cow mil%. The amount of the water soluble pigment 3lactoflavin4 in mil% varies with the breed and the individual cow or buffalo. #+ )holesterol The amount of cholesterol in mil% varies directly with the butter fat content of the mil%. 8holesterol is a yellow-white fatty chemical compound found in such foods as eggs, meat and dairy products. %+ Phospholipids The principal phospholids found in mil% is called BlecithinB. 2ecithin is a fat-li%e substance containing nitrogen and phosphorus. The lecithin is closely associated with mil% fat, and when mil% is separated, some of the lecithin remains in the butter, though most of it passes into butter mil%. &+ Enzymes n6ymes are comple* organic compounds secreted by living plant and animal cells which initiate chemical reactions but emerge unchanged. n6ymes are specific in their action. 2ipase,

for e*ample, acts only upon fat and proteins act only upon protein and its derivatives. They are relatively unstable. 1igh temperatures, unfavourable p1, light and certain chemicals tend to inactivate them. The following are the principal en6ymes in mil%. a+ )atalase This en6yme splits hydrogen pero*ide into water. Temperature of =EH 8 to =0H 8 for ,( minutes destroys this en6yme. b+ *mylase Amylase, a starch splitting en6yme, is considered to be the least variable in #uantity of all the en6ymes in normal mil%. c+ Phosphatase It is always present in mil%. Its concentration is fairly constant. It has the ability to free phenel from alcoholic esters of phosphoric acid. d+ 'ipase It is an en6yme in mil% which brea%s down mil% fat, thus causing a strong or BrancidB flavour. e+ Galactase -alactase slowly reduces mil% protein to simple compounds" hence it plays a part in the BripeningB of some cheeses. f+ 'actase 2actase hydrolyses lactose into simpler sugar li%e glucose and galactose. .+ Gases Mil% contains some gases such as carbon-dio*ide, o*ygen and nitrogen. 8arbon-dio*ide enters the mil% in the cowIs buffaloIs udder, while o*ygen, nitrogen and other gases get into the mil% during mil%ing. -ases also may be formed by the growth of certain bacteria. The gases in mil% have no special significance, they slowly pass from the mil% as it stands and almost entirely escape during pasteurisation. 0+ 1itrogenous substances :ther nitrogen containing compound besides the proteins, are found in mil% in very small #uantities. These are fibrin, mucoid protein and an alcohol soluble protein. :ther nitrogen containing substances which are not proteins but are found on mil% in very small amounts are uric acid, urea, creation, creatinine and traces of some amino acids.