Inside the Lactating Breast: The Latest Anatomy Research

Donna T. Geddes, PhD

Although it is well recognized that a thorough understanding of the anatomy of an organ is essential to enable
assessment of any abnormalities in that organ, there has been little investigation of the anatomy of the normal
lactating breast since Sir Astley Cooper performed detailed dissections of the anatomy of the breast more than
160 years ago. Many mothers recognize that breast milk provides the ultimate nutrition and protection for the
infant; however, a significant proportion of women experience difficulties breastfeeding, some of which lead
to weaning the infant. Recently, a small number of studies have focused on the gross anatomy of the breast,
and have found that the ductal system is comprised of fewer numbers of main ducts than previously thought.
In addition, the ducts are compressible and do not contain large amounts of milk, the amount of fatty tissue
in the breast is variable, and a proportion is situated within the glandular tissue. These findings add to our
understanding of both the physiology and pathology of the lactating breast. J Midwifery Womens Health
2007;52:556 –563 © 2007 by the American College of Nurse-Midwives.
keywords: anatomy, breast, breastfeeding, lactation, mammary gland

INTRODUCTION BREAST DEVELOPMENT

The human breast reaches its full functional capacity Fetal Development
during lactation with the production of breast milk. In
The human breast develops from a thickened ectodermal
order to diagnose and treat breastfeeding problems and
ridge (milk line) situated longitudinally along the ante-
pathologies that arise during lactation, it is essential to
rior body wall from the groin to the axilla at about 6
have an extensive understanding of the normal anatomy
weeks’ gestation. Regression of the ridge occurs except
and physiology of the breast. This review details the
for the pectoral region (2nd– 6th rib), which forms the
development of the breast and the most recent findings in
mammary gland. Supernumerary glands may develop
breast anatomy. The effect breast anatomy has upon
anywhere along the ectodermal ridges, and in 2% to 6%
clinical practice as well as the importance of milk
of women, these glands either mature into mammary
ejection is reviewed.
glands or remain as accessory nipples.4,5
During the 7th and 8th weeks of gestation, the mam-
DEVELOPMENT OF THE BREAST
mary parenchyma invades the stroma, which appears as a
While it is undisputed that breast milk provides the raised portion called the mammary disc. Between the
optimal nutrition for the developing infant, breast milk 10th and 12th weeks, epithelial buds form; parenchymal
also contains unique protective factors for the mother.1 It branching occurs during the 13th through 20th weeks.
has been hypothesized that the mammary gland first Between the 12th and 16th weeks of gestation, the
evolved from the innate immune system as an inflamma- smooth musculature of the areola and nipple are formed,
tory response to provide protection to the young, and that and at approximately 20 weeks’ gestation, between 15
nutritional factors developed later.2,3 To date, nutrition and 256 solid cords form in the subcutaneous tissue.
has assumed a position of dominance over the protective Branching continues, and canalization of the cords oc-
factors in considerations of the physiology of human curs, forming the primary milk ducts by 32 weeks’
lactation. gestation.6 At 32 weeks’ gestation the ducts open onto the
The human breast is a dynamic organ that does not go area, which develops into the nipple.7 The adipose tissue of
through all developmental stages unless a woman expe- the mammary gland develops from connective tissue that
riences pregnancy and childbirth. The course of breast has lost its capacity to form fibres, and it is considered
development can be described in distinct phases begin- necessary to further growth of the parenchyma.4
ning with the fetal phase and progressing through the Shortly after birth, colostrum can be expressed from
neonatal/prepubertal and postpubertal phases. Develop- the infant’s mammary glands. This is attributed to the
ment of the breast can then proceed through a number of pro-lactation hormones present in the fetal circulation at
lactation cycles (pregnancy, lactogenesis I, lactogenesis birth. Regression of the mammary gland usually occurs
II, and involution; (see review by Kent on page 564). by 4 weeks postpartum and coincides with a decrease in
the secretion of prolactin from the anterior pituitary
gland of the infant.4
Address correspondence to Donna T. Geddes, PhD, Department of Bio-
chemistry and Molecular Biology, School of Biomedical, Biomolecular Neonatal and Prepubertal Development
and Chemical Sciences, Faculty of Life and Physical Sciences, M310 –The
University of Western Australia, 35 Stirling Highway, Crawley WA 6009 The ducts in the newborn breast are rudimentary and
Australia. E-mail: [email protected] have small, club-like ends that regress soon after birth.

556 Volume 52, No. 6, November/December 2007

© 2007 by the American College of Nurse-Midwives 1526-9523/07/$32.00 • doi:10.1016/j.jmwh.2007.05.004
Issued by Elsevier Inc.
Before puberty, the growth of the breast is isometric.
Allometric growth of both the stroma and epithelium
begins with the onset of puberty (8 –12 years of age).
Although the impact of obesity on breast development at
this stage is unknown, it is of interest that ruminants fed
a diet that is higher than their energy requirements have
impaired mammary development and subsequent im-
paired lactation performance.8,9

Puberty
At puberty, the increase in breast size is mainly caused
by the increased deposition of adipose tissue within the
gland. However, progressive elongation and branching of
the ducts creates a more extensive ductal network.10 The
major site of growth is the bud-like structures at the end
of the ducts, and these form the terminal duct lobular
units or acini.11 Although knowledge of the hormonal
regulation of mammary growth during puberty is not
extensive, these maturational changes are associated with
increased plasma concentrations of oestrogen, prolactin, Figure 1. Artist’s impression of the ductal system of the human lactating
breast. The ductal system was injected with coloured wax and
luteinizing hormone, follicle stimulating hormone, and
dissected. (Reproduced from Cooper.16)
growth hormone.12,13

Menstrual Cycle Changes Although it is generally thought that each lobe is a single
entity, a recent study that created three-dimensional
During the follicular phase of the menstrual cycle, the
reconstructions of the entire ductal system (16 lobes) of
lobules are small, with few alveoli, and there is low
a mastectomized breast of a 69-year-old female was able
mitotic activity. During the luteal phase, the lobules and
to demonstrate two connections between lobes.19 It is
alveoli develop with open lumens and mitotic activity is
generally believed that 15 to 25 ducts drain the alveoli
at its greatest.14 From day 27 to menstruation, these
and merge into larger ducts that eventually converge into
changes regress. However, the degeneration of the epi-
one main milk duct which dilates slightly to form the
thelial growth is not complete,15 and some of the follic-
lactiferous sinus before narrowing as it passes through
ular growth remains until the next cycle. With increasing
the nipple and opens onto the nipple surface (Figure 2).
years, there is a relative decrease in mitotic activity until
Recent histologic sections of mastectomy nipples have
about 35 years of age, when breast development pla-
shown more than 17 ducts on average18,20; however, it is
teaus.4
not known whether these are all patent, and others
suggest the average number of ducts is lower (5–9).21
GROSS ANATOMY OF THE NON-LACTATING BREAST
The diameters of the main ducts in the non-lactating
For the past 160 years, the descriptions of the anatomy of breast as measured by ultrasound are between 1.2 mm
the breast have changed little since Sir Astley Cooper’s16 and 2.5 mm in diameter. Dilated ducts in the non-
meticulous dissections of breasts of women who were lactating breast may be caused by conditions such as
lactating when they died (Figure 1). polycystic ovarian disease22 or ductal ectasia. The nipple
The breast is composed of glandular (secretory) and pores are 0.4 mm to 0.7 mm in diameter and are
adipose (fatty) tissue, and is supported by a loose surrounded by circular muscle fibres.4,5
framework of fibrous connective tissue called Cooper’s The heterogeneous distribution of glandular and adi-
ligaments. Traditional descriptions of breast anatomy pose tissue in the breast has hindered measurement of
describe the glandular tissue as consisting of 15 to 20 these tissues. However, the ratio of glandular to adipose
lobes that are comprised of lobules containing between tissue estimated by mammography is 1:1 on average, and
10 and 100 alveoli that are approximately 0.12 mm in it is well documented that the proportion of glandular
diameter17 (Figure 2). The size of each lobe is extremely tissue declines with both advancing age23 and increasing
variable, and some lobes may differ by 20-to 30-fold.18 breast size.24

Arterial Supply
Donna T. Geddes, PhD, is a research associate in the School of Biomedical,
Biomolecular and Chemical Sciences at The University of Western The blood supply to the breast is provided mainly by the
Australia. anterior and posterior medial branches of the internal

Journal of Midwifery & Women’s Health • www.jmwh.org 557

Figure 2. A, Traditional schematic diagram of the anatomy of the breast. The main milk ducts below the nipple are depicted as dilated portions or lactiferous
sinuses, and the glandular tissue is deeper within the breast. B, Schematic diagram of the ductal anatomy of the breast based on the findings
of Ramsay et al.38 Milk ducts are shown to be small and branch a short distance from the base of the nipple. The ductal system is erratic and
glandular tissue is situated directly beneath the nipple. (Reproduced with permission from Medela AG.)

mammary artery (60%) and the lateral mammary branch Lymphatic Drainage
of the lateral thoracic artery (30%).25 Smaller sources of
The drainage of lymph from the breast has been exten-
arterial blood include the posterior intercostal arteries
sively investigated with particular reference to breast
and the pectoral branch of the thoracoacromial artery.5
carcinoma, and there are two main pathways by which
There is wide variation in the proportion of blood
lymph is drained from the breast. The first is to the
supplied by each artery between women,26 and little
axillary nodes; the second is to the internal mammary
evidence of symmetry between breasts. Moreover, the
nodes. The majority of the lymph from both the medial
course of the arteries does not appear to be associated
and lateral portions of the breast is drained to the axillary
with the ductal system of the breast.4
nodes (75%), whereas the internal mammary nodes
receive lymph from the deep portion of the breast.
Venous Drainage However, as expected, there is a wide variation in the
drainage of lymph from the breast, and less common
The venous drainage of the breast is divided into the deep pathways have been demonstrated.5
and superficial systems which are joined by short con-
necting veins. Both systems drain into the internal
thoracic, axillary, and cephalic veins. The deep veins are PREGNANCY
assumed to follow the corresponding mammary arteries, During the first half of pregnancy, extension and branch-
while the superficial plexus consists of subareolar veins ing of the ductal system occurs, along with intensified
that arise radially from the nipple and drain into the lobular–alveolar growth (mammogenesis). Growth of the
periareolar vein, which circles the nipple and connects mammary gland is influenced by a number of hormones,
the superficial and deep plexus. Symmetry of the super- including oestrogen, progesterone, prolactin, growth hor-
ficial venous plexus is not apparent.25 mone, epidermal growth factor, fibroblast growth factor,
insulin-like growth factor,28,29 and parathyroid hormone–
related protein.30 Growth of the glandular tissue is be-
Innervation
lieved to occur by invasion of the adipose tissue.4 By
Cooper16 showed that the 2nd to 6th intercostal nerves mid-pregnancy, there is some secretory development,
supply the breast. The distribution and course of these with colostrum present in the alveoli and milk ducts. In
nerves are complex and variable. The anterior nerves the last trimester, there is a further increase in lobular
take a superficial course in the subcutaneous tissues, size.
while the lateral nerves travel a deep course through the While these changes typically lead to a marked in-
breast. The nipple and areola are supplied by the anterior crease in breast size during pregnancy, the proportion of
and lateral cutaneous branches of the 3rd to 5th intercos- growth varies greatly between women, ranging from
tal nerves most commonly the 4th intercostal nerve.27 little or no increase to a considerable increase in size.

558 Volume 52, No. 6, November/December 2007

While the major increase in breast size is usually com- described the milk ducts as being significantly larger
pleted by week 22 of pregnancy, significant breast compared to those of the non-lactating breast. In contrast,
growth occurs during the last trimester of pregnancy in Cardenosa and Eklund35 have reported that the ducts do
some women, and some women undergo significant not enlarge during lactation.
breast growth postpartum. At the end of pregnancy, the To date, both computed tomography and magnetic
volume of breast tissue had increased by 145 ⫾ 19 ml resonance imaging have had little to offer in elucidating
(mean ⫾ standard error of the mean; n ⫽ 13; range, mammary anatomy. However, in two recent studies that
12–227 ml), with a further increase to 211 ⫾ 16 ml (n ⫽ used magnetic resonance imaging to image the breast,
12; range, 129 –320 ml) by 1 month of lactation. The rate one study was able to identify some central ducts in the
of growth of the mother’s breast during pregnancy is breasts of lactating women,36 and another attempted to
correlated with the increase in the concentration of quantify fatty and glandular tissue volumes in the breasts
human placental lactogen in the mother’s blood, which of non-lactating women.37 These findings suggest that
suggests that this hormone stimulates breast growth in this modality may offer some new insights into the
women.31 anatomy of the breast in future.
During pregnancy, mammary blood flow approxi- Our laboratory has recently reinvestigated the anatomy
mately doubles in volume. This increased blood flow is of the lactating breast using high-resolution ultrasound.38
concomitant with both the increased metabolic activity Ultrasound is non-invasive and allows the structures of
and temperature of the breast. This elevation in blood the breast to be examined without distortion. Compared
flow persists during lactation and appears to decline to with the quoted 15 to 25 ducts of conventional texts,4,5
prepregnancy levels about 2 weeks after weaning.32 fewer ducts were imaged with ultrasound (mean, 9;
range, 4 – 8), which concurs with both Love and Bar-
GROSS ANATOMY OF THE LACTATING BREAST sky’s21 observations of lactating women expressing milk
with a breast pump (mean, 5; range, 1–17) and Going and
The breast reaches its full functional capacity at lactation, Moffatt’s39 dissection of a nipple (4 patent ducts) from a
and as a result, several internal and external changes
woman who was lactating. Interestingly, these are in
occur. During pregnancy, the areola darkens in colour,
agreement with Cooper,16 who found 7 to 12 patent ducts
and the Montgomery glands, which are a combination of
in cadaver dissections of breast from a woman who was
sebaceous glands and mammary milk glands, increase in
lactating before death, although he could cannulate up to
size. The secretions of these glands, which number
22 ducts.
between 1 and 15,33 are thought to provide maternal
Ultrasound imaging has also elucidated other charac-
protection from both the mechanical stress of sucking
and pathogenic invasion. In addition, it is also suspected teristics of the milk ducts, in that they are small (mean, 2
the secretion may act as a means of communication with mm), superficial, and easily compressed. In addition,
the infant via odor. In this connection, a recent study they do not display the typical sac like appearance of the
demonstrated that increased numbers of Montgomery “lactiferous sinus” originally thought to exist (Figure 3).
glands is associated with increased infant weight gain in Instead, branches drain glandular tissue located directly
the first 3 days after birth, infant breastfeeding behaviour beneath the nipple and often merge into the main
(increased latching speed and sucking activity), and collecting duct very close to the nipple38 (Figure 3).
decreased time to onset of lactation in primiparous Furthermore, the milk ducts increase in diameter at milk
mothers,33 suggesting that there is indeed a functional ejection,40 leading to the conclusion that it is likely that
role of the Montgomery glands during lactation. the main function of the ducts is the transport rather than
The standard descriptions of the human breast are storage of milk. In addition, the actual course of the ducts
based on Cooper’s16 magnificent cadaver dissections of from the nipple into the breast is erratic, and they are
the breasts of women who were lactating at the time of intertwined much like the roots of a tree38 (Figure 1),
death. Although imaging modalities have become more making them difficult to separate surgically.5
sophisticated, research has focused extensively on abnor- It is widely believed that the lactating breast is
malities of the non-lactating breast. Mammography of predominantly composed of glandular tissue during lac-
the lactating breast is limited because of the increase in tation. Using a semi-quantitative ultrasound measure-
glandular tissue and the secretion of breast milk, which ment of the glandular and adipose tissues in the breast,
causes an increase in radiodensity, making images of the we have found there to be approximately twice as much
breast difficult to interpret.34 glandular tissue as adipose tissue in the lactating breast.
Galactography (the injection of radio-opaque contrast However, there is great variability, and in some women,
media into the duct orifice at the nipple and subsequent up to half of the breast is comprised of adipose tissue. In
radiography) has illustrated only a portion of the ductal addition, the amount of fat situated between the glandular
system, and few studies have examined lactating women. tissues is highly variable. At this stage, no relationship
Of those that have looked at lactating breasts, some have between the amount of glandular tissue in the breast and

Journal of Midwifery & Women’s Health • www.jmwh.org 559

 in mothers with very low milk supply, breast hypoplasia
(too little glandular tissue), or hyperplasia (overgrowth of
glandular tissue). With rising rates of obesity, there is
some concern about the effect of obesity on lactation,
particularly with increasing reports that obese women are
experiencing breastfeeding difficulties (see review by
Jevitt et al. on page 606). Only a few studies have been
carried out to investigate the effect of obesity on lacta-
tion, and they have been difficult to perform because of
known confounding factors such as the mode of delivery
and parity. These studies show that pregnant women with
a high body mass index are more likely to experience
delayed lactogenesis II.44 While the cause of delayed
lactation is not clear, hormonal influences on milk
production, increased difficulty attaining a successful
infant latch to the breast, and socio-cultural factors have
been suggested.45
Knowledge of the normal features of the ductal system
is integral to diagnosing ductal abnormalities such as
galactoceles and blocked ducts. A palpable lump and
ultrasonic features of non-compressible ducts is indica-
tive of a blocked duct and should not be considered
“normal” for the lactating breast. Furthermore, the ultra-
sound scan may identify the level of the blockage,
Figure 3. A and B, Ultrasound image of a main milk duct (Toshiba, Aplio). providing useful information for treatment with thera-
The nipple is the round hypoechoic (dark) structure in the left of peutic ultrasound.
the image (N). The main duct (M) branches into two ducts
Mothers of premature and sick infants rely on breast
(B) approximately 5 mm from the nipple. Note the small
diameter of the ducts (approximately 3 mm).
pumps to initiate and maintain lactation. Clinically, it has
been observed that larger shield sizes may optimize milk
removal for some mothers. It is therefore feasible that
either the storage capacity of the breast or milk produc- compression of superficial ducts within the breast by the
tion has been demonstrated.38 shield may indeed compromise milk flow. Further re-
Nerve fibres have been identified in association with search is required to determine the effect of ductal
the major duct system in the lactating breast; however, anatomy on pumping performance in women.
they are sparse in the region of the smaller ducts, areola, Many women who have breast reduction surgery may
and nipple.41 Although these nerves are sensory, apart be able to partially breastfeed their infant, but relatively
from a marked increase in areola and nipple sensitivity few are able to exclusively breastfeed.52 This is likely
within the first 24 hours postpartum,42 women tend not to because of the codistribution of glandular and fatty tissue
be particularly sensitive to breast changes associated within both the lactating38 and non-lactating breast,53
with some abnormal conditions. For example, women making it difficult to preferentially remove fatty tissue. In
with mastitis often experience influenza-like symptoms addition, milk outflow is probably disrupted, because
before becoming aware of breast tenderness or the there are fewer numbers of ducts than previously
presence of a blocked duct. In addition, the absence of thought.21,38 Furthermore, it is possible that the milk
motor innervation of the lactocyte and glandular tissue ejection reflex may be inhibited if the nerve supply to the
further supports that the production of milk is indepen- nipple is disturbed.
dent of neural stimulation.43 It should be noted, however, The absence of lactiferous sinuses or milk reservoirs
that as with the gross anatomy of the breast, the nerve leads one to reconsider the mechanism by which the
supply of the breast has not been extensively investigated infant removes milk from the breast. Generally, it is
recently. believed that the predominant action involved in remov-
These new findings with regard to breast anatomy ing milk from the breast is peristalsis or a stripping
have several clinical implications (Table 1). For exam- action.54 We have found that milk flows into the infant’s
ple, abnormalities of the lactating breast have not been mouth when its tongue is lowered and vacuum is applied
extensively investigated in comparison to the pathologi- to the breast. This finding suggests that the vacuum
cal, non-lactating breast. Although ultrasound imaging is applied by the breastfeeding infant is a major component
only semi-quantitative and can be subjective, it may of milk removal.55 Indeed, it is evident that correct
provide information on the proportion of glandular tissue positioning and attachment of the infant to the breast is

560 Volume 52, No. 6, November/December 2007

 Table 1. Summary of Some of the Common Breastfeeding Conditions and Symptoms and the Connection With Breast Anatomy
Clinical Condition Symptoms Anatomical Relationship
Glandular anomaly
Hypoplasia Low milk production Possible deficiency of glandular tissue
Hyperplasia Excessive breast growth, Excess glandular tissue
lymphoderma, possible necrosis
Breast surgery
Reduction Low milk production Large volume of glandular tissue removed, severing milk ducts (fewer in number than
mammoplasty previously thought); possible nerve damage inhibiting milk ejection reflex
Breast augmentation Low milk production Possible compression of milk ducts by implant
Possible deficiency in volume of glandular tissue
Palpable mass
Blocked duct Mass (small or large) ⫾ pain Compression of ducts: possible cause of blocked duct; if large lobe affected a significant
Possible reduction in milk reduction in milk production may occur; identification of the level of duct obstruction
production by ultrasound ensures treatment of entire affected area
Galactocele Mass (generally small) Possible ductal abnormality
Benign mass (cyst, Mass Possible compression of ducts causing blocked duct; possible obstruction of milk flow in
fibroadenoma) the area of attachment of the infant to the breast
Malignant mass Palpable non-resolving mass Irregular shaped mass that may be mistaken for a blocked duct or galactocele;
ultrasound ⫾ mammography needed for diagnosis
Infant sucking Ineffective suck Lack of milk sinuses and evidence that vacuum plays a major role in milk removal may
mechanism alter intervention
Milk expression Differences in efficiency of Theorized that women with large milk ducts or duct dilations at milk ejection express
pumping milk quickly
Differences in effectiveness of Poor shield fit may result in compression of superficial ducts and inhibit milk flow
pumping
Milk ejection Time of increased milk availability Small ducts lacking lactiferous sinuses do not store a large amount of milk; optimization
of milk removal during milk ejection will improve milk removal from the breast

important for successful breastfeeding; however, the though many women are able to sense the first milk
mechanism should be fully understood in order to diag- ejection, few are able to sense subsequent ones.
nose and manage infants with sucking abnormalities. While it is well known that stress can influence milk
Finally, the absence of the lactiferous sinuses further ejection—resulting in diminished amounts of milk re-
emphasises the critical nature of milk ejection for suc- moved by both the infant48 and breast pump51—it is
cessful breastfeeding, because only small amounts of often the subtle stress which affects maternal confidence
milk are available before the stimulation of milk ejec- and subsequently milk ejection that is overlooked. There-
tion.40,46 fore, it is important to provide positive support to the
mother during both breastfeeding and pumping. Another
MILK EJECTION factor that may influence milk ejection and milk removal
is the ductal anatomy of the breast. In a study of mothers
Milk ejection is critical for successful lactation, because
expressing with an electric breast pump, ultrasound was
only small volumes of milk (1–10 mL) can be either
used to image duct dilation in the breast that was not
expressed46 or removed by the breastfeeding infant40
pumped. It was found that mothers with larger ducts
before milk ejection. Failure to remove sufficient quan-
expressed more milk during milk ejection and had longer
tities of milk results in a decrease in milk production
milk ejections than mothers with smaller ducts.50 There-
because of local control mechanisms.47 Stimulation of
fore, the rate of milk removal for a mother may be
the nipple initiates milk ejection via initiation of nervous
influenced in part by her ductal anatomy.
impulses to the hypothalamus, which stimulates the
posterior pituitary gland to release oxytocin into the
CONCLUSION
bloodstream.48 Oxytocin causes the myoepithelial cells
surrounding the alveoli to contract, forcing milk into the New anatomy research has shown that the milk ducts of
ducts. This results in increased intraductal pressure,49 the breast are small, compressible, superficial, and
duct dilation40,50 (measured by ultrasound), and conse- closely intertwined. They do not display typical dilated
quently increased milk flow rate50 (measured by contin- “sinuses” and do not typically store large amounts of
uous weigh balance during breast expression). Multiple milk. In addition, the amount of adipose tissue in the
milk ejections almost always occur during breastfeed- breast is highly variable, particularly between the glan-
ing40 (mean, 2.5; range, 0 –9) and breast expression50 dular tissue. This fundamental knowledge of the anatomy
(mean, 3– 6 for 15-minute expression period), and al- of the breast—particularly when it is fully functional—

Journal of Midwifery & Women’s Health • www.jmwh.org 561

allows one to begin to understand and the myriad of 15. Longacre TA, Bartwo SA. A correlative morphologic study
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Orme, Green, Browne and Longmans, 1840.
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