OCCASIONAL REVIEW

The physiology of lactation

nursing or medical curricula, so many healthcare professionals

involved in the care of the mother and infant lack the skills to
manage problems with breastfeeding. Since the ability to
optimally breastfeed an infant depends on the support a mother
receives from those around her, all healthcare providers who
interact with women or infants should be knowledgeable about
the basics of lactation and the role that their specialty plays in
breastfeeding. Therefore, the aim of this article is to provide a
broad outline of the physiology of lactation and to discuss the
aetiology and management of breastfeeding, so that the clinical
issues in the newborn period can be addressed.

E Jones
SA Spencer

Abstract
The evidence is now overwhelming that prolonged breastfeeding is one
of the most important contributors to infant health. However, only 69%
of women initiate breastfeeding in the UK, and this declines rapidly to
42% at 6 weeks. The majority of problems could be avoided if mothers
were given adequate support; therefore, these figures represent a failure
by the healthcare system to provide women with the help they need to
sustain breastfeeding. Mothers of pre-term infants are especially
vulnerable to lactation failure, and require expert intervention if
breastfeeding is to succeed. The aim of this article is to provide
healthcare professionals responsible for assisting women with the
management of lactation and breastfeeding with an up-to-date knowledge of the anatomy and physiology of the breast, the mechanisms of
milk secretion and the principles of breastfeeding, so that clinicians are
able to address management issues during the critical period following
delivery.

Keywords breastfeeding;

insufficient

milk;

lactation;

Functional anatomy of the lactating breast

The breast is primarily composed of glandular (secretory) tissue
and adipose (fatty) tissue bound together by a loose framework
of connective tissue called Coopers ligaments.6 During lactation, there is a decrease in the amount of adipose tissue relative
to glandular tissue,7 and the size and weight of the breast
increases. The mammary gland is composed of between 15 and
20 sections called lobes, which consist of alveoli. Within a lobe,
small ducts drain the alveoli and coalesce forming larger ducts,
which eventually join to form a single duct to transport milk to
the nipple. Most textbooks describe a distension of the duct
located in the alveolar area of the breast as the lactiferous ducts
or sinuses, where milk is stored before and during a feed.
However, recent dynamic studies using ultrasound7 have given a
new insight into the functional anatomy. In the living tissue,
sinuses are not seen and the ducts only dilate during milk
ejection. The low number and size of ducts and the absence of
sinuses suggest that the ducts transport milk rather than store it.
Another important finding relates to the distribution of
glandular tissue, which is less abundant in the lateral aspects
of the breast than previously thought. Functional tissue is well
represented adjacent to the nipple and immediately below the
areola. In fact, there is minimal subcutaneous fat at the base of
the nipple (Figure 1).

mammary

physiology

Introduction
The epidemiological evidence is now overwhelming that breastfeeding is one of the most important contributors to infant health,
and that prolonged breastfeeding protects babies from common
childhood infections through mechanisms that are interactive,
adaptive and extend into childhood.1 Breastfeeding also offers a
range of health benefits for mothers, including a reduced risk of
ovarian2 and pre-menopausal breast cancer.3 Both the World
Health Organisation and the Department of Health recommend
that exclusive breastfeeding should continue for the first 6
months of an infants life. However, despite the proven
advantages of breastfeeding, only 69% of women initiate
breastfeeding in the UK, and the rate of exclusive breastfeeding
declines rapidly to 42% at 6 weeks following delivery.4 The
majority of the reasons given for ending breastfeeding relate
directly to problems that could have been avoided or solved with
adequate breastfeeding support both in the immediate postpartum period and following discharge from hospital.
Although the National Institute for Health and Clinical
Excellence5 suggests that maternity care providers should
implement breastfeeding training, it is not compulsory in all

Transition from pregnancy to lactation

The mammary gland develops the biochemical capacity to
secrete milk during pregnancy and this occurs in two separate
phases: mammogenesis and lactogenesis.8 Mammogenesis occurs early in pregnancy and is characterised by the proliferation
of the distal elements of the ductal tree, creating multiple alveoli
of variable size and shape. Initially, the development is a
continuum of the hypertrophy associated with the menstrual
cycle, and women notice an increase in breast tenderness and
nipple sensitivity early on in pregnancy. Subsequently, the
subcutaneous veins can be observed to enlarge, and the areola
becomes more deeply pigmented.
During mid-pregnancy, secretory differentiation begins with a
rise in mRNA for many milk proteins and enzymes important for
milk formation and secretion.9 This switch to secretory differentiation is classified as stage I lactogenesis10 and is the process
whereby the mammary gland becomes competent to secrete
milk. During this period, fat droplets begin to increase in size in
secretory cells, and become a major cell component at the end of
pregnancy. Other milk-specific components are also produced

E Jones MPhil RN RM ENB 405 is Research Midwife at the University Hospital of

North Staffordshire NHS Trust, Stoke on Trent, UK.
SA Spencer DM FRCPH is Consultant Paediatrician at the University Hospital
of North Staffordshire NHS Trust, Stoke on Trent, UK.

PAEDIATRICS AND CHILD HEALTH 17:6

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OCCASIONAL REVIEW

galactopoiesis or lactogenesis III. Under normal circumstances,

the breasts will continue to produce milk indefinitely as long as
milk removal continues.

Current research about milk synthesis

There is compelling evidence to suggest that extracellular storage
of milk allows biologically active substances in milk to influence
mammary function.14 A whey protein, termed feedback inhibitor
of lactation (FIL) regulates the rate of milk secretion acutely
according to frequency or completeness of milk removal in each
mammary gland. Therefore, milk production slows when milk
accumulates in the breast (and more FIL is present) and speeds
up when the breast is emptier (and less FIL is present). There is
also evidence that fat content is determined by how empty the
breast is (emptier breast higher fat milk), rather than the time
of day or stage of feed. Diurnal variations in milk content also
exist, which reflect maternal diet and daily hormonal fluctuations.
The storage capacity of a breast is a functional concept and is
defined as the maximum amount of milk that can be stored in the
breast between feeds.15 This can vary widely both between
mothers and also between breasts for the same mother. Storage
capacity is not necessarily determined by appearance, since it is
dependent on the ratio of glandular to adipose tissue in the
breasts. In practice, mothers with large or small storage capacities
can produce sufficient milk for their baby. A mother with a large
capacity may be able to go longer between feeds without
impacting on milk supply and infant growth. A mother with a
smaller capacity will need to feed more frequently to maintain
her milk supply since her breast will become full (slowing
production) more quickly. Since milk production directly reflects
the infants appetite, all mothers should be strongly encouraged
to feed their babies on demand.

Figure 1 Anatomy of the lactating breast. Changes to observe: ducts

branch closer to the nipple; lactiferous sinuses absent; glandular tissue
closer to the nipple; and subcutaneous fat minimal at the base of the
nipple. rMedela AG 2006, Baar/Switzerland.

such as lactose, casein and a-lactalbumin. Since these secretions

are not removed by infant suckling, they are re-absorbed into the
blood through the paracellular pathway. Full milk secretion is
held in check by high circulating levels of progesterone and
oestrogen, and the gland remains quiescent but poised to initiate
lactation around parturition.
At birth, the expulsion of the placenta results in a sudden drop
in progesterone, oestrogen and human placental lactogen levels.
This abrupt withdrawal of progesterone in the presence of high
prolactin levels cues lactogenesis II (or the onset of a copious
milk production) around 3040 hours after birth. However,
mothers do not typically begin to notice the sensation of breast
fullness until 5073 hours following delivery. Milk composition
also changes dramatically during this period, with a fall in the
sodium and chloride concentrations and an increase in the
lactose concentrations that start immediately after birth and are
largely complete by 72 hours post-partum.11 These changes are
explained by the closure of the tight junctions that block the
paracellular pathway, and precede the onset of the large increase
in milk volume by at least 24 hours. The concentrations of
immunoglobulin sIgA and lactoferrin rise dramatically during the
period following delivery, and although their concentrations fall
as milk volume increases, their secretion rate is still substantial
throughout the course of lactation.12 Oligosaccharides concentrations are also high during this early period and have a substantial
protective effect against a variety of pathogens. The sharp
increase in volume that occurs between 36 and 96 hours postpartum reflects a dramatic increase in the rates of synthesis for all
components of mature milk. This phase of lactation is hormonally driven, and milk removal is not needed for the changes that
bring about lactogenesis II in the majority of women.13 However,
it is clear that milk removal by day 3 is necessary for the
successful continuation of lactation.
After lactogenesis II, there is a switch from endocrine to
autocrine control, and milk removal is the primary control
mechanism for milk synthesis. This phase of lactation is termed

PAEDIATRICS AND CHILD HEALTH 17:6

Milk ejection
Milk removal from the breast is accomplished by the contraction
of the myoepithelial cells, whose processes form a basket-like
network around the alveoli where milk is stored.16 When the
infant is suckled, afferent impulses from sensory stimulation of
nerve terminals in the areolus travel to the central nervous
system where they promote the release of oxytocin from the
posterior pituitary. The oxytocin is carried through the blood
stream to the mammary gland where it interacts with specific
receptors on myoepithelial cells, initiating their contraction and
expelling milk from the alveoli into the ducts. The passage of
milk through the ducts is facilitated by longitudinally arranged
myoepithelial cell processes whose contraction shortens and
widens the ducts, allowing free flow of milk to the nipple. The
process by which milk is expelled from the alveoli is called the
milk ejection reflex and is essential to milk removal from the
lactating breast. When this reflex is inhibited, milk cannot be
removed from the breast and local mechanisms bring about an
inhibition in milk secretion. When milk removal stops completely, mammary involution is triggered and the gland loses its
capacity to secrete milk.
Initially, the milk ejection reflex is unconditioned and requires
a physical stimulus. Once it has become conditioned, the reflex

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poor attachment and positioning; ineffective infant suckling;

infrequent feeds; breast engorgement; impaired milk ejection;
inverted nipples; mastitis; and maternal/infant separation.

can be inhibited by physical stress such as pain, and by emotional

stress such as embarrassment and anxiety.17

Factors associated with impaired lactogenesis

How to overcome problems of insufficient milk

All breastfeeding mothers need comprehensive guidance with

respect to the initiation and continuation of breastfeeding. In
order to minimise feeding problems, breastfeeding education
should be available during the antenatal period so that parents
have a realistic idea about how breastfeeding works. In the
immediate period following delivery, mothers should be encouraged to have skin-to-skin contact with their baby. The separation
of a mother and her baby within the first hour of birth for routine
post-natal procedure should be avoided, whenever possible.
Initiation of breastfeeding should be encouraged as soon as
possible following birth, ideally within the first hour. From the
first feed, mothers should be offered skilled breastfeeding support
to enable comfortable positioning and to ensure that the baby
attaches correctly to the breast. This is necessary to establish
effective feeding and prevent problems such as sore nipples.
There is strong evidence that a stressful labour and delivery
are associated with delayed onset of lactation and/or infant
suckling difficulties. Delayed onset of lactation is defined as more
than 72 hours. Caesarean section and primiparity are two of the
most important risk factors.18 In a hospital-based study in Italy,
8% of exclusively breastfed newborns lost more than 10% of
their birth weight during the first 35 days.19 The investigators
determined that 26% of these cases were attributable to
inadequate maternal milk volume, and 74% were caused by
poor breastfeeding technique by either the mother or the infant.
Since a newborn who has experienced stress during labour and
delivery may be too weak or too sleepy to attach and suckle
effectively at the breast, additional support should be given to
mothers who have had a narcotic, a general anaesthetic or a
difficult birth. Furthermore, since early milk removal increases
the efficiency of milk synthesis, if a baby is reluctant to feed,
mothers should be encouraged to express milk on a regular basis
in order to stimulate stage II lactogenesis. A comprehensive list of
infant and maternal risk factors can be found in Table 1. Irregular
or incomplete removal of milk results from and/or contributes to:

The definitive indicator of an adequate milk supply is infant

weight gain in the early neonatal period. The evidence suggests
that when babies are breastfeeding well:
 maximum weight loss occurs by 72 hours
 maximum loss of body weight is less than 10% and
 birth weight is regained by 57 days following birth.

Other indicators of sufficient breast milk intake are frequent

wet nappies and pale, odourless urine from 4872 hours and a
change in stool colour from meconium at 024 hours to yellow at
7296 hours. Poor weight gain, static weight or weight loss at any
time after the first week is a cue that breastfeeding is not going
well.
As many women leave hospital in the early days following
delivery, breastfeeding problems often become apparent after the
clinical situation has become complicated by insufficient milk.
Since there is a direct relationship between correct attachment,
an unrestricted milk flow and the resultant lack of damage to the
nipple, correct attachment and positioning is one of the keys to
breastfeeding success. Mothers should feel strong tugging which
is not persistently painful. Figure 2 demonstrates correct
attachment. The babys mouth is wide open and the lower lip
is further from the base of the nipple than the top lip. The baby
has a mouth full of breast, including the nipple, much of the
areola and all the underlying tissue including the milk ducts. This
will cause a typical jaw action as the baby works on the breast.
The jaw muscles work rhythmically, and this action extends as
far back as the ears. If the cheeks are being sucked inwards, the
baby is not attached properly.20
Milk production works on a use it or lose it principle.
Therefore, sleepy babies should be encouraged to feed at least
eight to 12 times in 24 hours. Sleeping near the baby facilitates
breastfeeding because mothers are alerted by the signs of early

List of infant and maternal risk factors

For baby

For mother

 Birth by vacuum extraction

 Continued rooting after feeding

 Total labour more than 14 hours

 Caesarean section

 Infant irritable, restless or sleepy and refusing to feed

 Use of supplemental formula and dummies

 First-time mother
 Inverted nipples

 Poor attachment and positioning

 No visible or audible swallowing

 Sore nipples
 Prior breastfeeding problems
 Prior breast surgery

 No effective breastfeeding seen prior to discharge

 Tongue tie

 Type 1 diabetes
 Obesity

 Hyperbilirubinaemia
 Hypoglycaemia

 Multiple birth
 Smoking

Table 1

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encourage the mother to let the baby determine the length of a

feed. In cases of persistent poor or inadequate growth, specialist
facilities are required to determine whether underfeeding or an
intrinsic organic problem in the infant is the cause of the
problem.

Problems associated with pre-term lactation

In the extremely pre-term infant, there are particular difficulties
in establishing breastfeeding due to the earlier stage of breast
development at parturition and the inability of the newborn
infant to suckle. Some mothers may not have reached lactogenesis I prior to pre-term delivery, which means that the mammary
epithelium may not be sufficiently prepared by the hormones of
pregnancy to respond with efficient milk synthesis. Fortunately in
the vast majority of mothers of pre-term infants, compensatory
growth can be achieved, even if somewhat delayed, by efficient,
early and frequent milk removal.6 Consequently, during the first
post-delivery encounter, neonatal staff should discuss the
benefits of human milk, its role in the care of pre-term babies,
and the urgency to begin a milk expression schedule. A hospitalgrade double breast pump should be available to enable mothers
to express both breasts simultaneously. This method of milk
expression results in a higher milk yield, reduced pumping time
and may be an effective measure to trigger milk ejection.21 Breast
massage is another helpful technique that can be taught to
enhance the milk ejection reflex and improve milk yields. With
regular pumping and encouragement from staff, the majority of
these mothers can produce enough milk for their babies and go
on to establish full breastfeeding once the infant is ready to
suckle, which may be several weeks later.
Figure 2 Correct attachment and positioning. r Sue Saunders.

Conclusion
Healthcare professionals with responsibility for assisting women
with the management of lactation and breastfeeding should
possess an up-to-date knowledge of the anatomy of the breast,
the mechanisms of milk secretion and the principles of
breastfeeding. A sound understanding of what represents
appropriate attachment and positioning is also required in order
to address ineffective milk removal and nipple trauma. A flexible
pattern of feeds that is responsive to an infants needs should be
encouraged. Frequent feeds, unrestricted in length, will help the
baby to secure adequate nutrition and will help to promote milk
synthesis. Mothers of extremely pre-term infants should be
taught effective milk expression techniques as a matter of
urgency. Since early lactation difficulties are common even in
women who are highly motivated and receive breastfeeding
support while in hospital, it is essential that all breastfeeding
mothers and infants continue to receive professional support in
~
the immediate period following discharge from hospital.

feeding cues, thus helping to increase the frequency of feeds.

Breast massage and compression during a feed will help to
accelerate milk flow and ensure effective drainage. Since the use
of dummies may mask the early signs of hunger, dummy use
should be discouraged, particularly in mothers who are trying to
increase their milk supply. Formula supplementation will also
reduce milk synthesis, and should be avoided unless there is a
strong medical indication.
Thoroughly emptying the breast is more important than the
frequency of milk removal in stimulating milk production.
Consequently, one of the fastest ways to increase milk supply is
to manually express milk by hand or pump after each breastfeed.
The rate of milk synthesis is up to five times higher in a welldrained breast compared with a full breast. It is also of vital
importance to identify and eliminate factors that inhibit adequate
drainage such as an underwired bra or a breast pump shield with
an opening that compresses the nipple. A milk expression shield
with a small diameter can also increase nipple soreness.
Finally, in order to maximise the action of the milk ejection
reflex, it is important to remove distractions in order for mothers
to relax and enjoy feed times. Simple interventions such as taking
the phone off the hook, settling into a comfortable chair with a
drink on hand and listening to the radio can enhance the rate of
milk transfer during a breastfeed. It is also important to

PAEDIATRICS AND CHILD HEALTH 17:6

Acknowledgements
The authors would like to thank Dr Donna Ramsay and
Professor Peter Hartmann of the University of Western
Australia.

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PAEDIATRICS AND CHILD HEALTH 17:6

Practice points
Lactogenesis is not robust
Early and effective milk removal is crucial for the initiation of
lactation
 Triggering the milk ejection reflex is essential for effective milk
removal
 There is a direct relationship between correct attachment,
unrestricted milk flow and avoidance of damage to the nipple



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