Genetic Evidence
Genetic Evidence
Genetic Evidence
Lisa M. Diamond
To cite this article: Lisa M. Diamond (2021) The New Genetic Evidence on Same-Gender
Sexuality: Implications for Sexual Fluidity and Multiple Forms of Sexual Diversity, The Journal of
Sex Research, 58:7, 818-837, DOI: 10.1080/00224499.2021.1879721
The New Genetic Evidence on Same-Gender Sexuality: Implications for Sexual Fluidity
and Multiple Forms of Sexual Diversity
Lisa M. Diamond
Department of Psychology, University of Utah
ABSTRACT
In September of 2019, the largest-ever (N = 477,522) genome-wide-association study of same-gender
sexuality was published in Science. The primary finding was that multiple genes are significantly asso
ciated with ever engaging in same-gender sexual behavior, accounting for between 8–25% of variance in
this outcome. Yet an additional finding of this study, which received less attention, has more potential to
transform our current understanding of same-gender sexuality: Specifically, the genes associated with
ever engaging in same-gender sexual behavior differed from the genes associated with one’s relative
proportion of same-gender to other-gender behavior. I review recent research on sexual orientation and
sexual fluidity to illustrate how these findings speak to longstanding questions regarding distinctions
among subtypes of same-gender sexuality (such as mostly-heterosexuality, bisexuality, and exclusive
same-gender experience). I conclude by outlining directions for future research on the multiple causes
and correlates of same-gender expression.
In September, 2019, the largest-ever genome-wide-association than 6% of UK Biobank members and less than 2% of 23andMe
study (GWAS) of same-gender sexuality was published in members agreed to participate in the study (Saey, 2019), poten
Science (Ganna et al., 2019), reporting an analysis of 477,522 tially because of its focus on sexual orientation, which might
genomes (408,995 from the United Kingdom Biobank and introduce bias into the study. The sample was criticized for
68,527 from the United States genetic testing company, excluding individuals of non-European ancestry, although this
23andMe). The primary findings generated considerable scien exclusion was necessitated by the study’s methodology:
tific and media attention: Five autosomal loci were significantly Because gene frequencies and genotypic effects can vary across
associated with same-gender sexual behavior (defined as ever different populations, combining different populations in
engaging in same-gender sexual contact), and withstood repli a single GWAS study risks distorting the effects of interest
cation in independent samples. These five loci were simply (see Gebreyesus et al., 2019). In cases of population heteroge
ones that met their strict tests for individual statistical signifi neity, it is preferable to conduct separate GWAS studies within
cance, and overall their analyses indicated that thousands of different populations. Hence, the exclusion of non-European
other genetic variants, each with tiny effects, significantly con participants was methodologically appropriate, but obviously
tributed to the likelihood of ever engaging in same-gender limits the generalizability of the findings. Similarly, the exclu
sexual behavior. They estimated that 8–25% of the variance sion of transgender individuals (because their numbers were
in this outcome was genetically influenced, but found that the too small to yield meaningful group comparisons) may have
magnitude of measurable genetic influences was too small to been methodologically appropriate, but additionally constrains
yield reliable predictions of any individual’s same-gender sex the generalizability of the findings. Other limitations include
ual behavior based on their genome. Hence, their findings are the fact that most study participants were over the age of 40,
more useful for understanding the overarching construct of which may have affected the findings because such individuals
sexual orientation than for understanding the expression of were reared in social environments less accepting of same-
sexual orientation at an individual level. gender sexuality (and less conducive to its behavioral expres
As with most genetic research, media coverage oversimpli sion) than contemporary young adults (Lambert, 2019). This
fied the findings. Some headlines emphasized the failure of the may partially explain the fact that only 4% of the study parti
study to find a single “gay gene” (Chen, 2019; Davis, 2019; cipants reported ever having engaged in same-gender sexual
Ducharme, 2019; Harris, 2019; Khan, 2019; Saey, 2019; Scutti, behavior. This number is lower than some – although not all –
2019), whereas others trumpeted the study’s success in verify of the prevalence estimates of same-gender sexual behavior
ing genetic influences on same-gender sexuality (Abbot, 2019; that have emerged from population-based studies (Chandra
Belluck, 2019; Kaiser, 2019; Klelland, 2019; Tanner, 2019). et al., 2011; Copen et al., 2016; Kuyper & Vanwesenbeeck,
Methodological critiques also received well-justified attention: 2009; Twenge et al., 2015). Hence, the key phenotype of inter
Self-selection may have influenced the results, given that less est in the Ganna study et al. – same-gender sexual behavior –
CONTACT Lisa M. Diamond [email protected] Department of Psychology, University of Utah, 380 South 1530 East, Room 502, Salt Lake City, Utah,
84112-0251.
© 2021 The Society for the Scientific Study of Sexuality
THE JOURNAL OF SEX RESEARCH 819
may be underreported or underrepresented among the study’s sexual behavior in different populations (Frye, 1990; Randall &
participants. Collectively, these issues suggest that we interpret Byers, 2003; Sanders & Reinisch, 1999).
the findings with caution. Although it represents an important Ganna et al. did have a measure that more closely approx
step forward in our understanding of genetic contributions to imates the conventional construct of sexual orientation: One’s
same-gender sexuality (in the same way that early twin studies ratio of same-gender to other-gender sexual contact. The 4% of
represented important initial steps), it cannot provide defini participants who reported ever having same-gender sexual
tive answers to these questions. contact also reported the proportion of their sexual partners
The Ganna et al. study also received critiques for operatio that were of the same gender. Of course, “sexual partner” has
nalizing “same-gender sexual behavior” as ever engaging in its own definitional problems: Does a one-time experience with
same-gender sexual behavior. According to this definition, someone make that person a “sexual partner?” How much
someone with a single same-gender sexual experience occupies sexual contact needs to occur? Different participants might
the same category as someone with a lifetime of exclusively have adopted different interpretations of “sexual partner,”
same-gender sexual relationships. Such categorization conflicts introducing inevitable measurement error into the “propor
with scientific and lay conceptions of sexual orientation as an tionality” variable. Setting aside this complication for the
enduring and overarching sexual predisposition for one or moment, Ganna et al.’s analyses using the proportionality
more genders (Hamer, 2019; Reilly, 2019). Yet the unavoidable variable yielded a startling finding: The genetic effects differ
reality is that individuals pursue same-gender sexual contact entiating between individuals who “ever” versus “never”
for a variety of reasons. Some individuals report a consistent engaged in same-gender sexual behavior were not the same
and longstanding desire for same-gender sexual partners, genetic effects differentiating between individuals with varying
whereas others pursue same-gender behavior only in highly proportions of same-gender sexual behavior (i.e., mostly het
specific circumstances. Historically, this distinction has been erosexual vs. bisexual vs. exclusively same-gender).
described in terms of “constitutional” versus “facultative” The authors interpreted this pattern to suggest that there is
same-gender sexuality (Bell et al., 1981). Constitutional same- no genetic underpinning for the longstanding “single conti
gender sexuality has historically been attributed to a stable and nuum” model of sexual orientation, exemplified by the Kinsey
enduring same-gender orientation, whereas facultative same- scale, which posits exclusive heterosexuality as the opposite of
gender sexuality has been viewed as a situational manifestation exclusive same-gender sexuality. Ganna et al.’s conclusion on
of “play, exploration, lack of opposite-gender partners, hazing, this issue received little discussion in the media coverage of the
initiation rituals, intoxication, sexual frustration, prostitution, study, and only one mention among the peer-reviewed scien
boredom, opportunism, curiosity, and mistakes” (Muscarella, tific articles citing it (Jabbour et al., 2020). Yet the lack of
1999, p. 9). Alternatively, one might interpret the distinction genetic correlation between “ever/never” engaging in same-
between constitutional and facultative same-gender sexuality gender sexual behavior and one’s proportion of same-gender
in terms of the more familiar distinction between traits and sexual behavior is arguably the study’s most interesting result,
states. Some individuals are thought to possess the enduring with implications that go beyond the “single continuum”
“trait” of sexual orientation, whereas others are thought to model. Specifically, this finding bears on longstanding ques
occupy a temporary “state” of same-gender behavior. Yet tions about the difference between sexual orientation (i.e., an
there is no reliable basis on which to distinguish between overarching sexual predisposition for one or both genders) and
“state” and “trait” expressions of same-gender sexuality. For sexual fluidity (i.e., a capacity for situational variability in
example, what specific types and amounts of same-gender sexual responsiveness). This review elaborates potential impli
sexual behavior qualify someone as “oriented” versus “experi cations of Ganna et al.’s findings for these questions and out
menting?” Given these limitations, Ganna et al.’s “ever/never” lines directions for future research.
measure of same-gender sexual behavior (which they used to I begin by reviewing recent research on sexual fluidity and
maximize the study’s statistical power) cannot be interpreted as sexual orientation, highlighting unanswered theoretical and
an index of “sexual orientation,” and they straightforwardly empirical questions. Next, I describe the methods and results
acknowledged this fact. Although their study tests for genetic of the Ganna et al. study, focusing on the findings relevant to
influences on the phenotypic expression of same-gender sexual the single continuum model. I then offer my own speculations
behavior, it cannot differentiate between “trait” versus “state” regarding potential interpretations of these findings, and their
expressions of this behavior. implications for conceptualizing different subtypes of same-
Even if we restrict our interpretation of the study’s findings gender sexuality (such as bisexuality versus “fluid/flexible”
to “same-gender sexual behavior” instead of “sexual orienta heterosexuality). These speculations are offered in the spirit
tion,” definitional issues remain. For example, terms such as of hypothesis generation, and do not represent concrete theo
“same-gender sexual behavior” and “other-gender sexual beha retical proposals. There are many alternative interpretations of
vior” have ambiguous interpretations: One individual might Ganna et al.’s findings (including the possibility that their
“count” manual-genital stimulation as sexual contact, whereas results are artifactual), and we presently lack sufficient data to
another might only “count” oral-genital or genital-genital con yield definitive conclusions. Yet we do have sufficient data to
tact. Furthermore, these decisions may vary according to the rethink our own longstanding assumptions, and to consider
gender of each partner, the ages of the participants, and the most interesting and important possibilities for future
whether the contact was wanted or unwanted (reviewed in investigation. This is a chief goal of the present review.
Diamond & Savin-Williams, 2009). As a result, different stu Before proceeding, it bears noting that research on sexual
dies yield different estimates of same-gender and other-gender orientation has historically used restrictive definitions of sex
820 L. M. DIAMOND
and gender (reviewed in van Anders, 2015) that center and exclusive other-gender sexuality to exclusive same-gender sexu
privilege the experiences of cisgender individuals (i.e., those ality. This is the “single continuum” model discussed by Ganna
whose gender identities and/or expressions match their birth- et al., which Kinsey operationalized as a 0 to 6 scale (0 represent
assigned sex/gender). Accordingly, the term “same-gender” has ing exclusive other-gender behavior, 3 representing equal
been used to denote cisfemale-cisfemale or cismale-cismale degrees of same-gender and other-gender behavior, and 6 repre
pairings, while the term “other-gender” has been used to senting exclusive same-gender sexual behavior). Although
denote cisfemale-cismale pairings. Yet the terms “same- Kinsey originally developed his scale to represent sexual beha
gender” and “other-gender” fail to adequately describe pairings vior, it has been adapted over the years to assess sexual attraction,
involving transgender individuals (whose gender identities which is generally considered a more direct index of sexual
and/or presentations do not correspond to their birth- orientation (Mustanski, Van Wagenen et al., 2014).
assigned sex/gender) and nonbinary individuals (who may Notably, a core assumption of conventional models of sexual
experience and express their gender as both male and female, orientation (shared even by the Supreme Court in their 2015
neither male nor female, or fluid). Investigations of sexual decision legalizing same-gender marriage, “Obergefell
orientation have recently begun to wrestle with the full com v. Hodges,” 2015, p. 8) is that such orientations are fixed.
plexity, diversity, and fluidity of gender (Hyde et al., 2019; Hence, whether one is a “Kinsey 2” or a “Kinsey 6,” this status
Schudson et al., 2017; van Anders, 2015), but such nuanced is considered permanent. This view has been challenged over the
approaches remain underrepresented in extant research. This past several decades by scholars documenting that some indivi
underrepresentation is reflected in the present review. Hence, duals report unexpected changes in their sexual attractions, often
for the purposes of this discussion, the terms “gender,” “man,” prompted by situational or interpersonal factors. The capacity
“woman,” “same-gender,” and “other-gender” should be pre for such change has been denoted sexual fluidity (reviewed in
sumed to refer to cisgender individuals, on whom the vast Diamond, 2008, 2013; Diamond et al., 2020). Sexual fluidity has
majority of research on sexual orientation has been conducted. been postulated as an explanation for the fact that some indivi
Greater representation of the unique experiences of transgen duals who engage in same-gender sexual behavior do not per
der and nonbinary individuals remains a critical priority for ceive themselves as fundamentally oriented to do so, but rather
future research on sexual orientation. “open” to it under the right circumstances (Cassingham &
In addition, I want to guard against the impression that O’Neil, 1993; Diamond, 2008; Savin-Williams, 2017; Ward,
behavioral genetics holds the key to understanding different 2015). Some of these individuals describe themselves as “fluid,”
“types” of same-gender sexuality. Although I discuss Ganna “heteroflexible,” or “mostly heterosexual” (Kort, 2014; Savage,
et al.’s findings and their implications in detail, this should not 2002; Savin-Williams, 2017; Thompson & Morgan, 2008; Zane,
be taken to suggest that behavioral genetics provides inherently 2016) and the degree to which these individuals differ from self-
superior models of human variation than other scientific identified lesbian/gay/bisexual individuals remains debated
approaches (including biological models that focus on nonge (Savin-Williams et al., 2012; Savin-Williams & Vrangalova,
netic factors, as reviewed in Bailey et al., 2016; Wang et al., 2013; Vrangalova & Savin-Williams, 2010).
2019). Although contemporary Western culture tends to pri A key challenge for research on sexual fluidity and “mostly
vilege genetic “origin stories” (exemplified by the success of heterosexuality” concerns definitional ambiguity. The empiri
companies such as Ancestry and 23andMe), genetic influences cal evidence for sexual fluidity comes exclusively from self-
on complex human traits (especially traits that are subjected to report data, which introduces reporting biases. Just as indivi
culturally-specific interpretations) are obviously not determi duals may have different interpretations of “sexual behavior,”
nistic. Our genomes cannot tell us what social groups we they might also adopt different criteria for reporting whether
“truly” belong to, and using genetics for such aims runs the their sexual attractions have changed. Researchers, too, have
risk of reifying and entrenching social hierarchies (see different interpretations of “sexual fluidity,” and have investi
TallBear, 2007, 2013). Hence, although Ganna et al. provided gated a range of diverse phenomena under this general
important new data on sexual orientation, these data must be umbrella. For example, extensive evidence shows that some
considered in dialogue with what we already know and what we individuals show experimentally-elicited patterns of sexual
seek to learn. arousal that diverge from their self-described sexual orienta
tion, across diverse assessment methods including genital
photoplethysmography, functional magnetic brain imaging,
Sexual Fluidity and Sexual Orientation: Current Thinking
pupil dilation, and eye tracking (Chivers, 2017; Chivers &
and Unanswered Questions
Bailey, 2005; Chivers et al., 2004; Peterson et al., 2010; Rieger
Sexual orientation is typically conceptualized as “an enduring et al., 2012; Savin-Williams et al., 2017). This phenomenon has
pattern of or disposition to experience sexual or romantic desires been described as “non-specificity” (Chivers & Bailey, 2005),
for, and relationships with, people of one’s same sex, the other but obviously overlaps with the construct of sexual fluidity:
sex, or both sexes” (Institute of Medicine, 2011, p. 27). Although How exactly do these constructs differ? Evidence also shows
this conceptualization suggests three distinct categories (gay/ that some individuals engage in periodic sexual behavior that
lesbian, heterosexual, and bisexual, which also correspond to diverges from their self-described sexual orientation or identity
commonly used sexual identity labels), such categories are heur (Cramer et al., 2015; Matthews et al., 2014; Savin-Williams,
istics rather than natural types. Kinsey famously argued that 2017; Ward, 2015). Does this provide evidence for sexual
“The world is not to be divided into sheep and goats” (Kinsey fluidity, or is it simply attributable to bisexuality? Finally,
et al., 1948, p. 639), and postulated a continuum ranging from there is the matter of change in attractions over time.
THE JOURNAL OF SEX RESEARCH 821
Longitudinal studies have found that some individuals report than do exclusively-attracted individuals (Diamond, 2008,
changes in their patterns of sexual attraction over the course of 2016), less day-to-day stability in their sexual attractions
years (Charlton et al., 2016; Diamond, 2008; Ott et al., 2011, (Diamond et al., 2017), and more genital arousal to their less-
2012; Savin-Williams et al., 2012; Savin-Williams & Ream, preferred gender in the laboratory (Chivers et al., 2015).
2007; Spittlehouse et al., 2020) as well as days (Diamond The possibility that bisexuality and fluidity are fundamen
et al., 2017; Diamond & Wallen, 2011; Farr et al., 2014). Do tally related may help to address questions regarding gender
these changes stem from sexual fluidity? differences. The notion that women are more sexually fluid
The problem with ascribing all of these phenomena to than men has a long history (Baumeister, 2000; Blumstein &
“fluidity” is that “fluidity” does not seem to represent a single Schwartz, 1977; Cass, 1990; Diamond, 2008; Weinberg et al.,
overarching trait. Rather, different individuals appear to show 1994), but contemporary empirical data on this point is mixed,
different forms of sexual variability for potentially different further underscoring that “fluidity” as a single trait may not
reasons. For example, several studies have found that indivi exist. Rather, there appear to be varying degrees of gender
duals who appear more “fluid” on one dimension (such as difference for different types of sexual variability, and these
showing more day-to-day instability in their sexual attractions) gender differences may have different causes. If some forms of
are not necessarily more “fluid” on other dimensions, such as sexual variability are facilitated by bisexuality, then this pro
showing stronger arousal to their non-preferred gender in the vides a potential explanation for gender differences, given that
laboratory, having more sexual contact with non-preferred bisexual patterns of attraction are more common among
partners, or reporting more retrospective change in their women than among men (as shown by Chandra et al., 2011;
attractions (Diamond et al., 2020, 2017). Hence, although Copen et al., 2016; Gates, 2011; Mosher et al., 2005; Savin-
fluidity has been used as a “catch all” term for sexual variability, Williams et al., 2012). One way to disentangle the associations
scientific research must investigate different types of sexual among gender, bisexuality, and fluidity is to examine gender
variability (for example, longitudinal change in sexual attrac differences in different forms of sexual variability separately
tions versus unexpected patterns of laboratory-based arousal) among bisexual and non-bisexual individuals. With regard to
as distinct phenomena with potentially distinct causes, conse longitudinal change in attractions, such change appears to be
quences, and rates of prevalence in different populations. independently associated with both bisexuality and gender
For example, cultural factors often contribute to discrepan (Dickson et al., 2003; Mock & Eibach, 2012; Ott et al., 2011;
cies between individuals’ self-described attractions and beha Savin-Williams et al., 2012; Savin-Williams & Ream, 2007).
viors. Mustanski and colleagues (Mustanski, Birkett et al., Specifically, individuals with bisexual attractions show more
2014) pooled two national probability samples, yielding longitudinal change than those with exclusive attractions, yet
a combined sample size of over 50,000, to examine ethnic among those with exclusive attractions, women show more
differences in the degree of concordance between self- change than do men (reviewed in Diamond et al., 2017).
reported behavior and identity. Black and Hispanic youths Similarly, studies of day-to-day stability in sexual attractions
who described themselves as gay/lesbian showed more discor have found that both gender and bisexuality independently
dant patterns of sexual behavior than did Whites, potentially contribute to day-to-day instability in attractions (Diamond
due to variation across ethnic groups in the meanings and et al., 2017).
consequences of different expressions of same-gender sexual Gender and bisexuality may foster sexual variability through
behavior and identity. Some cultures may discount the rele different mechanisms. For example, gender differences in some
vance of same-gender attractions or behavior, and others may forms of sexual variability may reflect the different sociocul
provide no route for developing a lesbian, gay, or bisexual tural norms faced by women versus men, and the fact that
identity while maintaining one’s ethnic identity (Chan, 1995; women’s sexuality has been subjected to historically greater
Consolacion et al., 2004; Jamil et al., 2009; McLean, 2003; Parks social control than men’s sexuality (Baumeister & Twenge,
et al., 2004). Assessments of different forms of sexual variability 2002; Fine, 1988; Tolman, 2002, 2006). Notably, this social
must take these potential influences into account. control is not always enacted by men: As found by
Baumeister and Twenge (2002), women across cultures fre
quently restrict one another’s sexuality, enforcing cultural
Bisexuality as Fluidity?
norms and regulating the social currency associated with mar
Some forms of sexual fluidity might simply be outgrowths of riage and reproduction. Given the social and economic costs
bisexual orientations (Diamond et al., 2020, 2017). After all, that women have historically incurred by rejecting traditional
bisexuality (conceptualized as an enduring pattern of sexual roles as wives and mothers (Faderman, 1981; Kitzinger, 1987;
attractions to more than one gender) and sexual fluidity can Rich, 1980), women desiring same-gender sexual behavior may
both produce the same observable manifestation: a history of find it easier and safer to pursue such behavior alongside
sexual attraction/behavior with more than one gender. heterosexual behavior, rather than rejecting heterosexuality
Theoretically, bisexual orientations may foster sexual variabil altogether. Such dynamics could conceivably inflate rates of
ity because the simple existence of attractions to more than one bisexuality (relative to exclusive same-gender sexuality) among
gender grants situational and interpersonal factors more lati women. An additional influence might be the fact that
tude to shift one’s experiences across different times and con women’s expressions of bisexuality have been (and remain
texts (Ross et al., 2012; Weinberg et al., 1994). This could currently) viewed more positively than men’s expressions of
account for the fact that bisexually-attracted individuals report bisexuality (Dodge et al., 2016; Dyar & Feinstein, 2018; Yost &
more longitudinal change in their attractions and behaviors Thomas, 2012), which may further encourage greater
822 L. M. DIAMOND
expression of bisexuality among women than men. Finally, the reporting any same-gender attractions. As scientists, how can
greater social condemnation of gender nonconformity and we presume to understand anything about same-gender sexu
same-gender sexuality among men than among women plays ality if we do not understand its most common manifestation?
an additional role. In the contemporary West and around the The second reason to investigate mostly-heterosexual indi
world, men and women have more negative attitudes about viduals is that they raise core questions about the full spectrum
men’s same-gender sexuality than about women’s same-gender of exogenous and endogenous influences on same-gender
sexuality (Bettinsoli et al., 2019), which may reflect hegemonic attraction and behavior, and the degree to which sporadic
notions of masculinity that require strict adherence to the (versus consistent) same-gender desires are fundamentally
masculine gender role, especially in the domain of sexual con unique types of experiences, with distinct phenomenologies,
tact (Goodnight et al., 2014; Herek, 1986; Van Beusekom et al., etiologies, and developmental outcomes (i.e., whether same-
2019). As a result of these rigid norms, men with bisexual gender sexual behavior as a state differs fundamentally from
patterns of attraction may experience more social pressure to same-gender behavior as a trait). Investigating this question
maintain strictly heterosexual identities and relationships than has implications for our basic understanding of human same-
do bisexual women. gender sexuality, its development, and its evolved bases, which
remain longstanding topics of theoretical and empirical
inquiry (N. W. Bailey & Zuk, 2009; Barron & Hare, 2020; De
The Most Controversial Group: Mostly-Heterosexuals
Block & Adriaens, 2004; Burri et al., 2015; Cornil & Bakker,
Overlap between the constructs of sexual fluidity and bisexu 2019; Hyde & Durik, 2000; Jeffery, 2015; Kirkpatrick, 2000;
ality becomes particularly evident when considering “mostly Luoto et al., 2019; Luoto & Rantala, 2017; Radtke, 2013;
heterosexual” individuals. As with bisexuality, this group is Wunsch, 2010).
disproportionately female, with population prevalence esti A final reason to pursue these questions is their potential
mates ranging between 7.6% and 9.5% for women and between public health relevance. Extensive research has documented
3.6% and 4.1% for men (Savin-Williams & Vrangalova, 2013). that individuals reporting any same-gender attraction or beha
“Mostly heterosexuality” has grown in visibility in recent years: vior, regardless of self-identification, show dramatic disparities
In 2014 it was added as an identity category by the popular in physical and mental health compared to exclusively hetero
dating website OkCupid, and a growing number of young sexual individuals (Blosnich et al., 2014; Dyar et al., 2019;
celebrities openly claim “mostly heterosexual,” “fluid,” or “het Fredriksen-Goldsen et al., 2013; Hatzenbuehler et al., 2013;
eroflexible” identities (Renfro & Ahlgrim, 2020). A diverse Marshal et al., 2011; Mustanski, Van Wagenen et al., 2014;
range of experiences are subsumed under this umbrella: Potter & Patterson, 2019; La Roi et al., 2016). These disparities
Some individuals report feeling they are “straight-leaning” have been linked to the chronic stress, stigmatization, and
bisexuals; others report having fallen in love with only one victimization commonly experienced by sexually-diverse and
same-gender partner; others pursue sporadic sexual gratifica gender-diverse individuals (Bostwick et al., 2014; Burton et al.,
tion from same-gender partners when other-gender partners 2013; Friedman et al., 2014; Marshal et al., 2011; Meyer, 2003;
are unavailable (Diamond, 2008; Feinstein et al., 2020; Garvey Persson et al., 2015; Rosario et al., 2014). Notably, research has
et al., 2018; Kort, 2014; Redcay et al., 2018; Savin-Williams, reliably shown that individuals with both same-gender and
2017; Savin-Williams & Vrangalova, 2013; Thompson & other-gender sexual behavior have the poorest mental and
Morgan, 2008; Vrangalova & Savin-Williams, 2012, 2014; physical health of all sexually diverse individuals (Blosnich
Zane, 2016). Such diversity makes it difficult to interpret the et al., 2014; Bostwick et al., 2010; Dyar et al., 2019; Friedman
“mostly heterosexual” subgroup: Are they heterosexually- et al., 2014; Gorman et al., 2015; Marshal et al., 2011; Rosario
leaning bisexuals, “fluid” heterosexuals, or a mixture of both? et al., 2016). Scholars have ascribed this pattern to the specific
To answer such questions, researchers have investigated stigmatization and marginalization experienced by bisexuals
exactly how mostly-heterosexual individuals differ from self- (Brewster et al., 2013; Feinstein et al., 2019; Friedman et al.,
identified bisexual, queer, lesbian, gay, and heterosexual indi 2014; Katz-Wise, Mereish et al., 2017). Negative attitudes
viduals (Calzo et al., 2017; Jabbour et al., 2020; Kuyper & Bos, toward bisexuality have a long history in both mainstream
2016; Morandini et al., 2019; Savin-Williams, 2018; Talley society and the larger lesbian/gay community (Mulick &
et al., 2016). The results of these studies suggest that “mostly Wright, 2002; Weiss, 2003; Welzer-Lang & Tomolillo, 2008;
heterosexual” individuals resemble bisexually-attracted indivi Yost & Thomas, 2012), manifested in stereotypes of bisexuals
duals on some domains, but heterosexual individuals on other as confused, questioning, experimental, promiscuous, irre
domains. How much do such differences matter? Given that all sponsible, and unfaithful (Brewster & Moradi, 2010; Dyar &
sexual categories are human creations, parsing their bound Feinstein, 2018). As a result, bisexual individuals often experi
aries may seem trivial. Yet there are several reasons to investi ence marginalization from both heterosexual and lesbian/gay
gate whether mostly-heterosexuals represent a unique group. communities, restricting their access to social support and
First, representative studies reliably find that they comprise the validation (Friedman et al., 2019).
single largest subgroup of individuals with same-gender attrac The extent to which “mostly heterosexual” individuals share
tions or behavior (reviewed in Savin-Williams & Vrangalova, these experiences remains unclear. Some studies of health
2013). For example, the 2011–2013 data from the National disparities have specifically examined “mostly heterosexuals”
Survey of Family Growth (Copen et al., 2016) found that as a distinct group (Savin-Williams & Vrangalova, 2013), and
individuals reporting “mostly heterosexual” same-gender have found that they show rates of mental, physical, and sexual
attractions accounted for 72% of women and 57% of men health problems that are similar to self-identified bisexuals
THE JOURNAL OF SEX RESEARCH 823
(Corliss et al., 2009; Kuyper & Bos, 2016; Vrangalova & Savin- sexual behavior, represented as one’s proportion of same-
Williams, 2014; Zou et al., 2015). Heightened health disparities gender sexual partners (relative to total sexual partners). This,
have also been found among individuals reporting different of course, is not a direct measure of sexual orientation, given
types of sexual fluidity (Everett, 2015; Katz-Wise, Reisner et al., that sexual behavior is more constrained by social context and
2017; Katz-Wise, Williams et al., 2017). In order to address and opportunity than sexual attraction. Some individuals may pos
prevent these health disparities, we need a fuller understanding sess exclusively same-gender attractions, but may have bisexual
of their differential expression across the spectrum of sexual histories of sexual behavior; other individuals may possess
diversity, and the mechanisms responsible for these differ bisexual attractions, but engage in exclusively same-gender or
ences. A fuller understanding of potential genetic contribu exclusively other-gender sexual behavior. Gradations in sexual
tions to these differences may aid this goal, which adds to the attraction would obviously have provided a superior index of
relevance of Ganna et al.’s study. sexual orientation, but such data were not available for the full
sample (they were only available for the 23andMe subsample).
Before proceeding with their analysis of the “proportionality
Ganna et al. (2019): Basic Methods and Results
of sexual partners” measure, Ganna et al. examined how the
The GWAS by Ganna et al. (2019) is not the first GWAS proportionality measure correlated with other self-reported
focusing on sexual orientation (for reviews see Bailey et al., indices of sexual orientation that were available in the
2016; Dawood et al., 2009; Hill et al., 2013), but it is the largest 23andMe subsample, such as one’s ratio of same-gender to
and most statistically powerful. The authors analyzed the gen other-gender attractions and one’s identification as bisexual
omes of 477,522 individuals (408,995 from the United versus lesbian/gay. Each of these variables correlated around
Kingdom Biobank and 68,527 individuals from the United .98 with proportionality of same-gender sexual behavior.
States genetic testing company, 23andMe). A sample of such Hence, although proportionality of sexual behavior is not
size is necessitated by the low base rate of the key phenotype equivalent to the construct of sexual orientation, it provides
(ever engaging in same-gender sexual behavior), which was a reasonable “stand in” for the purposes of their analyses. The
reported by only 4% of their participants. As noted earlier, total number of individuals with same-gender sexual behavior
the low rate of self-reported same-gender sexual behavior is was too small to yield a GWAS estimate of genetic influence on
a limitation of the study. As with all studies using self-report “proportionality of same-gender sexual behavior,” but Ganna
assessments of same-gender sexuality, some participants may et al. calculated family-based heritability estimates for this
misrepresent their sexual history or decline participation in the variable, which ranged between .22 and .27.
study due to social stigmatization (and potentially the fear of
public identification). As summarized earlier, the primary find
“Ever/never” versus “Proportionality:” the Single
ing of the study was that between 8 and 25% of the variance in
Continuum Model
“ever versus never” engaging in same-gender sexual behavior
could be ascribed to genetic influence. The authors identified 5 This brings us to Ganna et al.’s findings regarding the “single
specific autosomal loci that were significantly associated with continuum” model. Among the 4% of individuals with same-
the “ever/never” phenotype: 2 among men only, 1 among gender sexual behavior, they calculated genetic correlations
women only, and 2 among both men and women. The genetic between the “ever/never” trait and the “proportionality” trait
correlation between men and women for “ever/never” (i.e., the (i.e., estimates of the proportion of variance that these two
extent to which the genes associated with “ever/never” for men phenotypes share due to genetic factors). If there were
overlap with the genes associated with “ever/never” for a single genetic continuum ranging from exclusive heterosexu
women) was .63, suggesting shared but not identical genetic ality to exclusive same-gender sexuality (with different degrees
influences. The authors also calculated traditional family-based of bisexuality in between), then the genes distinguishing
estimates of the heritability of “ever/never” using the subset of between the “ever” and “never” groups should also distinguish
dyads within the overall sample who had varying degrees of among individuals within the “ever” group (i.e., distinguishing
biological relatedness to one another (at the level of first cousin between those with lower versus higher proportions of same-
or closer): Family-based heritability was .32, which is quite gender sexual behavior). In other words, the genes associated
similar to the heritability estimates that have been calculated with being “not straight” should also be associated with one’s
from twin registry studies (reviewed in Bailey et al., 2016). degree of “not straightness.”
Ganna et al. noted that the difference between the GWAS Yet Ganna et al.’s results disconfirm this model. The genetic
heritability estimate (8–25%) and the family-based estimate correlation between the “ever/never” trait and the proportion
(32%) is similar to differences between GWAS and family- ality trait was nonsignificant (.03 for men and −.3 for women;
based estimates of the heritability of many other traits. Family- further, below I address the fact that the correlation for women
based estimates are uniformly higher, which may reflect the was actually negative, albeit nonsignificantly so). Ganna et al.
role of nonadditive genetic effects and phenotypic concluded that these two traits “capture aspects of sexuality
heterogeneity. that are distinct on the genetic level, which in turn suggests that
Although “ever/never” engaging in same-gender sexual there is no single continuum from opposite-sex to same-sex
behavior was the primary phenotype analyzed in the study, sexual behavior” (2019, p. 6). The word “single” is important
the authors conducted additional analyses of the 4% of parti here: Ganna et al.’s findings on the heritability of proportion
cipants within the “ever” category. For these analyses, the ality show that there is a meaningful genetic continuum differ
phenotype of interest was the specific degree of same-gender entiating individuals with low versus high ratios of same-
824 L. M. DIAMOND
gender to other-gender behavior. Yet clearly this is not the only This does not mean that there is no genetic underpinning to
genetic trait influencing the expression of same-gender sexu the traditional Kinsey scale; it simply suggests that the “0”
ality. The genes associated with “ever/never” make a separate rating – exclusive heterosexuality – might best be set aside
contribution. In essence, “Trait A” contributes to whether one and analyzed separately. The rest of the scale, from 1 (mostly
will ever pursue same-gender sexual behavior and “Trait B” heterosexual) to 6 (exclusive same-gender sexuality) might still
contributes to how much same-gender sexual behavior one will be conceptualized as a linear genetic continuum. Ganna et al.
pursue. This is a complex and counterintuitive conclusion, tested this possibility by dividing the “ever” group into four
which may explain why it received little scientific or media subgroups (approximating a Kinsey-type linear progression):
attention. Let us break it down, using an analogy familiar to (1) Those with less than a third of same-gender partners, (2)
students of genetic influence: flower color. those with between one third and two-thirds same-gender
Imagine a group of flowers that range from pure white partners, (3) those with over two-thirds (but not 100%) same-
to dark yellow. If “yellow versus white” is determined by gender partners, and (4) those with 100% same sex partners. If
a single set of genes, then these genes will distinguish the a single set of genes produces continuous variation in the
white flowers from all of the yellow flowers (whether those phenotype of proportionality, then groups that are more phe
flowers are light yellow or dark yellow). This is analogous notypically similar should be more genetically similar.
to the “ever/never” distinction, for which Ganna et al. This is exactly what Ganna et al. found: The genetic correla
found significant genetic influence. Now let us turn to tion between the “exclusively same-gender” group and the
subtypes of yellow flowers (light versus dark). One possibi “over two-thirds same-gender” group was .95; the correlation
lity is that the genes coding for “yellow versus white” also between the “exclusively same-gender” and the “between one-
determine the degree of a flower’s yellowness. Low genetic third and two-third same-gender” groups was .80. The correla
expression produces white flowers, medium genetic expres tion between the “exclusively same-gender” group and the “less
sion produces light yellow flowers, and high genetic expres than one third same-gender” group (the most phenotypically
sion produces dark yellow flowers. In this model, white different group) dropped precipitously, to a nonsignificant .13.
flowers and dark yellow flowers represent opposite expres In other words, the genetic correlation between the “100%
sions of the same underlying trait. same-gender” group and its phenotypically closest category
Ganna et al.’s findings disconfirmed this model. Let us (the “over two thirds same-gender” group) was 7 times greater
interpret Ganna et al.’s surprising results by thinking through than the genetic correlation between the “100% same-gender”
an alternative model: There is still a set of “yellowness” genes, group and the most phenotypically distinct category (the “less
but their only function is to distinguish between white flowers than one third” group). These differences are illustrated in
(i.e., the never group) and yellow flowers (the ever group). Figure 1, which uses Venn diagrams to show the genetic
Within the ever group, all of the flowers share a genetic pre correlations between the “exclusively same-gender” group and
disposition for yellowness, but this basic predisposition does the other three groups (Ganna et al. did not report correlations
not influence each flower’s degree of yellowness. Rather, the among all possible pairs of groups, so we do not know the
latter trait (let us call it “saturation”) is determined by genetic correlation between, for example, the “less than one
a separate set of genes. To understand the final appearance of third” group and the “one third to two thirds” group). One
flowers in the “yellow” group, we need to consider their yellow might interpret the pattern displayed in Figure 1 to suggest that
ness and their saturation. there is a meaningful genetic continuum ranging from
Figure 1. Venn diagrams representing the genetic correlations (reported in Ganna et al., 2019) between groups of individuals with different degrees of same-gender
sexual behavior. The left-most graph shows the genetic correlation between individuals with exclusive same-gender behavior and individuals with over two-thirds
same-gender behavior; the middle graph shows the correlation between individuals with exclusive same-gender behavior and those with one-third to two-thirds same-
gender behavior; the right-most graph shows the correlation between individuals with exclusive same-gender behavior and those with less than one-third same-gender
behavior.
THE JOURNAL OF SEX RESEARCH 825
moderate bisexuality (i.e., at least one-third same-gender sexual speculation, and I want to underscore the fact that these spec
behavior) to exclusive same-gender sexuality, but this conti ulations are offered to inspire and guide future research rather
nuum does not contain the mildest form of bisexuality (less than to support a single model. Although the phenotypic
than one third same-gender sexual behavior), which might expression of this trait is “ever engaging in same-gender sexual
represent “mostly heterosexuality.” This interpretation is con behavior,” this behavior might be an artifact of something else
sistent with the research reviewed above which has found that that has not been directly observed. Such a possibility is parti
“mostly heterosexual” individuals appear distinct from hetero cularly likely given that sexual behavior is a highly complex
sexual, lesbian, gay, and bisexual individuals, but also similar in human behavior involving a broad range of psychological
some ways (Vrangalova & Savin-Williams, 2012). Importantly, mechanisms. Might the “ever/never” trait represent
the fact that different genes contribute differently across differ a predisposition for sexual fluidity? After all, a predisposition
ent points along the Kinsey scale does not necessarily mean for sexual fluidity might account for the fact that some indivi
that there are fundamentally different processes at work along duals with low (or absent) same-gender interest end up enga
these different points. There may be different sets of genes ging in same-gender sexual behavior. Yet this framing does not
involved, but relatively similar neurological and/or psychologi seem supportable. Sexual fluidity (in its different manifesta
cal mechanisms, and the nature of these mechanisms remains tions) theoretically represents a capacity for variation, and
a critical unanswered question in this area. none of the data collected by Ganna et al. speak to this issue.
Also, pursuing sporadic and uncharacteristic sexual behavior is
only one form of sexual variability, and previous research
What Do “Ever/never” and “Proportionality” Represent?
shows that this form of variability does not correlate with
Let us now consider the meaning of the two distinct traits other forms of variability, such as the capacity to respond to
suggested by Ganna et al.’s findings (yellowness versus satura “non-preferred” sexual stimuli in the laboratory (Diamond
tion). “Proportionality” seems interpretable as a straightforward et al., 2020). Hence, although there may be individual differ
continuum of same-gender sexual interest, ranging from weak to ences in sexual fluidity “lurking” within Ganna et al.’s sample
strong. This trait does not presume that stronger same-gender (potentially within the “mostly heterosexual” group), we can
interests necessitate proportionally weaker other-gender inter not observe them.
ests (which is one of the long-critiqued shortcomings of the What other trait might underlie the distinction between the
traditional Kinsey scale). Rather, as suggested by others (Klein “ever” group and the “never” group? Unfortunately, we know
et al., 1985; Sell, 1996; Storms, 1980) and noted by Ganna et al., almost nothing about the diverse life experiences and desires of
same-gender and other-gender sexual interest might represent either group: All we know is that the “ever” group engaged in
orthogonal dimensions, such that one can be high on one same-gender sexual behavior at least once, and the “never”
dimension, high on both dimensions, or low on both dimen group did not. We also know that same-gender sexual behavior
sions (with the latter category potentially manifesting as is culturally stigmatized and relatively uncommon, whereas
asexuality). exclusive heterosexuality is both widespread and highly
It might seem contradictory to argue that “proportion of rewarded. Hence, we can say with certainty that all of the
same-gender sexual behavior” should be interpreted as individuals in the “ever” group (whether they had a single
a reasonable index of “same-gender interest” – what about other- same-gender experience or many such experiences) were will
gender interest? Wouldn’t one’s proportion of same-gender sex ing to engage in a new and unfamiliar behavior that exposed
ual behavior depend on one’s combination of same-gender and them to social and physical risks. The moment they stepped out
other-gender interests? Not necessarily: Society encourages and of the “never” category, they exposed themselves to potential
rewards other-gender behavior, but stigmatizes and punishes social stigmatization and victimization. In contrast, the “never”
same-gender sexual behavior. Hence, one’s proportion of same- group did not take this risk. We do not know whether they ever
gender sexual behavior should more reliably indicate one’s desired or had opportunities for same-gender sexual contact,
degree of same-gender sexual interest than one’s relative degree we only know that they never took this particular step.
of other-gender interest. As a result, we can reasonably presume Hence, we might characterize the “ever” group as (1) more
that individuals with more frequent same-gender sexual part open to unfamiliar and/or taboo experiences and/or (2) more
ners have stronger motivations to pursue such partners. The tolerant of risk. Both of these potential traits are theoretically
same cannot be said for other-gender behavior: Given its privi independent of same-gender interest, but could facilitate the
leged cultural status (along with the varying norms regulating behavioral expression of same-gender interest. In other
how often one is allowed to pursue other-gender sexual behavior words, perhaps one needs some degree of “openness” or
at different ages and in different social contexts), more frequent “risk tolerance” to pursue any same-gender sexual behavior
participation in other-gender relationships does not necessarily (i.e., to move from the “never” category to the “ever” cate
indicate stronger sexual interest in other-gender (versus same- gory), but once one arrives in the “ever” category, higher
gender) sexual partners – too many other factors are at play. levels of openness or risk tolerance do not make an individual
Hence, “proportionality of same-gender sexual behavior” pro more same-gender oriented. Rather, that form of variation
vides a reasonable index of gradations in same-gender sexual (from Kinsey 1 to Kinsey 6) is altogether separate. As with the
interest, independent of gradations in other-gender interest. flower analogy (yellowness + saturation), we would want to
Now let us turn to the “ever/never” trait: If this trait does not understand the combination of “same-gender interest” and
represent gradations in same-gender interest, then what might “risk tolerance” (or “openness”) to understand one’s final
it represent? This is where we tread cautiously into the realm of expression of same-gender sexuality. We already know (as
826 L. M. DIAMOND
shown in Figure 1) that individuals with the lowest propor If “ever/never” relates to a genetically influenced predispo
tions of same-gender sexual behavior (the mostly heterosex sition for “openness” or “risk tolerance,” we would expect
uals) are genetically distinct from individuals reporting higher positive genetic correlations with traits in the “risk behavior”
levels (moderate bisexuality and exclusive same-gender sexu group, with “number of sex partners” (which may reflect both
ality). Yet they are also genetically distinct from the “exclusive risk tolerance and an openness to new sexual opportunities)
heterosexuals.” Perhaps the interaction between the trait of and also with the personality trait of “openness to experience.”
same-gender sexual interest and additional traits such as risk This is what Ganna et al. observed, for both women and men.
tolerance or openness can help to explain this paradox. “Ever vs. never” engaging in same-gender sexuality was posi
Specifically, risk tolerance or openness should have more tively genetically correlated with number of total sex partners,
influence on same-gender sexual behavior among those with openness to experience, cannabis use, smoking, general risk
weaker same-gender interests than among those with stronger behavior, loneliness, major depression, and ADHD. These
same-gender interests. Among individuals with strong same- positive correlations indicate that the shared genetic variance
gender interests, the enduring strength of their motivation for between “ever/never” and these traits manifests as stronger trait
same-gender sexual contact might push them out of the expression among individuals who have ever engaged in same-
“never” category and into the “ever” category no matter gender sexual contact. Ganna et al. took care to point out that
how much they might fear unfamiliar or risky experiences. the potentially controversial correlations between “ever/never”
There are certainly many examples of this possibility, such as and mental health dimensions such as loneliness, depression,
prominent members of society who eventually risk every and ADHD should not be interpreted to suggest that same-
thing – including their jobs, families, and entire commu gender sexual behavior “leads” to mental health problems (or
nities – to pursue same-gender relationships. In essence, vice versa). They clarified that the shared genetic variance for
strong same-gender interests “win out” over fears of risk all of these traits could reflect social and environmental factors
and unfamiliarity. that expose sexually-diverse individuals to greater mental and
Yet for individuals with relatively weaker same-gender physical health risks, and this is also a potential explanation for
interests, this balance is different. An individual with milder the correlations with risk behaviors.
same-gender interests might only pursue same-gender sexual Yet what is most striking about Ganna et al.’s findings is the
behavior if such contact is rendered more familiar, safe, and difference between the pattern of genetic correlations found for
accessible (perhaps due to living in a more tolerant social “ever/never” versus “proportion of same-gender sexual part
environment) or if they are more comfortable with the asso ners.” Whereas “ever versus never” engaging in same-gender
ciated risks (perhaps due to their own high level of risk toler behavior was positively genetically correlated with risk beha
ance). Accordingly, individuals in the “mostly heterosexual” vior, substance use, and openness to experience, “proportion of
group (compared to other subgroups of nonheterosexual indi same-gender partners” was negatively correlated with all of
viduals) may contain a disproportionate number of individuals these traits. In other words, within the group of individuals
whose same-gender sexual behavior reflects high openness reporting any same-gender behavior, those with the strongest
and/or risk tolerance rather than an enduring same-gender genetic expression for risk-associated traits are those with the
predisposition. This may be why they are so genetically dis smallest proportions of same-gender sexual behavior (i.e.,
similar (as shown in Figure 1) from individuals with larger those with mostly-heterosexual or bisexual patterns of beha
proportions of same-gender sexual behavior. vior, as opposed to exclusive same-gender sexual behavior).
This overall pattern is presented in Figures 2 and 3. These
figures display the genetic correlations reported in Ganna
Links to Other Traits et al. Figure 2 displays the results for men and Figure 3 displays
the results for women. In both graphs, the black bars display
If “ever/never” and “proportionality” truly represent separate the magnitude of the genetic correlations between “ever/never”
traits, then they should show different patterns of genetic engaging in same-gender behavior and ADHD, depression,
correlation with other heritable traits. Additionally, if “ever/ loneliness, general risk behavior, smoking, cannabis, openness
never” is related to an openness to novelty and/or a tolerance to experience and number of sex partners. The fact that all of
for risk, then it should show high genetic correlations with the black bars extend to the right side of the graph indicates
traits associated with these characteristics. This is exactly what that all of the correlations are positive (i.e., greater genetic
Ganna et al.’s research found. They reported genetic correla expression of each trait among individuals in the “ever”
tions with 28 traits that previous GWAS studies found to be group versus the “never” group). The gray bars display the
genetically influenced. They selected these traits to represent magnitude of the genetic correlations between “proportion of
five domains: Risk behavior (alcohol use, overall risk behavior, same-gender partners” and the same traits. The fact that all of
smoking, cannabis use), mental health (subjective well-being, the gray bars extend to the left side of the graph indicates that
anorexia, anxiety, self-rated health, schizophrenia, autism, all of the correlations are negative (i.e., greater genetic expres
bipolar disorder, ADHD, depression), personality traits (neu sion of each trait among individuals with lower proportions of
roticism, loneliness, openness to experience), reproductive same-gender behavior.)
traits (age at first birth, age at menarche and menopause for To clarify the meaning of this pattern, it helps to walk
women, number of children, number of sex partners), and through a specific example: In this case, we will focus on
physical traits (height, waist-to-hip ratio, birth weight, and cannabis use. Keep in mind that the correlations computed
2D:4D digit ratio, an index of prenatal androgen exposure). with the “ever/never” variable contrast the “ever” group (the
THE JOURNAL OF SEX RESEARCH 827
Figure 2. Significant genetic correlations, in men, between various traits and (1) higher versus lower proportions of same-gender behavior, and (2) ever versus never
engaging in same-gender behavior, as reported in Ganna et al. (2019) . Values on the right side of the graph represent positive correlations and values on the left side of
the graph represent negative correlations.
Figure 3. Significant genetic correlations, in women, between various traits and (1) higher versus lower proportions of same-gender behavior, and (2) ever versus never
engaging in same-gender behavior, as reported in Ganna et al., (2019). Values on the right side of the graph represent positive correlations and values on the left side of
the graph represent negative correlations.
4% of participants reporting any same-gender sexual contact) partners,” suggesting that the genes associated with cannabis
with the “never” group (the 96% of participants with no such use overlap with those associated with smaller proportions of
contact). The analyses involving the proportionality variable same-gender partners (within the group of individuals report
focus only on the 4% of participants within the “ever” group, ing any same-gender partners). This pattern of divergent cor
contrasting individuals with higher versus lower proportions of relations for “ever/never” and “proportionality” presents
same-gender sexual behavior. As shown in the black bars of further evidence that they represent distinct traits with differ
Figures 2 and 3, cannabis use is positively genetically correlated ent implications for same-gender sexuality. The findings pre
with “ever/never” in both men and women, suggesting that the sented in Figures 2–3 suggest that individuals with different
genes associated with “ever” having same-gender sexuality combinations of these traits may show different patterns of
overlap with the genes associated with cannabis use. Notably, same-gender sexuality and different life experiences on other
both risk tolerance and openness to novelty might augment domains.
one’s likelihood of using cannabis, given that it is an illicit drug, Parsing such differences may be particularly relevant for
but one that is relatively widely available. Turning to the gray future research on health disparities: The pattern of correla
bars in Figures 2 and 3, we see that cannabis use is negatively tions shown in Figures 2–3 is particularly notable because it
genetically correlated with “proportion of same-gender suggests that the greatest expression of mental and behavioral
828 L. M. DIAMOND
risk factors may be found among individuals with some – but significant. Yet the genetic correlation between men and
relatively low – same-gender sexual behavior (i.e., “high” on women for “proportion of same-gender partners” was not
ever/never but “low” on proportionality). This is notable given significant for either the UK Biobank sample (r = .1) or the
the fact (reviewed earlier) that individuals reporting both 23andMe sample (r = −.2). This pattern of results yields
same-gender and other-gender behavior have greater mental interesting (albeit speculative) possibilities. For example, if
and physical health problems than individuals with exclusively “ever/never” shows significant genetic overlap between men
same-gender or exclusively other-gender behavior. Might this and women, but “proportionality” does not, then men and
reflect a shared genetic variance with risk tolerance or openness women whose same-gender sexual behavior reflects “ever/
to novelty, which might augment bisexual individuals’ expo never” influences might be more similar to one another
sure to environments that increase their mental and physical than women and men whose same-gender sexual behavior
health risks? In raising this possibility, I want to emphasize its reflects “proportionality” influences. To clarify this possibi
speculative nature: As pointed out by Ganna et al., we cannot lity, consider the following (oversimplified) example, based
infer causal pathways from these patterns of correlation. In on my speculation that the trait underlying “ever/never”
speculating about genetic contributions to the health risks might reflect risk tolerance or openness to novelty.
faced by sexually-diverse individuals, I am not suggesting that I suggested that such traits might prove particularly influen
“mostly heterosexual” and “bisexual” profiles cause mental/ tial on the same-gender behavior of individuals with relatively
physical health risks or that mental/physical health risks low same-gender interest (because for individuals with high
cause same-gender behavior. Rather, my chief interest lies same-gender interests, the strength of their same-gender
with the notable divergence between the correlations involving motivations might “win out” over fears of risk or unfamiliar
“ever/never” and the correlations involving “proportionality.” ity). Accordingly, we might expect that the “mostly hetero
This divergence should direct our attention to different con sexual” subgroup is more likely to contain individuals with
stellations of genetic and environmental factors (including high expression of “ever/never” genes (i.e., risk tolerance,
unmeasured “third variables”) that may characterize different openness to novelty) than the “bisexual” and “exclusively
subsets of sexually-diverse individuals and differentially aug same-gender” subtypes. If the genes associated with “ever/
ment their life experiences. Although I have suggested that never” are relatively similar among men and women, then
“risk tolerance” and/or “openness to novelty” are potential this would suggest that men and women within the “mostly
latent factors worth exploring in future studies, they are cer heterosexual” group are more genetically similar to one
tainly not the only ones. Other genetic studies using more another than men and women within the “bisexual” and
diverse populations, and perhaps assessing a broader range of “exclusively same-gender” groups.
sexual behaviors, may help to identify additional possibilities. What about gender differences within the “exclusive same-
gender sexuality” subgroup? Recall that “proportion of same-
gender sexual behavior” was genetically uncorrelated across
Revisiting Gender Differences
men and women. In other words, although “proportion of
Ganna et al.’s findings provide new perspectives on gender same-gender sexual partners” is heritable in both men and
differences in same-gender sexuality and in sexual variability. women, there appear to be different genes responsible for this
As reviewed earlier, reliable gender differences have been trait in each gender. This is why it is more accurate to say that
observed for certain types of sexual variability (such as sporadic women’s same-gender sexuality is differently – rather than
same-gender sexual behavior among self-identified heterosex less – genetically influenced than men’s same-gender sexuality.
uals) and in rates of bisexual attraction, but the extent to which How might these differing genetic influences help us under
these gender differences reflect inherent sex differences or stand gender differences in the expression of same-gender
sociocultural effects (or some combination thereof) remains sexuality at “higher ends” of the Kinsey scale (bisexuality and
unclear. Previous genetic research (reviewed in Bailey et al., exclusive same-gender sexuality)? As I have noted earlier,
2016) has found more consistent evidence for genetic contri nationally representative studies of same-gender sexuality con
butions to men’s same-gender sexuality than to women’s same- sistently find that women are more likely than men to report
gender sexuality, and this finding might suggest that there is bisexual patterns of attraction and behavior. Yet this broad
more “room” for nongenetic factors to influence same-gender statement does not provide a complete picture of this gender
sexuality among women than among men (Diamond, 2008). difference: Closer examination of the specific distribution of
Yet a close analysis of Ganna et al.’s findings suggests addi same-gender and other-gender attractions for women and men
tional possibilities. Specifically, it might be more accurate to shows that there are plenty of men with bisexual patterns of
characterize women’s same-gender sexuality as differently attraction, sometimes nearly as many as women (depending on
genetically influenced than less genetically influenced (com how different studies assess sexual attractions). Yet the consis
pared to men’s same-gender sexuality). tently largest gender difference concerns the “Kinsey 6” cate
Ganna et al. found more genetic overlap between men and gory, containing individuals reporting exclusive same-gender
women for “ever/never” than for “proportionality,” suggest attractions. In studies that specifically differentiate this form of
ing that gender differences may take different forms (and attraction from “mostly same-gender” and “bisexual” attrac
have different causes) within different subtypes of same- tions, men are between five and ten times more likely to
gender sexuality. The genetic correlation between men and endorse this category than women (Copen et al., 2016;
women for “ever/never” was .65 for the UK Biobank sample Mosher et al., 2005; Ott et al., 2011; Savin-Williams et al.,
and .34 for the 23andMe sample, and both were statistically 2012).
THE JOURNAL OF SEX RESEARCH 829
Hence, instead of describing women as “more bisexual,” it is Specifically, the negative direction of the correlation for
actually more informative to describe women as “less exclu women suggests that there is greater expression of “ever/
sively same-gender attracted.” As summarized by Bailey and never” genes among women with lower proportions of same-
colleagues over two decades ago, “Something about male sexual gender sexual behavior (i.e., “mostly heterosexuals”) than
orientation development makes extreme departures from het among women with higher proportions of same-gender sexual
erosexual development especially likely compared with female behavior. For men, this is not the case. If “ever/never” genes are
sexual orientation development” (Bailey et al., 2000, p. 533). associated with risk tolerance or openness to novelty, this
What is the “something” responsible for this difference? One suggests that “mostly heterosexual” women might have higher
possible explanation is the fact (noted earlier) that women’s risk tolerance/openness than “mostly heterosexual” men, and
historically subordinate status has made it more socially and in turn, higher levels of risk behavior. Is there any evidence for
economically risky for them to completely desist from hetero such a gender difference? In fact, there is: Vrangalova and
sexual behavior, compared to men. This could conceivably Savin-Williams conducted a comprehensive review of the men
suppress the expression of exclusive same-gender sexuality in tal and physical health of “mostly heterosexual” individuals,
women. Another possibility (also noted earlier) is that hege summarizing findings across 22 different samples from five
monic notions of rigid masculinity make it difficult or impos Western studies (Vrangalova & Savin-Williams, 2014).
sible for men to maintain their status as “authentic men” (and “Mostly heterosexuals” showed disproportionate mental/phy
hence worthy partners for women) if they violate the masculine sical health disparities across a wide range of outcomes, with
gender role by engaging in same-gender behavior, even if only effect sizes ranging from .2 to .5 for comparisons with exclusive
occasionally. These social pressures might have the effect of heterosexuals. There were minimal gender difference for effect
channeling bisexual men toward either exclusive heterosexu sizes involving mental/physical health disparities (such as
ality (in which case they stop violating the norms of hetero mood disorders, eating problems, weight status, and physical
sexual masculinity) or exclusive same-gender sexuality (in functioning). Yet for risk behaviors, all of the effects were
which they stop conforming to the norms of heterosexual substantially larger for women. Specifically, the mean and
masculinity altogether). median effect sizes for number of sexual partners, sexual and
Yet Ganna et al.’s findings provide new evidence for the non-sexual risk taking, rates of sexually-transmitted infections,
possibility that genetic factors might play an additional role. cannabis, other drug use, smoking, and alcohol use were up to
If different genes account for gradations in male same- three times greater for women than for men (p. 428–429). This
gender interest than female same-gender interest, perhaps is exactly what one might expect if a tolerance for risk or an
one of the consequences of this difference is that same- openness to new experiences is more characteristic of “mostly
gender interest takes a fundamentally different form in heterosexual” women than “mostly heterosexual” men.
women compared to men, manifested in lower rates of Such a gender difference might reflect the differential social
exclusive same-gender sexuality. Of course, in considering and cultural circumstances that face “mostly heterosexual”
this admittedly speculative possibility, we must keep in mind women versus men. As noted earlier, the category “mostly
that the overall genetic contributions to both “ever/never” heterosexual” subsumes individuals with diverse motives for
and “proportionality” were low, compared to nongenetic same-gender sexual contact. Some claim enduring, mildly
factors (heritability was .32 for “ever/never” and around bisexual patterns of attraction, some claim to have fallen unex
.25 for “proportionality”). Yet the point of this discussion pectedly in love with a single same-gender partner, some con
is to highlight the potential usefulness of investigating sider themselves “fluid,” some feel politically motivated to
whether gender differences have different magnitudes and challenge their own heterosexuality, and some are open to
different causes within different subgroups of sexually- a range of novel sexual experiences (Defries, 1976; Golden,
diverse individuals. 1994; Savin-Williams, 2017; Thompson & Morgan, 2008;
Ward, 2015). One possibility is that for men, “mostly hetero
sexuality” is more reflective of underlying bisexuality than
Different “Risk Tolerance” or “Openness” among Mostly
a generalized sexual openness or risk tolerance (consistent
Heterosexual Women versus Men?
with findings on “mostly heterosexual” men’s patterns of geni
One additional gender difference in the Ganna et al. findings tal arousal, Jabbour et al., 2020). Across the globe, men’s same-
warrants discussion. As noted earlier, the genes associated with gender sexuality is more stigmatized than women’s same-
“ever/never” were not significantly correlated with the genes gender sexuality (Bettinsoli et al., 2019), and women’s same-
associated with “proportionality.” Yet there was a gender dif gender sexuality is often eroticized for heterosexual men
ference in this regard: the genetic correlation for men hovered (Diamond, 2008; Fahs, 2009). Heterosexual women occupy
around zero, for both the UK Biobank and the 23andMe cultural environments replete with positive images of “occa
samples. The genetic correlation for women was nonsignifi sional” female-female eroticism, including film and television
cant, but negative (−.3 for the UK Biobank sample and −.4 for depictions of heterosexually-identified women experimenting
the 23andMe sample). Interpreting nonsignificant correlations with and enjoying sporadic same-gender sexual contact
is obviously risky (especially in light of the other weaknesses of (Diamond, 2005; Yost & McCarthy, 2012). Such experimental
this study’s sample), but the stark gender difference in this contact is not presumed to indicate an enduring same-gender
pattern (which Ganna et al. do not discuss) is curious, and orientation (Diamond, 2008; Fahs, 2009). Yet “occasional”
raises additional questions about gender differences in same- same-gender sexual contact among men is more often inter
gender sexuality. preted as potentially indicative of underlying bisexuality (for
830 L. M. DIAMOND
example, Kort, 2014; Zane, 2016), and these cultural reactions across three consecutive administrations, each 10 years apart,
may suppress expressions of “mostly heterosexuality” of the National Sexual Attitudes and Lifestyle study in the
among men. United Kingdom (1990, 1999, and 2010). Rates of same-
Accordingly, the overall population of “mostly heterosex gender sexual behavior were relatively stable among men
ual” women (which is larger than the population of “mostly (hovering around 7.5%) but increased linearly among women
heterosexual” men, as reviewed earlier) might contain (from 3.7% to 9.7% to 16%). Similarly, Gartrell et al. (2012)
a mixture of mildly bisexual individuals and “sexually open” compared rates of same-gender sexual contact reported by
individuals. In contrast, the population of “mostly heterosex 17 year olds in the 2002 and 2011 administrations of the
ual” men might contain predominantly “mild bisexual” indi National Survey of Family Growth. The percentage of 17 year
viduals (given that the greater stigmatization of same-gender old boys reporting same-gender sexual contact decreased from
contact among men than among women might make men less 6.6% to 1.4% during this period, but the percentage of girls
likely to express “sexual openness” through same-gender sex reporting same-gender sexual contact doubled (from 5% to
ual behavior). In turn, if sexual “openness” is related to risk 10%). Kuyper and Vanwesenbeeck (2009) compared the pre
tolerance, than we should observe greater overall risk behavior valence of same-gender sexual behavior in the Netherlands in
among “mostly heterosexual” women than among “mostly 1989 versus 2005. Women showed a 3-fold increase in same-
heterosexual” men, which is consistent with the findings gender sexual behavior, whereas men’s rates of same-gender
reviewed above. Importantly, the patterns suggested here sexual behavior did not change. Twenge et al. (2016) examined
demonstrate the critical importance of considering social and same-gender sexual behavior across multiple consecutive
cultural context when understanding different patterns of administrations of the United States General Social Survey.
same-gender expression. As long as cultures have different Between 1990 and 2010, the percentage of U.S. adults reporting
expectations, norms, incentives, and costs for women’s versus adult same-gender sexual contact increased from 4.5% to 8.2%
men’s same-gender versus other-gender sexuality (communi among men but from 3.6% to 8.7% among women.
cated through parents, peers, schools, governments, churches, These increases were concentrated among individuals with
and the 6–7 hours of visual media that children and adoles bisexual patterns of behavior: Rates of exclusive same-gender
cents consume each day, as reported by the APA Task Force on sexual behavior remained stable over time, consistent with the
the Sexualization of Girls, 2007), we cannot interpret genetic notion that greater familiarity with same-gender sexuality and
influences on sexual expression without considering how these lower “riskiness” of same-gender sexuality may prove more
influences interact with environmental factors to produce influential on individuals with relatively lower versus higher
diverse forms of sexual phenomenology. same-gender interests. As cultural acceptance of same-gender
sexuality continues to increase, individual differences in risk
tolerance may have a smaller and smaller influence on same-
Future Directions gender sexual behavior, perhaps leading to a broader uncou
pling between same-gender sexual behavior and other forms of
How Will Ongoing Cultural Changes Affect Same-gender
risky or novel behavior (especially among individuals with
Sexuality?
bisexual patterns of attraction). Investigating such possibilities
Ganna et al’s findings suggest new ways to understand subtypes has important implications for understanding and preventing
of sexual expression, and suggest new directions for study. In health disparities among bisexual and mostly-heterosexual
particular, the difference between the genetic underpinnings of individuals (who show greater disparities than do exclusively
“ever” engaging in same-gender sexuality and the genetic same-gender attracted individuals, as reviewed earlier).
underpinnings of one’s proportion of same-gender sexual part
ners suggests greater attention to the manifestations of these
Do We Have the Right Categories?
distinctions in different populations. If the “ever/never” trait
relates to risk tolerance or openness to novelty, as I have Another avenue for future research involves investigating the
speculated, then it might become less influential on same- degree to which observations of subtypes of same-gender
gender sexuality over time, as cultural attitudes about same- expression and their differing genetic/environmental influ
gender sexuality become more accepting and it becomes less ences depends on our conceptual framings of gender and
novel or socially risky to engage in same-gender sexual beha sexual orientation. Historically, laypeople and scientists have
vior. It is difficult to imagine a scenario in which same-gender conceptualized individuals as oriented toward the same gender
sexual behavior entails no risks for stigma or marginalization, or the other gender (or both genders), as opposed to being
but attitudes have changed rapidly over the past several dec oriented toward women or men (or both). This framing directly
ades, and research suggests that such attitudinal changes pro affects the type of gender differences we observe. Consider, for
mote increased rates of same-gender sexual behavior (Twenge example, one of the most robust gender differences in same-
et al., 2016), especially same-gender sexual behavior that is gender sexuality: the fact that women show more genital arou
pursued by individuals with relatively little same-gender sexual sal than do men when presented with sexual stimuli depicting
interest (Hawgood, 2020). their “less-preferred” gender (Chivers & Bailey, 2005; Chivers
Notably, historical reductions in the riskiness and novelty of et al., 2004, 2007). Early work suggested that this “nonspecific”
same-gender sexuality appear to have affected women more pattern of genital arousal (i.e., arousal that is not specific to
strongly than men. For example, Mercer and colleagues Mercer one’s preferred gender) characterized all women, but later
et al. (2013) compared rates of same-gender sexual behavior work showed that nonspecific genital arousal was most
THE JOURNAL OF SEX RESEARCH 831
pronounced among self-described heterosexual women represented by the Kinsey scale continues to dominate social
(reviewed in Chivers, 2017), and scholars have considered scientific research on this topic, perhaps reflecting the cultural
a range of social and evolutionary reasons for heterosexual dominance of this model of sexual orientation in Western cul
women’s uniqueness in this regard (Chivers, 2017; Diamond, ture (which necessarily feeds back to influence how sexually-
2017; Kuhle & Radke, 2013). diverse individuals come to perceive, understand, and experi
Yet the definition of heterosexual women as “unique” ence their own patterns of eroticism). Certainly, the same-
depends on the classification of sexual stimuli as preferred or gender/other-gender framing is useful for capturing the fact
non-preferred, according to participants’ self-described pat that heterosexuality is culturally valued and expected, whereas
terns of attraction. Within this framework, heterosexual same-gender sexuality is stigmatized and marginalized. The
women are unique because they show stronger genital arousal experience of stigma and marginalization is so relevant to the
to their non-preferred gender (i.e., women) than do all other life experiences of individuals with same-gender attractions (and
groups. But what if we re-classified the sexual stimuli as simply to the likelihood that they will express these attractions) that it
“men” versus “women?” Using this re-classification, exclusively seems naive to categorize attractions as “woman-oriented” or
gay men are suddenly the outlier group (Diamond, 2017). “man-oriented” without taking account of which type of attrac
Whereas heterosexual women, lesbian women, bisexual tions are socially permitted versus punished. Yet as we move
women, heterosexual men, and bisexual men all show some forward in trying to understand genetic influences on sexuality,
degree of genital arousal to sexual stimuli depicting women, we should remain mindful of the extent to which our framing of
gay men do not. core constructs (such as same/other versus woman/man) shapes
Hence, should we describe heterosexual women’s genital arou our observations and interpretations.
sal patterns as uniquely “fluid” or gay men’s genital arousal
patterns as uniquely “rigid?” How much do these patterns depend
Questions of Mechanism
on the mechanisms underlying genital versus subjective arousal,
given that these mechanisms are distinct (Chivers, 2017), and that Future research on sexual orientation, sexual fluidity, and their
concordance between genital and subjective arousal differs for genetic/environmental underpinnings may also benefit from
men versus women (Suschinsky et al., 2009)? Furthermore, what closer attention to the full range of conscious and noncon
is the role of aversion to same-gender versus other-gender stimuli scious processes through which different types of sexual stimuli
and/or partners (or male versus female stimuli/partners) in shap are attended to, neurologically processed, and responded to
ing subtypes of sexual diversity (see Dehlin et al., 2019; Freund, (Dickenson et al., 2020; Safron et al., 2007; Safron & Hoffmann,
Langevin, Chamberlayne et al., 1974; Freund, Langevin, Zajac 2017). Such process-oriented work is exemplified by Chivers’s
et al., 1974; Jabbour et al., 2020; Safron et al., 2007; Semon et al., (2017) nuanced and sweeping analysis of the potential contri
2017)? As reviewed earlier, the Kinsey-type “single continuum” bution of visual attention, implicit and explicit processing, and
model of sexual orientation (challenged by Ganna et al. 2019) incentive motivation to heterosexual women’s “nonspecific”
posits exclusive same-gender attractions and exclusive other- patterns of genital arousal. Given that environments fluctuate
gender attractions as polar opposites, but perhaps the true oppo over the lifespan, whereas genes remain fixed (setting aside for
site of exclusive same-gender attraction is same-gender aversion now the complications of epigenetics, Charney, 2012; Ngun &
or indifference. Models which account for aversion and/or indif Vilain, 2014; Rice et al., 2012; Richardson & Stevens, 2015), the
ference are better suited to including the experiences of asexual mechanisms underlying change in sexual experience and
individuals (Bogaert et al., 2018; Brotto & Yule, 2017) and those expression warrant particularly close study. As reviewed
who experience their own attractions as “gender neutral” above, sexual fluidity has been defined as a heightened sensi
(Diamond, 2008). Further integration of these nuances into tivity to situational change in sexual responsiveness (Diamond,
genetically-informed research would make a strong contribution 2008), but this definition leaves unspecified the process
to understanding the nature and development of different forms through which sexual responsiveness changes at all. There is
of sexual diversity. a growing body of rigorous research on the role of learning and
On this point, it bears noting that a growing body of sexuality conditioning in human sexual response (Hoffmann, 2012,
researchers now refer to sexual orientations as gynephilic (pre 2017; Hoffmann, Janssen, & Turner; Klucken et al., 2009),
ferring women), androphilic (preferring men) and biphilic (pre and this work should be more comprehensively integrated
ferring both genders) rather than “same-gender” and “other- into investigations of genetic and environmental influences
gender” (for example, Antfolk et al., 2017; Chivers, 2017; on same-gender expression.
Huberman & Chivers, 2015; Huberman et al., 2015; Petterson Of course, the notion of learned or conditioned sexual
et al., 2018; Semenyna et al., 2017; Skorska & Bogaert, 2020; responses may bring to mind the unfortunate history of beha
Snowden et al., 2020; Timmers et al., 2018; Vásquez-Amézquita vior-modification approaches to “extinguishing” undesirable
et al., 2019). There is an intuitive appeal to this approach, given sexual impulses (Hoffmann, 2017), which has had particularly
that most individuals describe themselves as desiring aspects of harmful effects on sexually-diverse individuals who have been
“women” and “men” rather than “sameness” and “otherness.” subjected to “conversion” and “reparative” therapies (APA
This approach is also better suited to describing the experiences Task Force on Appropriate Therapeutic Responses to Sexual
of transgender and nonbinary individuals, since it focuses on the Orientation, 2009). Perhaps because of this history, sexual
gender expression of sexual partners without making presump orientation is commonly (if inaccurately) described as funda
tions about one’s own or one’s partners’ birth-assigned sex/ mentally immutable (Diamond & Rosky, 2016). Yet from
gender. Yet the “same-gender/other-gender” framework a basic developmental perspective, the role of learning and
832 L. M. DIAMOND
exposure in human and nonhuman sexual development is well constructs such as “gay/lesbian,” “bisexual,” “fluid,” “polya
established (reviewed in Hoffmann, 2012, 2017). As Hoffman morous,” “nonbinary,” and “transgender”) gives rise to sexual
summarized, conditioning is quite simply “a process by which and gender diversity. Perhaps most important, assessing how
organisms, including humans, learn about the relationship such gene/environment interactions shape the life experiences
between events. Through conditioning, we can learn to predict of sexually-diverse individuals, including their mental and
events, we can learn signals for biologically significant stimuli, physical health and their susceptibility to social stress and
we can learn the value of stimuli, and we can learn the con marginalization, offers important possibilities for fostering
sequences of our actions. Hence, sexual conditioning can pre human flourishing across the entire spectrum of sexual
pare us to respond sexually and can contribute to our erotic expression.
preferences and to how we behave sexually” (Hoffmann, 2017,
p. 2213).
Positing a role for learning and experience in the expression ORCID
of same-gender sexuality does not invalidate the notion of
Lisa M. Diamond http://orcid.org/0000-0002-9875-9589
genetically influenced sexual predispositions. Rather, drawing
from Freund and Blanchard (1993), we might think of genetic
influences as differential sensitivities to certain classes of repro References
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