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O brother, where art thou? The fraternal birth-order effect

on male sexual orientation
David A. Puts, Cynthia L. Jordan, and S. Marc Breedlove*
Neuroscience Program and Psychology Department, Michigan State University, East Lansing, MI 48824-1101

igmund Freud has a lot to answer line or feminine as they like just by con- FBO effect on birth weight: each subse-
for. As the 20th century began, trolling how much T reaches the brain, quent baby a mother carries tends to
he convinced medical profession- especially early in life (7). weigh more, but boys whose mother car-
als that abnormal family relation- For men, those same putative markers ried brothers before them did not weigh
ships caused children to develop a of prenatal T do not paint as clear a as much as boys with older sisters (13).
homosexual orientation, which could be picture: some suggest gay men had seen The FBO effect on birth weight was
‘‘cured’’ by psychoanalysis. Not until the less prenatal T, some suggest they had greater among boys who turned out to
1970s would psychiatric and psychologi- seen more, and most suggest no differ- be homosexual than those who became
cal associations finally repudiate this ence between gay and straight men straight, suggesting that those boys who
idea that gay people have any illness (5, 6, 8). But, if T did not provide an are made gay by older brothers are also
that needs treating. Since then, a host of easy understanding of male sexual more susceptible to the FBO effect on
reports bolster the idea that prenatal orientation, several other findings still birth weight. Scouring the literature,
events, rather than family relations, af- implicated congenital origins for male these authors found an old report that
fect the likelihood that a person will homosexuality. Twin studies found that placenta weight, which also tends to get
grow up to be straight or gay. For exam- sexual orientation is heritable in both heavier with each subsequent child car-
ple, for each additional brother that pre- sexes (9), and a portion of the X chro- ried, weighs even more for boys if they
cedes him, a boy’s chance of growing up mosome was implicated in some cases have elder brothers rather than elder
to be gay increases by a third. If Freud of brothers who were both gay (10). sisters (14). So, elder brothers make
had known that, he might have sug- Then, 10 years ago, Ray Blanchard and subsequent brothers lighter than they
gested that the presence of older broth- Anthony Bogaert (11) made a startling, would have been and their placentas
ers shifts family dynamics, subjecting the counterintuitive finding: the more elder heavier. It is hard to see how older
youngest son to a social milieu that brothers a boy has, the more likely he is brothers could accomplish this socially,
leads to homosexuality. However, in this to grow up to be gay. There is no effect but neither do we know that these FBO
issue of PNAS, Anthony Bogaert (1) of older sisters and no effect of younger effects on birth weight and placenta
provides evidence that the social influ- siblings of either sex. Curiously, neither weight have anything to do with FBO
ence of an older brother is irrelevant to older nor younger siblings seem to have effects on orientation.
whether his younger brother will de- any effect on the sexual orientation of Most recently, Blanchard et al. (15)
velop a homosexual orientation. It is the females. So it is only older brothers af- collated several data sets and found an-
number of older biological brothers the fecting younger brothers. This fraternal other surprise: older brothers increase
mother carried, not the presence of birth-order (FBO) effect on male sexual the probability of a boy becoming gay
older brothers while growing up, that orientation has been replicated in sev- only if that boy is right-handed. Among
makes some boys grow up to be gay. eral data sets, including that gathered left-handed men, there’s no difference
Older stepbrothers in the home have no by Alfred Kinsey’s team in the 1940s in the incidence of homosexuality no
effect, although older biological brothers and 1950s, long before anyone had matter how many brothers they have.
raised apart still exert their influence. guessed there might be such an influ- What’s curious about this finding is that,
These data, by elimination, strengthen ence. How many men are gay because of by itself, left-handedness makes males
the notion that the common denomina- older brothers? One estimate is that ap- and females slightly more likely to be
tor between biological brothers, the proximately one in seven homosexual gay (16). So left-handedness makes
mother, provides a prenatal environ- men in North America are gay because males slightly more likely to be gay but
ment that fosters homosexuality in her of older brothers (12). That means that also negates the effects of older brothers
younger sons. about a million Americans are either on orientation. Note that, unless the
Since Simon LeVay’s 1991 (2) report gay men today or boys who are going to presence of older brothers affects the
that a brain nucleus that is larger in grow up to be gay because their mother development of handedness, this result
men than women is also smaller in gay had sons before them. If their mothers makes it seem unlikely that older broth-
men than straight men, circumstantial had carried only sisters before, the de- ers are affecting sexual orientation
evidence has accumulated suggesting mographics say, those males would be through social influences. Why would
that some people really are born to be- straight. It stretches the meaning of the boys with a brain organized in a right-
come gay. Most of these findings make word to suggest these boys and men handed fashion be affected by their
family relationships of the sort Freud ‘‘choose’’ to be gay. Who gets to choose older brothers’ antics when boys with a
scrutinized seem irrelevant. For women, how many brothers their mother had left-handed brain organization are not?
a bewildering array of body parts (ears, before them? Still, if we want to know whether the
fingers, eyes, arms) all indicate that les- But the question of mechanism re- social influence of older brothers is
bians were, on average, exposed to more mains. How do older brothers affect
fetal testosterone (T) than straight sexual development in their younger
women (3–6). The idea that the brains brothers? Is it the social influence of Conflict of interest statement: No conflicts declared.
of lesbians might have been masculin- the older brothers or something else? See companion article on page 10771.
ized by exposure to fetal T fits easily Several reports address this question *To whom correspondence should be addressed. E-mail:
with animal models, where researchers indirectly. For example, retrospective breedsm@msu.edu.
can make a mammalian brain as mascu- examination of records revealed an © 2006 by The National Academy of Sciences of the USA

www.pnas.org兾cgi兾doi兾10.1073兾pnas.0604102103 PNAS 兩 July 11, 2006 兩 vol. 103 兩 no. 28 兩 10531–10532

what is making some younger brothers nal blood, her immune system will now
gay, that hypothesis must be tested see proteins it has never seen before,
directly. This is what Bogaert (1) has including proteins encoded on her son’s
done, and the results are compelling. He Y chromosome. If she mounts an im-
replicates the oft-reported FBO effect, mune response to these proteins, then
but finds no effect of older brothers any subsequent sons will be exposed, via
who do not share the same mother, i.e., active transport across the placenta, to
stepbrothers and brothers with the same Fig. 1. Maternal immunization hypothesis. When maternal antibodies directed against the
father only. That failure to see an effect a mother is carrying her first son, the placental male-specific proteins. These maternal
of stepbrothers might be blamed on a barrier protects each from exposure to the other’s antibodies might then perturb develop-
modest sample size, but, on the other proteins. But inevitable mixing of blood upon de- ment of the younger son, decreasing
livery will expose the mother for the first time to birth weight and affecting his brain to
hand, an even smaller sample size shows male-specific proteins (blue triangles), including increase the probability that he will
that biological brothers (sharing the those encoded on the Y chromosome. If her im- grow up to be gay (13). Whether this is
same mother) who are raised apart still mune system produces antibodies to these pro- what is really happening for sexual ori-
significantly increase a younger brother’s teins, then the placenta may actively transport
entation remains to be seen, but it is a
likelihood of becoming gay. those antibodies (indeed, all IgGs) to subsequent
offspring in utero, potentially affecting develop-
provocative hypothesis. Freud thought
If the presence of the older brothers that a distant, emotionally cold father
ment of later-born sons, but not later-born
during development is not responsible might prevent a boy from identifying
for the FBO effect on orientation, what with Dad and steer him to homosexual-
is? One idea proposed by Gualtieri and ity. How much stranger it will be if,
Hicks (17) in 1985 for other traits found a first son has very little exposure to the instead of the father’s psychological re-
more commonly in males than in fe- proteins he is making because of the jection, it is the mother’s immunological
males is the ‘‘maternal immunization placental barrier. But upon delivery and rejection that inadvertently but actively
hypothesis’’ (Fig. 1). A mother carrying the inevitable mixing of fetal and mater- makes her son gay?

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10532 兩 www.pnas.org兾cgi兾doi兾10.1073兾pnas.0604102103 Puts et al.