Copulation (zoology)

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In zoology, copulation is animal sexual behavior in which a male introduces sperm into the female's body, especially directly into her reproductive tract. [1] [2] 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 reptiles, some fish, 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. [3] [4]


In spiders and insects

Spiders are often confused with insects, but they are not insects; instead, they are arachnids. [5] [6] 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. [7]

For primitive insects, the male deposits spermatozoa on the substrate, sometimes stored within a special structure; courtship involves inducing the female to take up the sperm package into her genital opening, but there is no actual copulation. [8] [9] 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. [10] 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 ) into the female's reproductive tract. [11]

In mammals

Sexual behavior can be classified into behavioral states associated with reward motivation ("wanting"), reward consummation aka pleasure ("liking"), and satiety ("inhibition"); [12] 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. [12] [13] These behavioral states correlate with the phases of the human sexual response cycle: motivation − excitement; consummation − plateau and orgasm; satiety − refraction. [12] 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. [12] [13] 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. [14] 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. [15] Female lordosis behaviour became secondary in hominidae and is non-functional in humans. [16] Mammals usually copulate in a dorso-ventral posture, though there are some primate species that copulate in a ventro-vental posture. [17]

Most mammals possess a vomeronasal organ that is involved in pheromone detection, including sex pheromones. [18] 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. [note 1] [18] 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. [12]

The duration of copulation varies significantly between mammal species, [22] and may be correlated with body mass, lasting longer in small mammals than in large mammals. [23]

See also


  1. In humans and other animals, trace amine-associated receptors (TAARs) that are expressed in the olfactory epithelium function as olfactory receptors that detect volatile amine odorants, including certain pheromones; [19] [20] these TAARs putatively function as a class of pheromone receptors involved in the olfactive detection of social cues. [19] [20]

    A review of studies involving non-human animals indicated that TAARs in the olfactory epithelium can mediate attractive or aversive behavioral responses to an agonist. [19] This review also noted that the behavioral response evoked by a TAAR can vary across species. [19] For example, TAAR5 mediates attraction to trimethylamine in mice and aversion to trimethylamine in rats. [19] In humans, hTAAR5 presumably mediates aversion to trimethylamine, which is known to act as an hTAAR5 agonist and to possess a foul, fishy odor that is aversive to humans; [19] [21] however, hTAAR5 is not the only olfactory receptor that is responsible for trimethylamine olfaction in humans. [19] [21] As of December 2015, hTAAR5-mediated trimethylamine aversion has not been examined in published research. [21]

Related Research Articles

Fear Basic emotion induced by a perceived threat

Fear is an emotion induced by perceived danger or threat, which causes physiological changes and ultimately behavioral changes, such as mounting an aggressive response or fleeing the threat. Fear in human beings may occur in response to a certain stimulus occurring in the present, or in anticipation or expectation of a future threat perceived as a risk to oneself. The fear response arises from the perception of danger leading to confrontation with or escape from/avoiding the threat, which in extreme cases of fear can be a freeze response or paralysis.

Pheromone Secreted or excreted chemical factor that triggers a social response in members of the same species

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 impact the behavior of the receiving individuals. There are alarm pheromones, food trail pheromones, sex pheromones, and many others that affect behavior or physiology. Pheromones are used from basic unicellular prokaryotes to complex multicellular eukaryotes. Their use among insects has been particularly well documented. In addition, some vertebrates, plants and ciliates communicate by using pheromones.

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 name is derived from the fact that it lies adjacent to the unpaired vomer bone in the nasal septum. It is present and functional in all snakes and lizards, and in many mammals, including cats, dogs, cattle, pigs, and some primates; in humans and horses it is present, but is vestigial and non-functional.

The mesolimbic pathway, sometimes referred to as the reward pathway, is a dopaminergic pathway in the brain. The pathway connects the ventral tegmental area in the midbrain to the ventral striatum of the basal ganglia in the forebrain. The ventral striatum includes the nucleus accumbens and the olfactory tubercle.

Nucleus accumbens Region of the basal forebrain

The nucleus accumbens is a region in the basal forebrain rostral to the preoptic area of the hypothalamus. The nucleus accumbens and the olfactory tubercle collectively form the ventral striatum. The ventral striatum and dorsal striatum collectively form the striatum, which is the main component of the basal ganglia. The dopaminergic neurons of the mesolimbic pathway project onto the GABAergic medium spiny neurons of the nucleus accumbens and olfactory tubercle. Each cerebral hemisphere has its own nucleus accumbens, which can be divided into two structures: the nucleus accumbens core and the nucleus accumbens shell. These substructures have different morphology and functions.

Motivational salience is a cognitive process and a form of attention that motivates or propels an individual's behavior towards or away from a particular object, perceived event or outcome. Motivational salience regulates the intensity of behaviors that facilitate the attainment of a particular goal, the amount of time and energy that an individual is willing to expend to attain a particular goal, and the amount of risk that an individual is willing to accept while working to attain a particular goal.

Trimethylamine (TMA) is an organic compound with the formula N(CH3)3. It is a colorless, hygroscopic, and flammable tertiary amine. It is a gas at room temperature but is usually sold as a 40% solution in water. (It is also sold in pressurized gas cylinders.) TMA is a nitrogenous base and can be readily protonated to give trimethylammonium cation. Trimethylammonium chloride is a hygroscopic colorless solid prepared from hydrochloric acid. Trimethylamine is a good nucleophile, and this reaction is the basis of most of its applications. TMA is widely used in industry: it is used in the synthesis of choline, tetramethylammonium hydroxide, plant growth regulators or herbicides, strongly basic anion exchange resins, dye leveling agents, and a number of basic dyes. At higher concentrations it has an ammonia-like odor, and can cause necrosis of mucous membranes on contact. At lower concentrations, it has a "fishy" odor, the odor associated with rotting fish.

Animal sexual behaviour Sexual behavior of non-human animals

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 reproductively motivated or non-reproductively motivated.

Trace amine-associated receptors (TAARs), sometimes referred to as trace amine receptors, are a class of G protein-coupled receptors that were discovered in 2001. TAAR1, the first of six functional human TAARs, has gained considerable interest in academic and proprietary pharmaceutical research due to its role as the endogenous receptor for the trace amines phenylethylamine, tyramine, and tryptamine – metabolic derivatives of the amino acids phenylalanine, tyrosine and tryptophan, respectively – ephedrine, as well as the synthetic psychostimulants, amphetamine, methamphetamine and methylenedioxymethamphetamine. In 2004, it was shown that mammalian TAAR1 is also a receptor for thyronamines, decarboxylated and deiodinated relatives of thyroid hormones. TAAR2–TAAR9 function as olfactory receptors for volatile amine odorants in vertebrates.

Reward system Group of neural structures responsible for motivation and desire

The reward system is a group of neural structures responsible for incentive salience, associative learning, and positively-valenced emotions, particularly ones involving pleasure as a core component. Reward is the attractive and motivational property of a stimulus that induces appetitive behavior, also known as approach behavior, and consummatory behavior. A rewarding stimulus has been described as "any stimulus, object, event, activity, or situation that has the potential to make us approach and consume it is by definition a reward". In operant conditioning, rewarding stimuli function as positive reinforcers; however, the converse statement also holds true: positive reinforcers are rewarding.

Putative trace amine-associated receptor 3 (TAAR3) is a human pseudogene with the gene symbol TAAR3P. In other species such as mice, TAAR3 is a functional protein-coding gene that encodes a trace amine-associated receptor protein.


Trace amine-associated receptor 5 is a protein that in humans is encoded by the TAAR5 gene. In vertebrates, TAAR5 is expressed in the olfactory epithelium.

Conditioned place preference

Conditioned place preference (CPP) is a form of Pavlovian conditioning used to measure the motivational effects of objects or experiences. By measuring the amount of time an animal spends in an area that has been associated with a stimulus, researchers can infer the animal's liking for the stimulus. This paradigm can also be used to measure conditioned place aversion with an identical procedure involving aversive stimuli instead. Both procedures usually involve mice or rats as subjects. This procedure can be used to measure extinction and reinstatement of the conditioned stimulus. Certain drugs are used in this paradigm to measure their reinforcing properties. Two different methods are used to choose the compartments to be conditioned, and these are biased vs. unbiased. The biased method allows the animal to explore the apparatus, and the compartment they least prefer is the one that the drug is administered in and the one they most prefer is the one where the vehicle is injected. This method allows the animal to choose the compartment they get the drug and vehicle in. In comparison, the unbiased method does not allow the animal to choose what compartment they get the drug and vehicle in and instead the researcher chooses the compartments.

Olfaction sense that detects odors

Olfaction, or the sense of smell, is the special sense through which smells are perceived. It occurs when an odor binds to a receptor within the nasal cavity, transmitting a signal through the olfactory system. Olfaction has many functions, including detecting hazards, and pheromones, and plays a role in taste.

Olfactory memory refers to the recollection of odors. Studies have found various characteristics of common memories of odor memory including persistence and high resistance to interference. Explicit memory is typically the form focused on in the studies of olfactory memory, though implicit forms of memory certainly supply distinct contributions to the understanding of odors and memories of them. Research has demonstrated that the changes to the olfactory bulb and main olfactory system following birth are extremely important and influential for maternal behavior. Mammalian olfactory cues play an important role in the coordination of the mother infant bond, and the following normal development of the offspring. Maternal breast odors are individually distinctive, and provide a basis for recognition of the mother by her offspring.

Addiction A disease resulting in compulsive engagement in rewarding stimuli despite adverse consequences

Addiction is a biopsychosocial disorder characterized by repeated use of drugs, or repetitive engagement in a behavior such as gambling, despite harm to self and others. According to the "brain disease model of addiction," while a number of psychosocial factors contribute to the development and maintenance of addiction, a biological process that is induced by repeated exposure to an addictive stimulus is the core pathology that drives the development and maintenance of an addiction. Many scholars who study addiction argue that the brain disease model is incomplete and misleading.

Vomeronasal receptor

Vomeronasal receptors are a class of olfactory receptors that putatively function as receptors for pheromones. Pheromones have evolved in all animal phyla, to signal sex and dominance status, and are responsible for stereotypical social and sexual behaviour among members of the same species. In mammals, these chemical signals are believed to be detected primarily by the vomeronasal organ (VNO), a chemosensory organ located at the base of the nasal septum.

Odour is sensory stimulation of the olfactory membrane of the nose by a group of molecules. Certain body odours are connected to human sexual attraction. Humans can make use of body odour subconsciously to identify whether a potential mate will pass on favourable traits to their offspring. Body odour may provide significant cues about the genetic quality, health and reproductive success of a potential mate. Body odour affects sexual attraction in a number of ways including through human biology, the menstrual cycle and fluctuating asymmetry. The olfactory membrane plays a role in smelling and subconsciously assessing another human's pheromones. It also affects the sexual attraction of insects and mammals. The major histocompatibility complex genes are important for the immune system, and appear to play a role in sexual attraction via body odour. Studies have shown that body odor is strongly connected with attraction in heterosexual females. The women in the study ranked body odor as more important for attraction than “looks”. Humans may not simply depend on visual and verbal senses to be attracted to a possible partner/mate.

Non-reproductive sexual behavior consists of sexual activities animals participate in that do not lead to the reproduction of the species. Although procreation continues to be the primary explanation for sexual behavior in animals, recent observations on animal behavior have given alternative reasons for the engagement in sexual activities by animals. Animals have been observed to engage in sex for social interaction, demonstration of dominance, aggression relief, exchange for significant materials, and sexual stimulation. Observed non-procreative sexual activities include non-copulatory mounting, oral sex, genital stimulation, anal stimulation, interspecies mating, and acts of affection. There have also been observations of animals engaging in homosexual behaviors, as well as sex with dead animals and sex involving juveniles.

Pavlovian-instrumental transfer (PIT) is a psychological phenomenon that occurs when a conditioned stimulus that has been associated with rewarding or aversive stimuli via classical conditioning alters motivational salience and operant behavior. Two distinct forms of Pavlovian-instrumental transfer have been identified in humans and other animals – specific PIT and general PIT – with unique neural substrates mediating each type. In relation to rewarding stimuli, specific PIT occurs when a CS is associated with a specific rewarding stimulus through classical conditioning and subsequent exposure to the CS enhances an operant response that is directed toward the same reward with which it was paired. General PIT occurs when a CS is paired with one reward and it enhances an operant response that is directed toward a different rewarding stimulus.


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  12. 1 2 3 4 5 Georgiadis JR, Kringelbach ML, Pfaus JG (September 2012). "Sex for fun: a synthesis of human and animal neurobiology". Nature Reviews. Urology. 9 (9): 486–98. doi: 10.1038/nrurol.2012.151 . PMID   22926422. S2CID   13813765. The sexual pleasure cycle adheres to the basic structure of pleasure cycles related to other rewards (such as food), and can therefore also be expressed in terms of motivation–consummation–satiety or wanting–liking–inhibition (Figure 1; Box 2).6,11,1 ... Similar to other forms of learning, sexual behaviour develops over time as people learn to associate stimuli such as bodily features, personality, and contextual cues with genitally-induced sexual pleasure.7 Adolescence is arguably the most critical phase in sexual development ... Popular belief holds that humans also respond to some distal sexual incentive stimuli (breasts, pheromones) in an unconditioned manner, but this has been difficult to evaluate empirically (Box 1) ... Sexual wanting in both rats and humans involves interaction between gonadal hormones and external stimuli that become sexual incentives through association with genitally-induced sexual reward; pleasurable genital stimulation is thus a major factor in sexual learning ...
     Unconditioned sexual stimuli (that is, those for which the pleasurable effect requires no learning) include proximal genital tactile stimulation in humans and distal stimuli such as pheromones, odours, and certain auditory vocalizations in rats.7,16
     Sexual inhibition involves similar brain mechanisms in rats and humans
     Rats show a similar pattern of brain activation to humans in response to cues related to sexual reward
     Cortical, limbic, hypothalamic, and cerebellar regions are activated by sex-related stimuli in both humans and rats
  13. 1 2 Schultz W (2015). "Neuronal reward and decision signals: from theories to data". Physiological Reviews. 95 (3): 853–951. doi:10.1152/physrev.00023.2014. PMC   4491543 . PMID   26109341. Sexual behavior follows hormonal imbalances, at least in men, but is also strongly based on pleasure. To acquire and follow these primary alimentary and mating rewards is the main reason why the brain's reward system has evolved in the first place. Note that “primary” reward does not refer to the distinction between unconditioned versus conditioned reward; indeed, most primary rewards are learned and thus conditioned (foods are primary rewards that are typically learnt). ... Pleasure is not only one of the three main reward functions but also provides a definition of reward. As homeostasis explains the functions of only a limited number of rewards, the prevailing reason why particular stimuli, objects, events, situations, and activities are rewarding may be pleasure. This applies first of all to sex (who would engage in the ridiculous gymnastics of reproductive activity if it were not for the pleasure) and to the primary homeostatic rewards of food and liquid, and extends to money, taste, beauty, social encounters and nonmaterial, internally set, and intrinsic rewards. ... Desire makes behavior purposeful and directs it towards identifiable goals. Thus desire is the emotion that helps to actively direct behavior towards known rewards, whereas pleasure is the passive experience that derives from a received or anticipated reward. Desire has multiple relations to pleasure; it may be pleasant in itself (I feel a pleasant desire), and it may lead to pleasure (I desire to obtain a pleasant object). Thus pleasure and desire have distinctive characteristics but are closely intertwined. They constitute the most important positive emotions induced by rewards. They prioritize our conscious processing and thus constitute important components of behavioral control. These emotions are also called liking (for pleasure) and wanting (for desire) in addiction research (471) and strongly support the learning and approach generating functions of reward. ... Some of the stimuli and events that are pleasurable in humans may not even evoke pleasure in animals but act instead through innate mechanisms. We simply do not know. Nevertheless, the invention of pleasure and desire by evolution had the huge advantage of allowing a large number of stimuli, objects, events, situations, and activities to be attractive. This mechanism importantly supports the primary reward functions in obtaining essential substances and mating partners.
  14. 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
  15. 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
  16. Dixson A.F. Primate sexuality: Comparative studies of the Prosimians, Monkeys, Apes, and Human Beings. Oxford University Press, 2nd edition, 2012.
  17. Dixson, Alan F. (2009-05-15). Sexual Selection and the Origins of Human Mating Systems. OUP Oxford. ISBN   978-0-19-156973-9.
  18. 1 2 Nei M, Niimura Y, Nozawa M (December 2008). "The evolution of animal chemosensory receptor gene repertoires: roles of chance and necessity". Nature Reviews. Genetics. 9 (12): 951–63. doi:10.1038/nrg2480. PMID   19002141. S2CID   11135361. OR genes are predominantly expressed in sensory neurons of the main olfactory epithelium (MOE) in the nasal cavity. Mammals detect many types of chemicals in the air and some in the water as odorants, whereas fishes recognize water-soluble molecules, such as amino acids, bile acids, sex steroids and prostaglandins. Some mammalian OR genes are known to be expressed in other tissues, including the testis, tongue, brain and placenta17. However, the functional significance of such ‘ectopic expression’ of OR genes is not definitively known. TAARs are also expressed in the MOE. These receptors were first identified as brain receptors for the trace amines, a collection of amines that are present at low concentrations in the central nervous system18. TAARs were originally suspected to be involved in psychiatric disorders19 but are now known to function as a second class of olfactory receptors10. Some mouse TAARs recognize volatile amines that are present in urine, and it seems that the TAARs function to detect ligands associated with social cues10. ... Most mammals possess an additional olfactory organ called the vomeronasal organ (VnO). ... The VnO was previously thought to be a specialized organ for pheromone detection, but it is now known that the VnO and MOE share some overlapping functions22. ... However, at least one of the five V1R genes is expressed in the human olfactory mucosa72. Furthermore, a recent study suggests that these five genes can activate an OR-like signal transduction pathway in a heterologous expression system73. It is therefore possible that the products of these genes function as pheromone or olfactory receptors. Adult humans do not have a VnO but seem to be sensitive to pheromones74. Another interesting observation is that chickens have no functional or non-functional V1R and V2R genes or a VnO75, although birds use pheromones for mate choice and other behaviours76. It is possible that some OR genes in the MOE are able to detect pheromones, as in humans74,77.
  19. Liberles SD (October 2015). "Trace amine-associated receptors: ligands, neural circuits, and behaviors". Curr. Opin. Neurobiol. 34: 1–7. doi:10.1016/j.conb.2015.01.001. PMC   4508243 . PMID   25616211. Roles for another receptor are supported by TAAR5-independent trimethylamine anosmias in humans [32]. ... Several TAARs detect volatile and aversive amines, but the olfactory system is capable of discarding ligand-based or function-based constraints on TAAR evolution. Particular TAARs have mutated to recognize new ligands, with almost an entire teleost clade losing the canonical amine-recognition motif. Furthermore, while some TAARs detect aversive odors, TAAR-mediated behaviors can vary across species. ... The ability of particular TAARs to mediate aversion and attraction behavior provides an exciting opportunity for mechanistic unraveling of odor valence encoding.
    Figure 2: Table of ligands, expression patterns, and species-specific behavioral responses for each TAAR
  20. "Trace amine receptor: Introduction". International Union of Basic and Clinical Pharmacology. Retrieved 15 February 2014. Importantly, three ligands identified activating mouse Taars are natural components of mouse urine, a major source of social cues in rodents. Mouse Taar4 recognizes β-phenylethylamine, a compound whose elevation in urine is correlated with increases in stress and stress responses in both rodents and humans. Both mouse Taar3 and Taar5 detect compounds (isoamylamine and trimethylamine, respectively) that are enriched in male versus female mouse urine. Isoamylamine in male urine is reported to act as a pheromone, accelerating puberty onset in female mice [34]. The authors suggest the Taar family has a chemosensory function that is distinct from odorant receptors with a role associated with the detection of social cues. ... The evolutionary pattern of the TAAR gene family is characterized by lineage-specific phylogenetic clustering [26,30,35]. These characteristics are very similar to those observed in the olfactory GPCRs and vomeronasal (V1R, V2R) GPCR gene families.
  21. Wallrabenstein I, Singer M, Panten J, Hatt H, Gisselmann G (2015). "Timberol® Inhibits TAAR5-Mediated Responses to Trimethylamine and Influences the Olfactory Threshold in Humans". PLOS ONE. 10 (12): e0144704. Bibcode:2015PLoSO..1044704W. doi:10.1371/journal.pone.0144704. PMC   4684214 . PMID   26684881. While mice produce gender-specific amounts of urinary TMA levels and were attracted by TMA, this odor is repellent to rats and aversive to humans [19], indicating that there must be species-specific functions. ... Furthermore, a homozygous knockout of murine TAAR5 abolished the attraction behavior to TMA [19]. Thus, it is concluded that TAAR5 itself is sufficient to mediate a behavioral response at least in mice. ... Whether the TAAR5 activation by TMA elicits specific behavioral output like avoidance behavior in humans still needs to be examined.
  22. Naguib, Marc (2020-04-19). Advances in the Study of Behavior. Academic Press. ISBN   978-0-12-820726-0.
  23. 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.