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Digit Ratio

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DOI: 10.1007/978-3-319-16999-6_3829-1

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D
Digit Ratio
John T. Manning1 and Bernhard Fink2
1
Applied Sports, Technology, Exercise and
Medicine (A-STEM), Swansea University,
Swansea, UK
2
University of Goettingen, Goettingen, Germany
Synonyms
2D:4D; Digit ratio; Estrogen; Finger length; Pre-
natal sex steroids; Testosterone
Definitions
Effects of prenatal sex steroids on fertility, behav-
ior, and health.
Introduction
Digit ratio (or 2D:4D), i.e., the relative length of
the 2nd digit (“index finger”; 2D) and the 4th digit
(“ring finger”; 4D) has attracted the attention of
scholars for many years (Baker 1888; Minami
1952; Phelps 1952). At the center of this interest
lies the sex difference and ethnic variation in
2D:4D, and the appearance of the sexual dimor-
phism in 2D:4D at a very early stage in ontogeny.
# Springer International Publishing AG 2018
T. K. Shackelford, V.A. Weekes-Shackelford (eds.), Encyclopedia of Evolutionary Psychological Science,
https://doi.org/10.1007/978-3-319-16999-6_3829-1
Digit ratio is sexually dimorphic with males
typically having longer 4th digits relative to 2nd
digits than do females (male 2D:4D < female
2D:4D). The sex difference in 2D:4D suggests
an involvement of sex steroids (primarily testos-
terone [T] and estrogen [E]) and a developmental
origin that predates puberty and may indeed
extend back to the 1st trimester of pregnancy. In
2D:4D, we may have a relatively fixed record of
the influences of T and E during a narrow devel-
opmental window in early pregnancy. If this is
so, we can use 2D:4D to address many questions
regarding the etiology of sexually dimorphic
abilities (e.g., speed, stamina, strength), behaviors
(e.g., aggression, criminality, risk-taking), dis-
orders of development (e.g., autism, attention
deficit hyperactivity disorder, dyspraxia), and
major diseases (e.g., heart disease, many
cancers, osteoarthritis). There are differences in
mean 2D:4D across large ethnic groupings
(Caucasians, Blacks, East-Asians) in the fetus
and adults (Minami 1952), and across smaller
groupings such as the nation-states of Europe
(Manning and Fink 2011a). They hint at in utero
hormonal variation in humans, which may be
driven by sexual selection and contribute to
group differences in, e.g., marriage systems and
susceptibility to sex-dependent diseases. Further-
more, the group and sex differences in 2D:4D
may have a reach beyond that of Homo sapiens
to other taxa within the primates (Nelson and
2 Digit Ratio

Shultz 2010) and could indeed map on to other the phalanges (e.g., in the mouse, Zheng and
mammalian groups including the rodents (Zheng Cohn 2011).
and Cohn 2011). Finger lengths grow rapidly in children. If
2D:4D is to be used as a correlate of prenatal sex
steroids, it is necessary to establish whether it
remains stable over periods of rapid growth. Man-
Measurement of Digit Ratio
ning et al. (1998, 2013) have measured sex differ-
ences in right and left 2D:4D (boys < girls) in a
The most accurate measure of digit length is from
sample of 680 Caucasian children from 2 years to
radiographs that include the three phalanges.
18 years. There was no significant change in mean
However, experimenters are often unable to take
2D:4D or in the sex difference across year-groups.
X-rays of the hand, and where there is a pre-
Trivers et al. (2006) have reported stability of right
existing archive of X-rays, it has often been of
2D:4D with growth in Afro-Caribbean children,
the left hand only. Soft-tissue measurements of
but less stability in left 2D:4D. McIntyre et al.
digit length are more common and are usually
(2005) found a tendency for radiographic left
taken from the ventral surface of the digit. There
2D:4D to increase with growth in Caucasian chil-
is a limitation with this approach as the proximal
dren, but sex differences in 2D:4D within year
measurement point excludes approximately half
groups remained stable. The stability of 2D:4D
of the proximal phalanx. Direct measurement of
is enhanced because it is underpinned by bone
digit length has acceptable repeatability, but is
(the phalanges; e.g., Robertson et al. 2008) and
more time-consuming than indirect measurement
its maintenance during rapid growth is a remark-
(e.g., from photocopies or scans). However, the
able example of the homeostasis of form.
latter method causes a reduction in 2D:4D, which
is influenced by sex (greater in males than
females). The effects of indirect digit measure-
ment are a matter of debate (Manning et al. Human Comparative Differences
2005), and there are concerns regarding the “clar-
ity of report” regarding these effects (Ribeiro There are significant differences in mean 2D:4D
et al. 2016). across the major ethnic groups such that Cauca-
sians have higher 2D:4D than Blacks or East-
Asians. This pattern has been noted in fetal
ratios (Minami 1952) and in adults (Manning
Sex differences in Digit Ratios and Their
2002). Within ethnic groups, there is also evi-
Stability
dence for between-nation variation in 2D:4D.
For example, in the BBC internet study (Reimers
Digit ratio shows sex differences (2D:4D males
2007) with n > 200,000 participants, the mean
<2D:4D females) with low-to-medium effect
2D:4D (across sex and hand) per ethnic group was
sizes (Cohen’s d from about 0.20 to 0.60;
0.978 for Chinese participants, 0.981 for Blacks,
Manning 2002). Some of this variation is
and 0.989 for Caucasians. Within the Caucasian
influenced by measurement protocol (greater for
sample, there were significant between-nation dif-
indirect than direct 2D:4D; Hoenekopp and
ferences that map on to variation in national
Watson 2010) and by variation in the strength of
values for personality scores (Manning and Fink
sexual selection across populations (Manning
2011a), consumption of alcohol and cigarettes
et al. 2014a). However, we must also bear in
(Manning and Fink 2011b), and gender inequal-
mind that 2D:4D is a composite trait and that
ities (Manning et al. 2014a).
effect sizes for sex differences may vary across
Digit Ratio
Digit Ratios and Sex Steroids
Manning et al. (1998) suggested that 2D:4D
correlates negatively with prenatal testosterone
(PT) and positively with prenatal estrogen
(PE). In support of this model, they reported
that (i) 2D:4D was sexually dimorphic
(males < females), (ii) the dimorphism was stable
across age groups 2 years to 25 years and was not
affected by puberty, and (iii) low 2D:4D
(particularly right 2D:4D) was associated with
efficient spermatogenesis. Since the publication
of Manning et al. (1998), it has become apparent
that the sex difference in 2D:4D is determined by
the end of the 1st trimester of pregnancy (Galis
et al. 2010; Malas et al. 2006; Szwed et al. 2017)
and is in general stable in postnatal growth.
Fetal Testosterone and Estrogen
There is evidence that PT levels correlate nega-
tively with postnatal 2D:4D. Thus, children with
congenital adrenal hyperplasia (CAH), a trait
associated with high PT, have been reported to
have lower 2D:4D than controls (Brown et al.
2002; Ciumas et al. 2009; Okten et al. 2002).
Another study reported no association (Buck
et al. 2003), but overall a meta-analysis found
significant associations between low 2D:4D and
CAH (Hoenekopp and Watson 2010). In contrast
to CAH, men with Klinefelter’s syndrome
(47, XXY; which is accompanied by low PT)
have higher 2D:4D than population norms
(Chang et al. 2015; Manning et al. 2013; Savic
2014). It is to be noted that insensitivity to
T should remove variation in 2D:4D that is related
to PT, but leave the variation that is linked to
PE. Berenbaum et al. (2009) and van Hemmen
et al. (2017) reported that T-insensitive individ-
uals had increased mean 2D:4D, but variation in
2D:4D was still present. Their result is supportive
of a PT:PE link with 2D:4D.
In studies that assessed hormonal concentra-
tions in amniotic fluid, PT:PE ratio and PT
were found to be negatively correlated with
2D:4D (Lutchmaya et al. 2004; Ventura et al.
2013), and T in maternal serum was reported to
3
be positively correlated with children’s 2D:4D
(Barona et al. 2015; Ventura et al. 2013). With
regard to perinatal cord blood, we should not
expect to find correlations between 2D:4D and
T and E because sex-dependent variation is
established some 6 months before birth. How-
ever, associations were reported between low
2D:4D and high-functioning T-producing prenatal
Leydig cells that have their origin early in gesta-
tion (Mitsui et al. 2015).
The Ancient Origin of 2D:4D
Manning (2002, 2008) suggested that the sexual
dimorphism of 2D:4D is an ancient trait, which
may have its origin in the evolution of four-limbed
animals. Sex differences in 2D:4D are present in
birds (Nagy et al. 2016) and mammals (Nelson
and Shultz 2010). In rodents, inactivation of the
androgen receptor (AR) gene increases 2D:4D,
whereas inactivation of the estrogen receptor
(ER) gene lowers 2D:4D. The administration of
exogenous PT or the blocking of ERs lowers
2D:4D, whereas administration of PE or blocking
ARs increases 2D:4D (Auger et al. 2013;
Talarovicová et al. 2009; Zheng and Cohn
2011). These experimental data confirm that
2D:4D is fixed by PT:PE in a narrow prenatal
window and it is linked to the activation of
“skeletogenic” genes by PT:PE. The genes,
some 20 in number, have multiple functions,
which include the morphogenesis of the skeleton,
reproductive organs, and the brain (Zheng and
Cohn 2011).
Digit Ratio and Sensitivity to Testosterone
A number of studies have investigated the link
between low 2D:4D and high sensitivity to
T. This includes studies of associations between
2D:4D and the structure of the AR and those that
have reported associations between 2D:4D and
administered T. Modifiers of AR function include
polymorphisms in its coding sequence (e.g., var-
iable numbers of CAG repeats), interaction with
cofactors, intracellular trafficking of the T-bound
complex, and interaction with general transcrip-
tion factors. Over 170 proteins possess AR
4 Digit Ratio
coactivating or corepressing characteristics, indi-
cating that a multitude of signals converge on the
AR to regulate its function (Heemers and Tindall
2007; Heinlein and Chang 2002). Some studies
have reported a negative relationship between
2D:4D and CAGn, and one study found that
2D:4D in neonates was dependent on CAGn and
circulating levels of T (see Hoenekopp 2013 for a
meta-analysis). However, most reports have not
found a correlation between CAGn and 2D:4D.
The importance of this null-finding for the puta-
tive association between 2D:4D and PT is a matter
of debate. Hoenekopp (2013) has suggested that
CAGn has a weak effect on AR transcription rates,
and no or little or inconsistent effects on traits that
are known to be strongly influenced by T. An
alternative view of the correlations between
2D:4D and the AR is given by Voracek (2014).
Clearly the link between 2D:4D and PT would
benefit from an experimental rather than a corre-
lational approach.
A more powerful experimental method of
considering sensitivity to T is to administer the
hormone and note whether 2D:4D modulates the
response. It has been reported that 2D:4D modu-
lates the effects of exogenous T application on
empathy, moral judgment, cooperation, trust,
preference for status goods, aggression, and
brain activity associated with mental rotation
task scores (Buskens et al. 2016; Montoya et al.
2013; Pintzka et al. 2016; van Honk et al. 2011,
2012 for women; Carré et al. 2015; Wu et al. 2017
for men). The direct manipulation of T itself
makes this experimental procedure more powerful
than correlational data. The link between 2D:4D
and sensitivity to T is potentially further amplified
by associations between low 2D:4D and produc-
tion of T in competition-induced circumstances
(Manning et al. 2014b). Thus, men with low
2D:4D, and those with lower right 2D:4D relative
to left 2D:4D, tend to produce high levels of T and
high T:C ratios when challenged with intense
exercise and/or aggressive stimuli (Crewther
et al. 2015; Kilduff et al. 2013a, 2013b).
Digit Ratios, Target Traits, and Effect
Sizes
Masculinized and feminized traits are generally
associated with low and high 2D:4D, respectively.
Effect sizes for associations vary. The strongest
correlations with 2D:4D are found for perfor-
mance in sport and developmental disorders and
the weakest among personality traits. Autism is a
severe developmental disorder that is strongly
sex-dependent such that the male-to-female sex
ratio is 4 to 1 across the full IQ range, and rising to
9 to 1 in children with normal IQ. Children with
autism are often unresponsive to people and lack
awareness of the feelings of others (“mind-blind-
ness”; Baron-Cohen 1995). The sex-dependent
pattern of autism has led to the suggestion that it
may result from the influence of high levels of PT,
the “extreme male brain theory” (Baron-Cohen
and Hammer 1997). Thus, it has been proposed
that PT effects of hypermasculinization of the
brain may cause autism.
Manning et al. (2001) considered the relation-
ship between 2D:4D and autism in 72 children
(Asperger’s Syndrome [AS]; of these, 23 par-
ticipants had normal IQ), 210 unaffected relatives
(88 fathers, 88 mothers, 34 siblings), and a
matched normative control sample. In comparison
to controls, children with autism and children with
AS had lower 2D:4D, and fathers, mothers, and
sibs from affected families had lower 2D:4D.
These findings have been the focus of a num-
ber of replications and two meta-analyses
(Hoenekopp 2012; Teatero and Netley 2013).
Both studies were supportive of a correlation
between low 2D:4D and autism spectrum disor-
der. More recently, Barona et al. (2015) reported
an association between low 2D:4D and emotional
and communication difficulties in a large
community-based sample. However, this re-
lationship was restricted to the lowest 10% of
male 2D:4D ratios. Currently, it is unknown
whether associations between low 2D:4D
(high PT) and symptoms of “mind-blindness”
are present throughout normative populations
Digit Ratio
(as predicted by the extreme male brain theory) or
whether these effects are found only among males
who have been exposed to very high PT.
Digit Ratio and Sports Performance
Manning and Taylor (2001) reported a negative
relationship between 2D:4D and performance
across a range of sports and extended this analysis
to a detailed consideration of professional soccer
players. Since this initial report, there have been
similar findings for other sports such as running,
skiing, surfing, rugby, rowing, basketball, and
Sumo wrestling, and for measures of fitness and
strength. A meta-analysis confirmed the initial
finding of Manning and Taylor, and found sub-
stantial relationships but with some heterogeneity
in effect sizes such that endurance sports tended
to show the strongest effects (Hoenekopp and
Schuster 2010).
Digit Ratio, Aggression, Risky/Criminal
Behavior
The associations between 2D:4D and aggression
and rule breaking show sex differences, with
higher effect sizes in men compared to women
(e.g., Hoenekopp and Watson 2010). There is
evidence that T influences such behavior, with
greater intensity of expression associated with
high T (Mehta et al. 2015). It is to be expected
that 2D:4D would be related to aggression, risky/
criminal behavior, and dominance because 2D:4D
modulates the action of T (Carré et al. 2015;
Montoya et al. 2013; van Honk et al. 2011,
2012). PT is associated with neuronal loss and
there is evidence of a positive relationship be-
tween 2D:4D and brain morphological features.
Darnai et al. (2016) reported significant positive
correlations between 2D:4D and volume mea-
surements such as total cerebral cortex, total cer-
ebellar white matter, and total cerebellar cortex in
men but not women. Gorka et al. (2015) found
low 2D:4D was associated with reduced anterior
cingulated cortex gray matter volume (a brain
region supporting conflict monitoring and behav-
ioral inhibition).
5
Aggression
There are reliable negative relationships between
2D:4D and sports performance and many of these
sports involve aggressive/competitive interac-
tions (Hoenekopp and Schuster 2010; Manning
and Taylor 2001). This pattern of significant
2D:4D effects in sports that provide intense chal-
lenge conditions is what we would expect if
2D:4D were a proxy for sensitivity to and ampli-
tude of T-spikes. Therefore, it seems reasonable to
assume that 2D:4D and aggression is negatively
associated. However, the effect size of the 2D:4D
and aggression relationship has proved difficult to
quantify.
A meta-analysis of studies concerning 2D:4D
and aggression revealed no significant rela-
tionship for women (n = 19 studies) and a
small but significant negative relationship for
men (n = 18 studies, r = 0.06;
Hoenekopp and Watson 2010). Subsequent
large-sample studies have reported male effect
sizes of about r = 0.10 (Butovskaya et al.
2013; Hoenekopp 2011).
Correlations of r = 0.06 to 10 for male
2D:4D associations with aggression are small.
However, aggression is not invariant across situ-
ations but is context-dependent (Millet 2011).
Thus, an interactionist perspective is necessary
to understand its relationship with T. Baseline
differences in T from individual to individual do
not correlate strongly with between-individual
variation in aggression. Rather, it is variation in
spikes of T in response to challenge situations that
correlate with levels of aggression (Carré and
Olmstead 2015). Low 2D:4D (particularly low
right-left 2D:4D) individuals tend to produce
marked T-spikes on challenge with aggressive
stimuli and/or intense exercise (Crewther et al.
2015; Kilduff et al. 2013a, 2013b). They are also
sensitive to high concentrations of T (Carré et al.
2015; Montoya et al. 2013; Pintzka et al. 2016;
van Honk et al. 2011, 2012) and show negative
correlations between 2D:4D and aggression in
challenge conditions (Kilduff et al. 2013b; Millet
and Dewitte 2007).
6 Digit Ratio
Therefore, it may be that the 2D:4D and
aggression correlation is significant in men when
in challenge conditions, and weak in contexts
other than challenges (Kilduff et al. 2013b;
Manning et al. 2014b; Millet and Dewitte 2007).
Support for this suggestion comes from real-
world studies. Thus, in professional soccer
players, there is a negative correlation between
the number of yellow and/or red cards per match
and 2D:4D in junior and elite players (Mailhos
et al. 2016; Perciavalle et al. 2013). Similarly,
patients with “boxer’s” fractures (of fourth or
fifth metacarpals) acquired during street-fighting
have lower 2D:4D ratios than controls (Joyce
et al. 2013).
Risky/Criminal Behavior
The relationships between 2D:4D and risky/crim-
inal behavior appear to show higher effect sizes
than those found for the relationship of 2D:4D and
aggression. For example, Hoskin and Ellis (2015)
reported a mean correlation of about r = 0.40
between 2D:4D and criminality. Hanoch et al.
(2012) found a correlation of r = 0.24 between
2D:4D and impulsivity among offenders and non-
offenders. Schwerdtfeger et al. (2010) reported a
correlation of r = 0.36 between 2D:4D and
traffic violations. Furthermore, large internet sam-
ples of 2D:4D enable us to consider risk avoid-
ance across nations rather than individuals. In the
BBC internet study, Manning and Fink (2011a)
reported significant correlations for mean male
2D:4D per nation and national values of uncer-
tainty avoidance (23 nations: right hand r = 0.58,
left hand r = 0.59).
Personality
It is generally thought that 2D:4D is only weakly
associated with aspects of personality such as the
“Big-Five” constructs (Austin et al. 2002; Fink
et al. 2004; Lippa 2006). This view may be cor-
rect. However, the field would benefit from a
consideration of the context in which personality
is examined. For example, in conditions such as
aggressive challenge, the relationships between
2D:4D and traits such as neuroticism may change
(Manning and Fink 2018).
Sexual Orientation
Interest in the relationships between 2D:4D and
sexual orientation came shortly after the publica-
tion of Manning et al. (1998). Unfortunately, the
first papers (Robinson and Manning 2000;
Williams et al. 2000) used indirect methods
(photocopies) to measure finger lengths. This
practice has continued such that most studies
show this limitation. A meta-analysis concluded
that lesbians showed lower mean 2D:4D than
heterosexual women, but the 2D:4D of gay men
did not differ from that of heterosexual men
(Grimbos et al. 2010). This is a surprising result
which should be treated with caution. Homosex-
uality in men is thought to be primarily deter-
mined by early genetic and hormonal influences
while homosexuality in women may be more
influenced by later-life effects. Finger lengths in
almost all the studies considered by Grimbos et al.
(2010) were measured indirectly. The BBC inter-
net study concluded that lesbians and heterosex-
ual women did not differ in their 2D:4D but gay
men had significantly higher 2D:4D than hetero-
sexual men (Manning et al. 2007). Further work
on 2D:4D and sexual orientation would benefit
from the use of direct digit measurement.
Digit Ratios and Health
There are possible applications for 2D:4D and
diagnosis, prognosis, and response to therapeutic
interventions in a number of diseases and devel-
opmental disorders.
Sperm Production
Manning et al. (1998) reported negative relation-
ships between 2D:4D and spermatogenesis. How-
ever, this was in a sample from a fertility clinic
and the participants included a proportion of
men who were producing few or no sperm. The
links between 2D:4D and spermatogenesis in the
normative population are more controversial and
need to be clarified (see Auger and Eustache 2010
for data and discussion).
Digit Ratio 7

Digit Ratio, Table 1 The relationships between national means of 2D:4D and age-standardized DALY’s by 100,000 in
29 countries and 16 malignant neoplasms. For cancer of the breast, cervix, uterus, and ovary, the mean of female right and
left 2D:4D was used, and for cancer of the prostate, the mean of male right and left 2D:4D was used
Malignant neoplasm r r2 p Adjusted p
Mouth 0.35 0.12 0.06
Esophagus 0.19 0.04 0.31
Stomach 0.57 0.32 0.001 0.02 (2-Tailed)
Colon/rectum 0.05 0.003 0.79
Liver 0.56 0.31 0.002 0.03 (2-Tailed)
Pancreas 0.23 0.06 0.22
Lung 0.51 0.26 0.005 0.03 (2-Tailed)
Melanoma 0.41 0.17 0.03
Breast 0.50 0.25 0.005
Cervix 0.48 0.23 0.008 0.04 (1-Tailed)
Uterus 0.11 0.01 0.57
Ovary 0.08 0.007 0.67
Prostate 0.53 0.28 0.003 0.02 (1-Tailed)
Bladder 0.47 0.22 0.01
Lymphoma 0.07 0.004 0.73
Leukemias 0.58 0.33 0.001 0.02 (2-Tailed)

Heart Disease years; Global Health Estimates 2015) of a number

There is accumulating evidence that low 2D:4D in
men is associated with protection against early
myocardial infarction and coronary heart disease
(e.g., Manning and Bundred 2001; Wu et al. 2013;
Zhenghao et al. 2012). However, the correlation
between 2D:4D and heart disease in women
requires clarification.
Cancers
There is now much interest in links between
2D:4D and cancers that are dependent on sex
steroids. We feel this is particularly important as
there is the possibility that 2D:4D may be useful
in prediction of susceptibility and also disease
progression where the inhibition of sex steroids
may be part of the treatment (e.g., Li et al. 2017).
We include here cancer of the breast (Muller
et al. 2011) and prostate (Rahman et al. 2010).
There are now 19 papers concerning 2D:4D and
susceptibility to various sex-dependent cancers
(Bunevicius 2018). In order to focus attention on
those cancers that may be linked to 2D:4D, we
present associations between overall mean 2D:4D
per nation and DALY’s (disability adjusted life
of cancers, including those for the stomach, liver,
lung, cervix, prostate, and various forms of leuke-
mia (Table 1).
Osteoarthritis
The relationship between low 2D:4D and risk for
osteoarthritis of the knees seems to be firmly
established (e.g., Ferraro et al. 2010). This corre-
lation appears to be independent of wear-and-tear
consequent on engagement in sport or other phys-
ical activity (Haugen et al. 2011). Further work is
indicated regarding 2D:4D and prognosis and
response to treatments.
Smoking and Alcohol Choices
Smoking and the consumption of alcohol are
major public health issues. In the BBC internet
study, Manning and Fink (2011b) reported corre-
lations with 2D:4D such that low 2D:4D was
linked to high alcohol intake and smoking to
high 2D:4D. These correlations applied to indi-
viduals and to mean 2D:4D per country and
national levels of consumption of cigarettes and
alcohol. Further work is necessary to identify the
8 Digit Ratio
relationships between 2D:4D, health (heart dis-
ease, lung cancer, respiratory conditions), and
consumption of tobacco and alcohol).
Conclusion
In humans, 2D:4D is a sexually dimorphic trait
(male 2D:4D < female 2D:4D) that is more or less
fixed by the end of the 1st trimester of pregnancy,
and which is little affected by puberty. It is a
composite trait such that the sexual dimorphism
may differ between the phalanges. The sex differ-
ence in 2D:4D is ancient among the four-limbed
vertebrates with similar sex-dependent ratios in
mammals (rodents and primates). There is exper-
imental and correlational evidence for the in-
fluence of prenatal sex steroids on mammalian
2D:4D. Low 2D:4D is associated with high PT
low PE and high 2D:4D is related to low PT
and high PE. Digit ratio is determined in a narrow
fetal time-window and the effect sizes of cor-
relations with target traits depending on whether
such traits are influenced in that time-window.
There are large effect sizes in the relationships
between 2D:4D and sport (particularly sports
that are dependent on endurance) and develop-
mental disorders. Effect sizes are smaller with
regard to complex sex-dependent behavioral traits
that are difficult to measure. A potentially impor-
tant aspect of 2D:4D are its links with sex-
dependent diseases such as cancers, heart disease,
and osteoarthritis.
Our understanding of 2D:4D has advanced
considerably since the late 1990s. However, the
speed of this advance has been affected by the
many methods that have been used to measure
finger length. The lack of clarity of report
concerning this issue means that we still do not
understand the properties of various indirect
methods of digit measurement. This apart, we
feel that the diverse program of research into
2D:4D will continue to reveal the importance of
the prenatal hormonal environment on behavior,
physiology, and predisposition to disease.
Cross-References
▶ Bernhard Fink
▶ Hormonal Effects on Human Fetal
Development
▶ John Manning
▶ Robert Trivers
▶ Sex Differences
▶ Sexual Dimorphism
▶ The Biology of Human Gender
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