Summary: Sexual union of a male and a female in non-human species.

Top Publications

  1. Olivier J, de Jong T, Jos Dederen P, van Oorschot R, Heeren D, Pattij T, et al. Effects of acute and chronic apomorphine on sex behavior and copulation-induced neural activation in the male rat. Eur J Pharmacol. 2007;576:61-76 pubmed
  2. Parker G, Lessells C, Simmons L. Sperm competition games: a general model for precopulatory male-male competition. Evolution. 2013;67:95-109 pubmed publisher
    ..A negative correlation between a and M is biologically plausible, and may buffer deviations from the previous models. There is some support for our predictions from comparative analyses across dung beetle species and frog populations. ..
  3. Li B, Zhao D. Copulation behavior within one-male groups of wild Rhinopithecus roxellana in the Qinling Mountains of China. Primates. 2007;48:190-6 pubmed
    ..Three post-conception copulations were also observed in this study. These results suggest a skewed sexual competition, with multiple females competing for a single male, which was shown by courtship attempts and female interference. ..
  4. Brennan P, Prum R. The limits of sexual conflict in the narrow sense: new insights from waterfowl biology. Philos Trans R Soc Lond B Biol Sci. 2012;367:2324-38 pubmed publisher
    ..We propose a return to a broader view of conflict where indirect selection and intrasexual selection are considered important in the evolution of conflict. ..
  5. Rezaval C, Pavlou H, Dornan A, Chan Y, Kravitz E, Goodwin S. Neural circuitry underlying Drosophila female postmating behavioral responses. Curr Biol. 2012;22:1155-65 pubmed publisher
    ..Transfer of male sex peptide (SP) during copulation mediates these postmating responses via sensory neurons that coexpress the sex-determination gene fruitless (fru)..
  6. Townsend S, Deschner T, Zuberbuhler K. Female chimpanzees use copulation calls flexibly to prevent social competition. PLoS ONE. 2008;3:e2431 pubmed publisher
    The adaptive function of copulation calls in female primates has been debated for years. One influential idea is that copulation calls are a sexually selected trait, which enables females to advertise their receptive state to males...
  7. Tseng H, Tang J, Gandhi P, Luo C, Ou C, Tseng C, et al. Mutual adaptation between mouse transglutaminase 4 and its native substrates in the formation of copulatory plug. Amino Acids. 2012;42:951-60 pubmed publisher
    ..Together with the properties of TGM4 and the nature of its disulfide-linked SVS protein substrates, male mice can easily transform the semen into a rigid and durable copulatory plug, which is an important advantage in sperm competition. ..
  8. Lizé A, Doff R, Smaller E, Lewis Z, Hurst G. Perception of male-male competition influences Drosophila copulation behaviour even in species where females rarely remate. Biol Lett. 2012;8:35-8 pubmed publisher response to the perceived intensity of sperm competition, reflected in Drosophila melanogaster by increased copulation duration following prior exposure to a rival...
  9. Addison J, Hart M. Spawning, copulation and inbreeding coefficients in marine invertebrates. Biol Lett. 2005;1:450-3 pubmed
    ..This result was robust to several different analytical approaches. We note several mechanisms that might contribute to this pattern, and appeal for more studies and ideas that might help to explain our observations. ..

More Information


  1. Mazzi D, Kesäniemi J, Hoikkala A, Klappert K. Sexual conflict over the duration of copulation in Drosophila montana: why is longer better?. BMC Evol Biol. 2009;9:132 pubmed publisher
    ..The reproductive behaviour of Drosophila montana suggests the occurrence of sexual conflict over the duration of copulation. During the last stages of copulation, females vigorously attempt to dislodge the mounting male, while males ..
  2. Brown E, Gay L, Vasudev R, Tregenza T, Eady P, Hosken D. Negative phenotypic and genetic associations between copulation duration and longevity in male seed beetles. Heredity (Edinb). 2009;103:340-5 pubmed publisher
    ..We find that the duration of a male's first copulation is negatively associated with subsequent male survival, phenotypically and genetically...
  3. Ho J, Murray J, Demas G, Goodson J. Vasopressin cell groups exhibit strongly divergent responses to copulation and male-male interactions in mice. Horm Behav. 2010;58:368-77 pubmed publisher
    ..the Fos responses of eight hypothalamic and three extra-hypothalamic AVP-immunoreactive (-ir) cell groups to copulation, nonaggressive male-male interaction, and aggressive male-male interaction in both dominant and subordinate ..
  4. Aluja M, Rull J, Sivinski J, Trujillo G, Perez Staples D. Male and female condition influence mating performance and sexual receptivity in two tropical fruit flies (Diptera: Tephritidae) with contrasting life histories. J Insect Physiol. 2009;55:1091-8 pubmed publisher
    ..b>Copulation duration also depended on both male and female nutritional condition, and the interaction between male diet and ..
  5. Apger McGlaughon J, Wolfner M. Post-mating change in excretion by mated Drosophila melanogaster females is a long-term response that depends on sex peptide and sperm. J Insect Physiol. 2013;59:1024-30 pubmed publisher
  6. Vitta A, Lorenzo M. Copulation and mate guarding behavior in Triatoma brasiliensis (Hemiptera: Reduviidae). J Med Entomol. 2009;46:789-95 pubmed
    ..of this species copulate with several males in short time intervals, and we examine whether the duration of copulation may vary in the presence of other males compared with what is observed with isolated pairs...
  7. Valtonen T, Viitaniemi H, Rantala M. Copulation enhances resistance against an entomopathogenic fungus in the mealworm beetle Tenebrio molitor. Parasitology. 2010;137:985-9 pubmed publisher
    ..A previous study on Tenebrio molitor showed that copulation decreases phenoloxidase activity in the haemolymph, and concluded that copulation corrupts immunity in this ..
  8. McFarlane S, Lane J, Taylor R, Gorrell J, Coltman D, Humphries M, et al. The heritability of multiple male mating in a promiscuous mammal. Biol Lett. 2011;7:368-71 pubmed publisher
  9. Hanin O, Azrielli A, Zakin V, Applebaum S, Rafaeli A. Identification and differential expression of a sex-peptide receptor in Helicoverpa armigera. Insect Biochem Mol Biol. 2011;41:537-44 pubmed publisher
    ..Males orient to the females by upwind anemotaxis which usually leads to a successful mating. During copulation insect males transfer seminal peptides, produced in Male Accessory Glands (MAGs) which are implicated in post-..
  10. Bono J, Matzkin L, Kelleher E, Markow T. Postmating transcriptional changes in reproductive tracts of con- and heterospecifically mated Drosophila mojavensis females. Proc Natl Acad Sci U S A. 2011;108:7878-83 pubmed publisher
    ..postmating transcriptional response of the lower reproductive tract of Drosophila mojavensis females following copulation with either conspecific or heterospecific (Drosophila arizonae) males at three time points postmating...
  11. Price T, Lizé A, Marcello M, Bretman A. Experience of mating rivals causes males to modulate sperm transfer in the fly Drosophila pseudoobscura. J Insect Physiol. 2012;58:1669-75 pubmed publisher
    ..This suggests that the evolution of parasperm in flies was not driven by sperm competition and adds to the increasing evidence that males can make extremely sophisticated responses to mating competition. ..
  12. Firman R, Simmons L. Experimental evolution of sperm quality via postcopulatory sexual selection in house mice. Evolution. 2010;64:1245-56 pubmed publisher
    ..Thus, we present significant evidence that sperm competition has profound fitness consequences for both male and female house mice. ..
  13. Klingerman C, Patel A, Hedges V, Meisel R, Schneider J. Food restriction dissociates sexual motivation, sexual performance, and the rewarding consequences of copulation in female Syrian hamsters. Behav Brain Res. 2011;223:356-70 pubmed publisher
    ..In Experiment 2, neural activation increased 1h after copulation in the NAc, and did not differ significantly between 10-day food-restricted and ad libitum-fed females in any ..
  14. Shan G, Walthall W. Copulation in C. elegans males requires a nuclear hormone receptor. Dev Biol. 2008;322:11-20 pubmed publisher
    ..We suggest that UNC-55 acts as an interface between genes involved in male tail pattern formation and those responsible for function. ..
  15. Diaz Fleischer F, Arredondo J, Aluja M. Enriching early adult environment affects the copulation behaviour of a tephritid fly. J Exp Biol. 2009;212:2120-7 pubmed publisher
    ..We expanded previous studies to evaluate how early experience in an enriched environment affects copulation behaviour when animals are confronted with a complex semi-natural environment...
  16. Garcia Gonzalez F. The relative nature of fertilization success: implications for the study of post-copulatory sexual selection. BMC Evol Biol. 2008;8:140 pubmed publisher
  17. Briceño R, Eberhard W, Robinson A. Copulation behaviour of Glossina pallidipes (Diptera: Muscidae) outside and inside the female, with a discussion of genitalic evolution. Bull Entomol Res. 2007;97:471-88 pubmed
    ..aspects of the morphology and behaviour of male genitalia should often function to stimulate the female during copulation. The morphology and behaviour of the complex, species-specific male genitalia of the tsetse fly, Glossina ..
  18. Han C, Jablonski P. Female genitalia concealment promotes intimate male courtship in a water strider. PLoS ONE. 2009;4:e5793 pubmed publisher
    ..Our results indicate that the apparent victory of coevolutionary arms race by one sex in terms of morphology may trigger evolution of a behavioral phenotype in the opposite sex...
  19. Liu Y, Lebeouf B, Guo X, Correa P, Gualberto D, Lints R, et al. A cholinergic-regulated circuit coordinates the maintenance and bi-stable states of a sensory-motor behavior during Caenorhabditis elegans male copulation. PLoS Genet. 2011;7:e1001326 pubmed publisher
    ..We demonstrate that coordination of sensory inputs and motor outputs used to initiate, maintain, self-monitor, and complete an innate behavior is accomplished via the coupling of a few circuit components...
  20. Fürtbauer I, Heistermann M, Schülke O, Ostner J. Concealed fertility and extended female sexuality in a non-human primate (Macaca assamensis). PLoS ONE. 2011;6:e23105 pubmed publisher wild Assamese macaques (Macaca assamensis), a species where females lack prominent anogenital swellings and copulation calls...
  21. Gowaty P, Kim Y, Rawlings J, Anderson W. Polyandry increases offspring viability and mother productivity but does not decrease mother survival in Drosophila pseudoobscura. Proc Natl Acad Sci U S A. 2010;107:13771-6 pubmed publisher
    ..To our knowledge, this is the first study to examine simultaneously in outbred WT Drosophila pseudoobscura the lifetime costs and benefits to females of polyandry, monogamy with a single copulation, and monogamy with repeat copulations.
  22. Kuijper B, Stewart A, Rice W. The cost of mating rises nonlinearly with copulation frequency in a laboratory population of Drosophila melanogaster. J Evol Biol. 2006;19:1795-802 pubmed
    ..Females that mated more often had substantially reduced lifetime fecundity, and importantly, the fitness cost from remating rapidly accelerated. ..
  23. Tayler T, Pacheco D, Hergarden A, Murthy M, Anderson D. A neuropeptide circuit that coordinates sperm transfer and copulation duration in Drosophila. Proc Natl Acad Sci U S A. 2012;109:20697-702 pubmed publisher
    ..Here we identify a male-specific neural pathway that coordinates the timing of SSFT with the duration of copulation behavior in Drosophila...
  24. Clay Z, Pika S, Gruber T, Zuberbuhler K. Female bonobos use copulation calls as social signals. Biol Lett. 2011;7:513-6 pubmed publisher
    During mating events, females of many primate species produce loud and distinct vocalizations known as 'copulation calls'...
  25. Weir L, Grant J, Hutchings J. The influence of operational sex ratio on the intensity of competition for mates. Am Nat. 2011;177:167-76 pubmed publisher
    ..By contrast, courtship rate decreases as the OSR becomes increasingly biased, whereas mate guarding and copulation duration increase...
  26. Naccarati C, Audsley N, Keen J, Kim J, Howell G, Kim Y, et al. The host-seeking inhibitory peptide, Aea-HP-1, is made in the male accessory gland and transferred to the female during copulation. Peptides. 2012;34:150-7 pubmed publisher
    ..of the MAG, occurs at high concentrations in the gland and is transferred to the female reproductive tract on copulation. Aea-HP-1 has a pyroglutamic acid at the N-terminus, an amidated carboxyl at the C-terminus and an unusual 4-..
  27. Prudic K, Jeon C, Cao H, Monteiro A. Developmental plasticity in sexual roles of butterfly species drives mutual sexual ornamentation. Science. 2011;331:73-5 pubmed publisher
    ..Thus, reciprocal selection through time may result in mutual sexual ornamentation. ..
  28. Tan M, Jones G, Zhu G, Ye J, Hong T, Zhou S, et al. Fellatio by fruit bats prolongs copulation time. PLoS ONE. 2009;4:e7595 pubmed publisher
    ..Female bats often lick their mate's penis during dorsoventral copulation. The female lowers her head to lick the shaft or the base of the male's penis but does not lick the glans penis ..
  29. Pischedda A, Stewart A, Little M. Male × female interaction for a pre-copulatory trait, but not a post-copulatory trait, among cosmopolitan populations of Drosophila melanogaster. PLoS ONE. 2012;7:e31683 pubmed publisher
    ..Thus, the evolutionary mechanisms promoting male-female genotype interactions for pre-copulatory mating traits in D. melanogaster warrant further investigation. ..
  30. Tachikawa K, Yoshihara Y, Kuroda K. Behavioral transition from attack to parenting in male mice: a crucial role of the vomeronasal system. J Neurosci. 2013;33:5120-6 pubmed publisher
    ..Thus, the downregulation of pup pheromone-induced activation of the vomeronasal system might be important for the behavioral transition from attack to parenting in male mice. ..
  31. Koene J, ter Maat A. Coolidge effect in pond snails: male motivation in a simultaneous hermaphrodite. BMC Evol Biol. 2007;7:212 pubmed
    ..hermaphroditic pond snail, Lymnaea stagnalis, can mate in the male and female role, but within one copulation only one sexual role is performed at a time...
  32. Yapici N, Kim Y, Ribeiro C, Dickson B. A receptor that mediates the post-mating switch in Drosophila reproductive behaviour. Nature. 2008;451:33-7 pubmed
    ..SPR is highly conserved across insects, opening up the prospect of new strategies to control the reproductive and host-seeking behaviours of agricultural pests and human disease vectors. ..
  33. Boerjan B, Tobback J, De Loof A, Schoofs L, Huybrechts R. Fruitless RNAi knockdown in males interferes with copulation success in Schistocerca gregaria. Insect Biochem Mol Biol. 2011;41:340-7 pubmed publisher
    ..gene (Fru(M)) codes for a set of transcription factors that are involved in the regulation of male courtship and copulation. Fru(M) is expressed in an interconnected neuronal circuit containing central and sensory neurons as well as ..
  34. Rönn J, Katvala M, Arnqvist G. Coevolution between harmful male genitalia and female resistance in seed beetles. Proc Natl Acad Sci U S A. 2007;104:10921-5 pubmed
    ..We show that more spiny male genitalia causes more harm to females during copulation and provide comparative evidence for the correlated evolution between these antagonistic adaptations in males ..
  35. Paoli T, Palagi E, Tacconi G, Tarli S. Perineal swelling, intermenstrual cycle, and female sexual behavior in bonobos (Pan paniscus). Am J Primatol. 2006;68:333-47 pubmed
    ..We suggest that GG-rubbing, which is generally the most frequent female sexual interaction, is a tool for social assessments among females. ..
  36. de Freitas J, Paula D, Cariello M. The influence of self-fertilization performance and copulation behaviour in reproduction by cross-fertilization in groups of Biomphalaria tenagophila (Mollusca, Planorbidae). Mem Inst Oswaldo Cruz. 1997;92:739-43 pubmed
    ..the process of self-fertilization do not increase their reproductive success after the end of grouping; (b) the copulation behaviour and the presence of one snail whose eggs have a low viability rate influence the partner's ..
  37. Emmons S, Lipton J. Genetic basis of male sexual behavior. J Neurobiol. 2003;54:93-110 pubmed
    ..of genes and circuits that govern behaviors ranging from motivation and mate-searching to courtship and copulation. Some worm and fly genes have counterparts with related functions in higher animals and many more such ..
  38. Hori T, Yamsaard T, Ueta Y, Harakawa S, Kaneko E, Miyamoto A, et al. Exposure of C57BL/6J male mice to an electric field improves copulation rates with superovulated females. J Reprod Dev. 2005;51:393-7 pubmed
    It is well-known that there are considerable strain differences in the relative copulation rates between male and superovulated female mice. In particular, the C57BL/6J strain of mice has a lower rate of successful copulation...
  39. Perez Staples D, Aluja M. Sperm allocation and cost of mating in a tropical tephritid fruit fly. J Insect Physiol. 2006;52:839-45 pubmed
    ..Ejaculate size was not related to male mating history; time elapsed since the last mating, copulation duration, female thorax length or head width...
  40. Lagoda G, Muschamp J, Vigdorchik A, Hull E. A nitric oxide synthesis inhibitor in the medial preoptic area inhibits copulation and stimulus sensitization in male rats. Behav Neurosci. 2004;118:1317-23 pubmed
    Dopamine in the medial preoptic area (MPOA) facilitates copulation in male rats, and nitric oxide (NO) regulates basal and female-stimulated MPOA dopamine release...
  41. Ravi Ram K, Ji S, Wolfner M. Fates and targets of male accessory gland proteins in mated female Drosophila melanogaster. Insect Biochem Mol Biol. 2005;35:1059-71 pubmed
    ..gland proteins (Acps) in Drosophila are components of the seminal fluid and are transferred to females during copulation. In mated females, Acps enhance egg production, augment sperm storage, induce refractory mating behaviors, and ..
  42. Park S, Mann K, Lin H, Starostina E, Kolski Andreaco A, Pikielny C. A Drosophila protein specific to pheromone-sensing gustatory hairs delays males' copulation attempts. Curr Biol. 2006;16:1154-9 pubmed
    ..Together, these data indicate that CheB42a is required for a normal gustatory response to female cuticular hydrocarbon pheromones that modulate male courtship. ..
  43. Pizzari T, Cornwallis C, Løvlie H, Jakobsson S, Birkhead T. Sophisticated sperm allocation in male fowl. Nature. 2003;426:70-4 pubmed
    ..Our results indicate that female promiscuity leads to the evolution of sophisticated male sexual behaviour. ..
  44. Carvalho G, Kapahi P, Anderson D, Benzer S. Allocrine modulation of feeding behavior by the Sex Peptide of Drosophila. Curr Biol. 2006;16:692-6 pubmed
    ..Here, we show that copulation strongly stimulates female food intake...
  45. Arnqvist G, Kirkpatrick M. The evolution of infidelity in socially monogamous passerines: the strength of direct and indirect selection on extrapair copulation behavior in females. Am Nat. 2005;165 Suppl 5:S26-37 pubmed
  46. Sato S, Braham C, Putnam S, Hull E. Neuronal nitric oxide synthase and gonadal steroid interaction in the MPOA of male rats: co-localization and testosterone-induced restoration of copulation and nNOS-immunoreactivity. Brain Res. 2005;1043:205-13 pubmed the medial preoptic area (MPOA) has been implicated in various physiological functions, including male rat copulation. Based on their apparent sensitivity to gonadal steroid manipulation, we hypothesized that nNOS cells contain ..
  47. Hanlon R, Naud M, Shaw P, Havenhand J. Behavioural ecology: transient sexual mimicry leads to fertilization. Nature. 2005;433:212 pubmed
    ..These results are surprising, given the high rate at which females reject copulation attempts by males, the strong mate-guarding behaviour of consort males, and the high level of sperm competition ..
  48. Mossinson S, Yuval B. Regulation of sexual receptivity of female Mediterranean fruit flies: old hypotheses revisited and a new synthesis proposed. J Insect Physiol. 2003;49:561-7 pubmed
    ..Duration of a female's first copulation did not affect her subsequent receptivity...
  49. Schindelman G, Whittaker A, Thum J, Gharib S, Sternberg P. Initiation of male sperm-transfer behavior in Caenorhabditis elegans requires input from the ventral nerve cord. BMC Biol. 2006;4:26 pubmed
    ..The definition of circuit components is a crucial first step toward understanding how genes specify the neural circuit and hence the behavior. ..
  50. Eberhard W. The function of female resistance behavior: intromission by male coercion vs. female cooperation in sepsis flies (Diptera: Sepsidae). Rev Biol Trop. 2002;50:485-505 pubmed
    Female resistance behavior that occurs prior to intromission does not by itself imply forced copulation. Such behavior may function instead as a test of the male in order to favor some males over others, or to induce the male to desist...
  51. Rolff J, Siva Jothy M. Copulation corrupts immunity: a mechanism for a cost of mating in insects. Proc Natl Acad Sci U S A. 2002;99:9916-8 pubmed
  52. Carosi M, Heistermann M, Visalberghi E. Display of proceptive behaviors in relation to urinary and fecal progestin levels over the ovarian cycle in female tufted capuchin monkeys. Horm Behav. 1999;36:252-65 pubmed
    ..Furthermore, the ejaculations observed always occurred within proceptive periods. The results clearly indicate that female behavior is a good indicator of the periovulatory phase and can enhance female attractivity. ..
  53. Ward I, Bennett A, Ward O, Hendricks S, French J. Androgen threshold to activate copulation differs in male rats prenatally exposed to alcohol, stress, or both factors. Horm Behav. 1999;36:129-40 pubmed
    ..Such deficits may be concealed when behavior is evaluated in gonadally intact animals. ..