Thursday, January 16, 2020

Same sex sexual attraction evolved as just one of a suite of traits responding to strong selection for prosocial behavior (reduced reactive aggression, increased social affiliation, social communication, & ease of social integration)

Prosociality and a Sociosexual Hypothesis for the Evolution of Same-Sex Attraction in Humans. Andrew B. Barron and Brian Hare. Front. Psychol., 16 January 2020. https://doi.org/10.3389/fpsyg.2019.02955

Abstract: Human same-sex sexual attraction (SSSA) has long been considered to be an evolutionary puzzle. The trait is clearly biological: it is widespread and has a strong additive genetic basis, but how SSSA has evolved remains a subject of debate. Of itself, homosexual sexual behavior will not yield offspring, and consequently individuals expressing strong SSSA that are mostly or exclusively homosexual are presumed to have lower fitness and reproductive success. How then did the trait evolve, and how is it maintained in populations? Here we develop a novel argument for the evolution of SSSA that focuses on the likely adaptive social consequences of SSSA. We argue that same sex sexual attraction evolved as just one of a suite of traits responding to strong selection for ease of social integration or prosocial behavior. A strong driver of recent human behavioral evolution has been selection for reduced reactive aggression, increased social affiliation, social communication, and ease of social integration. In many prosocial mammals sex has adopted new social functions in contexts of social bonding, social reinforcement, appeasement, and play. We argue that for humans the social functions and benefits of sex apply to same-sex sexual behavior as well as heterosexual behavior. As a consequence we propose a degree of SSSA, was selected for in recent human evolution for its non-conceptive social benefits. We discuss how this hypothesis provides a better explanation for human sexual attractions and behavior than theories that invoke sexual inversion or single-locus genetic models.


In most contemporary human cultures that have been studied individuals who self-identify as exclusively homosexual are rare (Ward et al., 2014; Bailey et al., 2016), but a larger minority of the population report some homosexual sexual behavior and experience and a degree of same sex sexual attraction (SSSA) (Bagley and Tremblay, 1998; Savin-Williams and Vrangalova, 2013; Bailey et al., 2016). While estimates of the population prevalence and distribution of SSSA vary (Bailey et al., 2016) contemporary studies support Kinsey et al.’s (1948, 1953) conclusion that in human populations there is continuous variation in the expression of homosexuality. The variation forms a smooth cline from a large majority who report exclusive or mostly heterosexual attraction and/or behavior, through groups who report degrees of both homosexual and heterosexual attractions and/or behavior to a small minority who report exclusive homosexual attractions and behavior (Savin-Williams and Vrangalova, 2013; Bailey et al., 2016).
For evolutionary biologists SSSA and associated homosexual sexual orientation has presented somewhat of a conundrum. SSSA persists both within and across cultures (Witham and Mathy, 1985; Crompton, 2006) and within families, since sexual orientation has high heritability (Pillard and Bailey, 1998; Mustanski et al., 2005; Santtila et al., 2008; Bailey et al., 2016). Evidence from human twin studies and genome-wide genetic association studies suggest that about one third of the variation in sexual orientation can be attributed to additive genetic factors (Santtila et al., 2008; Bailey et al., 2016; Ganna et al., 2019). For evolutionary biologists the puzzle is typically posed like this: how can a heritable SSSA persist in a population when homosexual sex of itself is non-reproductive and homosexual people have fewer offspring on average than heterosexual people (Bell et al., 1981; King et al., 2005; Wrangham, 2019). There is expected to be strong selection against genetic factors that contribute to SSSA: how, therefore, can heritable homosexual attractions persist in a population (Kirkpatrick, 2000; Gavrilets and Rice, 2006; Bártová and Valentová, 2012; Rice et al., 2012; Jeffery, 2015)?
Various explanations to this puzzle have been proposed. The prevalence of SSSA is certainly too high for the trait to be maintained by recurrent random mutation (Moran, 1972). Models have consequently been proposed to explain how SSSA could be maintained in a population as a stable genetic polymorphism, but presently there is scant or no evidence to support these theories.
The theory of sexually antagonistic selection proposes that genetic factors contributing to SSSA in one sex could persist in populations if they conferred a strong selective advantage when expressed in the other sex, and various models of this kind have been posed to explain human SSSA (Gavrilets and Rice, 2006; Camperio-Ciani et al., 2008; Rice et al., 2012). Camperio-Ciani et al. (2008) explored whether the female relatives of homosexual males had more offspring than female relatives of heterosexual males, which could be indirect evidence for antagonistic sexual selection. There is some evidence that females with male homosexual relatives have more children than females with no male homosexual relatives in a Western European population (Camperio-Ciani et al., 2008; Lemmola and Camperio-Ciani, 2009), but this finding is at best only weakly supported in other populations or cultures (Vasey et al., 2007; Bailey et al., 2016; Semenyna et al., 2017), and so overall there is little evidence for sexually antagonistic selection as an explanation for SSSA in human males. No study has yet explored whether this theory might apply to human females.
An alternative hypothesis proposes that SSSA and homosexual behavior could be maintained in a population if genetic factors contributing to these traits had pleiotropic effects that conferred a reproductive advantage. Zietsch et al. (2008) explored a version of this hypothesis and reported that psychologically masculine females and psychologically feminine men typically identified as non-heterosexual, but if did self identify as heterosexual they also self-reported a greater number of sexual partners than average for heterosexuals. We note that number of sexual partners is a long way from a measure of reproductive success or fitness. We also note that a critical test of this hypothesis was whether heterosexuals with a non-heterosexual identical twin have more sexual partners than members of heterosexual identical twin pairs. Here, there was a trend in the hypothesized direction but no statistically significant difference in number of sexual partners (Zietsch et al., 2008). Zietsch et al.’s (2008) study is certainly intriguing. The data they present are compatible with a relationship between SSSA, homosexual behavior and increased socio-sexuality – a point we develop later.
Kin-selection theories propose genes promoting SSSA could persist in a population if people expressing SSSA enhanced the reproduction of relatives (Bailey and Zuk, 2009). It is assumed the indirect fitness benefit of more relatives would compensate for the presumed fitness costs associated with SSSA and same-sex sexual behavior. Invoking kin-selection theory to explain human SSSA seems a little odd. The many examples of social and reproductive traits in animals that have evolved as a consequence of kin selection emphasize the evolution of non-reproductives, not same-sex sexual behavior (Kirkpatrick, 2000). In human societies there is very little evidence homosexual people increase the reproductive output of relatives (Bobrow and Bailey, 2001; Rahman and Hull, 2005; Vasey and VanderLaan, 2012; Abild et al., 2014; Prum, 2017) offering weak empirical support to kin selection theories for SSSA. But several studies by Vasey and others have emphasized the avuncularity (defined as altruistic uncle-like behavior) and generosity of transgender males expressing SSSA (Vasey et al., 2007; Vasey and VanderLaan, 2012, 2015; Abild et al., 2014) perhaps indicating a relationship between SSSA and affiliative behavior.
Here we propose a sociosexual hypothesis for the evolution of SSSA that explores possible adaptive social functions of same-sex directed attractions and behavior. Benefits of SSSA and same sex sexual behavior for the development and maintenance of same sex social bonds and group affiliation have been proposed previously, most notably by Kirkpatrick (2000) and later by Bártová and Valentová (2012). But here we link the evolution of human SSSA to the suite of traits that evolved as a consequence of selection for ease of social integration (prosocial behavior), within-group tolerance and social affiliation. This has been described as an evolutionary process of human self-domestication (Eisenberg et al., 1983; Clay and Zuberbühler, 2011; Gleeson, 2016; Hare, 2017; Theofanopoulou et al., 2017; Niego and Benítez-Burraco, 2019; Wrangham, 2019).
Assessing SSSA in non-human animals is not easy, but what is clear is that homosexual behavior is not a human innovation. It is widespread in primates (Sommer and Vasey, 2006) and other animals (Bagemihl, 1999; Bailey and Zuk, 2009), and is certainly ancestral to hominids. Analyses of the contexts of occurrence of homosexual sexual behavior in primate societies suggest the behavior has various diverse functions. These include appeasement, pacification, reinforcement of social dominance structures, juvenile play, social tolerance, stress reduction, and barter (Sommer and Vasey, 2006; Clay and de Waal, 2015). Heterosexual sexual behavior shows a similar diversity of expression across primate societies (Sommer and Vasey, 2006). It appears there has been an expansion of the social functions of sexual interactions (both homosexual and heterosexual) as more complex societies evolved in primates (Werner, 2006). As a consequence, sexual behavior in primates has been subject to selection for adaptive social functions as well as the obvious reproductive functions.
Social evolutionary processes have been a major driver of recent human cognition and behavior (Eisenberg et al., 1983; Dos Santos and West, 2005); particularly selection for increased intra-group tolerance and reduced intra-group aggression (Bowles and Gintis, 2013; Hare, 2017). Prosocial individuals would have more readily accessed the fitness benefits of cooperative group living (Hare, 2017), and would have gained both greater reproductive success and social mobility (Bowles and Gintis, 2013). Enhanced tolerance also would allow for smoother integration of juveniles that moved from their natal group to a new group – bringing new ideas and technology with them. Selection for prosociality is thought to have driven the recent evolution of bonobos from their chimp-like ancestor, and proto-dogs from their wolf-like ancestor also (Hare, 2017).
In humans, dogs, and bonobos, a common suite of traits has evolved as a consequence of selection for prosociality. These are juvenilization of facial features, extended cognitive developmental periods, reduced social threat responses, reduced aggression, reduced aggressive reactivity, cooperative play behavior, and increased cooperative-communicative capacity and engagement (Hare, 2017; Theofanopoulou et al., 2017; Wrangham, 2019). This set of traits is very similar to those that have arisen from artificial selection on species for reduced aggression and fear of humans in order to domesticate them (Belyaev et al., 1985; Hare, 2017). Consequently, recent human evolution has been described as a process of self-domestication arising from natural selection for prosocial behavior (Gleeson, 2016; Hare, 2017; Niego and Benítez-Burraco, 2019; Wrangham, 2019).
Across both domesticated species and self-domesticated species it is common to see an increase in expression of same-sex sexual behavior. This is part of the expansion of the contexts of sexual behavior (same-sex oriented and heterosexual) into adult play, usually interpreted as part of an adult affiliative function for sex (Dagg, 1984; Poiani, 2011). For example, in the evolution of dogs from wild dogs, and wild dogs from wolves both self-domestication and domestication have increased expression of adult sexual play and homosexual sexual behavior relative to their wild relatives (Dagg, 1984). While domestication of livestock has not always increased rates of homosexual behavior, there are several well studied examples where domestication has yielded high levels of same-sex sexual behavior among adults (Dagg, 1984; Perkins and Roselli, 2007).
The bonobo has experienced a parallel process to humans of prosocial evolution from a chimp-like ancestor (Hare, 2017; Tan et al., 2017). Like humans, bonobos show a suite of features associated with self-domestication (Hare, 2017). Bonobos exhibit higher levels than chimpanzees of same-sex sexual behavior in contexts of adult play and social affiliation also (Clay and Zuberbühler, 2011; Dixon, 2011; Woods and Hare, 2011; Bailey et al., 2016; Hare and Yamamoto, 2017).
Same-sex sexual attraction, homosexual behavior and same sex affiliations are distinct dimensions of sexuality (Bolin and Whelehan, 2009; Jordan-Young, 2010; Greenberg et al., 2016; Valentova and Varella, 2016), but they are related. SSSA is a motivator of homosexual behavior, and sexual behavior is a strong motivator of social bonds and affiliations. Sex is a strong reinforcer of pair bonds in all social mammals studied (Young and Wang, 2004). Sexual behavior in social contexts functions as a reinforcer of social bonds also (Kirkpatrick, 2000). Same-sex social bonds are likely to be as important as heterosexual social bonds for any individual operating within a social group (Kirkpatrick, 2000). A degree of SSSA could therefore reasonably confer a selective advantage, by facilitating engagement in sociosexual behavior with the associated benefits of social reinforcement, affiliation, play, appeasement, and conflict resolution (Kirkpatrick, 2000; Bártová and Valentová, 2012). Selective benefits for SSSA could be increased ease of social bonding and reduced intragroup conflict through a willingness to engage in or initiate homosexual sexual play. Human ethnographic evidence points to an adaptive benefit for SSSA in alliance formation and maintenance (Kirkpatrick, 2000; Muscarella et al., 2005).
Mechanistic analyses indicate links between increased prosociality and SSSA. Raghanti et al. (2018) have argued that the neurochemical profile of the human striatum is unique among primates with elevated dopamine, serotonin, and neuropeptide Y signaling. They argue this feature evolved early in hominid evolution and increased sensitivity to social cues to promote empathy and affiliative behavior (Raghanti et al., 2018). Self domestication in both dogs and humans is believed to have caused evolutionary changes in serotonin, oxytocin and androgen systems that regulate affiliative, threat, and aggressive behavior (Hare, 2017), and are involved in chimpanzee social affiliation (Samuni et al., 2017). These are the same endocrine systems that have been implicated in the development of human SSSA and homosexual behavior (Mustanski et al., 2002; Balthazart, 2011; Fleischman et al., 2015). In domesticated sheep changes in these neurochemical systems have been considered causal of increased levels of homosexual behavior (Perkins and Roselli, 2007). Taken together, these studies suggest an overlap between the neurochemical systems involved in affiliation and prosocial behavior and those involved in an increased incidence of same-sex sexual behavior in animals. Such a relationship is expected given that sex is itself a mechanism of social bonding in mammals (Young and Wang, 2004; Young et al., 2005).
Prosociality, increased in-group tolerance and increased social affiliation: these are extremely complex traits involving widespread changes in behavior, anatomy, and neurophysiology (Hare, 2017; Theofanopoulou et al., 2017; Raghanti et al., 2018). Genetic changes underlying the evolution of such traits are likely to be complex and highly polygenic. Presently not much is known about the genetic basis of human SSSA, but as we learn more about it, it is clear human SSSA is also highly polygenic and a complex multicomponent trait (Mustanski et al., 2005; Prum, 2017; Sanders et al., 2017; Ganna et al., 2019; Swift-Gallant et al., 2019). The high heritability of human SSSA is caused by a large number of genes each with individually small effect. These genes likely contribute to different aspects of sexuality which can assort independently (Mustanski et al., 2005; Sanders et al., 2017; Ganna et al., 2019). Genetic models for the evolution of human SSSA should therefore reflect this complexity and be polygenic and multicomponent, rather than positing individual genes of large effect, as has occurred previously (Gavrilets and Rice, 2006; Rice et al., 2012).
A polygenic and additive genetic model of SSSA is compatible with the nature and distribution of SSSA in human populations, which features continuous variation in the degree of SSSA from a majority reporting exclusively heterosexual attractions to a small minority reporting exclusively homosexual attractions (Bailey et al., 2016). Along this cline of variation individuals expressing degrees of both homosexual and heterosexual attractions are stable sexualities and not transitional forms (Bailey et al., 2000; Diamond, 2008; Rosenthal et al., 2012; Savin-Williams and Vrangalova, 2013). We propose this pattern of variation could have arisen from selection for prosociality increasing the frequency of alleles in a population across multiple loci that contribute to prosocial behavior. This would include alleles contributing to SSSA because of the benefits of sociosexual same-sex behavior for same-sex social bonding and affiliation. If a trait is highly polymorphic and polygenic [as sexual orientation seems to be (Sanders et al., 2017; Ganna et al., 2019)] the random recombination of genes in sexual reproduction would result in a spectrum of heritable variation for strength of SSSA in a population (Prum, 2017).
Given this argument one might ask why SSSA is not more common in human populations. Indeed, Kirkpatrick (2000) wondered that bisexuality might be an adaptive optimum since it would allow for sociosexual affiliative behavior with members of both sexes. Kirkpatrick (2000) proposes that any reproductive disadvantage from a low level of same-sex sexual behavior could be minor or negligible, irrespective of the degree of SSSA associated with the behavior.
To this point we note simply that while individuals reporting exclusive SSSA are rare in most contemporary human populations, SSSA is not. While specific measures vary all studies recognize that males and females reporting some degree of SSSA are relatively common, and not rare (Kinsey et al., 1948, 1953; Kirkpatrick, 2000; Mustanski et al., 2002; Bailey et al., 2016). Bisexuality is more common than homosexuality, but the nature of variation in SSSA is often not well appreciated since experimentalists are prone to force a binary dichotomy across what is in reality continuous and multivariate variation in sexuality (Jordan-Young, 2010). There may also be cultural reasons why the degree of SSSA in populations may go under-reported.
We emphasize that our hypothesis is not that homosexual people are domesticated, or even more prosocial than the population average. Rather, we recognize that self-domestication has been an important process in the recent evolution of our species as a whole. SSSA has increased in frequency in humans as a consequence of the self-domestication syndrome experienced by our species. If correct, this sociosexual hypothesis comprehends the phenomenon of human SSSA as part of broader adaptive prosocial changes in recent human cognitive and social evolution (Burkart et al., 2009, 2014; Hare, 2017).
Two other authors have remarked on a link between SSSA and selection for prosociality: Prum (2017) and Wrangham (2019). Prum (2017) argues that for humans female dispersal was the ancestral condition, with females rather than males leaving their natal group. He proposes that female SSSA could evolve as part of selection for female prosociality to aid female introgression into a new social group and strengthen female-female social bonds (Prum, 2017, p. 508). He further argues that male SSSA and homosexual behavior could have evolved through female mate choice (Prum, 2017). Females may have preferred males that show a degree of SSSA since this male trait would lessen the intensity and investment of males in sexual and social control of females, and would subsequently have fostered the evolution of prosocial males and more cooperative male–male and female–male relationships (Prum, 2017, p. 509).
Wrangham (2019) has also recognized an association between prosociality and homosexuality, but Wrangham proposes a very different hypothesis for why this association might be so. Wrangham (2019, p. 189) suggests human homosexuality is a maladaptive by product of selection against reactive aggression in humans. Wrangham (2019) argues that selection for reduced reactive aggression reduced prenatal testosterone levels in males, which resulted in a maladaptive expression of homosexuality in a minority of males.
Models of human evolution are naturally hard, if not impossible, to prove or disprove, but here we note that Wrangham’s explanation for an association between homosexuality and prosociality does not, and cannot, explain homosexuality in women. By contrast, prosocial benefits of SSSA would be expected to apply to both female–female social relationships and male–male social relationships (Kirkpatrick, 2000). Prum’s (2017) evolutionary argument is interesting in many ways, not least of which is because it proposes different (but interacting) selective pressures for the evolution of male and female SSSA in humans. Here we have argued a link between prosocial evolution and SSSA. Prum (2017) recognizes this selective force for females, but considers female mate choice the primary driver of human male SSSA, with prosociality in human males an outcome of female mate choice. This hypothesized evolutionary scenario is perhaps more complex than ours, but that does not mean it is less likely. If non-prosocial species could be found in which female mate choice had lead to the evolution of male SSSA this would lend strong support to Prum’s (2017) model for social evolution.
Wrangham’s reasoning and evidence draw on the endocrine hypothesis for human homosexuality, which has been strongly refuted (Jordan-Young, 2010). There are many variants of the endocrine hypothesis, but they all propose that SSSA is caused by some malfunction or gendered misexpression of endocrine systems considered responsible for establishing gender-typical behavioral differences between heterosexual males and heterosexual females (Mustanski et al., 2002; Balthazart, 2011; Rice et al., 2012; Bailey et al., 2016). Hypotheses vary as to when or how in development a change in endocrine systems could result in SSSA. Arguments in support of the endocrine hypothesis come from a range of experimental manipulations of mammals, including primates, which demonstrate a role for androgens in the organization and development of male and female typical sexual and social behavior, and also show that severe manipulations of endocrine systems in early development can result in males showing female-typical sexual behavior and vice-versa (Balthazart, 2011; Poiani, 2011).
The endocrine hypothesis does not, however, fit well to features of human SSSA (Jordan-Young, 2010). The example of female congenital adrenal hyperplasia (CAH) is often cited as evidence supporting an endocrine basis of human SSSA (Balthazart, 2011). This disorder causes prenatal hypertrophy of the adrenal gland, and consequently the developing fetus is exposed to higher than normal levels of testosterone. Females with CAH report a higher incidence of adult homosexual orientation than that of the population as a whole, but most females with CAH report exclusively heterosexual attraction (Meyer-Bahlburg et al., 2008; Jordan-Young, 2010). This would suggest that for women there is not a simple relationship between elevated prenatal testosterone and SSSA. Further, in both animal studies and the human cases of CAH pre- or perinatal endocrine manipulations have consequences for the development of anatomical secondary sexual characteristics and genital morphology. Female rhesus monkeys given testosterone postnatally develop an enlarged clitoris (Pfaff, 1999; Dixson, 2013) and some females with CAH also develop partially masculinized genitalia (Bailey et al., 2016). There is no evidence that homosexual people (male or female) have intersex genital development (Jordan-Young, 2010; Bailey et al., 2016) suggesting it is unlikely an endocrine imbalance pre or perinatally is causal of human SSSA.
Rice and Gavrilets (Rice et al., 2012) argued that a misexpressed epigenetic modifier of testosterone sensitivity or insensitivity that affected development of the brain only and not the body and genitals might possibly explain why homosexual people show SSSA but do not have intersex bodies. This is an interesting theory, but there is currently no evidence such a precise epigenetic modifier of testosterone sensitivity exists in either humans or other animals.
However, it is proposed, the endocrine hypothesis effectively categorizes homosexuals as partially intersex: homosexual men as partially feminized and homosexual women as partially masculinized (Mustanski et al., 2002; Balthazart, 2011). Such a portrayal of homosexuality perpetuates discredited ideas of homosexuality as sexual inversion (Ellis and Symonds, 1896), and the historic medical and psychological view of homosexuality as pathological. These views of homosexuality have long since been rejected by clinical and social psychology because in clinical psychology they have been found to be inaccurate, unsupported, and unconstructive (Haumann, 1995; Jordan-Young, 2010; Bailey et al., 2016). We argue that it is time for evolutionary psychology to also question the veracity of the endocrine hypothesis for human homosexuality.
Our proposed hypothesis for human SSSA has no requirement for sexual inversion. It would not require that SSSA be masculine-like for females or feminine-like for males. Rather, consideration of both an additive genetic model for SSSA and selection on SSSA in prosocial contexts would predict a diversity of expression of SSSA in both males and females.
We have argued that SSSA evolves as part of selection for increased prosociality. This hypothesis is testable. If it is correct there should be a detectable benefit to SSSA in contexts of within-group cohesion or cooperative tasks. Some evidence already points to a relationship between affiliation and SSSA in humans. Kirkpatrick (2000) documents ethnographic examples of SSSA and homosexual behavior strengthening important social affiliations in both males and females and SSSA supporting long term supportive social bonds. Human males self-reported a higher level of homoerotic motivation if they were primed with words related to friendship than if they were primed with words related to sex (Fleischman et al., 2015). This suggests that for males social affiliation is a greater releaser of SSSA than a sexual context (Fleischman et al., 2015). Whether within-group SSSA enhances cooperation and group performance to provide individual selective benefits remains to be tested, however.
Animal models could provide a powerful resource to explore these questions. We have described how homosexual behavior is more common in highly prosocial species than non-prosocial close relatives. We would predict homosexual behavior to enhance cooperation, group cohesion and performance and ultimately increase the reproductive success of individuals that are part of a high-functioning group in animals also. Comparing the consequences of homosexual behavior in bonobos and chimpanzees for group function would be a test of this hypothesis (Moscovice et al., 2019).
If the sociosexual hypothesis of SSSA evolution is correct we would expect to see an introgression of systems causal of human SSSA and social and affiliative behavior at both genetic and physiological levels of analysis. As we have discussed above, current evidence is compatible with this hypothesis, but significant gaps remain in our understanding of the genomic and neurophysiological basis of human sexual orientation and much work remains to be done.
Exploration of human SSSA has thus far been dominated by assumptions that the trait must be maladaptive (Bell et al., 1981; King et al., 2005; Wrangham, 2019). It may be timely and beneficial to explore alternatives that consider the sociosexual adaptive functions of same sex attraction and sexual behavior, and the full spectra of expression of SSSA.

Wednesday, January 15, 2020

Male Urogenital System Mapped Onto the Sensory Cortex Thru Functional Magnetic Resonance Imaging: The nipple was found to project to the same cortical region as the genitals

Allen K, Wise N, Frangos E, et al. Male Urogenital System Mapped Onto the Sensory Cortex: Functional Magnetic Resonance Imaging Evidence. J Sex Med 2020;XX:XXX–XXX. https://doi.org/10.1016/j.jsxm.2019.12.007

Abstract
Introduction The projection of the human male urogenital system onto the paracentral lobule has not previously been mapped comprehensively.

Aim To map specific urogenital structures onto the primary somatosensory cortex toward a better understanding of sexual response in men.

Methods Using functional magnetic resonance imaging, we mapped primary somatosensory cortical responses to self-stimulation of the penis shaft, glans, testicles, scrotum, rectum, urethra, prostate, perineum, and nipple. We further compared neural response with erotic and prosaic touch of the penile shaft.

Main Outcome Measure We identified the primary mapping site of urogenital structures on the paracentral lobule and identified networks involved in perceiving touch as erotic.

Results We mapped sites on the primary somatosensory cortex to which components of the urogenital structures project in men. Evidence is provided that penile cutaneous projection is different from deep penile projection. Similar to a prior report in women, we show that the nipple projects to the same somatosensory cortical region as the genitals. Evidence of differential representation of erotic and nonerotic genital self-stimulation is also provided, the former activating sensory networks other than the primary sensory cortex, indicating a role of “top-down” activity in erotic response.

Clinical Implications We map primary sites of projection of urogenital structures to the primary somatosensory cortex and differentiate cortical sites of erotic from nonerotic genital self-stimulation.

Strength & Limitations To our knowledge, this is the first comprehensive mapping onto the primary somatosensory cortex of the projection of the components of the urogenital system in men and the difference in cortical activation in response to erotic vs nonerotic self-stimulation. The nipple was found to project to the same cortical region as the genitals. Evidence is provided that superficial and deep penile stimulation project differentially to the cortex, suggesting that sensory innervation of the penis is provided by more than the (pudendal) dorsal nerve.

Conclusion This study reconciles prior apparently conflicting findings and offers a comprehensive mapping of male genital components to the paracentral lobule. We provide evidence of differential projection of light touch vs pressure applied to the penile shaft, suggesting differential innervation of its superficial, vs deep structure. Similar to the response in women, we found nipple projection to genital areas of the paracentral lobule. We also provide evidence of differential representation of erotic and nonerotic genital self-stimulation, the former activating sensory networks other than the primary sensory cortex, indicating a role of top-down activity in erotic response.

Rolf Degen summarizing: People with a nice personality were more ready to fall for mundane bullshit

Expanding the bullshit research out of pseudo-transcendental domain. Vladimíra Čavojová, Ivan Brezina, Marek Jurkovič. Current Psychology, January 15 2020. https://link.springer.com/article/10.1007/s12144-020-00617-3

Abstract: The ability to distinguish bullshit from factual, but less appealing, information is becoming a crucial skill in present-day society. The aim of this paper was to extend the research conducted by Pennycook et al. (Judgment and Decision Making, 10(6), 549–563, 2015) by developing and validating the General Bullshit Receptivity Scale (GBRS), designed to measure bullshit in more general and non-transcendental context, than the Bullshit Receptivity Scale (BSR). In this paper we assessed the psychometric properties of the GBRS on representative sample of Slovak participants (N = 458) and explored the relation between the GBRS and original BSR scale, epistemically suspect beliefs, ontological confusion, spirituality, personality and analytical thinking. People who thought the randomly generated transcendental statements were more profound were more susceptible to accepting more general bullshit and other epistemically suspect beliefs, and this tendency was accompanied with a low level of analytical thinking. Non-cognitive factors (agreeableness, spirituality) also contributed to perceptions the bullshit was profound and truthful. The most “impressive” bullshit was bullshit that did not contain obscure vocabulary but seemingly provided recipients with intuitive, though untruthful insights. We believe we have succeeded in constructing and refining a new measure for detecting other kind of bullshit that enables us to better understand the underlying cognitive mechanisms and personality variables.

Keywords: Bullshit receptivity General bullshit receptivity scale Epistemically suspect beliefs Ontological confusion Analytical thinking

Women who reported to have sex weekly during the study period were 28% less likely to experience menopause than women who had sex less than monthly

Sexual frequency is associated with age of natural menopause: results from the Study of Women's Health Across the Nation. Megan Arnot and Ruth Mace. Royal Society Open Science, Volume 7, Issue 1, January 15 2020. https://doi.org/10.1098/rsos.191020

Abstract: It is often observed that married women have a later age of natural menopause (ANM) than unmarried women; however, the reason for this association is unknown. We test an original hypothesis that sexual frequency acts as a bio-behavioural mediator between marital status and ANM. We hypothesize that there is a trade-off between continued ovulation and menopause based on the woman's chances of becoming pregnant. If a woman is sexually inactive, then pregnancy is impossible, and continued investment in ovulation would not be adaptive. In addition, we test an existing hypothesis that the observed relationship is because of the exposure to male pheromones. Data from 2936 women were drawn from 11 waves of the Study of Women's Health Across the Nation, which is a longitudinal study conducted in the United States. Using time-varying Cox regression, we found no evidence for the pheromone hypothesis. However, we did observe that women who reported to have sex weekly during the study period were 28% less likely to experience menopause than women who had sex less than monthly. This is an indication that ANM may be somewhat facultative in response to the likelihood of pregnancy.

4. Discussion

While women often stop reproducing many years prior to the menopause [63], the permanent reproductive cessation resulting from the menopause means a woman is no longer physically able to increase her direct fitness. Many lifestyle factors have been found to associate with ANM, and these are seldom discussed from an evolutionary perspective. Here, we have focused specifically on the relationship between marital status and ANM, testing an original hypothesis that sexual frequency acts as a bio-behavioural mediator between ANM and marital status, in addition to the existing hypothesis that married women enter menopause later owing to male pheromones [7]. To test the latter hypothesis, three measures of male–female cohabitation were used as a proxy of exposure to male pheromones, and we found no evidence to suggest that menopause timing was responsive to living with a male, and therefore possibly male pheromones. It should be noted that this hypothesis could be fundamentally flawed, as there is no conclusive evidence either that humans produce pheromones, or that they are capable of detecting them [64]. Nonetheless, this is, to our knowledge, the first study addressing the pheromonal hypothesis since it was originally proposed, and while male household presence is merely a proxy of pheromones—and it may be that the hypothesis is moot owing to the absence of evidence for human pheromones—it is an indication that the relationship between marital status and ANM is not capturing the effect of male pheromones on the menstrual cycle.
This study did not replicate findings from previous research that married women enter menopause later. In fact, following complete adjustment, the converse was found, with women who were married or in a relationship having an increased risk of entering menopause compared to divorced, separated and single women. Conflicting results regarding marital status' effect on ANM have been found elsewhere (e.g. [65]), and one reason for this may be the way in which the researcher chooses to code the variable. In this analysis, romantic partnerships that may not have been acknowledged in previous studies owing to having not been formalized by a marriage ceremony (e.g. cohabiting but unmarried) were taken into account. In addition, some prior studies have not included marital status as time-variant and dichotomized the variable as ‘ever married’ or ‘never married’ (e.g. [66]). Hence, the responsive way in which this study coded marital status may account for the difference in results.
Another reason for this difference may be the cultural setting of previous studies. For example, research originating from Iran found that ever married women experience a later menopause than those who never married [8]. However, in the case of Iran where dowry is still common practice, it means marriage is contingent upon family wealth [67]. Therefore, the effect of marital status on ANM would be confounded by a woman's socioeconomic position, which itself would relate to other aspects of her health and life history that have been associated with menopause timing—such as BMI and age of menarche—therefore resulting in a significant difference in ANM between those who have and have not been married. Within Iran, sex outside of marriage is prohibited both legally and socially, meaning marital status would be highly correlated with sexual behaviour [68]. Hence, it may be that previous studies identifying married women enter menopause later are simply capturing the effect of health and lifestyle patterns that themselves associate with both marital status and menopause timing, rather than demonstrating that marital status itself is a cross-cultural correlate of ANM. Future research should aim to address the cultural setting in which the data were collected when interpreting the results.
Evidence supporting the notion that ANM associated with sexual frequency during the pre- and peri-menopause was found. Even following complete adjustment, results still indicated that women who engage in sexual activity weekly or monthly have a lower risk of entering menopause relative to those who report having some form of sex less than monthly. If we interpret these results from a fitness-maximizing framework, it may be the physical cues of sex signal to the body that there is a possibility of becoming pregnant, and therefore an adaptive trade-off may occur between continued energetic investment in ovulation and reproductive cessation. During ovulation, the woman's immune function is impaired making the body more susceptible to disease [28,29]. Hence, if a pregnancy is unlikely owing to a lack of sexual activity, then it would not be beneficial to allocate energy to a costly process, especially if there is the option to invest resources into existing kin [30,31]. The idea that women cease fertility in order to invest in kin is known as the Grandmother Hypothesis, which predicts that the menopause originally evolved in humans to reduce reproductive conflict between different generations of females, and allow women to increase their inclusive fitness through investing in their grandchildren [30,69,70]. It may be costly for a woman to cease ovulatory function if the chances of her becoming pregnant are still high. In other words, if she is still able to increase her direct fitness, then it may be better to maintain the function of her menstrual cycle for slightly longer.
It should be noted that there may be a bidirectional relationship between the physical condition of the woman when approaching the menopause and sexual engagement. As oestrogen levels decline, women are more likely to experience vaginal dryness and discomfort, making them less inclined to engage in sex [71]. This study has attempted to control for this factor through adjusting for both oestradiol levels and the woman's self-perceived overall health, with the association between sexual frequency and ANM still persisting following this adjustment. This suggests that—even when controlling for the complicated relationship between health, hormonal fluctuations and desire for sex—the menopause may be somewhat facultative in response to sexual behaviour, rather than being solely the result of a physiological constraint (e.g. degrading oocyte quality).

Firms with higher H-1B visa lottery win rates are more likely to receive additional venture capital funding, to have a successful exit via an IPO or acquisition, & to receive more patents & patent citations

Give Me Your Tired, Your Poor, Your High-Skilled Labor: H-1B Lottery Outcomes and Entrepreneurial Success. Stephen G. Dimmock, Jiekun Huang, Scott J. Weisbenner. NBER Working Paper No. 26392, October 2019. https://www.nber.org/papers/w26392

Abstract: We study how access to high-skill labor affects the outcomes of start-up firms. We obtain exogenous variation in firms’ ability to access skilled labor by using win rates in H-1B visa lotteries. Relative to other firms that also applied for H-1B visas, firms with higher lottery win rates are more likely to receive additional venture capital funding and to have a successful exit via an IPO or acquisition. H-1B visa lottery winners also subsequently receive more patents and patent citations. Overall, our results show that access to high-skill labor is a critical determinant of success for start-up firms.




Approval & regulation processes for pharmaceutical agents have increased in complexity; FDA review times have shortened, but total time from the authorization of clinical testing to approval has remained at approx 8 years

FDA Approval and Regulation of Pharmaceuticals, 1983-2018. Jonathan J. Darrow, Jerry Avorn, Aaron S. Kesselheim. JAMA. 2020;323(2):164-176. doi:10.1001/jama.2019.20288

Key Points
Question  How has the regulation of prescription drugs evolved from the 1980s to 2018, and what trends have occurred in drug approvals?

Findings  Approvals of new generic drugs have increased over time, leading to greater competition. Technological advances have been reflected in increased approvals of biologics over time. The number of expedited development and approval programs has expanded greatly since 1983, reducing the amount of evidence available at the time of approval and increasing uncertainty about the existence or amount of clinical benefit. These regulatory innovations have not clearly led to an increase in new drug approvals or to reduced total development times.

Meaning  While retaining policies that encourage efficient review, Congress and other government officials should also consider the implications of less rigorous clinical outcome requirements and whether the current complex array of regulatory programs should be simplified.

Abstract
Importance  US law requires testing of new drugs before approval to ensure that they provide a well-defined benefit that is commensurate with their risks. A major challenge for the US Food and Drug Administration (FDA) is to achieve an appropriate balance between rigorous testing and the need for timely approval of drugs that have benefits that outweigh their risks.

Objective  To describe the evolution of laws and standards affecting drug testing, the use of new approval programs and standards, expansions of the role and authority of the FDA, and changes in the number of drugs approved from the 1980s to 2018.

Evidence  Sources of evidence included principal federal laws and FDA regulations (1962-2018) and FDA databases of approved new drugs (1984-2018), generic drugs (1970-2018), biologics (1984-2018), and vaccines (1998-2018); special development and approval programs (Orphan drug [1984-2018], Fast-Track [1988-2018], Priority Review and its predecessors [1984-2018], Accelerated Approval [1992-2018], and Breakthrough Therapy [2012-2018]); expanded access (2010-2017) and Risk Evaluation and Mitigation Strategies (2008-2018); and user fees paid to the FDA by industry (1993-2018).

Findings  From 1983 to 2018, legislation and regulatory initiatives have substantially changed drug approval at the FDA. The mean annual number of new drug approvals, including biologics, was 34 from 1990-1999, 25 from 2000-2009, and 41 from 2010-2018. New biologic product approvals increased from a median of 2.5 from 1990-1999, to 5 from 2000-2013, to 12 from 2014-2018. The median annual number of generic drugs approved was 136 from 1970 to the enactment of the Hatch-Waxman Act in 1984; 284 from 1985 to the enactment of the Generic Drug User Fee Act in 2012; and 588 from 2013-2018. Prescription drug user fee funding expanded from new drugs and biologics in 1992 to generic and biosimilar drugs in 2012. The amount of Prescription Drug User Fee Act fees collected from industry increased from an annual mean of $66 million in 1993-1997 to $820 million in 2013-2017, and in 2018, user fees accounted for approximately 80% of the salaries of review personnel responsible for the approval of new drugs. The proportion of drugs approved with an Orphan Drug Act designation increased from 18% (55/304) in 1984-1995, to 22% (82/379) in 1996-2007, to 41% (154/380) in 2008-2018. Use of Accelerated Approval, Fast-Track, and Priority Review for new drugs has increased over time, with 81% (48/59) of new drugs benefiting from at least 1 such expedited program in 2018. The proportion of new approvals supported by at least 2 pivotal trials decreased from 80.6% in 1995-1997 to 52.8% in 2015-2017, based on 124 and 106 approvals, respectively, while the median number of patients studied did not change significantly (774 vs 816). FDA drug review times declined from more than 3 years in 1983 to less than 1 year in 2017, but total time from the authorization of clinical testing to approval has remained at approximately 8 years over that period.

Conclusions and Relevance  Over the last 4 decades, the approval and regulation processes for pharmaceutical agents have evolved and increased in complexity as special programs have been added and as the use of surrogate measures has been encouraged. The FDA funding needed to implement and manage these programs has been addressed by expanding industry-paid user fees. The FDA has increasingly accepted less data and more surrogate measures, and has shortened its review times.

Just How Important Is Breast Size in Attraction? Across cultures, men do not consistently prefer large breasts

Just How Important Is Breast Size in Attraction? Across cultures, men do not consistently prefer large breasts. Robert D. Martin, Jan 14, 2020. https://www.psychologytoday.com/intl/blog/how-we-do-it/202001/just-how-important-is-breast-size-in-attraction

As the previous blog piece reported, a woman’s capacity for milk production has no connection with breast size before pregnancy. What matters is the increase in size during pregnancy. Any evolutionary explanation for men’s responses to a woman’s breasts in terms of reproductive capacity must hence lie elsewhere.

However, one key finding from a host of studies of female breast size is that men predominantly prefer medium size, not large. This bears directly on the widespread demand for cosmetic breast augmentation. The major motivation seems to be a woman’s own perception of her body image rather than male preferences.

Various attempts have been made to link male preferences for female breast size to fertility indicators with some evolutionary function. But such preferences are too inconsistent across cultures to permit clear recognition of an evolutionary basis. Moreover, multiple complicating factors, such as BMI, breast firmness and shape, features of the nipple and surrounding areola, and men’s marital status, preclude any simple explanation.

Check also Waist to hip ratio and breast size modulate the processing of female body silhouettes: An EEG study. Farid Pazhoohi, Joana Arantes, Alan Kingstone, Diego Pinal. Evolution and Human Behavior, January 7 2020. https://www.bipartisanalliance.com/2020/01/waist-to-hip-ratio-and-breast-size.html


Psychological impact of positive & negative information in memory: Positive content is often favored & the affect prompted is more temporally persistent than the affect prompted by memories of negative events

In Human Memory, Good Can Be Stronger Than Bad. Constantine Sedikides, John J. Skowronski. Current Directions in Psychological Science, January 14, 2020. https://doi.org/10.1177/0963721419896363

Abstract: Some researchers assert that the psychological impact of negative information is more powerful than that of positive information. This assertion is qualified in the domain of human memory, in which (a) positive content is often favored (in the strength of memories for real stimuli or events and in false-memory generation) over negative content and (b) the affect prompted by memories of positive events is more temporally persistent than the affect prompted by memories of negative events. We suggest that both of these phenomena reflect the actions of self-motives (i.e., self-protection and self-enhancement), which instigate self-regulatory activity and self-relevant processes.

Keywords: memory, self, self-motives, self-enhancement, self-protection


Check also The evaluative information ecology: On the frequency and diversity of “good” and “bad”. Christian Unkelbach, Alex Koch & Hans Alves. European Review of Social Psychology, Volume 30, 2019 - Issue 1, Pages 216-270. Nov 24 2019. https://www.bipartisanalliance.com/2019/11/positive-good-information-is-more.html

Tuesday, January 14, 2020

Contrary to findings from previous correlational studies, we do not find any impact of social media usage on well-being and academic success

Collis, Avinash, and Felix Eggers. 2020. “Effects of Restricting Social Media Usage.” SocArXiv. January 14. doi:10.31235/osf.io/udgxt

Abstract: Recent research has shown that social media services create large consumer surplus. Despite their positive impact on economic welfare, concerns are raised about the negative association between social media usage and performance or well-being. However, causal empirical evidence is still scarce. To address this research gap, we conduct a randomized controlled trial among students in which we track participants’ digital activities over the course of three quarters of an academic year. In the experiment, we randomly allocate half of the sample to a treatment condition in which social media usage is restricted to a maximum of 10 minutes per day. We find that participants in the treatment group substitute social media for instant messaging and do not decrease their total time spent on digital devices. Contrary to findings from previous correlational studies, we do not find any impact of social media usage on well-being and academic success. Our results also suggest that antitrust authorities should consider instant messaging and social media services as direct competitors before approving acquisitions.

People valorize the unproductive efforts of others in part because they believe such efforts reflect one’s inner virtues

Celniker, Jared, Andrew Gregory, Hyunjin Koo, Paul K. Piff, Peter Ditto, and Azim Shariff. 2020. “The Moralization of Unproductive Effort.” PsyArXiv. January 14. doi:10.31234/osf.io/nh9ax

Abstract: People believe that effort is valuable, but what kind of value does it confer? We find that displays of effort signal moral character. Importantly, we focus on displays of unproductive or unnecessary effort to highlight the heuristic nature of these intuitions—even “useless” effort is deemed virtuous. We conducted five studies to demonstrate the nature of these effects. In the domains of paid employment and charitable giving, the exertion of effort is deemed morally admirable (Studies 1-3) and is monetarily rewarded (Studies 1, 3, and 4), even when that effort results in no additional product. We test and find convergent patterns in a cross-cultural replication (Study 1b) and using a “big data” analysis of naturalistic donation behaviors (Study 4). We consider cultural and evolutionary accounts of effort moralization and discuss the implications of these effects for social welfare policy, automation, and the future of work.

General Discussion

Five studies, using multiple methodologies and cross-cultural samples, found that people ascribe greater moral value to greater exertions of effort, even when that effort is unproductive. Displays of effort serve as signals of one’s moral character, and these judgments inform decisions about how to allocate scarce monetary resources. People valorize the efforts of others in part because they believe such efforts reflect one’s inner virtues.
Our investigation refines previous research on effort evaluations and advances it in important ways. First, by explicitly controlling for productivity in our studies, we extend prior research (Amos, Zhang & Read, 2019) by showing that effort is valued even when it produces no value. Second, we provide the first discriminative evidence that effort cues affect moral evaluations specifically rather than positive character ascriptions more generally. Across four preregistered experiments, manipulating effort produced consistent differences in assessments of moral traits but not assessments of warmth or competence. These findings support theorizing that places moral character judgments at the center of impression formation (Goodwin, 2015; Uhlmann, Pizzaro & Diermeier, 2015). Finally, we broaden research on the martyrdom effect (Olivola & Shafir, 2013)—which has focused on manipulating one’s own commitment to a cause—by conceptually replicating it in paradigms focused on interpersonal moral judgments and real-world donation behaviors.

Unpacking Explanations of Effort Moralization
In addressing methodological limitations of prior work, limitations of a cultural explanation for effort moralization were also revealed. If PWE beliefs moralize effort, then inefficient effort should be denigrated because it is wasteful. Yet across our studies, participants consistently viewed unproductive effort as morally virtuous. Furthermore, individual differences
in participants’ work ethic beliefs did not moderate these effects, implying a limited role of PWE in explaining effort moralization. Finally, replicating one of our experiments in South Korea (including the failure of PWE beliefs to moderate valuations of effort) intimates that people moralize effort outside the West as well.
There is, in fact, some evidence that individuals in modern hunter-gatherer societies also moralize hard work (Smith & Apicella, 2019), suggesting that effort moralization may rest on more fundamental, and potentially evolutionary, origins. In the collaborative, group-living environments in which our species evolved, focusing on displays of costly signaling, like displays of effort, may have been an efficient and adaptive way to assess the cooperative intent of others (Gintis, Smith & Bowles, 2001). Partner choice markets can explain the use of competitive altruism as a signal of one’s value as a cooperation partner (Barclay, 2013), and effort may serve a similar function. This signaling account may provide a more parsimonious framework for conceptualizing effort moralization as a basic social heuristic. Rather than directly causing people to moralize effort, PWE beliefs may be scaffolded upon and exaggerate shared intuitions about the moral value of effort.
Cross-cultural replication of the current findings would seem a crucial next step in disentangling universal and culture-specific accounts of effort moralization. While South Korea is rooted in a distinct cultural tradition that has traditionally eschewed the overtly individualistic institutions and values of the U.S. (Hofstede, 1983), it is still a highly industrialized nation with some of the longest working hours in the OECD (OECD, 2019). Replications in societies that differ from the U.S. and South Korea on other dimensions would provide a more rigorous test of the role of culture in effort valuations.