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Concealed ovulation

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Concealed ovulation or hidden estrus in a species is the lack of any perceptible change in an adult female (for instance, a change in appearance or scent) when she is "in heat" and near ovulation. Some examples of such changes are swelling and redness of the genitalia in baboons and bonobos Pan paniscus, and pheromone release in the feline family. In contrast, the females of humans and a few other species[1] have few external signs of fecundity, making it difficult to for the male to consciously deduct, by means of external signs only, whether or not a female is near ovulation.

While women can be taught to recognize their own level of fertility (fertility awareness), whether men can detect fertility in women is highly debated. Several small studies have found that fertile women (compared to women in infertile portions of the menstrual cycle, or using hormonal contraception) appear more attractive to men.[2][3] It has also been suggested that a woman's voice may become more attractive to men during this time.[4] Two small studies of monogamous human couples found that women initiated sex significantly more frequently when fertile, but male-initiated sex occurred at a constant rate, without regard to the woman's phase of menstrual cycle.[5] It may be that a woman's awareness of men's courtship signals[6] increases during her highly fertile phase due to an enhanced olfactory awareness of chemicals specifically found in men's body odor.[7][8]

Analyses of data provided by the post-1998 U.S. Demographic and Health Surveys found no variation in the occurrence of coitus in the menstrual phases (except during menstruation itself).[9] This is contrary to other studies, which have found female sexual desire and extra-pair copulations ("EPC's") to increase during the midfollicular to ovulatory phases (that is, the highly fertile phase).[10] These findings of differences in woman-initiated versus man-initiated sex are likely caused by the woman’s subconscious awareness of her ovulation cycle (because of hormone changes causing her to feel increased sexual desire), contrasting with the man’s inability to detect ovulation because of its being “hidden”.

In 2008, researchers announced the discovery in human semen of hormones usually found in ovulating women. They theorized that follicle stimulating hormone, luteinising hormone, and estradiol may encourage ovulation in women exposed to semen. These hormones are not found in the semen of chimpanzees, suggesting this phenomenon may be a human male counter-strategy to concealed ovulation in human females. Other researchers are skeptical that the low levels of hormones found in semen could have any effect on ovulation.[11] One group of authors has theorized that concealed ovulation and menstruation were key factors in the development of symbolic culture in early human society.[12][13]

Evolutionary hypotheses

Concealed (or “hidden”) ovulation means the period of fertility is not detectable in women, whereas female chimpanzees "advertise" ovulation by an obvious swelling of the genitals. Women can be partly aware of their ovulation along the menstrual phases, but men are essentially unable to detect ovulation in women. Most primates have semiconcealed ovulation, which could be taken as evidence that the common ancestor had semiconcealed ovulation (still found today in gorillas), and that subsequently fully concealed ovulation evolved in humans, while fully "advertised" ovulation evolved in chimpanzees (see Homininae and "Why is Sex Fun?").

Evolutionary psychologists have advanced a number of different possible explanations for concealed ovulation (and elements of several hypotheses may be true[14]). Some posit that the lack of signaling in some species is a trait retained from evolutionary ancestors, not something that existed previously and later disappeared. If signaling is supposed to have existed and was lost, then it could have been merely due to reduced adaptive importance and lessened selection,[15] or due to direct adaptive advantages for the concealment of ovulation. Yet another possibility (regarding humans specifically) is that while highly specific signaling of ovulation is absent, human female anatomy evolved to mimic permanent signaling of fertility.[16]

Paternal investment hypothesis

The paternal investment hypothesis is strongly supported by many evolutionary biologists.[14] Several hypotheses regarding human evolution integrate the idea that women increasingly required supplemental paternal investment in their offspring. The shared reliance on this idea across several hypotheses concerning human evolution increases its significance in terms of this specific phenomenon.

This hypothesis suggests that women concealed ovulation to obtain men's aid in rearing offspring. Schoroder[14] summarizes this hypothesis outlined in Alexander and Noonan’s 1979 paper: if women no longer signaled the time of ovulation, men would be unable to detect the exact period in which they were fecund. This led to a change in men's mating strategy: rather than mating with multiple women in the hope that some of them, at least, were fecund during that period, men instead chose to mate with a particular woman repeatedly throughout her menstrual cycle. A mating would be successful in resulting in conception when it occurred during ovulation, and thus, frequent matings, necessitated by the effects of concealed ovulation, would be most evolutionarily successful.

Continuous female sexual receptivity suggests human sexuality is not solely defined by reproduction; a large part of it revolves around conjugal love and communication between partners. Copulations between partners while the woman is pregnant or in the infertile period of her menstrual cycle do not achieve the original purpose of sex, that is, conception, but do strengthen the bond between these partners. Therefore, the increased copulations because of concealed ovulation are thought to have played a role in fostering pair bonds in humans.[17]

The pair bond would be very advantageous to the reproductive fitness of both partners throughout the period of pregnancy, lactation, and rearing of offspring. Pregnancy, lactation and caring for post-lactation offspring require vast amounts of energy and time on the part of the woman. She must at first consume more food, then provide food to her offspring, while her ability to forage is reduced throughout. Supplemental male investment in the mother and her offspring is advantageous to all parties. While the man supplements the woman’s limited gathered food, the woman is enabled to devote the necessary time and energy to the care of their offspring. The offspring benefits from the supplemental investment, in the form of food and defense from the father, and receives the full attention and resources of the mother. Through this shared parental investment, both man and woman would increase their offspring’s chances for survival, thereby increasing their reproductive fitness. In this way natural selection would favor the establishment of pair bonds in humans. To the extent that concealed ovulation strengthened pair bonding, selective pressure would favor concealed ovulation as well.

Another, more recent, hypothesis is that concealed ovulation is an adaptation in response to a promiscuous mating system, similar to that of our closest evolutionary relatives, bonobos and chimpanzees. The theory is that concealed ovulation evolved in women to lessen paternity certainty, which would both lessen the chances of infanticide (as a father is less likely to kill offspring that might be his), and potentially increase the number of men motivated to assist her in caring for her offspring (partible paternity). This is supported by the fact that all other mammals with concealed ovulation, such as dolphins and gray langurs, are promiscuous, and that the only other ape species that have multi-male communities, as humans do, are promiscuous. It is argued that evidence such as the Coolidge effect, showing that a man does not seem to be naturally geared towards sexual mate-guarding behavior (that is, preventing other males from having access to his sexual partner), supports the conclusion that sexual monogamy (though perhaps not social monogamy and/or pair bonding) was rare in early modern humans.[18]

Reduced infanticide hypothesis

This hypothesis suggests the adaptive advantage for women who had hidden estrus would be a reduction in the possibility of infanticide by men, as they would be unable to reliably identify, and kill, their rivals’ offspring.[14] This hypothesis is supported by recent studies of wild Hanuman langurs, documenting concealed ovulation, and frequent matings with males outside their fertile ovulatory period.[19] Heistermann et al. hypothesize that concealed ovulation is used by women to confuse paternity and thus reduce infanticide. He explains that, as ovulation is always concealed in women, so that a men can only determine paternity (and thus decide on whether to kill the woman’s child) probabilistically, based on his previous mating frequency with her, he would be unable to escape the possibility that the child might be his own, even if he were aware of promiscuous matings on the woman's part.

Sex and reward hypothesis

Schroder[14] reviews a hypothesis by Symons and Hill, that after hunting, men exchanged meat for sex with women. Women who continuously mimicked estrus may have benefited from more meat than those that did not. If this occurred with enough frequency, then a definite period of estrus would have been lost, and, with it, sexual signaling specific to ovulation would have disappeared.

Social-bonding hypothesis

Schroder[14] presents the idea of a “gradual diminution of mid-cycle estrus and concomitant continuous sexual receptivity in human women” because it facilitated orderly social relationships throughout the menstrual cycle by eliminating the periodic intensification of male-male aggressiveness in competition for mates.[14] The extended estrous period of the bonobo (reproductive-age females are in heat for 75% of their menstrual cycle) has been said to have a similar effect to the lack of a "heat" in women. While concealed human ovulation may have evolved in this fashion, extending estrus until it was no longer a distinct period, as paralleled in the bonobo, this theory of why concealed ovulation evolved has frequently been rejected. Schroder outlines the two objections to this hypothesis: (1) natural selection would need to work at a level above the individual, which is difficult to prove; and (2) selection, because it acts on the individuals with the most reproductive success, would thus favor greater reproductive success over social integration at the expense of reproductive success.

However, since 1993 when that was written, group selection models have seen a resurgence.[20][21][22] (see group selection, reciprocal altruism, and kin selection)

Cuckoldry hypothesis

Schroder in his review writes that Benshoof and Thornhill hypothesized that estrus became hidden after monogamous relationships became the norm in Homo erectus.[14] Concealed ovulation allowed the woman to mate secretly at times with a genetically superior man, and thus gain the benefit of his genes for her offspring, while still retaining the benefits of the pair bond with her usual sexual partner. Her usual sexual partner would have little reason to doubt her fidelity, because of the concealed ovulation, and would have high, albeit unfounded, paternity confidence in her offspring. His confidence would encourage him to invest his time and energy in assisting her to care for the child, even though it was not his own. Again, the idea of a man's investment being vital to the child’s survival is a central fixture of a hypothesis regarding concealed ovulation, even as the evolutionary benefits accrue to the child, the woman, and her clandestine partner, and not to her regular sexual partner.

Concealed ovulation as a side effect of bipedalism

Pawlowski[23] presents the importance of bipedalism to the mechanics and necessity of ovulation signaling. The more open savannah environment inhabited by early humans (as made available by bipedalism) brought greater danger from predators. This would have caused humans to live in denser groups, and, in such a scenario, the long-distance sexual signaling provided by female genital swellings would have lost its function. Concealed ovulation is thus argued to be a loss of function evolutionary change rather than an adaptation. Thermoregulatory systems were also modified in humans with the move to the savannah to conserve water. It is thought that female genital swellings would have incurred added cost because of ineffective evaporation of water from the area. Pawlowski continues by saying the change to bipedalism in early hominins changed both the position of female genitals and the line of vision of males. Since males could no longer constantly see the female genitals, swelling of them during estrus as a mode of signaling would have become useless. Also, anogenital swelling at each ovulatory period may have interfered with the mechanics of bipedal locomotion, and selection may have favored females who were less hindered by this occurrence. This hypothesis ultimately concludes that bipedalism, which was strongly selected for, caused the physiological changes and a loss of function of sexual signaling through female genital swelling, leading to the concealed ovulation we now observe.

Pawlowski’s paper offers views that differ from the other hypotheses regarding concealed ovulation in that it pinpoints physiological changes in early humans as the cause of concealed ovulation rather than social or behavioral ones.[23] One of the strengths of this is derived from the other hypotheses’ weaknesses – it is difficult to track the evolution of a behavior as it leaves no verifiable evidence in the form of bone or DNA. However, the fact that the Hanuman langurs also display some concealed ovulation and that it is not directly caused by a physiological change to bipedalism may suggest bipedalism was not, at least, the sole cause of concealed ovulation in humans. As stated earlier, it is possible for many elements of different hypotheses to be true regarding the selective pressures for concealed ovulation in humans.

See also

References

  1. ^ Sandy J. Andelman (June 1987). "Evolution of Concealed Ovulation in Vervet Monkeys (Cercopithecus aethiops)". The American Naturalist. 129 (6): 785–799. doi:10.1086/284675.
  2. ^ S.C. Roberts, J. Havlicek, J. Flegr, M. Hruskova, A.C. Little, B.C. Jones, D.I. Perrett and M. Petrie (August 2004). "Female facial attractiveness increases during the fertile phase of the menstrual cycle". Proceedings of the Royal Society B. 271 (Suppl 5): S270–S272. doi:10.1098/rsbl.2004.0174. PMC 1810066. PMID 15503991.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  3. ^ Geoffrey Miller, Joshua M. Tybur and Brent D. Jordan (June 2007). "Ovulatory cycle effects on tip earnings by lap dancers: economic evidence for human estrous?" (PDF). Evolution and Human Behavior. 28 (6): 375–381. doi:10.1016/j.evolhumbehav.2007.06.002. Retrieved 2008-01-21.
  4. ^ Pipitone, R. (2008-05-18). "Women's voice attractiveness varies across the menstrual cycle". Evolution and Human Behavior. 29 (4): 268–274. doi:10.1016/j.evolhumbehav.2008.02.001. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  5. ^ Susan B. Bullivant; Sarah A. Sellergren; Kathleen Stern; et al. (February 2004). "Women's sexual experience during the menstrual cycle: identification of the sexual phase by noninvasive measurement of luteinizing hormone". Journal of Sex Research. 41 (1): 82–93. doi:10.1080/00224490409552216. PMID 15216427. {{cite journal}}: Unknown parameter |author-separator= ignored (help)
  6. ^ Lee Ann Renninger, T. Joel Wade, Karl Grammer "Getting that female glance: Patterns and consequences of male nonverbal behavior in courtship contexts" Evolution and Human Behavior 25 (2004) 416–431
  7. ^ Randy Thornhill and Steven W. Gangestad "The Scent of Symmetry: A Human Sex Pheromone that Signals Fitness?" Evolution and Human Behavior 20: 175–201 (1999)
  8. ^ Brooksbank, B.W.L. Urinary excretion of androst-16-en-3 alpha-ol in human male axillary sweat. Experientia 30:864–865, 1962.
  9. ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1086/430016, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi=10.1086/430016 instead.
  10. ^ Regan, P.C. Rhythms of desire: the association between menstrual cycle phases and female sexual desire. Canadian Journal of Human Sexuality 5:145–156, 1996.
  11. ^ Motluk, Alison (2006-08-05). "The secret life of semen". New Scientist (2563). Retrieved 2008-06-28. {{cite journal}}: External link in |journal= (help)
  12. ^ Chris Knight (1991). Blood relations: menstruation and the origins of culture. New Haven, Conn: Yale University Press. ISBN 0-300-04911-0.
  13. ^ Knight, Chris (1995). "The Human Symbolic Revolution: A Darwinian Account" (PDF). Cambridge Archaeological Journal. 5 (1): 75–114. doi:10.1017/S0959774300001190. Retrieved 2006-12-13. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  14. ^ a b c d e f g h Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1016/0162-3095(93)90026-E, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi=10.1016/0162-3095(93)90026-E instead.
  15. ^ Burt, Austin (June 1992). "'Concealed ovulation' and sexual signals in primates". Folia Primatologica. 58: 1–6. doi:10.1159/000156600.
  16. ^ Frederick S. Szalay and Robert K.Costello (June 1991). "Evolution of permanent estrus displays in hominids". Journal of Human Evolution. 20 (6): 439–464. doi:10.1016/0047-2484(91)90019-R.
  17. ^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 19281665, please use {{cite journal}} with |pmid=19281665 instead.
  18. ^ Sex at Dawn. HarperCollins. 2012. ISBN 978-0-06-220794-4. {{cite book}}: |access-date= requires |url= (help); Unknown parameter |authors= ignored (help)
  19. ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1098/rspb.2001.1833, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi=10.1098/rspb.2001.1833 instead.
  20. ^ Koeslag, J.H. (1997). Sex, the prisoner's dilemma game, and the evolutionary inevitability of cooperation. J. theor. Biol. 189, 53--61
  21. ^ Koeslag, J.H. (2003). Evolution of cooperation: cooperation defeats defection in the cornfield model. J. theor. Biol. 224, 399-410
  22. ^ Wilson, D. S.; Wilson, E. O. (2008). "Evolution "for the good of the group". [Article]". American Scientist. 96 (5): 380–389.
  23. ^ a b Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1086/200017, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi=10.1086/200017 instead.

Further reading

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