Biology of Lactation 342-460B

Mammary Gland

The Mammary Gland

What is a mammary gland?

The mammary gland is the milk secreting structure, which includes a teat, a duct system and
lobes (lobules) of secretory tissue drained by the duct system. Mammary glands are modified
sweat (sudoriferous) glands, which secretes milk (exocrine gland) and serve as accessory glands
to the reproductive system. Due to its economic importance, the following discussion will focus
on the mammary gland of the dairy cow.

The udder
The udder is a complex organ made up of a series of systems, which includes
- A supportive system.
- A secretory system composed of epithelial cells.
- A duct system for storage and conveyance of milk.
- Blood, lymph, and nerve systems.

In the cow, the udder is separated into two halves by a longitudinal groove, the intermammary
groove. It consists of four separate mammary glands (quarters), each gland has one teat and each
teat has one opening. The right and left halves are entirely separate. The rear quarters are
normally larger than the forequarters and secrete about 60% of the daily milk yield. The milk
from each gland is emptied through the teat. Rear teats are usually shorter than the fore teats.
About 25-50% of all cows have extra teats. Extra teats should be removed during calfhood.

The weight of the udder varies with the age of the cow, the stage of lactation, the amount of milk
in the udder, and inherited differences among cows. The weight of the empty udder varies
between 12-30 kg with an average capacity of about 35 kg. The weight and capacity of the
udder increases with age of the cow up to six years, with the greatest increases occurring
between the first and the second lactation. It is very important for the dairy producer to have
cows with udders that are large enough to produce large volume of milk and in the same time not
to have them excessively large or with poor udder attachment.

Udder of other species

- Sheep and goats: Two glands and two teats.
- Sow: 12-14 teats and two glands per teat.
- Mare: Four glands and only two teats.

Supportive System
A stronger udder supportive (suspensory) system is required to maintain proper attachments of
the gland to the body. The udder of the cow is supported by two major groups of ligaments.

1- Median Suspensory Ligaments (MSL): The most important part of the suspensory system in
cows. The MSL divide the udder into two distinct halves and provide the primary support for the
udder. When the cow is viewed from behind, a distinct midline groove marks the position of

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median suspensory ligament. The MSL is composed of elastic and fibrous connective tissues that
arise from the midline of the abdominal wall and extent between the udder halves. The elastic
tissues of the MSL provide the necessary elasticity needed by the udder as it fills with milk to
expand away from the body.

2- Lateral Suspensory Ligaments (LSL): These structures contain more collagen than elastin
and thus provide support but not elasticity to the udder. The LSL arise from the tendons above
and posterior to the udder and extend along both sides of the udder providing support to the
interior of the udder. The LSL extend to the midline on the floor of the udder joining the MSL at
the base of the udder. However, they don’t join under the bottom of the udder.

The skin provides the udder with protection but with little support. The udder is attached to the
skin by a fine connective tissue and a coarse connective tissue attaches the forequarters to the
abdominal wall.

Teat Structure
Cows have four teats (one teat drains on gland). The skin of the teat is characterized by the
absence of hair and glands (sweat or sebaceous glands). Teat size and shape is independent of the
size, shape or milk production of the udder. Average size for the fore teats is ~6.6 cm long and ~
2.9 cm in diameter. The corresponding values for the rear teats are 5.2 and 2.6 cm, respectively.

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Teat (streak) canal: About 7 to 16 mm long with a diameter of 0.08 mm. It is closed by circular
sphincter muscles. The streak canal retains the milk in the udder against the pressure developed
by the accumulation of milk. It is also the main barrier against infection and is lines with skin-
like epidermis cells that produce a bacteriostatic secretion.

Furstenburg’s rosette: Mucosal folds of the streak canal lining at the internal end of the canal.
It folds over the canal opening due to pressure when the udder is full, which aids in he retention
of milk. It may be a major point of entry for leukocytes leaving the teat lining and entering the
into the teat cistern.

Annular (cricoid) rings: Located at the proximal end of the teat cistern, marking the boundary
between the teat cistern and the gland cistern

Teat cistern: The cavity within the teat and is located just below the gland cistern. It is
continuous with the gland cistern. It is lined with numerous longitudinal and circular mucosal
folds, which from pocket on the inner lining of the teat.

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Mammary Gland

Interior Anatomy of the Mammary Gland

The mammary gland is a complex organ that proceeds through the early embryonic stages, into
pregnancy, lactation and finally regression.
Structure of the mammary gland

1- Connective tissue
2- Ductular system
3- Secretory tissue

Connective tissue (stroma)

The function of the connective tissue is to support the ductular and the secretory tissues. A
desirable udder should contain a minimal amount of connective tissue and maximal amounts of
serectory tissue.

Ductular System
The ductular system of the mammary gland starts at the alveoli and ends at the streak canal. No
modification of the milk composition occurs once it has moved into the duct. Milk is first
collected in the lumen of the alveoli and secretory (terminal) ductules. It then drains into larger
ducts until it enters the primary (mammary) ducts attached to the gland cistern. The gland
cistern is the largest collecting point of milk. The gland cistern is linked to the teat cistern. The
two cisterns are partially separated by an annular fold. Usually 10-12 ducts lead into each gland
cistern, but as many as 20 ducts can be found.

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Milk in the teat cistern is prevented from escaping to the outside by a sphincter muscle which
contracts around the teat (streak) canal through which milk is drawn to the outside.

The size and the shape of the gland cistern of each quarter vary considerably. The capacity of the
gland cistern varies from 100-400 ml of milk. No relationship exists between the size of the
cistern and the amount of milk secreted by the quarters. The gland and teat cisterns are
continuos. However, in most cows, a definite circular construction (annular folds) is located
between the two cavities.

Secretory Tissue
The secretory epithelial cells comprise about 1/2 of the total cells in the secretory tissue during
lactation. Secretory tissue is arranged into lobes with each lobes consists of many lobules. Each
lobule containing clusters or groups of alveoli (150-220 alveoli) which are surrounded by a
network of blood vessels. Each alveolus has all the components required to produce and deliver
milk to the duct system. Each alveolus consists of a single layer of epithelial cells (secretory
cells) surrounding the central lumen into which the epithelial cells eject the milk they
synthesize.

Alveoli (acini): Alveoli are sack-like structure where milk is synthesized and secreted. The
lumen of the alveolus is lined by a single layer of secretory epithelial cells. The epithelial lining
is surrounded by contractile epithelial cells, which contact in response to the hormone oxytocin,
resulting in milk being squeezed out of the alveolar lumen and into the small ducts. Outside the
myoepithelial cells, the alveolus is surrounded by a connective tissue basement membrane. The
capillary bed on the outside the alveolus is part of the stromal tissue between alveoli.

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Secretory Cells
A lactating secretory cell is the basic unit of milk synthesis. Milk precursors are taken from the
blood into the cell through the basal and the lateral membrane and milk is discharged into the
lumen through the apical membrane. Individual cells are joined to their neighbor cells on all
sides by tight junctional complex structure located around the apical portion of the apical
portion that forms a tight barrier, which prevents the passage of materials between cells under
normal conditions. Secretory cells are probably also bound to adjacent cells through gap
junction, which allow low molecular weight materials to pass from one cell to another. The
intracellular exchange may help a given alveolus to synchronized milk discharge from various
secretory cells into the lumen.

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Major Components of a Secretory Epithelial Cell

1- Nucleus The most prominent feature of the cell. It contains the genetic
materials that control the functions of cell as well as the genetic
information for the synthesis of milk protein and several enzymes
2- Endopalsmic Reticulum Membrane-bound channels that begin with the nucleus and extent
through the cytoplasm. They are the site of lipid synthesis and part
of the flow of membranes that form inner vesicles. The ER has two
parts;
A- Rough ER: Covered with ribosomes, and are the sites of milk
protein (export protein) biosynthesis.
B- Smooth ER: Devoid of ribosomes and serve as intracellular
channels for transferring materials within the cell.
3- Golgi Apparatus Site of protein packaging (addition of Ca & P to casein, formation
casein micelles). It is also the site of lactose biosynthesis. Other
non-fat milk constituents are also packed in the Golgi apparatus
before being secreted.
4- Secretory vesicles Originate in the Golgi apparatus to carry non-fat constituents and
vesicular membranes to the apical membrane
5- Lysosomes Contain enzymes which degrade unwanted materials in the cell.
The enzymes are released during involution (e.g. dry period or
during mastitis).
6- Cytoplasm The fluid matrix of the secertory cell. It includes enzymes,
macromolecules, nutrients. It is also the site of anaerobic
breakdown of glucose.
7- Junctions: Structures that join adjacent cells.
a- Tight junctions: Located around the apical portion of the cell. Join adjacent cells
and form tight barriers, which stop the passage of materials
between cells.
b- Gap junctions: Join adjacent cells and allow materials of low molecular weights to
move between cells.

8- Membranes:
a- Basal and lateral membranes: Milk precursors are taken from blood into secretory
cells through these membranes.
b- Apical membrane: Milk is discharged through this membrane into the
lumen.
c- Basement membrane: Milk precursors coming from the blood through this
membrane and are located in the interstitial space below the
secretory cell.

Unattached ribosomes exist in the cytosol in the areas surrounding the endoplasmic reticulum
and the Golgi, where synthesis of cellular proteins (structural and enzymatic) takes place.

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A secretory cell from a lactating mammary gland.

Precursors of Milk
The single-thick layer is some times refer to as the mammary barrier as this cell layer separates
the materials in the blood and the milk. Precursors of milk come from the bloodstream. The
primary substrates extracted from blood by the secretory cell include glucose, amino acids (both
essential and nonessential), fatty acids, and minerals. For ruminants, acetate and β-
hydroxybutyrate are also major substrates extracted from the blood. Glucose is the precursor for
lactose, citric acid and most of the glycerol portion of the triacylglycerol.

The major carbohydrate found in the milk is lactose and only glucose is found in the blood.
Some fatty acids are typical of milk fat in some species and most of milk proteins are not found
in the blood. Several materials in milk come unchanged from the blood, including minerals,
hormones, and several proteins especially immunoglobulins. In ruminants, immunoglobulins
pass the mammary barrier in large amounts into colostrum just prior to parturition.

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Blood Supply
The rate at which blood flow to the mammary gland is one of the key-factors in determining milk
synthetic rate. It takes from 400 to 500 liters of blood flowing the mammary glands to produce
one liter of milk. For a cow with an average milk production level, this translate to about 20% of
the entire output by the heart. There is a 2-6 fold increase in blood flow in the mammary gland
staring 2-3 days prepartum. However, the decrease in production with advancing lactation is not
due to decreased blood flow. This suggests that the efficiency of extraction of the components
from the blood while it passes through the udder is very important.

Arterial blood: Most of the blood is supplied to the udder by the two external pudenal (external
pudic) arteries, one for each half of the udder. The arteries enter the mammary gland from the
abdominal cavity through the inguinal canal. As they enter the mammary gland, the arteries form
a sigmoid flexure, which probably allows for the lowering of the udder when it becomes filled
with milk. The mammary artery branches soon after it penetrates the mammary gland into
cranial mammary artery (supplies the front side of the udder) and the caudal mammary artery
(supplies the rear part of the udder). Numerous branches from these arteries provide oxygenated
to all parts of the udder. The arteries in the teat are part of these and are known as the papillary
arteries.

Venous blood: The blood from each half of the udder leaves by two veins, the external pudenal
and the subcutaneous abdominal (milk vein) vein. Veins leave the mammary gland anti-parallel
to the arteries. Two routes of venous blood carry CO2 back to the heart. Many papillary veins
course upward from the teat and join, making several large mammary veins the venous circle.
After the venous circle, blood can leave the udder by one of two routes;

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1- Two external pudic veins which will eventually join the vena cava to deliver blood to the
heart (see figures).

2- Two subcutaneous abdominal veins (milk veins). These are anterior extension of the large
mammary veins and run forward a long the ventral abdominal wall just under the skin. They
penetrate the thoracic cavity at the milk well and eventually join the anterior vena cava.

The exact route is probably dependent upon the position of the animal. When a cow is in the
standing position, most blood returns to the heart via the milk veins. However, when a cow lies
down and shuts off blood flow through the milk veins, milk production is not affected because of
the other possible exists for blood.

Nervous System
The efferent innervation of the mammary gland is entirely sympathetic in origin. Stimulation of
the nerves leading to the mammary gland causes a vasoconstriction of the blood vessels, which
has a significant impact on milk secretion because the blood flow to the mammary gland is
decreased. The efferent nerves innervate the muscle fibres within the connective tissue
surrounding the lobules, lobes, and the blood vessels. However, the nerves do not pierce the
alveoli. The efferent fibers nerves apparently play an important role in innervating the smooth
muscles within the tea, especially those around the teat streak to keep it closed between milkings

- Innervation of the udder is sparse compared with other tissues.

- Sensory (afferent) nerves are involved in milk ejection and found in the teats and skins.

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- Similar to other skin glands, there is no parasympathetic innervation to the gland.

- Sympathetic nerves are associated with the arteries but not with alveoli.
- There is no innervation of the secretory system. Myoepithelial cells contract in response to
oxytocin and not in response to direct innervation.
- Few nerves go to the interior of the udder.

The primary functions of the sympathetic nerve fibers to the udder are control of the blood
supply to the udder and innervation of smooth muscle surrounding the milk collecting ducts and
the sphincter muscles within the teat. Stimulation of the sympathetic nervous system causes a
vasoconstriction of the blood vessels and this has an inhibiting effect on milk secretion.

Lymphatic System
Lymph is a colorless tissue fluid that is drained from tissue spaces by thin-walled lymph vessels.
Lymph originates as a filtrate of the blood serum and is similar in composition to it except that
lymph has no red blood serum and about ½ the protein content. Lymph flows from the udder to
the thoracic duct and is eventually discharged into the blood system. The lymph nodes of the
udder and the other lymph nodes of the body are important for disease resistance in the cow. The
lymph nodes form lymphocytes which are involved in immunity. In response to bacterial
infection (e.g. mastitis), lymph nodes increase their output of lymphocytes which are transported
to the udder to compact infection.
Around the time of parturition, the filtration of lymph out of the blood capillaries in the udder
may exceed the drainage back into the blood. This causes accumulation of fluid in the
intercellular tissue spaces. This disorder is known as udder edema. This is more serious in first-
calf heifers and in older cows with pendulous udders.

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