Sperm competition is a term used to refer to the competitive process between spermatozoa of two or more different males to fertilize an egg of a lone female. Competition can occur when females have multiple potential mating partners. Greater choice and variety of mates increases a female's chance to produce more viable offspring. However, multiple mates for a female means an individual male has decreased chances of producing offspring. Sperm competition is an evolutionary pressure on males, and has led to the development of adaptations to increase males' chance of reproductive success. Sperm competition results in a sexual conflict of interest between males and females. Males have evolved several defensive tactics including: mate-guarding, mating plugs, and releasing toxic seminal substances to reduce female re-mating tendencies to cope with sperm competition. Offensive tactics of sperm competition involve direct interference by one male on the reproductive success of another male, for instance by physically removing another male's sperm prior to mating with a female.
Sperm competition is often compared to having tickets in a raffle; a male has a better chance of winning (i.e. fathering offspring) the more tickets he has (i.e. the more sperm he inseminates a female with). However, sperm are not free to produce, and as such males are predicted to produce sperm of a size and number that will maximize their success in sperm competition. By making many spermatozoa, males can buy more "raffle tickets", and it is thought that selection for numerous sperm has contributed to the evolution of anisogamy with very small sperm (because of the energy trade-off between sperm size and number). Alternatively males may evolve faster sperm to enable their sperm to reach and fertilize the ovum first. Dozens of adaptations have been documented in males that help them succeed in sperm competition.
Mate-guarding is a defensive behavioral trait that occurs in response to sperm competition; males try to prevent other males from approaching the female (and/or vice versa) thus preventing their mate from engaging in further copulations. Precopulatory and postcopulatory mate-guarding occurs in birds, lizards, insects and primates. Mate-guarding also exists in the fish species Neolamprologus pulcher, as some males try to "sneak" matings with females in the territory of other males. In these instances the males guard their female by keeping them in close enough proximity so that if an opponent male shows up in his territory he will be able to fight off the rival male which will prevent the female from engaging in extra-pair copulation with the rival male.
Strategic mate-guarding occurs when the male only guards the female during her fertile periods. This strategy can be more effective because it may allow the male to engage in both extra-pair paternity and within-pair paternity.
Copulatory plugs are frequently observed in insects, reptiles, some mammals, and spiders. Copulatory plugs are inserted immediately after a male copulates with a female, which reduce the possibility of fertilization by subsequent copulations from another male, by physically blocking the transfer of sperm. The Indian Meal Moth (Plodia interpunctella) are restricted in engaging in further mating activities because the spermatacore serves as a copulatory plug immediately after copulation. Bumblebee mating plugs, in addition to providing a physical barrier to further copulations, contain linoleic acid, which reduces re-mating tendencies of females.
Similarly, Drosophila melanogaster males release toxic seminal fluids, known as ACPs (accessory gland proteins), from their accessory glands to impede the female from participating in future copulations. These substances act as an anti-aphrodisiac causing a dejection of subsequent copulations, and also stimulate ovulation and oogenesis. Seminal proteins can have a strong influence on reproduction, sufficient to manipulate female behavior and physiology.
Another strategy, known as sperm partitioning, occurs when males conserve their limited supply of sperm by reducing the quantity of sperm ejected. In Drosophila, ejaculation amount during sequential copulations is reduced; this results in half filled female sperm reserves following a single copulatory event, but allows the male to mate with a larger number of females without exhausting his supply of sperm. To facilitate sperm partitioning, some males have developed complex ways to store and deliver their sperm. In the blue headed wrasse, Thalassoma bifasciatum, the sperm duct is sectioned into several small chambers that are surrounded by a muscle that allows the male to regulate how much sperm is released in one copulatory event.
A strategy common among insects is for males to participate in prolonged copulations. By engaging in prolonged copulations a male has an increased opportunity to place more sperm within the female's reproductive tract and prevent the female from copulating with other males.
It has been found that some male mollies (Poecilia) have developed deceptive social cues to combat sperm competition. Focal males will direct sexual attention toward typically non-preferred females when an audience of other males is present. This encourages the males that are watching to attempt to mate with the non-preferred female. This is done in an attempt to decrease mating attempts with the female that the focal male prefers, hence decreasing sperm competition.
Offensive adaptation behavior differs from defensive behavior because it involves an attempt to ruin the chances of another male's opportunity in succeeding in copulation by engaging in an act that tries to terminate the fertilization success of the previous male. A male on the offensive side of mate-guarding may terminate the guarding male's chances at a successful insemination by brawling with the guarding male to gain access to the female. In Drosophila, males release seminal fluids that contain additional toxins like pheromones and modified enzymes that are secreted by their accessory glands intended to destroy the sperm that have already made their way into the female's reproductive tract from a recent copulation. Based on the "last male precedence" idea, some males can remove sperm from previous males by ejaculating new sperm into the female; hindering successful insemination opportunities of the previous male.
The "good sperm hypothesis" is very common in polyandrous mating systems. The "good sperm hypothesis" suggests that a male's genetic makeup will determine the level of his competitiveness in sperm competition. When a male has "good sperm" he is able to father more viable offspring than males that do not have the "good sperm" genes. Females may select males that have these superior "good sperm" genes because it means that their offspring will be more viable and will inherit the "good sperm" genes which will increase their fitness levels when their sperm competes.
One evolutionary response to sperm competition is the variety in penis morphology of many species. For example, the shape of the human penis has been selected through sperm competition. The human penis has been selected to displace seminal fluids that were implanted within the female reproductive tract by a rival male. The thrusting action which occurs during sexual intercourse manually removes seminal fluid out of the cervix area from a previous mating.
Evolution to increase their ejaculate size in the presence of sperm competition has a consequence on testis size. Large testes can produce more sperm required for larger ejaculates, and can be found across the animal kingdom when sperm competition occurs  Males with larger testes have been documented to achieve higher reproductive success rates than males with smaller testes in male yellow pine chipmunks. Male yellow pine chipmunks that had large testes fathered more offspring than males with smaller testes.
In some insects and spiders, for instance Nephila fenestrate, the male copulatory organ breaks off or tears off at the end of copulation and remains within the female to serve as a copulatory plug. This broken genitalia is believed to be an evolutionary response to sperm competition. This damage to the male genitalia means that these males can only mate once.
Female choice for males with competitive sperm
Female factors can influence the result of sperm competition through a process known as "sperm choice". Proteins present in the female reproductive tract or on the surface of the ovum may influence which sperm succeeds in fertilizing the egg. During sperm choice females are able to discriminate and differentially use the sperm from different males. One instance where this is known to occur is inbreeding; females will preferentially use the sperm from a more distantly related male than a close relative.
It has been found that because of female choice (see sexual selection), morphology of sperm in many species occurs in many variations to accommodate or combat (see sexual conflict) the morphology and physiology of the female reproductive tract. However, it is difficult to understand the interplay between female and male reproductive shape and structure that occurs within the female reproductive tract after mating that allows for the competition of sperm. Polyandrous females mate with many male partners. Females of many species of arthropod, mollusk and other phyla have a specialized sperm-storage organ called the spermatheca in which the sperm of different males sometimes compete for increased reproductive success.
Evidence exists that illustrates the ability of genetically similar spermatozoa to cooperate so as to ensure the survival of their counterparts thereby ensuring the implementation of their genotypes towards fertilization. Cooperation confers a competitive advantage by several means, some of these include incapacitation of other competing sperm and aggregation of genetically similar spermatozoa into structures that promote effective navigation of the female reproductive tract and hence improve fertilization ability. Such characteristics lead to morphological adaptations that suit the purposes of cooperative methods during competition. For example, spermatozoa possessed by the Wood mouse (Apodemus sylvaticus) possess an apical hook which is used to attach to other spermatozoa to form mobile trains that enhance motility through the female reproductive tract. Spermatozoa that fail to incorporate themselves into mobile trains are less likely to engage in fertilization. Other evidence suggests no link between sperm competition and sperm hook morphology.
Selection to produce more sperm can also select for the evolution of larger testes. Relationships across species between the frequency of multiple mating by females and male testis size are well documented across many groups of animals. For example, among primates, female gorillas are relatively monogamous, so gorillas have smaller testes than humans, which in turn have smaller testes than the highly promiscuous bonobos. Male chimpanzees that live in a structured multi-male, multi-female community, have large testicles to produce more sperm, therefore giving him better odds to fertilize the female. Whereas the community of gorillas consist of one alpha male and two or three females, when the female gorillas are ready to mate, normally only the alpha male is their partner.
Regarding sexual dimorphism among primates, humans falls into an intermediate group with moderate sex differences in body size but relatively large testes. This is a typical pattern of primates where several males and females live together in a group and the male faces an intermediate amount of challenges from other males compared to exclusive polygyny and monogamy but frequent sperm competition.
The male black-winged damselfly provides a striking example of an adaptation to sperm competition. Female black-winged damselflies are known to mate with several males over the span of only a few hours and therefore possess a receptacle known as a spermatheca which stores the sperm. During the process of mating the male damselfly will pump his abdomen up and down using his specially adapted penis which acts as a scrub brush to remove the sperm of another male. This method proves quite successful and the male damselfly has been known to remove 90-100 percent of the competing sperm.
A similar strategy has been observed in the Dunnock, a small bird. Before mating with the polyandrous female, the male dunnock pecks at the female's cloaca in order to peck out the sperm of the previous male suitor.
A notion emerged in 1996 that in some species, including humans, a significant fraction of sperm specialize in a manner such that they cannot fertilize the egg but instead have the primary effect of stopping the sperm from other males from reaching the egg, e.g. by killing them with enzymes or by blocking their access. This type of sperm specialization became known popularly as "kamikaze sperm" or "killer sperm", but most follow-up studies to this popularized notion have failed to confirm the initial papers on the matter. While there is also currently little evidence of killer sperm in any non-human animals certain snails have an infertile sperm morph ("parasperm") that contains lysozymes, leading to speculation that they might be able to degrade a rivals' sperm.
Sperm competition has led to other adaptations such as larger ejaculates, prolonged copulation, deposition of a copulatory plug to prevent the female re-mating, or the application of pheromones that reduce the female's attractiveness. The adaptation of sperm traits, such as length, viability and velocity might be constrained by the influence of cytoplasmic DNA (e.g. mitochondrial DNA); mitochondrial DNA is inherited from the mother only and it is thought that this could represent a constraint in the evolution of sperm.
- Parker, Geoffrey A. 1970. Sperm competition and its evolutionary consequences in the insects, Biological Reviews 45: 525-567.
- Stockley, P (1997). "Sexual conflict resulting from adaptations to sperm competition". Trends Ecol. Evol 12 (4): 154–159. doi:10.1016/S0169-5347(97)01000-8. PMID 21238013.
- Wedell, Nina; Gage, Matthew J.G.; Parker, Geoffrey A. (2002). "Sperm competition, male prudence and sperm-limited females". Trends in Ecology & Evolution 17 (7): 313–320. doi:10.1016/S0169-5347(02)02533-8.
- Simmons, L.W (2001). Competition and its Evolutionary Consequences in the Insects. Princeton University Press. ISBN 0691059888.
- Birkhead, T.R; Pizzari, Tommaso (2002). "Postcopulatory sexual selection". Nat. Rev. 3 (4): 262–273. doi:10.1038/nrg774.
- Olsson et al., 1997; Wedell et al., 2002
- Jiang-Nan Yang (2010). "Cooperation and the evolution of anisogamy". Journal of Theoretical Biology 264 (1): 24–36. doi:10.1016/j.jtbi.2010.01.019. ISSN 0022-5193. PMID 20097207.
- Kokko, H; Monaghan (2001). "Predicting the direction of sexual selection". Ecology Letters 4 (2): 159–165. doi:10.1046/j.1461-0248.2001.00212.x.
- Hasselquist, D; Bensch, Staffan (1991). "Trade-off between mate guarding and mate attraction in the polygynous great reed warbler". Behav Ecol Sociobiol 28 (3): 187–193. doi:10.1007/BF00172170.
- Baer, B.; Morgan, E. D.; Schmid-Hempel, P. (2001). "A nonspecific fatty acid within the bumblebee mating plug prevents females from remating". Proceedings of the National Academy of Sciences 98 (7): 3926–3928. doi:10.1073/pnas.061027998.
- McGraw, Lisa A.; Gibson, Greg; Clark, Andrew G.; Wolfner, Mariana F. (2004). "Genes Regulated by Mating, Sperm, or Seminal Proteins in Mated Female Drosophila melanogaster". Current Biology 14 (16): 1509–1514. doi:10.1016/j.cub.2004.08.028. PMID 15324670.
- Clark, Nathaniel L.; Swanson, Willie J. (2005). "Pervasive Adaptive Evolution in Primate Seminal Proteins". PLoS Genetics 1 (3): e35. doi:10.1371/journal.pgen.0010035. PMC 1201370. PMID 16170411.
- Hunter, F. M.; Harcourt, R.; Wright, M.; Davis, L. S. (2000). "Strategic allocation of ejaculates by male Adelie penguins". Proceedings of the Royal Society B: Biological Sciences 267 (1452): 1541–1545. doi:10.1098/rspb.2000.1176.
- Rasotto, M.B; Shapiro, D. Y. (1998). "Morphology of gonoducts and male genital papilla, in the bluehead wrasse: implications and correlates on the control of gamete release". J. Fish Biol. 52 (4): 716–725. doi:10.1111/j.1095-8649.1998.tb00815.x.
- Schöfl, G; Taborsky, Michael (2002). "Prolonged tandem formation in firebugs (Pyrrhocoris apterus) serves mate-guarding". Behavioral Ecology and Sociobiology. 52 (5): 426–433. doi:10.1007/s00265-002-0524-9.
- Plath, Martin; Richter, Stephanie; Tiedemann, Ralph; Schlupp, Ingo (2008). "Male Fish Deceive Competitors about Mating Preferences". Current Biology 18 (15): 1138–1141. doi:10.1016/j.cub.2008.06.067. PMID 18674912.
- Birkhead, T.R.; Martinez, J. G.; Burke, T.; Froman, D. P. (1999). "Sperm mobility determines the outcome of sperm competition in the domestic fowl". Proc. R. Soc. Lond. B 266 (1430): 1759–1764. doi:10.1098/rspb.1999.0843.
- Hosken, D. J.; Garner, T. W. J.; Tregenza, T.; Wedell, N.; Ward, P. I. (2003). "Superior sperm competitors sire higher-quality young". Proceedings of the Royal Society B: Biological Sciences 270 (1527): 1933–1938. doi:10.1098/rspb.2003.2443.
- Searcy, W.A.; Andersson, Malte (1986). "Sexual selection and the evolution of song". Annual Review of Ecology and Systematics 17: 507–533. doi:10.1146/annurev.es.17.110186.002451. JSTOR 2097007.
- Birkhead, T.R (2000). "Defining and demonstrating postcopulatory female choice". Evolution 54 (3): 1057–1060. doi:10.1554/0014-3820(2000)054[1057:dadpfc]2.3.co;2. PMID 10937281.
- Shackelford, T.K; Goetz, Aaron T. (2007). "Adaptation to sperm competition in humans". Current Directions in Psychological Science 16: 47–50. doi:10.1111/j.1467-8721.2007.00473.x.
- Schulte-Hostedde, AI; Millar, John S. (2004). "Intraspecific variation of testis size and sperm length in the yellow-pine chipmunk". Behav. Ecol. Sociobiol 55 (3): 272–277. doi:10.1007/s00265-003-0707-z.
- Fromhage, L (2006). "Emasculation to plug up females: the significance of pedipalp damage in Nephila fenestrata". Behav. Ecol. 17 (3). doi:10.1093/beheco/arj037.
- Christenson, T.E (1989). "Sperm depletion in the golden orb-weaving spider, Nephila clavipes". J. Arachnol 17: 115–118.
- Kempenaers, B (1999). "Extra-pair paternity and egg hatchability in tree swallows: evidence for the genetic compatibility hypothesis". Behavioral Ecology 10 (3): 304–311. doi:10.1093/beheco/10.3.304.
- Snook, R (2005). "Sperm in competition: not playing by the numbers". Trends in Ecology & Evolution 20 (1): 46–53. doi:10.1016/j.tree.2004.10.011.
- Pitnick, S, Markow, T, & Spicer, G. (1999). "Evolution of multiple kinds of female sperm-storage organs in Drosophila". Evolution 53 (6): 1804–1822. doi:10.2307/2640442. JSTOR 2640442.
- Pai, Aditi; Bernasconi, Giorgina (2008). "Polyandry and female control: the red flour beetleTribolium castaneum as a case study". Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 310B (2): 148–159. doi:10.1002/jez.b.21164.
- Weigensberg, I; D.J. Fairbairn (1994). "Conflicts of interest between the sexes: a study of mating interactions in a semiaquatic bug". Animal Behaviour 48 (4): 893–901. doi:10.1006/anbe.1994.1314.
- Moore, Harry; Dvoráková, Katerina; Jenkins, Nicholas; Breed, William (2002). "Exceptional sperm cooperation in the wood mouse". Nature 418 (6894): 174–177. doi:10.1038/nature00832. PMID 12110888.
- FIRMAN, R. C.; CHEAM, L. Y.; SIMMONS, L. W. (2011). "Sperm competition does not influence sperm hook morphology in selection lines of house mice". Journal of Evolutionary Biology 24 (4): 856–862. doi:10.1111/j.1420-9101.2010.02219.x. PMID 21306461.
- Harcourt, A.H., Harvey, P.H., Larson, S.G., & Short, R.V. 1981. Testis weight, body weight and breeding system in primates, Nature 293: 55-57
- Low, Bobbi S. (2007). "Ecological and socio-cultural impacts on mating and marriage". In R.I.M. Dunbar & L. Burnett. Oxford handbook of evolutionary psychology. Oxford: Oxford University Press. pp. 449–462. ISBN 0198568304.
- Birkhead, T.R.; Hunter, F.M. (1990). "Mechanisms of sperm competition". Trends in Ecology & Evolution 5 (2): 48–52. doi:10.1016/0169-5347(90)90047-H.
- Alcock 1998
- Barrie Heather and Hugh Robertson (2005). The Field Guide to the Birds of New Zealand (Revised ed.). Auckland: Viking. ISBN 0143020404.[page needed]
- Baker 1996
- Swallow, John G; Wilkinson, Gerald S. (2002). "The long and short of sperm polymorphisms in insects". Biological Reviews of the Cambridge Philosophical Society 77 (2): 153–182. doi:10.1017/S1464793101005851. PMID 12056745.
- Buckland-Nicks, John (1998). "Prosobranch parasperm: Sterile germ cells that promote paternity?". Micron 29 (4): 267–280. doi:10.1016/S0968-4328(97)00064-4.
- Dowling, D. K.; Nowostawski, A. Larkeson; Arnqvist, G. (2007). "Effects of cytoplasmic genes on sperm viability and sperm morphology in a seed beetle: implications for sperm competition theory?". Journal of Evolutionary Biology 20 (1): 358–368. doi:10.1111/j.1420-9101.2006.01189.x. PMID 17210029.
- Alcock, John 1998. Animal Behavior. Sixth Edition. 429-519.
- Baker, Robin 1996. Sperm Wars: The Science of Sex ISBN 0-7881-6004-4.
- Eberhard, William 1996 Female Control: Sexual Selection by Cryptic Female Choice ISBN 0-691-01084-6
- Freeman, Scott; Herron, Jon C.; (2007). Evolutionary Analysis (4th ed.). Pearson Education, Inc. ISBN 0-13-227584-8.
- Olsson, M., Madsen, T. & Shine, R. 1997. Is sperm really so cheap? Costs of reproduction in male adders, Vipera berus. Proceedings of the Royal Society of London B 264: 455-459.
- Ryan, Christopher & Jethá, Calcilda. Sex at Dawn: The prehistoric origins of modern sexuality. New York: Harper, 2010.
- Shackelford, T. K. & Pound, N. 2005. Sperm Competition in Humans : Classic and Contemporary Readings ISBN 0-387-28036-7.
- Shackelford, T. K., Pound, N., & Goetz, A. T. (2005). Psychological and physiological adaptations to sperm competition in humans. Review of General Psychology, 9, 228-248. Full text
- Simmons, Leigh W. 2001. Sperm competition and its evolutionary consequences in the insects. Princeton University Press, ISBN 0-691-05988-8 and ISBN 0-691-05987-X
- Singh S R, Bashisth N. Singh and Hugo F. Hoenigsberg (2002).Female remating, sperm competition and sexual selection in Drosophila.Genet. Mol. Res. 1 (3): 178-215
- Snook, Rhonda R. Postcopulatory reproductive strategies. Encyclopedia of Life Sciences http://www.els.net