Lactation is the process by which milk is synthesized and secreted from the mammary

glands of the postpartum female breast in response to an infant sucking at the nipple. Breast
milk provides ideal nutrition and passive immunity for the infant, encourages mild uterine
contractions to return the uterus to its pre-pregnancy size (i.e., involution), and induces a
substantial metabolic increase in the mother, consuming the fat reserves stored during
pregnancy.

Structure of the Lactating Breast

Mammary glands are modified sweat glands. The non-pregnant and non-lactating female
breast is composed primarily of adipose and collagenous tissue, with mammary glands
making up a very minor proportion of breast volume. The mammary gland is composed of
milk-transporting lactiferous ducts, which expand and branch extensively during pregnancy
in response to estrogen, growth hormone, cortisol, and prolactin. Moreover, in response to
progesterone, clusters of breast alveoli bud from the ducts and expand outward toward the
chest wall. Breast alveoli are balloon-like structures lined with milk-secreting cuboidal cells,
or lactocytes, that are surrounded by a net of contractile myoepithelial cells. Milk is secreted
from the lactocytes, fills the alveoli, and is squeezed into the ducts. Clusters of alveoli that
drain to a common duct are called lobules; the lactating female has 12–20 lobules organized
radially around the nipple. Milk drains from lactiferous ducts into lactiferous sinuses that
meet at 4 to 18 perforations in the nipple, called nipple pores. The small bumps of the areola
(the darkened skin around the nipple) are called Montgomery glands. They secrete oil to
cleanse the nipple opening and prevent chapping and cracking of the nipple during
breastfeeding.

The Process of Lactation

The pituitary hormone prolactin is instrumental in the establishment and maintenance of
breast milk supply. It also is important for the mobilization of maternal micronutrients for
breast milk.

Near the fifth week of pregnancy, the level of circulating prolactin begins to increase,
eventually rising to approximately 10–20 times the pre-pregnancy concentration. We noted
earlier that, during pregnancy, prolactin and other hormones prepare the breasts anatomically
for the secretion of milk. The level of prolactin plateaus in late pregnancy, at a level high
enough to initiate milk production. However, estrogen, progesterone, and other placental
hormones inhibit prolactin-mediated milk synthesis during pregnancy. It is not until the
placenta is expelled that this inhibition is lifted and milk production commences.

After childbirth, the baseline prolactin level drops sharply, but it is restored for a 1-hour spike
during each feeding to stimulate the production of milk for the next feeding. With each
prolactin spike, estrogen and progesterone also increase slightly.

When the infant suckles, sensory nerve fibers in the areola trigger a neuroendocrine reflex
that results in milk secretion from lactocytes into the alveoli. The posterior pituitary releases
oxytocin, which stimulates myoepithelial cells to squeeze milk from the alveoli so it can
drain into the lactiferous ducts, collect in the lactiferous sinuses, and discharge through the
nipple pores. It takes less than 1 minute from the time when an infant begins suckling (the
latent period) until milk is secreted (the let-down). Figure 1 summarizes the positive feedback
loop of the let-down reflex.

Fig 1. Let-Down Reflex. A positive feedback loop ensures continued milk production as long
as the infant continues to breastfeed.
The prolactin-mediated synthesis of milk changes with time. Frequent milk removal by
breastfeeding (or pumping) will maintain high circulating prolactin levels for several months.
However, even with continued breastfeeding, baseline prolactin will decrease over time to its
pre-pregnancy level. In addition to prolactin and oxytocin, growth hormone, cortisol,
parathyroid hormone, and insulin contribute to lactation, in part by facilitating the transport
of maternal amino acids, fatty acids, glucose, and calcium to breast milk.

Changes in the Composition of Breast Milk

In the final weeks of pregnancy, the alveoli swell with colostrum, a thick, yellowish
substance that is high in protein but contains less fat and glucose than mature breast milk
(Table 3). Before childbirth, some women experience leakage of colostrum from the nipples.
In contrast, mature breast milk does not leak during pregnancy and is not secreted until
several days after childbirth.

*Cow’s milk should never be given to an infant. Its composition is not suitable and its
proteins are difficult for the infant to digest.
Compositions of Human Colostrum, Mature Breast Milk, and Cow’s Milk (g/L) (Table
3)
Human colostrum Human breast milk Cow’s milk*
Total protein 23 11 31
Immunoglobulins 19 0.1 1
Fat 30 45 38
Lactose 57 71 47
Calcium 0.5 0.3 1.4
Phosphorus 0.16 0.14 0.90
Sodium 0.50 0.15 0.41

Colostrum is secreted during the first 48–72 hours postpartum. Only a small volume of
colostrum is produced—approximately 3 ounces in a 24-hour period—but it is sufficient for
the newborn in the first few days of life. Colostrum is rich with immunoglobulins, which
confer gastrointestinal, and also likely systemic, immunity as the newborn adjusts to a
nonsterile environment.

After about the third postpartum day, the mother secretes transitional milk that represents an
intermediate between mature milk and colostrum. This is followed by mature milk from
approximately postpartum day 10 (see Table 3). As you can see in the accompanying table,
cow’s milk is not a substitute for breast milk. It contains less lactose, less fat, and more
protein and minerals. Moreover, the proteins in cow’s milk are difficult for an infant’s
immature digestive system to metabolize and absorb.

The first few weeks of breastfeeding may involve leakage, soreness, and periods of milk
engorgement as the relationship between milk supply and infant demand becomes
established. Once this period is complete, the mother will produce approximately 1.5 liters of
milk per day for a single infant, and more if she has twins or triplets. As the infant goes
through growth spurts, the milk supply constantly adjusts to accommodate changes in
demand. A woman can continue to lactate for years, but once breastfeeding is stopped for
approximately 1 week, any remaining milk will be reabsorbed; in most cases, no more will be
produced, even if suckling or pumping is resumed.

Mature milk changes from the beginning to the end of a feeding. The early milk, called
foremilk, is watery, translucent, and rich in lactose and protein. Its purpose is to quench the
infant’s thirst. Hindmilk is delivered toward the end of a feeding. It is opaque, creamy, and
rich in fat, and serves to satisfy the infant’s appetite.

During the first days of a newborn’s life, it is important for meconium to be cleared from the
intestines and for bilirubin to be kept low in the circulation. Recall that bilirubin, a product of
erythrocyte breakdown, is processed by the liver and secreted in bile. It enters the
gastrointestinal tract and exits the body in the stool. Breast milk has laxative properties that
help expel meconium from the intestines and clear bilirubin through the excretion of bile. A
high concentration of bilirubin in the blood causes jaundice. Some degree of jaundice is
normal in newborns, but a high level of bilirubin—which is neurotoxic—can cause brain
damage. Newborns, who do not yet have a fully functional blood–brain barrier, are highly
vulnerable to the bilirubin circulating in the blood. Indeed, hyperbilirubinemia, a high level of
circulating bilirubin, is the most common condition requiring medical attention in newborns.
Newborns with hyperbilirubinemia are treated with phototherapy because UV light helps to
break down the bilirubin quickly.

Chapter Review
The lactating mother supplies all the hydration and nutrients that a growing infant needs for
the first 4–6 months of life. During pregnancy, the body prepares for lactation by stimulating
the growth and development of branching lactiferous ducts and alveoli lined with milk-
secreting lactocytes, and by creating colostrum. These functions are attributable to the actions
of several hormones, including prolactin. Following childbirth, suckling triggers oxytocin
release, which stimulates myoepithelial cells to squeeze milk from alveoli. Breast milk then
drains toward the nipple pores to be consumed by the infant. Colostrum, the milk produced in
the first postpartum days, provides immunoglobulins that increase the newborn’s immune
defenses. Colostrum, transitional milk, and mature breast milk are ideally suited to each stage
of the newborn’s development, and breastfeeding helps the newborn’s digestive system expel
meconium and clear bilirubin. Mature milk changes from the beginning to the end of a
feeding. Foremilk quenches the infant’s thirst, whereas hindmilk satisfies the infant’s
appetite.

colostrum
thick, yellowish substance secreted from a mother’s breasts in the first postpartum
days; rich in immunoglobulins
foremilk
watery, translucent breast milk that is secreted first during a feeding and is rich in
lactose and protein; quenches the infant’s thirst
hindmilk
opaque, creamy breast milk delivered toward the end of a feeding; rich in fat; satisfies
the infant’s appetite
lactation
process by which milk is synthesized and secreted from the mammary glands of the
postpartum female breast in response to sucking at the nipple
let-down reflex
release of milk from the alveoli triggered by infant suckling
prolactin
pituitary hormone that establishes and maintains the supply of breast milk; also
important for the mobilization of maternal micronutrients for breast milk