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In
zoology Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and ...
, copulation is
animal sexual behavior Animal sexual behaviour takes many different forms, including within the same species. Common mating or reproductively motivated systems include monogamy, polygyny, polyandry, polygamy and promiscuity. Other sexual behaviour may be reproduc ...
in which a male introduces
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...
into the female's body, especially directly into her reproductive tract. This is an aspect of
mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...
. Many animals that live in water use
external fertilization External fertilization is a mode of reproduction in which a male organism's sperm fertilizes a female organism's egg outside of the female's body. It is contrasted with internal fertilization, in which sperm are introduced via insemination and then ...
, whereas
internal fertilization Internal fertilization is the union of an egg and sperm cell during sexual reproduction inside the female body. Internal fertilization, unlike its counterpart, external fertilization, brings more control to the female with reproduction. For inte ...
may have developed from a need to maintain
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s in a liquid medium in the
Late Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. ...
epoch. Internal fertilization with many vertebrates (such as all reptiles, some
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
, and most
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s) occurs via cloacal copulation, known as cloacal kiss (see also
hemipenis A hemipenis (plural hemipenes) is one of a pair of intromittent organs of male squamates (snakes, lizards and worm lizards). Hemipenes are usually held inverted within the body, and are everted for reproduction via erectile tissue, much like tha ...
), while
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 ...
s copulate
vagina In mammals, the vagina is the elastic, muscular part of the female genital tract. In humans, it extends from the vestibule to the cervix. The outer vaginal opening is normally partly covered by a thin layer of mucosal tissue called the hymen ...
lly, and many basal
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s reproduce sexually with external fertilization.


In spiders and insects

Spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s are often confused with
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, but they are not insects; instead, they are
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroon ...
s. Spiders have separate male and female sexes. Before mating and copulation, the male spider spins a small web and ejaculates on to it. He then stores the sperm in reservoirs on his large
pedipalp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") and ...
s, from which he transfers sperm to the female's genitals. The females can store sperm indefinitely. For primitive insects, the male deposits
spermatozoa A spermatozoon (; also spelled spermatozoön; ; ) is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote. (A zygote is a single cell, with a complete set of chromosomes, ...
on the substrate, sometimes stored within a special structure;
courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private m ...
involves inducing the female to take up the sperm package into her genital opening, but there is no actual copulation. In groups that have reproduction similar to spiders, such as
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
, males extrude sperm into secondary copulatory structures removed from their genital opening, which are then used to inseminate the female. In dragonflies, it is a set of modified
sternites The sternum (pl. "sterna") is the ventral portion of a segment of an arthropod thorax or abdomen. In insects, the sterna are usually single, large sclerites, and external. However, they can sometimes be divided in two or more, in which case the ...
on the second abdominal segment. In advanced groups of insects, the male uses its aedeagus, a structure formed from the terminal segments of the abdomen, to deposit sperm directly (though sometimes in a capsule called a ''
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
'') into the female's reproductive tract.


In mammals

Sexual behavior can be classified into behavioral states associated with
reward Reward may refer to: Places * Reward (Shelltown, Maryland), a historic home in Shelltown Maryland * Reward, California (disambiguation) * Reward-Tilden's Farm, a historic home in Chestertown Maryland Arts, entertainment, and media * "Rewa ...
motivation (" wanting"), reward consummation aka pleasure ("liking"), and
satiety Satiety ( ) is a state or condition of fullness gratified beyond the point of satisfaction, the opposite of hunger. It is a state which induces meal termination.Hetherington, M.Sensory-specific satiety and its importance in meal termination ''Neuro ...
("inhibition"); these behavioral states are regulated in mammals by reward-based sexual learning, fluctuations in various neurochemicals (i.e.,
dopamine Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic compound, organic chemical of the catecholamine and phenethylamine families. Dopamine const ...
sexual desire Sexual desire is an emotion and motivational state characterized by an interest in sexual objects or activities, or by a drive to seek out sexual objects or to engage in sexual activities. It is an aspect of sexuality, which varies significantly ...
aka "wanting";
norepinephrine Norepinephrine (NE), also called noradrenaline (NA) or noradrenalin, is an organic chemical in the catecholamine family that functions in the brain and body as both a hormone and neurotransmitter. The name "noradrenaline" (from Latin '' ad'', ...
sexual arousal Sexual arousal (also known as sexual excitement) describes the physiological and psychological responses in preparation for sexual intercourse or when exposed to sexual stimuli. A number of physiological responses occur in the body and mind 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 ...
and melanocortins
sexual attraction Sexual attraction is attraction on the basis of sexual desire or the quality of arousing such interest. Sexual attractiveness or sex appeal is an individual's ability to attract other people sexually, and is a factor in sexual selection or mat ...
), and gonadal hormone cycles and further influenced by
sex pheromone Sex pheromones are pheromones released by an organism to attract an individual of the same species, encourage them to mate with them, or perform some other function closely related with sexual reproduction. Sex pheromones specifically focus on ind ...
s and
motor reflex Tendon reflex (or T-reflex) may refer to: *The stretch reflex or muscle stretch reflex (MSR), when the stretch is created by a blow upon a muscle tendon. This is the commonly used definition of the term. Albeit a misnomer, in this sense a common ...
es (i.e., lordosis behaviour) in some mammals. These behavioral states correlate with the phases of the
human sexual response cycle The human sexual response cycle is a four-stage model of physiological responses to sexual stimulation, which, in order of their occurrence, are the excitement, plateau, orgasmic, and resolution phases. This physiological response model was fir ...
: motivation − excitement; consummation − plateau and orgasm; satiety − refraction. Sexual learning (a form of
associative learning Learning is the process of acquiring new understanding, knowledge, behaviors, skills, values, attitudes, and preferences. The ability to learn is possessed by humans, animals, and some machines; there is also evidence for some kind of learnin ...
) occurs when an animal starts to associate bodily features, personality, contextual cues, and other stimuli with genitally-induced sexual pleasure. Once formed, these associations in turn impinge upon both sexual wanting and sexual liking. In most female mammals, the act of copulation is controlled by several innate neurobiological processes, including the motor sexual reflex of
lordosis Lordosis is historically defined as an ''abnormal'' inward curvature of the lumbar spine. However, the terms ''lordosis'' and ''lordotic'' are also used to refer to the normal inward curvature of the lumbar and cervical regions of the human spi ...
.PFAFF Donald W. , SCHWARTZ-GIBLIN Susan, MACCARTHY Margareth M. , KOW Lee-Ming : Cellular and molecular mechanisms of female reproductive behaviors, in KNOBIL Ernest, NEILL Jimmy D. : The physiology of reproduction, Raven Press, 2nd edition, 1994 In males, the act of copulation is more complex, because some learning is necessary, but the innate processes (retrocontrol of penis intromission in the vagina, rhythmic movement of the pelvis, detection of female pheromones) are specific to copulation. These innate processes direct heterosexual copulation.MEISEL Robert L. , SACHS Benjamin D. : The physiology of male sexual behavior. in KNOBIL Ernest, NEILL Jimmy D. The physiology of reproduction, Raven Press, 2nd edition, 1994 Female lordosis behaviour became secondary in
hominidae The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the ea ...
and is non-functional in humans.Dixson A.F
Primate sexuality: Comparative studies of the Prosimians, Monkeys, Apes, and Human Beings
Oxford University Press, 2nd edition, 2012.
Mammals usually copulate in a dorso-ventral posture, though there are some primate species that copulate in a ventro-vental posture. Most mammals possess a
vomeronasal organ The vomeronasal organ (VNO), or Jacobson's organ, is the paired auxiliary olfactory (smell) sense organ located in the soft tissue of the nasal septum, in the nasal cavity just above the roof of the mouth (the hard palate) in various tetrapods. T ...
that is involved in pheromone detection, including
sex pheromone Sex pheromones are pheromones released by an organism to attract an individual of the same species, encourage them to mate with them, or perform some other function closely related with sexual reproduction. Sex pheromones specifically focus on ind ...
s. Despite the fact that humans do not possess this organ, adult humans appear to be sensitive to certain mammalian pheromones that putative
pheromone receptor A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
proteins in the
olfactory epithelium The olfactory epithelium is a specialized epithelial tissue inside the nasal cavity that is involved in smell. In humans, it measures 9 cm2 and lies on the roof of the nasal cavity about 7 cm above and behind the nostrils. The olfactory ...
are capable of detecting. While sex pheromones clearly play a role in modifying sexual behavior in some mammals, the capacity for general pheromone detection and the involvement of pheromones in regulating human sexual behavior has not yet been determined. The duration of copulation varies significantly between mammal species, and may be correlated with body mass, lasting longer in small mammals than in large mammals.Stallmann, Robert R., and A. H. Harcourt.
Size matters: the (negative) allometry of copulatory duration in mammals
" Biological Journal of the Linnean Society 87.2 (2006): 185-193.


See also

*
Pelvic thrust The pelvic thrust is the thrusting motion of the pelvic region, which is used for a variety of activities, such as dance or sexual activity. Sexual activity The pelvic thrust is used during copulation by many species of mammals, including hum ...


Notes


References


Bibliography

* * Møller, A. P., and T. R. Birkhead.
Copulation behaviour in mammals: evidence that sperm competition is widespread
" Biological Journal of the Linnean Society 38.2 (1989): 119–131. * Birkhead, Timothy R., L. Atkin, and A. P. Møller.
Copulation behaviour of birds
" Behaviour 101.1 (1987): 101–138. * * * * * *Carlson, Debra A
Reproductive biology of the coyote (Canis latrans): integration of behavior and physiology
Utah State University, 2008. *Castro, Ana Mafalda Lopes Sardica Velez
Mexican gray wolf courtship and mating: behavior & basic endocrinology during breeding season
Diss. Universidade de Lisboa. Faculdade de Medicina Veterinária, 2016. * Szykman, Micaela, et al.
Courtship and mating in free-living spotted hyenas.
Behaviour 144.7 (2007): 815–846. * Dixson, Alan F.
Baculum length and copulatory behavior in primates.
American Journal of Primatology 13.1 (1987): 51–60. {{Animal sexual behavior Animal sexuality Ethology Mammalian sexuality Physiology Sex positions