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Uterine contractions are
muscle contraction Muscle contraction is the activation of Tension (physics), tension-generating sites within muscle cells. In physiology, muscle contraction does not necessarily mean muscle shortening because muscle tension can be produced without changes in musc ...
s of the uterine smooth muscle that can occur at various intensities in both the non-pregnant and pregnant uterine state. The non-pregnant uterus undergoes small, spontaneous contractions in addition to stronger, coordinated contractions during the
menstrual cycle The menstrual cycle is a series of natural changes in hormone production and the structures of the uterus and ovaries of the female reproductive system that makes pregnancy possible. The ovarian cycle controls the production and release of eg ...
and orgasm. Throughout
gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregn ...
, the uterus enters a state of uterine quiescence due to various neural and hormonal changes. During this state, the uterus undergoes little to no contractions, though spontaneous contractions still occur for the uterine myocyte cells to experience
hypertrophy Hypertrophy is the increase in the volume of an organ or tissue due to the enlargement of its component cells. It is distinguished from hyperplasia, in which the cells remain approximately the same size but increase in number. Although hypertro ...
. The pregnant uterus only contracts strongly during orgasms, labour, and in the postpartum stage to return to its natural size.


Throughout menstrual cycle

Uterine contractions that occur throughout the
menstrual cycle The menstrual cycle is a series of natural changes in hormone production and the structures of the uterus and ovaries of the female reproductive system that makes pregnancy possible. The ovarian cycle controls the production and release of eg ...
, also termed ''endometrial waves'' or ''contractile waves'', appear to involve only the sub-
endometrial The endometrium is the inner epithelium, epithelial layer, along with its mucous membrane, of the mammalian uterus. It has a basal layer and a functional layer: the basal layer contains stem cells which regenerate the functional layer. The funct ...
layer of the
myometrium The myometrium is the middle layer of the uterine wall, consisting mainly of uterine smooth muscle cells (also called uterine myocytes) but also of supporting stromal and vascular tissue. Its main function is to induce uterine contractions. Stru ...
.


Follicular and luteal phase

In the early
follicular phase The follicular phase, also known as the preovulatory phase or proliferative phase, is the phase of the estrous cycle (or, in primates for example, the menstrual cycle) during which follicles in the ovary mature from primary follicle to a full ...
, uterine contractions in the non-pregnant woman occur 1–2 times per minute and last 10–15 seconds with a low intensity of usually 30
mmHg A millimetre of mercury is a manometric unit of pressure, formerly defined as the extra pressure generated by a column of mercury one millimetre high. Currently, it is defined as exactly , or approximately 1 torr =  atmosphere = &nb ...
or less. This sub-
endometrial The endometrium is the inner epithelium, epithelial layer, along with its mucous membrane, of the mammalian uterus. It has a basal layer and a functional layer: the basal layer contains stem cells which regenerate the functional layer. The funct ...
layer is rich in
estrogen Estrogen (also spelled oestrogen in British English; see spelling differences) is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three ...
and
progesterone Progesterone (; P4) is an endogenous steroid and progestogen sex hormone involved in the menstrual cycle, pregnancy, and embryogenesis of humans and other species. It belongs to a group of steroid hormones called the progestogens and is the ma ...
receptors. The frequency of contractions increases to 3–4 per minute towards
ovulation Ovulation is an important part of the menstrual cycle in female vertebrates where the egg cells are released from the ovaries as part of the ovarian cycle. In female humans ovulation typically occurs near the midpoint in the menstrual cycle and ...
. During the
luteal phase The menstrual cycle is on average 28 days in length. It begins with Menstruation, menses (day 1–7) during the follicular phase (day 1–14), followed by ovulation (day 14) and ending with the luteal phase (day 14–28). While historically, medi ...
, the frequency and intensity decrease, possibly to facilitate any implantation.


Menstruation

If implantation does not occur, the frequency of contractions remains low; but at
menstruation Menstruation (also known as a period, among other colloquial terms) is the regular discharge of blood and Mucous membrane, mucosal tissue from the endometrium, inner lining of the uterus through the vagina. The menstrual cycle is characterized ...
the intensity increases dramatically to between 50 and 200
mmHg A millimetre of mercury is a manometric unit of pressure, formerly defined as the extra pressure generated by a column of mercury one millimetre high. Currently, it is defined as exactly , or approximately 1 torr =  atmosphere = &nb ...
producing labor-like contractions. These contractions are sometimes termed ''menstrual cramps'',medicinenet.com > Menstrual Cramps
Retrieved January 2011
although that term is also used for menstrual pain in general. These contractions may be uncomfortable or even painful, but they are generally significantly less painful than contractions during labour. Painful contractions are called
dysmenorrhea Dysmenorrhea, also known as period pain, painful periods or menstrual cramps, is pain during menstruation. Its usual onset occurs around the time that menstruation begins. Symptoms typically last less than three days. The pain is usually in th ...
.


Directionality of contractions

A shift in the
myosin Myosins () are a Protein family, family of motor proteins (though most often protein complexes) best known for their roles in muscle contraction and in a wide range of other motility processes in eukaryotes. They are adenosine triphosphate, ATP- ...
expression of the uterine smooth muscle has been hypothesized as arising for changes in the directions of uterine contractions during the menstrual cycle.


Labour and pregnancy

Uterine contractions are a vital part of natural
childbirth Childbirth, also known as labour, parturition and delivery, is the completion of pregnancy, where one or more Fetus, fetuses exits the Womb, internal environment of the mother via vaginal delivery or caesarean section and becomes a newborn to ...
, which occur during the process of labour and delivery, (typically this excludes
caesarean section Caesarean section, also known as C-section, cesarean, or caesarean delivery, is the Surgery, surgical procedure by which one or more babies are Childbirth, delivered through an incision in the mother's abdomen. It is often performed because va ...
). These labour contractions are characterized by their rhythmic tightening and relaxation of the
myometrium The myometrium is the middle layer of the uterine wall, consisting mainly of uterine smooth muscle cells (also called uterine myocytes) but also of supporting stromal and vascular tissue. Its main function is to induce uterine contractions. Stru ...
, the most prominent uterine muscle. Labour contractions primarily serve the purpose of opening and dilating the
cervix The cervix (: cervices) or cervix uteri is a dynamic fibromuscular sexual organ of the female reproductive system that connects the vagina with the uterine cavity. The human female cervix has been documented anatomically since at least the time ...
, which leads to the assisting of the passage of the baby through the vaginal canal during the first stage of labour. Throughout pregnancy, the uterus experiences motor denervation, thus inhibiting spontaneous contractions. The remaining contractions are predominantly hormonally controlled. The decrease in the coordination of uterine smooth muscles cells reduces the effectiveness of contractions, causing the uterus to enter a state of uterine quiescence. During the beginning of labour, contractions may initially be intermittent and irregular, but will transition into a more coordinated pattern as the labour progresses. This transition is governed by various myogenic, neurogenic, and hormonal factors working together. As labour progresses, contractions will typically increase in frequency and intensity, which leads to a significant rise in intrauterine pressure. Otherwise, not all contractions experienced by pregnant individuals are indications of the beginning of labour. Some women experience what are commonly called Braxton Hicks contractions before their initial due date, which are characterized as “false labour." Though similar to labour uterine contractions, these contractions do not play a prominent role in cervical dilation or the progression of childbirth.


Oxytocin

The
hormone A hormone (from the Ancient Greek, Greek participle , "setting in motion") is a class of cell signaling, signaling molecules in multicellular organisms that are sent to distant organs or tissues by complex biological processes to regulate physio ...
oxytocin has been identified as inducing uterine contractions, and labour in general. Oxytocin is produced by the body naturally and since the 1950s has also been available in
synthetic Synthetic may refer to: Science * Synthetic biology * Synthetic chemical or compound, produced by the process of chemical synthesis * Synthetic elements, chemical elements that are not naturally found on Earth and therefore have to be created in ...
pharmaceutical form. In either form, oxytocin stimulates uterine contractions to accelerate the process of
childbirth Childbirth, also known as labour, parturition and delivery, is the completion of pregnancy, where one or more Fetus, fetuses exits the Womb, internal environment of the mother via vaginal delivery or caesarean section and becomes a newborn to ...
. Production and secretion of oxytocin is controlled by a
positive feedback Positive feedback (exacerbating feedback, self-reinforcing feedback) is a process that occurs in a feedback loop where the outcome of a process reinforces the inciting process to build momentum. As such, these forces can exacerbate the effects ...
mechanism, where its initial release, either naturally or in pharmaceutical form, stimulates production and release of further oxytocin. For example, when oxytocin is released during a contraction of the uterus at the start of childbirth, this stimulates production and release of more oxytocin and an increase in the duration, intensity and frequency of contractions. This process compounds in intensity and frequency and continues until the triggering activity ceases.


Prostaglandins

The concentration of prostaglandins in the
blood plasma Blood plasma is a light Amber (color), amber-colored liquid component of blood in which blood cells are absent, but which contains Blood protein, proteins and other constituents of whole blood in Suspension (chemistry), suspension. It makes up ...
and
amniotic fluid The amniotic fluid is the protective liquid contained by the amniotic sac of a gravid amniote. This fluid serves as a cushion for the growing fetus, but also serves to facilitate the exchange of nutrients, water, and biochemical products betwee ...
increases during labor. These inflammatory mediators encourage myometrial contractions to induce labor. Prostaglandins are also related to the changes in
gap junction Gap junctions are membrane channels between adjacent cells that allow the direct exchange of cytoplasmic substances, such small molecules, substrates, and metabolites. Gap junctions were first described as ''close appositions'' alongside tight ...
formation and connexin-43 expression during labor.


In orgasm

Uterine and vaginal contractions usually take place during female
sexual stimulation Sexual stimulation is anything that leads to sexual arousal or orgasm. This thing can be physical or of other senses, and is known as a stimulus. Sexual stimulation is a broad term, usually understood to mean physical touching of the genita ...
, including
sexual arousal Sexual arousal (also known as sexual excitement) describes the Physiology, physiological and psychological responses in preparation for sexual intercourse or when exposed to Sexual stimulation, sexual stimuli. A number of physiological response ...
, and
orgasm Orgasm (from Greek , ; "excitement, swelling"), sexual climax, or simply climax, is the sudden release of accumulated sexual excitement during the sexual response cycle, characterized by intense sexual pleasure resulting in rhythmic, involu ...
.


Monitors

Uterine contractions can be monitored by
cardiotocography Cardiotocography (CTG) is a technique used to monitor the fetal heartbeat and uterine contractions during pregnancy and labour. The machine used to perform the monitoring is called a cardiotocograph. Fetal heart sounds were described as early ...
, in which a device is affixed to the skin of the mother or directly to the fetal scalp. The pressure required to flatten a section of the uterine wall correlates with the internal pressure, thereby providing an estimate of it. A type of monitoring technology under development at
Drexel University Drexel University is a private university, private research university with its main campus in Philadelphia, Pennsylvania, United States. Drexel's undergraduate school was founded in 1891 by Anthony Joseph Drexel, Anthony J. Drexel, a financier ...
embeds conductive threads in the knitted fabric of a bellyband. When the fibers stretch in response to a contraction, the threads function like an antenna, and send the signals they pick up to an embedded RFID (
radio-frequency identification Radio-frequency identification (RFID) uses electromagnetic fields to automatically Automatic identification system, identify and Tracking system, track tags attached to objects. An RFID system consists of a tiny radio transponder called a tag, ...
device) chip that reports the data.


Mechanism


Resting state

The
resting membrane potential The relatively static membrane potential of quiescent cells is called the resting membrane potential (or resting voltage), as opposed to the specific dynamic electrochemical phenomena called action potential and graded membrane potential. The re ...
(Vrest) of uterine smooth muscle has been recorded to be between −35 and −80 mV. As with the resting membrane potential of other cell types, it is maintained by a Na+/K+ pump that causes a higher concentration of Na+ ions in the extracellular space than in the
intracellular space Intracellular space is the interior space of the plasma membrane The cell membrane (also known as the plasma membrane or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and prote ...
, and a higher concentration of K+ ions in the intracellular space than in the extracellular space. Subsequently, having K+ channels open to a higher degree than Na+ channels results in an overall efflux of positive ions, resulting in a negative potential. This resting potential undergoes rhythmic oscillations, which have been termed ''slow waves'', and reflect intrinsic activity of slow wave potentials. These slow waves are caused by changes in the distribution of Ca2+, Na+, K+ and Cl ions between the intracellular and extracellular spaces, which, in turn, reflects the permeability of the plasma membrane to each of those ions. K+ is the major ion responsible for such changes in
ion flux Flux describes any effect that appears to pass or travel (whether it actually moves or not) through a surface or substance. Flux is a concept in applied mathematics and vector calculus which has many applications in physics. For transport pheno ...
, reflecting changes in various K+ channels.


Excitation-contraction

As the uterus becomes essentially denervated during gestation, it is unlikely that any coordinated nervous regulation of the myometrium is centrally orchestrated.


Excitation

The excitation-contraction coupling of the uterine smooth muscle is also very similar to that of other
smooth muscle Smooth muscle is one of the three major types of vertebrate muscle tissue, the others being skeletal and cardiac muscle. It can also be found in invertebrates and is controlled by the autonomic nervous system. It is non- striated, so-called bec ...
s in general, with intracellular increase in
calcium Calcium is a chemical element; it has symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to it ...
(Ca2+) leading to contraction. Nitric oxide (NO) is particularly effective in relaxing the myometrium and in fact has a lower inhibitory concentration 50% (Ki) in human than guinea pig or non-human primate myometrium.


Restoration to resting state

Uterine smooth muscle mechanisms of relaxation differ significantly from those of other human smooth muscles. Removal of Ca2+ after contraction induces relaxation of the smooth muscle, and restores the molecular structure of the sarcoplasmic reticulum for the next contractile stimulus.


Measuring uterine contractility ''ex vivo''

Ethically donated human uterine tissues can be used to measure uterine contractility '' ex vivo''. In these experiments, sections of
myometrium The myometrium is the middle layer of the uterine wall, consisting mainly of uterine smooth muscle cells (also called uterine myocytes) but also of supporting stromal and vascular tissue. Its main function is to induce uterine contractions. Stru ...
are set up in an organ bath system that to measure changes in isometric force production. Following functional checks to ensure the tissue is physiologically active, compounds can be added to the organ bath in increasing concentrations to create a cumulative concentration-response curve (CCRC). A key advantage of measuring uterine contractility ''ex vivo'' is the ability to eliminate species differences. For example, while magnesium reduces myometrial contractility in
animal studies Animal studies is a recently recognised field in which animals are studied in a variety of cross-disciplinary ways. Scholars who engage in animal studies may be formally trained in a number of diverse fields, including art history, anthropology ...
and ''
in vitro ''In vitro'' (meaning ''in glass'', or ''in the glass'') Research, studies are performed with Cell (biology), cells or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in ...
'', it does not demonstrate the same effect in
clinical studies Clinical trials are prospective biomedical or behavioral research studies on human participants designed to answer specific questions about biomedical or behavioral interventions, including new treatments (such as novel vaccines, drugs, dietar ...
. And while the peptide hormone relaxin has been shown to inhibit uterine contractility in rats, mice, and pigs, it does not prevent uterine contractility in humans.


See also

* Involution (medicine)


References

{{Authority control Obstetrics Midwifery Uterus