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Mechanisms and For the purposes of this review, we will outline the
biochemical, hormonal and mechanical phases of normal spon-
management of normal taneous labour and vaginal delivery, before describing how these
may be influenced at different points by intervention, or by the
OBSTETRICS, GYNAECOLOGY AND REPRODUCTIVE MEDICINE 30:3 84 Crown Copyright Ó 2020 Published by Elsevier Ltd. All rights reserved.
REVIEW
Oestrogen
CRH
Early in pregnancy, oestrogen augments the effect of progester-
one through gene regulation. This role shifts near term as the
oestrogen:progesterone ratio increases. At this stage oestrogen
enhances oxytocin receptor expression.
Oxytocin
Oe
Oxytocin plays a role in the activation of myometrial smooth
muscle, and in cervical remodelling. Response to oxytocin de-
pends on gestation and phase of parturition, reflective of the
Ox
progesterone:oestrogen balance at the time. Oxytocin receptor
concentration increases significantly as term approaches, and is
at its highest in early labour. Oxytocin receptors are distributed
in the highest concentration at the uterine fundus and in the
decidua, followed by the lower segment, and finally by the
R cervix.
PG Oxytocin has both direct and indirect actions. It acts directly
on the myometrium to produce regular, effective contractions,
P and indirectly on the decidua to increase production of
prostaglandins.
CRH, Corticotropin-releasing hormone; Oe, Oestrogen;
Ox, Oxytocin; PG, Prostaglandins; R, Relaxin; P, Progesterone Relaxin
Relaxin is produced by the corpus luteum, the placenta and
Figure 1 decidua. Levels of relaxin peak at the end of the first trimester,
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REVIEW
fall, and then rise again immediately prior to delivery. Relaxin The net result of the changes detailed above is increased
acts on G protein-coupled receptors, similar to those of Lutei- cervical compliance and cervical response to an increase in intra-
nising Hormone and Follicle Stimulating Hormone. Relaxin tar- uterine pressure, due to progressive uterine contractions. The
gets the cervix, myometrium, endometrium and decidua. It acts cervix becomes progressively effaced and dilated to 10 cm, to
mainly to suppress uterine contractility and later plays a role in allow delivery of the fetus and placenta.
remodelling cervical connective tissue in preparation for labour. The cervix may be manipulated both chemically and me-
chanically in induction of labour. Common methods include
Mechanical processes of labour vaginal prostaglandins and cervical balloon catheters.
Rosa stated that “delivery can occur only when the uterine
Uterus
contraction, the axis of engagement, the descending fetus and
the pelvic diameters all fit together”. With this in mind, one The uterus is predominantly composed of smooth muscle and
thinks of the three Ps e Power, the Passenger and the Passage. changes shape from the first few weeks of pregnancy to accom-
The intimate relationship between these components is key to modate the growing fetus and prepare for its eventual expulsion.
a competent understanding of the mechanical process of As outlined in further detail below, processes within the
labour. myometrium generate electrical activity which leads to the con-
We will first explore the preparatory mechanical changes tractile forces required for labour. These processes are controlled
before looking at the three Ps in more detail. by hormonal, metabolic and mechanical factors.
Although the uterus receives both parasympathetic and
Cervix sympathetic innervation, it is a myogenic organ, i.e. it can con-
tract without nervous stimulation. Uterine myocytes are rela-
The cervix is composed of fibrous connective tissue, collagen
tively large, in keeping with the powerful contractile forces
(types I, III and IV), elastin, vasculature, fibroblasts and minimal
required during labour. They are connected by gap junctions,
smooth muscle. The proportion of collagen decreases with
which increase in number as pregnancy progresses and thereby
increasing gestation.
increase the capacity of the uterus to contract. Uterine contrac-
Cervical ripening is a product of increased vascularity, stro-
tions occur due to the generation of action potential within the
mal and glandular hypertrophy, and changes in the cervical
myocyte. When threshold potential has been reached, voltage-
extracellular matrix by matrix metallo-proteases, proteases and
dependent channels are activated, increasing intracellular cal-
collagenases. Cervical nerve density is sustained or increased at
cium (Ca2þ) and therefore action potential. Opening of voltage-
term, and alterations in this signalling pathway have been noted
dependent potassium (Kþ) channels leads to outward Kþ cur-
to delay cervical ripening and reduce immune cell infiltration in
rent, inactivating L-type Ca2þ channels, and leading to repolar-
animal studies.
ization. It is thought that myocyte resting membrane potential
The structural integrity of the cervix is reduced following a
becomes more depolarized as gestation advances, facilitating
progressive decrease in the number of crosslinks between
Ca2þ influx and increasing the frequency of myometrial con-
collagen helices and a switch from straight to wavy fibres, which
tractions. Reductions in Kþ channels near term also prolong
are further disorganized by the infiltration of macrophages and
myocyte action potentials and enhance contractility.
neutrophils and an increase in hyaluronic acid production. This
Intracellular Ca2þ forms a complex with calmodulin and ac-
effect is compounded by an influx of water, which occurs due
tivates myosin light-chain kinase (MLCK), leading to phosphor-
to an increase in decorin expression. Concurrently, there is
ylation of myosin and cross-bridge cycling with actin
decreased expression of collagen synthesis enzymes and pro-
myofilaments, generating a contraction. In this way, myometrial
teins, such as lysyl hydroxylase, lysyl oxidase, thrombospondin
contractions may be suppressed by blocking voltage-dependent
2, and tenascin C. Genetic mutations in this synthetic pathway
L-type Ca2þ channels (dihydropyridine receptors); this is the
are thought to be associated with cervical insufficiency, itself a
mechanism for current tocolytics, such as Nifedipine.
major player in miscarriage and pre-term birth. Infection, lead-
Myocytes are noted to have proportionately large Sarco-
ing to premature cervical remodelling, is associated with 25
plasmic reticula (SR). SR not only store Ca2þ, but also affect
e40% of preterm birth. Cervical length surveillance is another
smooth muscle excitability. Ca2þ released by SR augments
focus in an effort to identify those at increased risk of pre-term
cytosolic Ca2þ, and contributes to the actions of oxytocin,
delivery.
vasopressin and PG, thereby strengthening uterine contractions.
Inflammatory processes continue throughout labour.
SR Ca2þ release is regulated by intracellular pH and metabolites.
Increased activity of mediators such as IL-1 and IL-6. IL-6 stim-
Interestingly, during spontaneous labour, Ca2þ entry and efflux
ulates prostaglandin and leukotriene production, leading to cer-
across the plasma membrane is responsible for uterine contrac-
vical vessel dilatation and further inflammatory cell
tions, and there the SR does not contribute.
extravasation.
Oxytocin binds to a receptor in the cell membrane, thereby
Calcitonin gene-related peptide (CGRP) acts as a vasodilator,
inducing an increase in cytosolic Ca2þ, depolarizing the mito-
and as sensitivity to CGRP rises during pregnancy, it is thought to
chondrial membrane potential and enhancing contractility. Whilst
contribute not only to vascular remodelling in early pregnancy,
the importance of Ca2þ is apparent, Kþ conductance is the main
but also towards uterine quiescence, and eventually to vasodi-
determinant of resting membrane potential in myocytes. Manipu-
lation during cervical ripening. Substance P has a significant role
lation of Kþ channels affects the duration of the action potential
in the promotion of processes required for cervical ripening, and
and repolarizsation. Kþ channel activity is associated with uterine
can also induce uterine contractions.
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REVIEW
quiescence, and downregulation of KATP occurs in preparation for Perineum and vagina
labour. TREK-1 K channels respond to myometrial stretching by
Perineal and vaginal muscle vascularity increases throughout
increasing outward Kþ current. However, their activity is limited
pregnancy. Perineal connective tissue softens and smooth muscle
until gene expression increases close to term, to allow for uterine
cells hypertrophy to enable distension during labour.
distension and accommodation of fetal growth.
A subunit of Kþ voltage dependent channels is thought to be
expressed in lower levels in obese women. This may be com- Pelvis
pounded by the fact that elevated cholesterol levels also increase The ischial spines are a landmark for assessing descent of the
outward Kþ current. Both pathways result in increased Kþ fetal presenting part. Table 1 and Figure 2 summarize the di-
conductance and supposedly poorer myometrial activity, which mensions of the pelvis.
may contribute to the fact that obese women are more likely to
require delivery by caesarean section. Power
It has been suggested that uterine myocytes have inherent
pacemaking activity, however, to date, no anatomical or histo- The mechanical principles behind contraction production on a
logical pacemaking structure has been identified. cellular level have already been detailed. Contractions alter the in-
Contractions increase in amplitude and frequency as labour trauterine pressure, thereby exerting force on both the fetus and the
progresses. In the process myometrial vessels are compressed and more pliable soft tissues, leading initially to cervical dilation, and
occluded. The phasic nature of uterine contractions is therefore then combining with maternal effort to produce expulsive force.
important, so as to prevent prolonged myometrial hypoxia, leading
to fetal asphyxia. As contractions reach their peak, lactic acid builds Passage
up within the myocytes, affecting the Ca2þ current and decreasing
the intensity of the contraction to baseline. The following rest period Primarily the cervix must ripen and dilate. Maintaining an up-
right position during the first stage of labour is thought to
is essential in order to reverse hypoxic metabolic changes and
restore pH, before the action potential is generated again. shorten this process, however, the mechanics of this have not
Short cycles of hypoxia are thought to maintain the progressive been thoroughly elicited. Thereafter there are three bony
augmentation of contractions required for labour. This is often landmarks (Table 1), which may present a challenge to the fetal
referred to as hypoxia-induced force increase (HIFI). Parturition presenting part. Maternal position may also assist in this re-
requires significant preparation, not only with respect to myometrial gard, for example the use of McRobert’s position to drop the
hypertrophy and increases in contractile proteins, but also glycogen sacral promontory and thereby assist delivery in cases of
and free fatty acid storage ahead of the hypoxic challenges of labour. shoulder dystocia.
Strict control of Ca2þ and pH is necessary for uterine function. Pelvic floor soft tissue must also adequately stretch to accom-
Oxidative phosphorylation provides the energy required for con- modate delivery. There are mixed opinions as to whether high
tractions, while anaerobic metabolism supports ionic regulation. pelvic tone aids the fetal passage, or obstructs it. It is thought that
it may guide presentation, as well as passively stretching as
Lactate is produced by the myometrium even in the presence
of sufficient oxygen and despite transportation out of the myo- required. Unsurprisingly, it has been demonstrated that the peri-
cyte, continues to have an inhibitory effect on contractility. neal muscle force on the fetus increases as it descends through the
Accumulation of extracellular lactate is thought to contribute pelvis. Consequently, those who deliver vaginally have been
towards dystocia, and a recent trial has proposed that ingestion observed to have lower levels of pelvic tone than those who were
of oral bicarbonate may have a therapeutic role in this regard. delivered by Caesarean Section due to failed induction.
The changes detailed above are rapidly reversed in involution,
where progressive remodelling of the uterine collagen and elastin Passenger
content occurs. The uterus is back to near pre-pregnancy size in The diameter of the fetal head poses the greatest mechanical
about 6 weeks, due to changes in endocrine levels and removal challenge, and so a cephalic, fully-flexed presentation is optimal.
of mechanical stretch. If an infant’s head is successfully delivered, the healthcare
Inlet Sacral promontory, linea 12.7 (Longest distance between 11.5 (Iliopectineal eminence -> 10 (Obstetric conjugate: sacral
terminalis, superior rami of the the bilateral linea terminalis) sacroiliac joint) promontory -> symphysis pubis)
pubic bones, symphysis pubis
Mid Ischial spines 11.5 11.5 (lower sacroiliac joint -> 11.5 (S3 ->midpoint of pubic
midpoint of obturator membrane) symphysis)
Outlet Base at level of line drawn 10 (between ischial tuberosities) 11.5 12.7 (Pubis -> sacrococcygeal
between the ischial tuberosities, joint)
tip of sacrum, subpubic area
Table 1
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The relationship of the pelvic dimensions Diameter of presenting part and important landmarks of
the fetal skull
Inlet Presenting part of fetal skull Diameter
Outlet Table 2
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Variations of the fetal head position when the Descent and delivery of the fetal head during
presenting part is cephalic labour
Advancement
– to the ischial spines,
ROT LOT rotation and flexion
ROP DOP LOP of the fetal head
– concurrently the
fetal shoulders enter
the pelvis
Delivery
– of the fetal head
following extension
Figure 3
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REVIEW
- Following delivery, the infant should be dried and stimulated provider, it is important to respect and acknowledge an indi-
as appropriate, and transferred to the mother for skin-to-skin vidual woman’s choices, whilst providing her with evidence-
contact. based information and advice. Robust knowledge of the mecha-
- Obstetric review should be sought if the third stage is pro- nisms and management of normal labour is key to our under-
longed, at 30 min for active management, and at 1 h for standing of when to clinically intervene, and recognize areas
physiological management. where maternal and fetal morbidity can be reduced. A
Monitoring
FURTHER READING
NICE do not recommend continuous monitoring in low risk Buhimschi I, Malinow A, Saade GR, Garfield R, Weiner C. The forces of
women, due to increased rates of intervention, without a recip- labour. Fetal Matern Med Rev 2003; 14: 273e307. https://doi.org/
rocal reduction in adverse outcome. For this reason, it is rec- 10.1017/S0965539503001128.
ommended that low risk women receive intermittent Desseauve D, Fradet L, Lacouture P, Pierre F. Position for labor and
auscultation of the fetal heart, for at least 1 min following each birth: state of knowledge and biomechanical perspectives. Eur J
contraction. Fetal concerns that indicate escalation to continuous Obstet Gynecol Reprod Biol 2017 Jan; 208: 46e54. https://doi.org/
monitoring include rising baseline or audible decelerations. 10.1016/j.ejogrb.2016.11.006. Epub 2016 Nov 15.
Maternal concerns that indicate escalation of monitoring are Dodd JM, Crowther CA, Huertas E, Guise J-M, Horey D. Planned
varied, and include hypertension, proteinuria, vaginal bleeding, elective repeat caesarean section versus planned vaginal birth for
constant abdominal pain, pyrexia or suspected sepsis, contrac- women with a previous caesarean birth. [CD004224]. Cochrane
tions lasting longer than 60 s, or more than five contractions per Database Syst Rev, 2013.
minute, and dystocia. Inducing labour: NICE Clinical Guideline 70. et. NICE, 2008.
Intrapartum care for healthy women and babies. NICE Clinical
Particular circumstances Guideline 190. et. NICE, 2017.
Irani RA, Foster S. Overview of the mechanisms of induction of labour.
Induction of labour
Semin Perinatol 2015 Oct; 39: 426e9. https://doi.org/10.1053/j.
Induction of labour is indicated in certain cases where shortening
semperi.2015.07.001. Epub 2015 Sep. 11.
the pregnancy has been shown to reduce maternal and fetal
Kamel RM. The onset of human parturition. Arch Gynecol Obstet 2010
morbidity. Although not strictly considered ‘normal’, induction
Jun; 281: 975e82. https://doi.org/10.1007/s00404-010-1365-9.
of labour may often be the most realistic means of achieving a
Epub 2010 Feb 3.
‘normal’ vaginal delivery. The process is commenced with
Olson DM, Ammann C. Role of the prostaglandins in labour and
exogenous prostaglandin pessaries, or mechanical dilatation de-
prostaglandin receptor inhibitors in the prevention of preterm la-
vices, to encourage cervical ripening. Endogenous prostaglandin
bour. Front Biosci 2007 Jan; 12: 1329e43.
release is then stimulated by amniotomy, and once the cervix is
World Health Organization Human Reproduction Programme, 10 April
sufficiently ‘ripe’, synthetic oxytocin infusion is commenced to
2015. WHO statement on caesarean section rates. Reprod Health
stimulate regular, strong contractions. Concurrently, oxytocin
Matters 2015; 23: 149e50.
also binds to PG receptors in the decidua and cervix to activate
Wray S, Prendergast C. The myometrium: from excitation to con-
further prostaglandin release.
tractions and labour. In: Hashitani H, Lang R, eds. Smooth muscle
Vaginal birth after caesarean section spontaneous activity. Advances in experimental medicine and
With increasing rates of caesarean section across the world, biologyvol. 1124. Singapore: Springer, 2019.
vaginal birth after caesarean section (VBAC, also known as
TOLAC) is now considered a normal means of delivery in many Practice points
centres. A review conducted in 2013 compared planned
caesarean section to VBAC, and concluded the presence of risks C Labour is dependent on the interplay between biochemical, hor-
and benefits with both options. During VBAC, there is a small monal and mechanical factors which prepare for parturition far in
risk of uterine rupture and thus this necessitates continuous advance of established labour.
monitoring throughout labour, which in turn increases the rate of C Successful vaginal delivery requires cervical effacement and
intervention in labouring women. dilatation, progressive and effective uterine contractions and
concordance between the diameters of the fetal presenting part
Conclusion and the maternal pelvis.
C Knowledge of the rate of progress expected during normal labour
Labour is a varied experience for women and there is often an is essential for the correct diagnosis of dystocia, and as such,
overwhelming amount of information available from different, when escalation of care and intervention are required.
and sometimes contradicting sources. As the clinical care
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