Editorial

VOLUME 9 ISSUE 6 2013 201

inf ant

REVIEW

Oxytocin – a system activator

xytocin, a small peptide of just nine

amino acids, is normally associated

with labour and the milk ejection reflex.

However, oxytocin is not only a hormone

but also a neurotransmitter and a

paracrine substance in the brain

1,2

. During

breastfeeding it is released into the brain of

both mother and infant where it induces a

great variety of functional responses.

Through three different release pathways

(

FIGURE 1), oxytocin functions rather like a

system activator and often influences the

release of other signalling substances such

as opioids, serotonin, dopamine and

noradrenaline. Through these activations,

different behavioural and physiological

effects are facilitated and coordinated into

adaptive patterns, which are influenced by

the type of stimuli and environmental

factors

Through the endogenous release of

oxytocin or its administration, for example

by nasal spray in humans, various types of

socially interactive behaviours can be

stimulated. These include maternal and

sexual behaviours as well as the develop-

ment of bonding and attachment

Oxytocin stimulates well-being, it induces

anti-stress effects, decreases sensitivity to

pain, decreases inflammation and

stimulates processes related to growth and

healing. In addition, repeated exposure to

oxytocin may give rise to long-term effects

by influencing the production or function

of other signalling systems. For example,

noradrenergic activity in the brain

may decrease as a consequence of the

Oxytocin effects in mothers and infants

during breastfeeding

Oxytocin integrates the function of several body systems and exerts many effects in mothers

and infants during breastfeeding. This article explains the pathways of oxytocin release and

reviews how oxytocin can affect behaviour due to its parallel release into the blood circulation

and the brain. Oxytocin levels are higher in the infant than in the mother and these levels are

affected by mode of birth. The importance of skin-to-skin contact and its association with

breastfeeding and mother-infant bonding is discussed.

Kerstin Uvnäs Moberg

MD, PhD

Professor of Physiology

Swedish University of Agriculture

[email protected]

Danielle K. Prime

PhD

Breastfeeding Research Associate

Medela AG, Baar, Switzerland

Keywords

oxytocin; breastfeeding; skin-to-skin;

anti-stress; milk ejection; bonding

Key points

Uvnäs Moberg K., Prime D.K. Oxytocin

effects in mothers and infants during

breastfeeding. Infan t 2013; 9(6): 201-06.

1. Oxytocin is released in the mother and

infant during breastfeeding and skin-to-

skin contact.

2. Milk ejection patterns vary between

women.

3. Oxytocin is released into circulating

blood and brain structures, in parallel.

4. Oxytocin levels are higher in the infant

than in the mother and differ with

mode of birth.

increased function of inhibitory alpha-2

adrenoceptors

The regulation of the release of oxytocin

is complex and can be affected by different

types of sensory inputs, by hormones such

as oestrogen and even by the oxytocin

molecule itself. This article will focus on

four major sensory input nervous

pathways (

FIGURES 2 and 3) activated by:

1. Sucking of the mother’s nipple, in which

the sensory nerves originate in the

breast.

2. Sucking in the infant, in which the

sensory nerves originate in the infant

oral mucosa.

3. The presence of food in the

gastrointestinal tract, affecting vagal

sensory nerves.

4. Skin-to-skin contact in both mothers

and infants, in which the sensory nerves

that originate in the skin respond to

warmth, touch, stroking and light

pressure.

The mother

Milk ejection

Oxytocin is critical for milk removal in,

perhaps, its most renowned role: the milk

ejection reflex. Following sucking, the

release of oxytocin causes the contraction

of myoepithelial cells in the breast pushing

milk from the alveoli, through the milk

ducts and toward the nipple.

In general, it takes around a minute of

infant sucking or stimulation with a breast

pump before milk ejection occurs

5,6

Interestingly, the milk ejection reflex can be

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202 VOLUME 9 ISSUE 6 2013

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easily conditioned and many women can

experience spontaneous milk ejections

between feeds

7,8

as the sight, thought,

sound and smell of the infant (or even a

breast pump in pump-dependent mothers)

can cause milk ejection and dripping of

milk from the nipples. Indeed, it is

common that milk ejection occurs prior to

the physical attachment of the infant or a

breast pump

7,9

Milk ejection can be associated with

different sensations that vary dramatically

between women. These may be localised to

the breast such as a ‘drawing’ pain or

tingling, to more systemic sensations such

as nausea, thirst, fainting or even mental

anxiety and depression

Sucking- and pumping-induced

oxytocin release has been described as

pulsatile

7,10

. On average, oxytocin pulses

occur with 90-second intervals over the

first 10 minutes of breastfeeding in the first

few days after birth

11,12

. The amount of

oxytocin that is released within a 10

minute breastfeed is much higher at four

months when compared to four days after

FIGURE 1 The release of oxytocin from the supraoptic (SON) and paraventricular (PVN) nuclei

of the hypothalamus. Magnocellular neurons in the SON and PVN release oxytocin centrally

and peripherally from the posterior pituitary, resulting in effects that include milk ejection and

uterine contraction. Parvocellular neurons, located in the PVN, project axons directly into brain

structures involved in the regulation of social interaction, anxiety, the activity of the

hypothalamic-pituitary-adrenal axis, well-being and the autonomic nervous system (eg the

amygdala, hypothalamus, anterior pituitary, nucleus accumbens, nucleus tractus solitarius and

the dorsal vagal motor nucleus).

Magnocellular neurons

Central oxytocin release:

Dendritic extracellular diffusion

Parvocellular neurons

Central oxytocin release:

Axons project into brain

structures and spinal cord

Axons of

parvocellular neurons

Paraventricular

nucleus (PVN)

Supraoptic

nucleus (SON)

pituitary

Axons of

magnocellular

neurons

Posterior

pituitary

Capillary

Oxytocin

Peripheral oxytocin release:

Axonal release into blood to

target distant organs

birth, but there is still a strong correlation

between the amount of oxytocin released

on the two occasions within individual

women

One pulse of oxytocin is generally

associated with one milk ejection and there

is a relationship between the amount of

oxytocin released and the number of

oxytocin pulses during the first 10 minutes

of breastfeeding

. The number of milk

ejections during breastfeeding ranges from

one to 17 over a period of up to 25

minutes

6,7

. Similarly, pulsatile patterns of

two to 14 milk ejections are seen during a

15 minute breast expression

. The simil-

arity between the milk ejection pattern

resulting from breastfeeding and breast

pumping is demonstrated in

FIGURE 4.

Tracking an individual mother over

time, it appears that her pattern of milk

ejection repeats throughout the first nine

months of lactation

. If a mother has two

or three milk ejections she will continue

that pattern, removing all her milk in these

brief episodes. Other mothers may follow a

more constant pulsing profile in that they

have continuous milk ejections throughout

the milk removal session. Both of these

patterns can remove similar volumes of

milk successfully, but the second scenario

would require more time to reach that

same level of milk removal

. Similar

repeatability has been observed in

individual infants at successive breast-

feeds

; these mother-derived patterns may

explain why some infants are quick feeders

and others take a bit more time.

Oxytocin-induced effects in the brain

It is not known why mothers have such

varied milk ejection patterns but, since

oxytocin is involved in so many functions,

these release patterns may not only impact

on milk removal. Systemic oxytocin release

and its release into the brain may result in

many other outcomes involving

breastfeeding and mother-infant

interactions. For example, breastfeeding in

mothers is associated with physiological

and psychological adaptations including:

■

Increased social interaction

■

Decreased anxiety

■

Decreased cortisol levels

■

Decreased blood pressure

■

Increased gastrointestinal tract

function

15-17

These effects are exerted in the brain but

are associated with circulating oxytocin

levels, supporting the concept that

oxytocin is released systemically and into

the brain in parallel, as has previously been

demonstrated in sheep in response to

suckling

The infant

Infants produce oxytocin too and in fact,

the production of oxytocin begins in the

fetus. An understanding of the infant

oxytocin system is complicated by

difficulties in collecting repeat blood

samples and measuring circulating

oxytocin levels in the newborn.

Nevertheless, some studies exist in which

oxytocin levels have been measured in the

newborn while other studies have

demonstrated the release of oxytocin

indirectly via expression of oxytocin-linked

effect patterns. Information can also be

drawn from animal experiments, as

oxytocin exerts similar effect patterns in all

mammals.

Infant oxytocin levels are affected by the

mode of delivery

In infants born by vaginal delivery,

oxytocin levels in umbilical arterial blood

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were actually higher than in infants

born by caesarean section: 69pg/mL

(range 20-315pg/mL) vs 33pg/mL

(range 9-195pg/mL), respectively. Mothers

also had higher oxytocin levels if they

delivered vaginally, compared to caesarean

section. In the first 30 minutes after birth,

infants (born both vaginally and by

caesarean) had higher oxytocin levels than

those of mothers: approximately 30pg/mL

(range 13-158pg/mL). Over subsequent

hours and days the differences due to

mode of birth diminish, yet infants will

maintain a higher oxytocin level than their

mothers (

FIGURE 5). In addition, oxytocin

levels recorded postpartum in infants born

vaginally correlate inversely with fetal

arterial pH and also with the duration of

labour. This suggests that the elevated

oxytocin levels recorded postpartum are

linked to the stress of being born (Bystrova

et al 2013, unpublished results)

Ingestion of oxytocin from maternal milk

Human milk contains many different

hormones and growth factors and it also

contains small amounts of oxytocin. The

concentration of oxytocin in maternal milk

is approximately 8pg/mL in the first few

days after birth and then decreases with

increased milk production. Even if the

ingested oxytocin was able to survive the

acid milieu of the stomach and was

absorbed from the small intestine of the

fetus, the dilution within the circulation

would limit any significant rise of oxytocin

levels. It would also be unlikely that

transport to the brain would occur because

of the blood-brain-barrier.

Oxytocin release by sucking

While the intake of oxytocin from human

milk has negligible effects, sucking in

newborns is associated with infant

oxytocin release. In calves, the act of

sucking at the udder is associated with a

rise in oxytocin levels but not when

drinking/lapping milk from a bucket

. This

effect is caused by activation of sensory

nerves in the oral mucosa during sucking.

Furthermore, oxytocin is released when

milk reaches the gastrointestinal tract

(

FIGURE 3). Food intake is linked to release

of the gut hormone cholecystokinin (CCK)

which, via activation of the afferent

(sensory) vagal nerve fibres, triggers

oxytocin release

. In support of this, infant

plasma levels of CCK have been shown to

rise during breastfeeding

FIGURE 3 The ingestion of food into the infant’s stomach can trigger oxytocin release. Food

intake results in the release of the gut hormone cholecystokinin (CCK), which, via activation of

sensory vagal nerve fibres, results in central and peripheral oxytocin release. (Image © Medela.)

FIGURE 2 Different kinds of sensory nerves can release oxytocin during mother-infant

interaction. While various sensory nerves can initiate this pathway, the nucleus tractus

solitarius (NTS) acts as a common relay station for sensory input to the oxytocin-producing

paraventricular (PVN) and supraoptic (SON) nuclei. (Image © Medela.)

Skin-to-skin contact

Mother’s breast

Infant oral mucosa

Systemic

oxytocin

release

Oxytocin release into

brain structures

PVN

SON

NTS

Sensory nerves enter the

spinal cord/brainstem and

connect to the NTS

Oxytocin release into

brain structures

PVN

SON

Systemic

oxytocin

release

NTS

Milk entering the stomach

Vagal sensory nerve

Cholecystokinin (CCK)

The mother and infant

Skin-to-skin contact

Oxytocin can be released by activation of

several types of sensory nerves originating

from the skin, nipples, gastrointestinal tract

and urogenital tract. Light pressure,

warmth and stroking contribute to

oxytocin release caused by ‘pleasant’ or

‘non-noxious’ sensory stimulation of the

skin

When newborn infants are put on their

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204 VOLUME 9 ISSUE 6 2013

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FIGURE 4 Examples

of the pulsatile

patterns of milk

ejection during

breastfeeding and

breast expression.

Graph A

demonstrates four

pulses measured

during a breastfeed

by monitoring the

size of the milk duct

with ultrasound; as

milk is ejected

through the ductal

system the milk duct diameter expands (image courtesy of Dr Donna Geddes, Australia). Graph

B demonstrates four pulses measured during breast expression by monitoring the rate of flow

of milk; as milk is ejected through the ductal system the milk flow rate increases.

mother’s chest immediately after birth,

oxytocin is released into the maternal

circulation (

FIGURE 2). The effect is, in part,

linked to massage of the breasts by the

infant

. However, the oxytocin release that

is induced by skin-to-skin contact does not

occur in short pulses in the same way as

oxytocin induced by sucking, but rather in

a few protracted pulses. It is not associated

with milk ejection, but may instead ‘prime’

the ensuing breastfeeding interaction.

It has not yet been demonstrated that

peripheral oxytocin levels rise in response

to skin-to-skin contact in human

newborns. Even if circulating oxytocin

levels do not rise, it may be assumed that

oxytocin levels increase in the infant brain

in response to these types of stimuli. In the

following sections a number of effect

patterns, both short- and long-term will be

described. These can be attributed to

oxytocin release and/or function in the

brain and are triggered in both the mother

and the infant during breastfeeding and

skin-to-skin contact.

Social interaction, well-being and

stress levels

When the mother and the newborn infant

are placed in skin-to-skin contact after

birth, the infant expresses an inborn

breast-seeking behaviour – a ‘social

approach behaviour’

. Both mother and

infant become more socially interactive

and synchronise their interactions

Mother and infant become calmer, the

infant cries less, the pain threshold

increases, cortisol levels decrease and skin

temperature of the mother’s breast and of

the infant increases

27,28

. As discussed below,

these effects are likely to involve oxytocin

release in the brain.

Oxytocin and inhibition of stress

It is well known that cortisol release is

controlled by the hypothalamic-pituitary-

adrenal (HPA) axis. In this mechanism of

interactions, corticotrophin releasing factor

(CRF) released from the hypothalamus,

stimulates the release of adrenocortico-

trophic hormone (ACTH) from the

anterior pituitary, which in turn releases

cortisol into the circulation from the

adrenal cortex. Oxytocin can inhibit the

function of the HPA axis at each of these

levels

What is not well-known is that

noradrenergic neurons originating in the

locus coeruleus (LC) and the nucleus

tractus solitarius (NTS) in the brainstem,

exert a powerful influence on the activity

of the HPA axis by stimulating production

of CRF in the hypothalamus and activity in

the sympathetic nervous system. Oxytocin

may decrease stress levels by counteracting

the activity in these noradrenergic

pathways: administration of oxytocin or

the release of oxytocin from oxytocinergic

nerves that terminate in the LC and the

NTS, increases the number or function of

inhibitory alpha-2 adrenoceptors located

on the noradrenergic neurons

30,31

This type of oxytocin-linked, anti-stress

pattern is facilitated in certain situations,

for example, when the skin is exposed to

touch, warmth and light pressure, which

explains why mothers and newborn infants

experiencing skin-to-skin contact exhibit a

marked anti-stress pattern. The high levels

of oxytocin seen in both mothers and

infants after vaginal birth (

FIGURE 5) may

play an important role.

Bonding and attachment

The act of sucking may also enhance

bonding between mothers and newborn

infants and the infant’s attachment to their

mother. In support of this, newborn lambs

that are allowed to suck at the udder soon

after birth recognise and follow their

mothers more quickly than those that are

separated from their mothers in the early

days after birth

. It seems that neural

reflexes induced from the oral mucosa by

the touching of the nipple (

FIGURE 2) and

also the presence of colostrum in the

gastrointestinal tract are involved in the

creation of this primitive ‘attachment’

behaviour (

FIGURE 3).

Long-term effects

Research has shown that mothers who

have breastfed for several weeks have lower

basal, systolic and diastolic blood pressure

and also lower stress reactivity. This

supports the existence of a long-term, anti-

stress influence of oxytocin in humans.

The finding of a reduced risk for certain

kinds of cardiovascular disease and type 2

diabetes in mothers who have breastfed, is

further support of the connection between

repeated exposure to endogenous oxytocin

and long-term, anti-stress effects

The act of skin-to-skin contact between

mother and infant during the first two

hours after birth has been associated with

long-term outcomes. These include

enhanced interaction between mother and

infant and a better ability to handle stress

0246810

24 6 810

Milk ejections during breast expression

Minutes

Milk ejections during breastfeeding

Minutes

Milk duct diameter increase (mm)

Milk flow rate (g/s)

0.6

0.4

0.2

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inf ant

in the infant – effects which may be

measured as long as one year after birth

34,35

As an example of a mechanism behind

these long-term effects, decreased levels of

anxiety and increased social interaction

seen in rats exposed to extra sensory stim-

ulation after birth has been attributed to an

increased production/function of oxytocin

receptors in the amygdala. The anti-stress

effects have been attributed to a decreased

function of the HPA axis

Medical interventions during birth

It can be concluded that oxytocin

stimulates socially interactive behaviour

and induces anti-stress effects and that

these effects may become long lasting if

induced early in life. Many medical

interventions during birth interfere with

spontaneous oxytocin release. For example:

■

Caesarean section may be linked to a

reduced oxytocin release during labour

(or none at all if there is no labour)

■

Epidural analgesia may be linked to

reduced oxytocin release as the Ferguson

reflex is partly blocked

■

Infusions of oxytocin may influence

spontaneous oxytocin release via a feed-

back inhibitory mechanism.

Recent data suggest that such effects can be

documented at two days after birth

11,37

Development of secure attachment

Oxytocin release is easily conditioned and,

after a while, its release and the consequent

oxytocin-related effects will be triggered in

the infant by just the sight, voice or smell

of the mother. With time, the infant may

learn to hold the ‘image’ of its mother even

when she is not present and in this way the

infant may remain calm and happy even

when alone. Only when the infant becomes

fearful does it need to return to the secure

base, or in physiological terms, receive

activation of sensory nerves from the skin,

when being held close by the mother (or

another caregiver) to become happy and

calm again. In the long-term, the function

of the oxytocin system may become well

established in those infants receiving

closeness and friendly encounters by

primary caregivers: a chronic state of

satisfaction and calm develops – secure

attachment.

Conclusion

Oxytocin is an integral component of

many body systems with long-term

implications for both mother and baby. It

is not only involved in milk ejection from

the mother, but is also a key hormone for

the infant and can be influenced by skin-

to-skin contact and birthing practices.

Consolidation of current understanding

of oxytocin should encourage consider-

ation of the value of the natural

interaction between mother and infant,

at and after birth.

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37. Nissen E., Gustavsson P., Widstrom A.M., Uvnäs-

Moberg K. Oxytocin, prolactin, milk production and

their relationship with personality traits in women

after vaginal delivery or Cesarean section.

J Psychosom Obstet Gynaecol 1998;19:49-58.

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