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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, an ...
, copulation is animal sexual behavior in which a male introduces sperm into the female's body, especially directly into her reproductive tract. This is an aspect of mating. Many animals that live in water use external fertilization, whereas internal fertilization may have developed from a need to maintain gametes in a liquid medium in the Late Ordovician epoch. Internal fertilization with many vertebrates (such as all reptiles, some
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
, and most birds) occurs via cloacal copulation, known as cloacal kiss (see also hemipenis), while mammals copulate vaginally, and many basal vertebrates reproduce sexually with external fertilization.


In spiders and insects

Spiders are often confused with insects, but they are not insects; instead, they are arachnids. 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 pedipalps, from which he transfers sperm to the female's genitals. The females can store sperm indefinitely. For primitive insects, the male deposits spermatozoa 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, 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 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. Spermatophore ...
'') into the female's reproductive tract.


In mammals

Sexual behavior can be classified into behavioral states associated with reward motivation (" wanting"), reward consummation aka pleasure ("liking"), and satiety ("inhibition"); these behavioral states are regulated in mammals by reward-based sexual learning, fluctuations in various neurochemicals (i.e., dopaminesexual desire aka "wanting"; norepinephrinesexual arousal; oxytocin and melanocortinssexual attraction), and gonadal hormone cycles and further influenced by sex pheromones and motor reflexes (i.e., lordosis behaviour) in some mammals. These behavioral states correlate with the phases of the human sexual response cycle: motivation − excitement; consummation − plateau and orgasm; satiety − refraction. Sexual learning (a form of associative learning) 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.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 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 tetrapo ...
that is involved in pheromone detection, including sex pheromones. Despite the fact that humans do not possess this organ, adult humans appear to be sensitive to certain mammalian pheromones that putative pheromone receptor proteins in the olfactory epithelium 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


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