Klatt, Wilhelm K.

Lobmaier, Janek S.

Lory, Vanda

Probst, Fabian

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© The Author 2015. Published by Oxford University Press on behalf  of  the International Society for Behavioral Ecology. All rights reserved. For permissions, please e-mail: journals.permissions@oup.comThe official journal of  theISBEInternational Society for Behavioral EcologyBehavioral EcologyBehavioral Ecology (2015), 26(5),1261–1267Invited CommentariesHuman sexuality and the menstrual cycle: a comment on Havlíček et al.Alan DixsonSchool of Biological Sciences, Victoria University of Wellington, BP 600, Wellington 5024, New ZealandHavlíček et al. (2015) have provided an excellent critique of  the cur-rent status of  the “ovulatory shift” hypothesis of  women’s sexual attractiveness, and the “human estrus” hypothesis of  female mate choice. The first of  these hypotheses posits that women become sig-nificantly more sexually attractive (e.g., due to changes in visual or vocal traits) during the periovulatory phase of  the menstrual cycle. The second hypothesis proposes that women are significantly more attracted to certain masculine traits (e.g., masculinized faces) during the fertile phase of  the cycle and that female mate choices are signifi-cantly affected on this basis. Yet, as Havlíček et al. point out, “physi-cal changes in women’s appearance around ovulation are almost beyond our capacity to measure.” Likewise, “even detailed acoustic analysis precludes unequivocal identification of  the follicular phase.” Such facts confirm that men are unable to determine when women are likely to ovulate, and that “shifts” in attractiveness around ovu-lation are biologically trivial, if  they exist at all. As for increased responsiveness to masculine traits during the fertile phase of  the cycle, these are likewise very small, if  they exist at all (e.g., see Peters et al. 2009; Harris 2011, 2013), and probably have no relevance to human sexual behavior in the real world (Dixson 2009, 2012).Havlíček et  al. rightly emphasize that hormones influence human sexual attractiveness via effects that are expressed over the long term, rather than during individual menstrual cycles. Thus, women who have consistently higher levels of  estradiol have more attractive faces (Law Smith et al. 2006). Women who have a healthy distribution of  body fat, and an “hourglass” body shape, have higher circulating levels of  estrogen and are more fertile (Jasieñska et  al. 2004; Singh et  al. 2010). The reason why cues relating to underlying reproductive health and fertility do not fluctuate signifi-cantly during menstrual cycles, relates to the long-term nature of  reproductive processes and mate-choice decisions that have char-acterized the course of  human evolution. Long-term monogamous or polygynous relationships between women and men occur in the majority of  recent human cultures around the world, and would likely have been present in our African ancestors (Dixson 2009). We reproduce slowly, and human offspring are dependent on their parents for many years. Where masculine mate choices are con-cerned, cues that advertise long-term female health and fertility are thus adaptive; likewise, female mate choice is focused on long-term reproductive outcomes, rather than on periovulatory, extrapair copulations in quest of  “better genes.”Perhaps, as Havlíček et  al. suggest, women who exhibit gener-ally higher levels of  circulating estradiol might also show some greater propensity to display small changes in attractiveness (and behavior) during the menstrual cycle. These small changes, if  they exist, would constitute side effects, and epiphenomena, rather than being reproductive adaptations. In which case, surely it is time to acknowledge that the “ovulatory shift” and “human estrus” hypotheses are flawed, and to accept that they should be discarded.Address correspondence to A. Dixson. E-mail: alan.dixson@vuw.ac.nz.Received 8 May 2015; accepted 17 May 2015; Advance Access publication 15 June 2015.doi:10.1093/beheco/arv085Editor-in-Chief: Leigh SimmonsREFERENCESDixson AF. 2009. Sexual selection and the origins of  human mating sys-tems. Oxford: Oxford University Press.Dixson AF. 2012. Primate sexuality: comparative studies of  the prosimians, monkeys, apes, and humans. 2nd ed. Oxford: Oxford University Press.Harris CR. 2011. Menstrual cycle and facial preferences reconsidered. Sex Roles. 64:669–681.Harris CR. 2013. Shifts in masculinity preferences across the menstrual cycle: still not there. Sex Roles. 69:507–515.Havlíček J, Cobey K, Barrett L, Klapilova K, Roberts C. 2015. The span-drels of  Santa Barbara? A  new perspective on the peri-ovulation para-digm. Behav Ecol. 26:1249–1260.Jasieñska G, Ziomkiewicz A, Ellison TP, Lipson FS, Thune I. 2004. Large breasts and narrow waist indicate high reproductive potential in women. Proc Biol Sci. 271:1213–1217.Law Smith MJL, Perrett DI, Jones BC, Cornwall RE, Moore FR, Feinberg DR, Boothroyd LG, Durrani SJ, Stirrat MR, Whiten S, et al. 2006. Facial appearance is a cue to oestrogen levels in women. Proc Roy Soc Lond B. 273:135–140.Peters M, Simmons LW, Rhodes G. 2009. Preferences across the menstrual cycle for masculinity and symmetry in photographs of  male faces and bodies. PLoS One. 4(1):e4138.Singh D, Dixson BJ, Jessop TS, Morgan BM, Dixson AF. 2010. Cross-cultural consensus for waist-to-hip ratio and women’s attractiveness. Evol Hum Behav. 31:176–181.The many sides of  the periovulational coin: a comment on Havliček et al.Janek S. Lobmaier,a,b Wilhelm K. Klatt,a,b  Vanda Lory,a,b and Fabian Probsta,baInstitute of Psychology, University of Bern, Fabrikstrasse 8, 3012 Bern, Switzerland and bCenter for Cognition Learning and Memory, University of Bern, Fabrikstrasse 8, 3012 Bern, SwitzerlandHavliček et  al. (2015) discuss the possibility that men’s increased attraction to periovulatory women and women’s cyclic shifts in Behavioral Ecologypreference for certain male traits may not constitute adaptations per se, but might instead be inevitable by-products of  putative adaptations related to between-individual differences in reproduc-tive potential. This is an interesting approach and has prompted us to reflect on aspects of  our own studies. In the following, we outline some thoughts that have emerged from our reflections, which call for a more differentiated view on what may be spandrels and what may be adaptations. We then propose an alternative explanation for why women might appear to look more attractive during the peri-ovulatory phase of  their menstrual cycle. The periovulatory paradigm is a 2-sided coin: any observable change between 2 specific cycle phases could be driven by the one or the other phase (baseline problem). If, for example, women report to be more convivial during the periovulatory phase com-pared with the luteal phase, this could be because women are more outgoing around ovulation. Alternatively, women might be more reserved and safety seeking during the luteal phase. Given that dur-ing the luteal phase a woman’s body prepares for potential preg-nancy, any risk-avoiding behavior would seem adaptive while there might be no direct advantage in being more sociable around ovula-tion. What might seem a spandrel when looking at one phase might make perfect adaptational sense when looking at the other.Furthermore, we see the need to differentiate between what may or may not be adaptations for men versus what may or may not be adaptations for women. From a man’s perspective, a healthy and feminine (i.e., attractive) appearance will always be important because cues to health and femininity putatively signal reproduc-tive potential. Insofar we agree with the authors that most men will readily pick out attractive women in a busy room. We also agree that the task would be much more difficult when asked to pick out ovulating women in the same busy room. While it is highly adap-tive for men to recognize cues to potential fertility (irrespective of  cycle phase), there is arguably no need for an adaptation that enables men to discriminate ovulating from nonovulating women in a group of  women they meet for the first time. It may however be of  adaptive value for men to be able to track the fertility win-dow of  their own (long term) partner (minimize cuckoldry risk, maximize reproduction). But such ability may rest on behavioral rather than on purely physical cues, as we will suggest below. For women on the other hand, it is not always of  equal importance to be attracted to the most masculine men. It may instead be advanta-geous to be attracted to healthy and strong men while fertile and to seek more feminine traits in a partner during the luteal phase. Such opportunistic mating strategies afford that a woman adjusts her behavior to the situational circumstances. Behavioral adjust-ments in turn require that a woman is (unconsciously) aware of  her menstrual cycle.In a series of  studies from our own lab (e.g., Bobst and Lobmaier 2012, 2014), we found that men preferred the ovulatory woman over women in their luteal phase in a forced-choice paradigm (very similar to Roberts et  al. 2004). Notwithstanding the justified criticism that forced-choice paradigms in no way resemble situations in the real world, we note that we found no evidence that differences in estra-diol may explain why ovulatory women appear to be more attrac-tive. This is in conflict with Havliček et  al.’s suggestion that men’s preference for portraits of  ovulatory women may be a by-product of  a general preference of  women with high estradiol levels. Shifts in apparent attractiveness may instead result from subtle behavioral changes: during ovulation, women may have been flirting with the camera more, resulting in more charismatic portraits. We suggest that women’s changes in attractiveness, preferences, and behavior across the menstrual cycle are not necessarily spandrels, but instead may originate in subtle appetitive changes in the woman. Because a woman can reproduce only during the fertile window of  her cycle, it is conceivable that her appetite for sex may increase subtly around ovulation, and this increased appetite may indeed be an adaptation.Address correspondence to J.S. Lobmaier. E-mail: janek.lobmaier@psy.unibe.ch.Received 3 June 2015; accepted 8 June 2015; Advance Access publication 29 June 2015.doi:10.1093/beheco/arv097Editor-in-Chief: Leigh SimmonsREFERENCESBobst C, Lobmaier JS. 2012. Men’s preference for the ovulating female is triggered by subtle face shape differences. Horm Behav. 62(4):413–417.Bobst C, Lobmaier JS. 2014. Is preference for ovulatory female’s faces asso-ciated with men’s testosterone levels? Horm Behav. 66(3):487–492.Havlíček J, Cobey KD, Barrett L, Klapilová K, Roberts SC. The spandrels of  Santa Barbara? A new perspective on the peri-ovulation paradigm. Behav Ecol. 26:1249–1260.Roberts SC, Havliček J, Flegr J, Hruskova M, Little AC, Jones BC, Perrett DI, Petrie M. 2004. Female facial attractiveness increases dur-ing the fertile phase of  the menstrual cycle. Proc Biol Sci. 271(Suppl 5):S270–S272.Are within-cycle variations in women’s sexual interests mere by-products? A comment on Havliček et al.Steven W. Gangestad and Nicholas M. Grebe Department of Psychology, University of New Mexico, MSC 03-2220, 1 University of New Mexico, Albuquerque, NM 87131, USAHavliček et al. (2015) (hereafter, HCBKR) propose that cyclic shifts in women’s mate preferences are by-products of  adaptations that adjust preferences to individual reproductive potential, as reflected by hormone levels. Similarly, men prefer hormone-related cues because they discriminate between individual women’s reproduc-tive potential, generating as by-product cyclic variation in women’s attractiveness. In their view, then, adaptations that drive cyclic vari-ation were not shaped by benefits pertaining to cyclic variation. These proposals may fruitfully inspire new theory, critical analy-sis, and research designed to test alternative conceptualizations of  cycle shifts. Nonetheless, HCBKR vastly overstate the case that researchers have neglected particular kinds of  explanations (e.g., by-product hypotheses) and insufficiently acknowledge substantial empirical and theoretical challenges their proposals face. We focus on two (see also Roney et al. 2015).We largely attend to HCBKR’s claims regarding women’s mate preferences and sexual interests. Indeed, in our view, selection has led to suppression of  incidental cues of  the fertile phase and men’s abilities to detect these cues are very poor. Moreover, men should value women for long-term reproductive potential, which we agree men discriminate better than fertile phases (though we do not see that men’s discrimination of  women’s fertile phase is solely by-product of  the latter; e.g., Puts et al. 2013). See Thornhill and Gangestad (2008).THE ROLE OF PROGESTERONEEstradiol levels peak just prior to ovulation and, hence, predict within-cycle fertility status. But as HCBKR note, mean levels also, 1262

The many sides of the periovulational coin: comment on Havliček et al

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The many sides of the periovulational coin: comment on Havliček et al

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