The estrous cycle (, originally ) is the set of recurring physiological changes that are induced by
reproductive 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 by complex biological processes to regulate physiology and beh ...
s in most
mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
ian
theria
Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes.
Ch ...
n
female
Female (Venus symbol, symbol: ♀) is the sex of an organism that produces the large non-motile ovum, ova (egg cells), the type of gamete (sex cell) that fuses with the Sperm, male gamete during sexual reproduction.
A female has larger gamet ...
s. Estrous cycles start after
sexual maturity
Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...
in females and are interrupted by anestrous phases, otherwise known as "rest" phases, or by
pregnancies
Pregnancy is the time during which one or more offspring develops ( gestates) inside a woman's uterus (womb). A multiple pregnancy involves more than one offspring, such as with twins.
Pregnancy usually occurs by sexual intercourse, but can ...
. Typically, estrous cycles repeat until death. These cycles are widely variable in duration and frequency depending on the species.
[Bronson, F. H., 1989. Mammalian Reproductive Biology. University of Chicago Press, Chicago, IL, USA.] Some animals may display bloody vaginal discharge, often mistaken for
menstruation
Menstruation (also known as a period, among other colloquial terms) is the regular discharge of blood and mucosal tissue from the inner lining of the uterus through the vagina. The menstrual cycle is characterized by the rise and fall of hor ...
. Many mammals used in commercial agriculture, such as cattle and sheep, may have their estrous cycles artificially controlled with hormonal medications for optimum productivity.
The male equivalent, seen primarily in ruminants, is called
rut.
Differences from the menstrual cycle
Mammals share the same reproductive system, including the regulatory
hypothalamic
The hypothalamus () is a part of the brain that contains a number of small nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrine system via the pituitary gland. The hypothalamus i ...
system that produces
gonadotropin-releasing hormone
Gonadotropin-releasing hormone (GnRH) is a releasing hormone responsible for the release of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary. GnRH is a tropic peptide hormone synthesized and released ...
in pulses, the
pituitary gland
In vertebrate anatomy, the pituitary gland, or hypophysis, is an endocrine gland, about the size of a chickpea and weighing, on average, in humans. It is a protrusion off the bottom of the hypothalamus at the base of the brain. The ...
that secretes
follicle-stimulating hormone
Follicle-stimulating hormone (FSH) is a gonadotropin, a glycoprotein polypeptide hormone. FSH is synthesized and secreted by the gonadotropic cells of the anterior pituitary gland and regulates the development, growth, pubertal maturation, an ...
and
luteinizing hormone
Luteinizing hormone (LH, also known as luteinising hormone, lutropin and sometimes lutrophin) is a hormone produced by gonadotropic cells in the anterior pituitary gland. The production of LH is regulated by gonadotropin-releasing hormone (GnRH) ...
, and the ovary itself that releases
sex hormone
Sex hormones, also known as sex steroids, gonadocorticoids and gonadal steroids, are steroid hormones that interact with vertebrate steroid hormone receptors. The sex hormones include the androgens, estrogens, and progestogens. Their effects ar ...
s, including
estrogen
Estrogen or oestrogen is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three major endogenous estrogens that have estrogenic hormonal acti ...
s 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 m ...
.
However, species vary significantly in the detailed functioning. One difference is that animals that have estrous cycles
resorb the
endometrium
The endometrium is the inner 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 functional laye ...
if conception does not occur during that cycle. Animals that have
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 make pregnancy possible. The ovarian cycle controls the production and release of eggs a ...
s shed the endometrium through
menstruation
Menstruation (also known as a period, among other colloquial terms) is the regular discharge of blood and mucosal tissue from the inner lining of the uterus through the vagina. The menstrual cycle is characterized by the rise and fall of hor ...
instead. Another difference is
sexual activity
Human sexual activity, human sexual practice or human sexual behaviour is the manner in which humans experience and express their sexuality. People engage in a variety of sexual acts, ranging from activities done alone (e.g., masturbation) ...
. In species with estrous cycles, females are generally only sexually active during the
estrus phase of their cycle. This is also referred to as being "in heat". In contrast, females of species with menstrual cycles can be sexually active at any time in their cycle, even when they are not about to
ovulate
Ovulation is the release of eggs from the ovaries. In women, this event occurs when the ovarian follicles rupture and release the secondary oocyte ovarian cells. After ovulation, during the luteal phase, the egg will be available to be fertilized ...
.
Humans have menstrual cycles rather than estrous cycles. They, unlike most other species, have
concealed ovulation Concealed ovulation or hidden œstrus in a species is the lack of any perceptible change in an adult female (for instance, a change in appearance or scent) when she is fertile and near ovulation. Some examples of perceptible changes are swelling an ...
, a lack of obvious external signs to signal estral receptivity at
ovulation
Ovulation is the release of eggs from the ovaries. In women, this event occurs when the ovarian follicles rupture and release the secondary oocyte ovarian cells. After ovulation, during the luteal phase, the egg will be available to be fertilized ...
(i.e., the ability to become pregnant). Animals with estrous cycles often have unmistakable outward displays of receptivity, ranging from engorged and colorful
genitals
A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...
to behavioral changes like
mating call
A mating call is the auditory signal used by animals to attract mates. It can occur in males or females, but literature is abundantly favored toward researching mating calls in females. In addition, mating calls are often the subject of mate choic ...
s.
Etymology and nomenclature
''Estrus'' is derived via Latin ' ('
frenzy
''Frenzy'' is a 1972 British thriller film directed by Alfred Hitchcock. It is the penultimate feature film of his extensive career. The screenplay by Anthony Shaffer was based on the 1966 novel ''Goodbye Piccadilly, Farewell Leicester Squar ...
', '
gadfly'), from Greek ''oîstros'' (literally 'gadfly', more figuratively 'frenzy', 'madness', among other meanings like 'breeze'). Specifically, this refers to the gadfly in
Ancient Greek mythology
A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities of de ...
that
Hera
In ancient Greek religion, Hera (; grc-gre, Ἥρα, Hḗrā; grc, Ἥρη, Hḗrē, label=none in Ionic and Homeric Greek) is the goddess of marriage, women and family, and the protector of women during childbirth. In Greek mythology, she ...
sent to torment
Io, who had been won in her
heifer
Heifer may refer to:
* Heifer (cow), a young cow before she has had her first calf
* Frank Heifer (1854–1893), American outfielder and first baseman
* '' The Heifer'' (''La vaquilla''), 1985 Spanish comedy film
* Heifer International
Heifer ...
form by
Zeus
Zeus or , , ; grc, Δῐός, ''Diós'', label=Genitive case, genitive Aeolic Greek, Boeotian Aeolic and Doric Greek#Laconian, Laconian grc-dor, Δεύς, Deús ; grc, Δέος, ''Déos'', label=Genitive case, genitive el, Δίας, ''D ...
.
Euripides
Euripides (; grc, Εὐριπίδης, Eurīpídēs, ; ) was a tragedian
Tragedy (from the grc-gre, τραγῳδία, ''tragōidia'', ''tragōidia'') is a genre of drama based on human suffering and, mainly, the terrible or sorrowful e ...
used ''oestrus'' to indicate 'frenzy', and to describe madness.
Homer
Homer (; grc, Ὅμηρος , ''Hómēros'') (born ) was a Greek poet who is credited as the author of the ''Iliad'' and the ''Odyssey'', two epic poems that are foundational works of ancient Greek literature. Homer is considered one of the ...
used the word to describe panic.
Plato
Plato ( ; grc-gre, Πλάτων ; 428/427 or 424/423 – 348/347 BC) was a Greek philosopher born in Athens during the Classical period in Ancient Greece. He founded the Platonist school of thought and the Academy, the first institution ...
also used it to refer to an irrational drive and to describe the
soul
In many religious and philosophical traditions, there is a belief that a soul is "the immaterial aspect or essence of a human being".
Etymology
The Modern English noun ''soul'' is derived from Old English ''sāwol, sāwel''. The earliest attes ...
"driven and drawn by the gadfly of desire". Somewhat more closely aligned to current meaning and usage of ''estrus'',
Herodotus
Herodotus ( ; grc, , }; BC) was an ancient Greek historian and geographer from the Greek city of Halicarnassus, part of the Persian Empire (now Bodrum, Turkey) and a later citizen of Thurii in modern Calabria ( Italy). He is known f ...
(''Histories'', ch. 93.1) uses ''oîstros'' to describe the desire of fish to
spawn
Spawn or spawning may refer to:
* Spawn (biology), the eggs and sperm of aquatic animals
Arts, entertainment, and media
* Spawn (character), a fictional character in the comic series of the same name and in the associated franchise
** '' Spawn: ...
.
The earliest use in English was with a meaning of 'frenzied passion'. In 1900, it was first used to describe '
rut in animals; heat'.
[
In ]British
British may refer to:
Peoples, culture, and language
* British people, nationals or natives of the United Kingdom, British Overseas Territories, and Crown Dependencies.
** Britishness, the British identity and common culture
* British English, ...
and most Commonwealth English
The use of the English language in current and former Member states of the Commonwealth of Nations, member countries of the Commonwealth of Nations was largely inherited from British Empire, British colonisation, with some exceptions. English s ...
, the spelling is ''oestrus'' or (rarely) ''œstrus''. In all English spellings, the noun ends in ''-us'' and the adjective in ''-ous''. Thus in North American English
North American English (NAmE, NAE) is the most generalized variety of the English language as spoken in the United States and Canada. Because of their related histories and cultures, plus the similarities between the pronunciations (accents), v ...
, a mammal may be described as "in estrus" when it is in that particular part of the estrous cycle.
Four phases
A four-phase terminology is used in reference to animals with estrous cycles.
Proestrus
One or several follicles of the ovary
The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
start to grow. Their number is species-specific. Typically this phase can last as little as one day or as long as three weeks, depending on the species. Under the influence of estrogen, the lining of the uterus (endometrium
The endometrium is the inner 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 functional laye ...
) starts to develop. Some animals may experience vaginal secretions that could be bloody. The female is not yet sexually receptive; the old corpus luteum
The corpus luteum (Latin for "yellow body"; plural corpora lutea) is a temporary endocrine structure in female ovaries involved in the production of relatively high levels of progesterone, and moderate levels of estradiol, and inhibin A. It is t ...
degenerates; the uterus and the vagina distend and fill with fluid, become contractile and secrete a sanguinous fluid; the vaginal epithelium
The vaginal epithelium is the inner lining of the vagina consisting of multiple layers of (squamous) cells. The basal membrane provides the support for the first layer of the epithelium-the basal layer. The intermediate layers lie upon the basal ...
proliferates and the vaginal cytology shows a large number of non-cornified nucleated epithelial cells. Variant terms for ''proestrus'' include ''pro-oestrus'', ''proestrum'', and ''pro-oestrum''.
Estrus
Estrus or oestrus refers to the phase when the female is sexually receptive ("in heat" American English, or "on heat" British English). Under regulation by gonadotropic hormones, ovarian follicle
An ovarian follicle is a roughly spheroid cellular aggregation set found in the ovaries. It secretes hormones that influence stages of the menstrual cycle. At the time of puberty, women have approximately 200,000 to 300,000 follicles, each with th ...
s mature and estrogen secretions exert their biggest influence. The female then exhibits sexually receptive behavior, a situation that may be signaled by visible physiologic changes. Estrus is commonly seen in the mammalian species, including primates.
In some species, the labia
The labia are part of the female genitalia; they are the major externally visible portions of the vulva. In humans, there are two pairs of labia: the ''labia majora'' (or the outer labia) are larger and thicker, while the ''labia minora'' are fol ...
are reddened. Ovulation
Ovulation is the release of eggs from the ovaries. In women, this event occurs when the ovarian follicles rupture and release the secondary oocyte ovarian cells. After ovulation, during the luteal phase, the egg will be available to be fertilized ...
may occur spontaneously in others. Especially among quadrupeds
Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
, a signal trait of estrus is the lordosis reflex, in which the animal spontaneously elevates her hindquarters.
Controlled internal drug release
Controlled internal drug release (CIDR) devices are used in livestock for the synchronization of estrus. They are T-shaped devices with a silicone-coated nylon core. The silicone coating is impregnated with progesterone. CIDRs are inserted intra- ...
devices are used in livestock
Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to animals ...
for the synchronization of estrus.
Metestrus or diestrus
This phase is characterized by the activity of the corpus luteum
The corpus luteum (Latin for "yellow body"; plural corpora lutea) is a temporary endocrine structure in female ovaries involved in the production of relatively high levels of progesterone, and moderate levels of estradiol, and inhibin A. It is t ...
, which produces progesterone. The signs of estrogen stimulation subside and the corpus luteum
The corpus luteum (Latin for "yellow body"; plural corpora lutea) is a temporary endocrine structure in female ovaries involved in the production of relatively high levels of progesterone, and moderate levels of estradiol, and inhibin A. It is t ...
starts to form. The uterine
The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the hollow organ, organ in the female reproductive system, reproductive system of most female mammals, including humans that accommodates the embryonic development, embryonic an ...
lining begins to appear. In the absence of pregnancy
Pregnancy is the time during which one or more offspring develops ( gestates) inside a woman's uterus (womb). A multiple pregnancy involves more than one offspring, such as with twins.
Pregnancy usually occurs by sexual intercourse, but ca ...
the diestrus phase (also termed pseudo-pregnancy
High-dose estrogen (HDE) is a type of hormone therapy in which high doses of estrogens are given. When given in combination with a high dose of a progestogen, it has been referred to as pseudopregnancy. It is called this because the estrogen and p ...
) terminates with the regression of the corpus luteum. The lining in the uterus is not shed, but is reorganized for the next cycle. Other spellings include ''metoestrus'', ''metestrum'', ''metoestrum'', ''dioestrus'', ''diestrum'', and ''dioestrum''.
Anestrus
Anestrus refers to the phase when the sexual cycle rests. This is typically a seasonal event and controlled by light exposure through the pineal gland
The pineal gland, conarium, or epiphysis cerebri, is a small endocrine gland in the brain of most vertebrates. The pineal gland produces melatonin, a serotonin-derived hormone which modulates sleep, sleep patterns in both circadian rhythm, circ ...
that releases melatonin
Melatonin is a natural product found in plants and animals. It is primarily known in animals as a hormone released by the pineal gland in the brain at night, and has long been associated with control of the sleep–wake cycle.
In vertebrates ...
. Melatonin may repress stimulation of reproduction in long-day breeders and stimulate reproduction in short-day breeders. Melatonin is thought to act by regulating the hypothalamic
The hypothalamus () is a part of the brain that contains a number of small nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrine system via the pituitary gland. The hypothalamus i ...
pulse activity of the gonadotropin-releasing hormone
Gonadotropin-releasing hormone (GnRH) is a releasing hormone responsible for the release of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary. GnRH is a tropic peptide hormone synthesized and released ...
. Anestrus is induced by time of year, pregnancy
Pregnancy is the time during which one or more offspring develops ( gestates) inside a woman's uterus (womb). A multiple pregnancy involves more than one offspring, such as with twins.
Pregnancy usually occurs by sexual intercourse, but ca ...
, lactation
Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The proces ...
, significant illness
A disease is a particular abnormal condition that negatively affects the structure or function of all or part of an organism, and that is not immediately due to any external injury. Diseases are often known to be medical conditions that ar ...
, chronic energy deficit, and possibly age. Chronic exposure to anabolic steroids may also induce a persistent anestrus due to negative feedback on the hypothalamus/pituitary/gonadal axis. Other spellings include ''anoestrus'', ''anestrum'', and ''anoestrum''.
After completion (or abortion) of a pregnancy, some species have ''postpartum estrus'', which is ovulation and corpus luteum production that occurs immediately following the birth of the young. For example, the mouse
A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
has a fertile postpartum estrus that occurs 14 to 24 hours following parturition.
Cycle variability
Estrous cycle variability differs among species, but cycles are typically more frequent in smaller animals. Even within species significant variability can be observed, thus cat
The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s may undergo an estrous cycle of 3 to 7 weeks. Domestication can affect estrous cycles due to changes in the environment. For most species, vaginal smear cytology
Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living and ...
may be used in order to identify estrous cycle phases and durations.
Frequency
Some species, such as cat
The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s, cow
Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult ma ...
s and domestic pig
The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus s ...
s, are polyestrous, meaning that they can go into heat several times per year. Seasonally polyestrous animals or seasonal breeder
Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
s have more than one estrous cycle during a specific time of the year and can be divided into short-day and long-day breeders:
* Short-day breeders, such as sheep
Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus ''Ovis'', in everyday usage it almost always refers to domesticated s ...
, goat
The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
s, deer
Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...
and elk
The elk (''Cervus canadensis''), also known as the wapiti, is one of the largest species within the deer family, Cervidae, and one of the largest terrestrial mammals in its native range of North America and Central and East Asia. The common ...
are sexually active in fall or winter.
* Long-day breeders, such as horse
The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million y ...
s, hamster
Hamsters are rodents (order Rodentia) belonging to the subfamily Cricetinae, which contains 19 species classified in seven genera.Fox, Sue. 2006. ''Hamsters''. T.F.H. Publications Inc. They have become established as popular small pets. The b ...
s and ferret
The ferret (''Mustela furo'') is a small, Domestication, domesticated species belonging to the family Mustelidae. The ferret is most likely a domesticated form of the wild European polecat (''Mustela putorius''), evidenced by their Hybrid (biol ...
s are sexually active in spring and summer.
Species that go into heat twice per year are diestrous. Canines are diestrous.
Monestrous species, such as canids
Canidae (; from Latin, ''canis'', "dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found within th ...
and bear
Bears are carnivoran mammals of the family Ursidae. They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats throughout the Nor ...
s, have only one breeding season
Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
per year, typically in spring to allow growth of the offspring
In biology, offspring are the young creation of living organisms, produced either by a single organism or, in the case of sexual reproduction, two organisms. Collective offspring may be known as a brood or progeny in a more general way. This ca ...
during the warm season to aid survival during the next winter.
A few mammalian species, such as rabbit
Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit speci ...
s and cats, do not have an estrous cycle, instead being induced to ovulate by the act of mating and are able to conceive at almost any arbitrary moment.
Generally speaking, the timing of estrus is coordinated with seasonal availability of food and other circumstances such as migration, predation etc., the goal being to maximize the offspring's chances of survival. Some species are able to modify their estral timing in response to external conditions.
Specific species
Cats
The female cat
The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
in heat has an estrus of 14 to 21 days and is generally characterized as an induced ovulator
Induced ovulation is when a female animal ovulates due to an externally-derived stimulus during, or just prior to, mating, rather than ovulating cyclically or spontaneously. Stimuli causing induced ovulation include the physical act of coitus or m ...
, since coitus induces ovulation. However, various incidents of spontaneous ovulation have been documented in the domestic cat and various non-domestic species. Without ovulation, she may enter interestrus, which is the combined stages of diestrus and anestrus, before reentering estrus. With the induction of ovulation, the female becomes pregnant or undergoes a non-pregnant luteal phase
The corpus luteum (Latin for "yellow body"; plural corpora lutea) is a temporary endocrine structure in female ovaries involved in the production of relatively high levels of progesterone, and moderate levels of estradiol, and inhibin A. It is t ...
, also known as pseudopregnancy
In mammalian species, pseudopregnancy is a physical state whereby all the signs and symptoms of pregnancy are exhibited, with the exception of the presence of a fetus, creating a false pregnancy. The corpus luteum (the remains of an ovulated ovar ...
. Cats are polyestrous but experience a seasonal anestrus in autumn and late winter.
Dogs
A female dog
The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
is usually diestrous (goes into heat typically twice per year), although some breeds typically have one or three cycles per year. The proestrus is relatively long at 5 to 9 days, while the estrus may last 4 to 13 days, with a diestrus of 60 days followed by about 90 to 150 days of anestrus. Female dogs bleed during estrus, which usually lasts from 7–13 days, depending on the size and maturity of the dog. Ovulation occurs 24–48 hours after the luteinizing hormone peak, which occurs around the fourth day of estrus; therefore, this is the best time to begin breeding. Proestrus bleeding in dogs is common and is believed to be caused by diapedesis
Leukocyte extravasation (also commonly known as leukocyte adhesion cascade or diapedesis – the passage of cells through the intact vessel wall) is the movement of leukocytes out of the circulatory system and towards the site of tissue damage or ...
of red blood cell
Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek ''erythros'' for "red" and ''kytos'' for "holl ...
s from the blood vessels due to the increase of the estradiol-17β hormone.
Horses
A mare
A mare is an adult female horse or other equine. In most cases, a mare is a female horse over the age of three, and a filly is a female horse three and younger. In Thoroughbred horse racing, a mare is defined as a female horse more than four ...
may be in heat for 4 to 10 days, followed by approximately 14 days in diestrus. Thus, a cycle may be short, totaling approximately 3 weeks. Horses mate in spring and summer; autumn is a transition time, and anestrus occurs during winter.
A feature of the fertility
Fertility is the capability to produce offspring through reproduction following the onset of sexual maturity. The fertility rate is the average number of children born by a female during her lifetime and is quantified demographically. Fertili ...
cycle of horses and other large herd animals is that it is usually affected by the seasons. The number of hours daily that light enters the eye of the animal affects the brain, which governs the release of certain precursors and hormones. When daylight hours are few, these animals "shut down", become anestrous, and do not become fertile. As the days grow longer, the longer periods of daylight cause the hormones that activate the breeding cycle to be released. As it happens, this benefits these animals in that, given a gestation period of about eleven months, it prevents them from having young when the cold of winter would make their survival risky.
Rats
Rats are polyestrous animals that typically have rapid cycle lengths of 4 to 5 days. Although they ovulate spontaneously, they do not develop a fully functioning corpus luteum unless they receive coital stimulation. Fertile mating leads to pregnancy in this way, but infertile mating leads to a state of pseudopregnancy lasting about 10 days. Mice and hamsters have similar behavior. The events of the cycle are strongly influenced by lighting periodicity.
A set of follicles starts to develop near the end of proestrus and grows at a nearly constant rate until the beginning of the subsequent estrus when the growth rates accelerate eightfold. Ovulation occurs about 109 hours after the start of follicle growth.
Estrogen
Estrogen or oestrogen is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three major endogenous estrogens that have estrogenic hormonal acti ...
peaks at about 11 am on the day of proestrus. Between then and midnight there is a surge in 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 m ...
, luteinizing hormone
Luteinizing hormone (LH, also known as luteinising hormone, lutropin and sometimes lutrophin) is a hormone produced by gonadotropic cells in the anterior pituitary gland. The production of LH is regulated by gonadotropin-releasing hormone (GnRH) ...
and follicle-stimulating hormone
Follicle-stimulating hormone (FSH) is a gonadotropin, a glycoprotein polypeptide hormone. FSH is synthesized and secreted by the gonadotropic cells of the anterior pituitary gland and regulates the development, growth, pubertal maturation, an ...
, and ovulation occurs at about 4 am on the next estrus day. The following day, metestrus, is called early diestrus or diestrus I. During this day, the corpora lutea
The corpus luteum (Latin for "yellow body"; plural corpora lutea) is a temporary endocrine structure in female ovaries involved in the production of relatively high levels of progesterone, and moderate levels of estradiol, and inhibin A. It is th ...
grow to a maximal volume, achieved within 24 hours of ovulation. They remain at that size for three days, halve in size before the metestrus of the next cycle and then shrink abruptly before estrus of the cycle after that. Thus the ovaries of cycling rats contain three different sets of corpora lutea at different phases of development.
Bison
Buffalo have an estrous cycle of about 22 to 24 days. Buffalo are known for difficult estrus detection. This is one major reason for being less productive than cattle
Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult mal ...
. During four phases of its estrous cycle, mean weight of corpus luteum
The corpus luteum (Latin for "yellow body"; plural corpora lutea) is a temporary endocrine structure in female ovaries involved in the production of relatively high levels of progesterone, and moderate levels of estradiol, and inhibin A. It is t ...
has been found to be 1.23±0.22g (metestrus), 3.15±0.10g (early diestrus), 2.25±0.32g (late diestrus), and 1.89±0.31g (proestrus/estrus), respectively. The plasma progesterone concentration was 1.68±0.37, 4.29±0.22, 3.89±0.33, and 0.34±0.14 ng/ml while mean vascular density (mean number of vessels/10 microscopic fields at 400x) in corpus luteum was 6.33±0.99, 18.00±0.86, 11.50±0.76, and 2.83±0.60 during the metestrus, early diestrus, late diestrus and proestrus/estrus, respectively.
Cattle
Female cattle, also referred to as "heifer
Heifer may refer to:
* Heifer (cow), a young cow before she has had her first calf
* Frank Heifer (1854–1893), American outfielder and first baseman
* '' The Heifer'' (''La vaquilla''), 1985 Spanish comedy film
* Heifer International
Heifer ...
s" in agriculture, will gradually enter standing estrus, or "standing heat," starting at puberty between 9 and 15 months of age. The cow estrous cycle typically lasts 21 days. Standing estrus is a visual cue which signifies sexual receptivity for mounting by male cattle. This behavior lasts anywhere between 8 and 30 hours at a time. Other behaviors of the female during standing estrus may change, including, but not limited to: nervousness, swollen vulva
The vulva (plural: vulvas or vulvae; derived from Latin for wrapper or covering) consists of the external sex organ, female sex organs. The vulva includes the mons pubis (or mons veneris), labia majora, labia minora, clitoris, bulb of vestibu ...
, or attempting to mount other animals. While visual and behavioral cues are helpful to the male cattle, estrous stages cannot be determined by the human eye. Rather, the stage can be estimated from the appearance of the corpora lutea or follicle composition.
Estrous control
Due to the widespread use of bovine animals in agriculture, cattle estrous cycles have been widely studied, and manipulated, in an effort to maximize profitability through reproductive management. Much estrous control in cattle is for the purpose of synchronization, a practice or set of practices most often used by cattle farmers to control the timing and duration of estrus in large herds.
There is variation between the available methods of cattle estrous synchronization. Treatment depends on herd size, specific goals for control, and budget. Some of the FDA
The United States Food and Drug Administration (FDA or US FDA) is a federal agency of the Department of Health and Human Services. The FDA is responsible for protecting and promoting public health through the control and supervision of food ...
-approved drugs and devices used to mimic natural hormones of the estrous cycle include, but are not limited to, the following classes:
* Gonadorelin
Gonadorelin is a gonadotropin-releasing hormone agonist (GnRH agonist) which is used in fertility medicine and to treat amenorrhea and hypogonadism. It is also used in veterinary medicine. The medication is a form of the endogenous GnRH and is i ...
: There are currently five available gonadorelin products that are FDA-Approved. Usually, gonadorelin is used in conjunction with another estrous control drug (typically, prostaglandin). This drug is used to mimic gonadotropin releasing hormone (GnRH) and may also be used to treat ovarian cysts.
* Prostaglandin
The prostaglandins (PG) are a group of physiologically active lipid compounds called eicosanoids having diverse hormone-like effects in animals. Prostaglandins have been found in almost every tissue in humans and other animals. They are derive ...
: Mimics the prostaglandin F2-alpha hormone released when no pregnancy has occurred and regresses the corpus luteum. This drug is used to achieve more consistent results in artificial insemination.
* Progestin
A progestogen, also referred to as a progestagen, gestagen, or gestogen, is a type of medication which produces effects similar to those of the natural product, natural female sex hormone progesterone in the body. A progestin is a ''synthetic co ...
: Used to suppress estrus and/or block ovulation. Most commonly, it is administered via an intravaginal insert comparable to an IUD
An intrauterine device (IUD), also known as intrauterine contraceptive device (IUCD or ICD) or coil, is a small, often T-shaped birth control device that is inserted into the uterus to prevent pregnancy. IUDs are one form of long-acting rever ...
, which used in controlling menstrual periods. It is also available as a medicated feed, but this method is not yet approved for cattle crop synchronization.
There is variation between the available methods of cattle estrous synchronization. Treatment depends on herd size, specific goals for control, and budget.
Bovine estrous cycles may also be impacted by other bodily functions such as oxytocin
Oxytocin (Oxt or OT) is a peptide hormone and neuropeptide normally produced in the hypothalamus and released by the posterior pituitary. It plays a role in social bonding, reproduction, childbirth, and the period after childbirth. Oxytocin ...
levels. Additionally, heat stress
Hyperthermia, also known simply as overheating, is a condition in which an individual's body temperature is elevated beyond normal due to failed thermoregulation. The person's body produces or absorbs more heat than it dissipates. When extreme ...
has been linked to impairment of follicular development, especially impactful to the first-wave dominant follicle. Future synchronization programs are planning to focus on the impact of heat stress on fertilization and embryonic death rates after artificial insemination
Artificial insemination is the deliberate introduction of sperm into a female's cervix or uterine cavity for the purpose of achieving a pregnancy through in vivo fertilization by means other than sexual intercourse. It is a fertility treatme ...
.
Additionally, work has been done regarding other mammalian females, such as in dogs, for estrous control; However, there are yet to be any approved medications outside of those commercially available.[Kutzler MA. Estrous Cycle Manipulation in Dogs. Vet Clin North Am Small Anim Pract. 2018 Jul;48(4):581-594. . Epub 2018 Apr 27. PMID 29709316.]
Others
Estrus frequencies of some other notable mammals:
* Sheep
Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus ''Ovis'', in everyday usage it almost always refers to domesticated s ...
: 18 days
* Pig
The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus ...
: 21 days
* Goat
The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
: 21 days
* Rabbit
Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit speci ...
: variable
* Donkey
The domestic donkey is a hoofed mammal in the family Equidae, the same family as the horse. It derives from the African wild ass, ''Equus africanus'', and may be classified either as a subspecies thereof, ''Equus africanus asinus'', or as a ...
: 13 to 31 days (average 23)
* Elephant
Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae an ...
: 16 weeks
* Wolf
The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly u ...
: 9 days
See also
* Mating system
A mating system is a way in which a group is structured in relation to sexual behaviour. The precise meaning depends upon the context. With respect to animals, the term describes which males and females mating, mate under which circumstances. Reco ...
* Musth
Musth or must (from Persian, )''The Oxford Dictionary and Thesaurus: American edition'', published 1996 by Oxford University Press; p. 984 is a periodic condition in bull (male) elephants characterized by aggressive behavior and accompanied by ...
* Neutering
Neutering, from the Latin ''neuter'' ('of neither sex'), is the removal of an animal's reproductive organ, either all of it or a considerably large part. The male-specific term is castration, while spaying is usually reserved for female animals. ...
* Progesterone-releasing intravaginal device
* Reproductive cycle
In biology, a biological life cycle (or just life cycle or lifecycle when the biological context is clear) is a series of changes in form that an organism undergoes, returning to the starting state. "The concept is closely related to those of the ...
* Rut (mammalian reproduction)
The rut (from the Latin ''rugire'', meaning "to roar") is the mating season of certain mammals, which includes ruminants such as deer, sheep, camels, goats, pronghorns, bison, giraffes and antelopes, and extends to others such as skunks and elepha ...
* Sexual swelling
Sexual swellings are enlarged areas of genital and perineal skin occurring in some female primates that vary in size over the course of the menstrual cycle. Thought to be an honest signal of fertility, male primates are attracted to these swell ...
References
Further reading
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External links
Systematic overview
Etymology
Horse estrous cycle
Dogs in Heat - FAQ
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{{Authority control
Mammalian sexuality
Reproduction in mammals
Theriogenology