Neuropsychologia 184 (2023) 108523

Contents lists available at ScienceDirect

Neuropsychologia
journal homepage: www.elsevier.com/locate/neuropsychologia

Review article

Handedness in twins reared apart: A review of the literature and new data
Gareth Richards a, *, Nancy L. Segal b
a
School of Psychology, Faculty of Medical Sciences, Newcastle University, Newcastle upon Tyne, Tyne and Wear, NE2 4DR, UK
b
Department of Psychology, California State University, Fullerton, CA, 92831, USA

A R T I C L E I N F O A B S T R A C T

Keywords: Reared-apart twin studies are a powerful means for identifying the relative contributions of heredity and envi­
Behavioural genetics ronment to variation in human physical and behavioural traits. One such characteristic is handedness, for which
Handedness it has long been noted that approximately 20% of twin pairs are comprised of one right-handed cotwin and one
Review
left-handed cotwin. Reared-together twin studies suggest a slightly greater concordance in monozygotic (MZT)
Twins
than dizygotic (DZT) twins, implying that genetics influences hand preference. We report here two studies of
Twins reared apart
handedness in reared-apart twins. Study 1 synthesizes the available data and estimates that at least N = 560
same-sex reared-apart twin pairs (for which zygosity is known with reasonable confidence) have been identified.
Of these, handedness data are available for both members of n = 415 pairs. We observed similar levels of
concordance/discordance for reared-apart monozygotic (MZA) and dizygotic (DZA) twins. However, although
direction of handedness (right or left) has frequently been examined, strength of handedness (strong or weak) has
not. Study 2 examined strength of hand preference and relative hand skill, as well as right- and left-hand speed,
information available for participants in the Minnesota Study of Twins Reared Apart (MISTRA). We provide
evidence of heritability for right-hand and left-hand speed. We also found hand preference strength was more
alike than chance in DZA, but not MZA, twins. Findings are discussed in relation to genetic and environmental
influences on human handedness.

1. Introduction included mention of handedness.

Monozygotic (MZ) twins share 100% of their genes identical by
descent, having formed when a single fertilised egg, or zygote, divided
during the first two weeks post conception. In contrast, dizygotic (DZ)
twins are no more alike than other full siblings in sharing, on average,
50% of their genes identical by descent. The classic twin method com­
pares the similarity (or concordance) of MZ and DZ twins, with a greater
resemblance in the former relative to the latter indicating that genetic
factors affect the phenotype under study. However, as twins are typi­
cally raised together, they may share very similar early environments;
this assumption is explicated in the paragraph that follows. One of the
most powerful methods through which behaviour geneticists can
ascertain the relative contributions of heredity and environment is to
study twins who have been reared apart from infancy (Segal, 2012). In
such cases, MZ and DZ twins still share 100% and 50% of their genes
identical by descent, respectively, but genetic effects are no longer
confounded by those of the shared environment. Although not typically
their primary focus, case reports of reared-apart twins have often
The equal environments assumption (EEA) is important to explain as
violation can affect the findings and interpretations of twin-based ana­
lyses. The EEA specifies that environmental factors relevant to traits of
interest are “equal” for MZ and DZ twins. Some critics of twin research
assert that since MZ twins generally share more activities (e.g., activities
related to the trait of sociability) than do DZ twins, this renders twin
study findings invalid. However, as pointed out by Segal (2012, p. 3),
“this difference is problematic only if (1) MZ twins are required to share
… activities to a greater degree than DZ twins, and (2) sharing activities
affects sociability in a meaningful way. If sharing activities increases
sociability, then the environments of the two types of twins would not be
equivalent with respect to this behavior. The equal environments
assumption would have been compromised in this hypothetical
example.” In fact, such objections have been tested empirically and have
been dismissed in most cases.
Other concerns of twin research involve twins’ sharing of prenatal
environments. It is not widely appreciated that the prenatal circum­
stances and adversities of MZ twins (e.g., twin-to-twin transfusion

* Corresponding author. School of Psychology, Newcastle University, 4.32 Dame Margaret Barbour Building, Wallace Street, Newcastle upon Tyne, UK.
E-mail address: gareth.richards@ncl.ac.uk (G. Richards).

https://doi.org/10.1016/j.neuropsychologia.2023.108523
Received 13 September 2022; Received in revised form 14 February 2023; Accepted 20 February 2023
Available online 12 April 2023
0028-3932/© 2023 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
G. Richards and N.L. Segal
syndrome [TTTS]; positioning in the womb) lead mostly to differences,
not similarities, in newborns. In contrast, DZ twins do not experience
TTTS, but are subject to the vagaries of differential positioning in the
womb and related effects. Again, a classic comparison of MZ and DZ
twins would be a violation of the EEA only if the effects were relevant to
the trait in question. Direction of handedness may be one such trait. That
is because only MZ twins result from zygotic division and could be
affected by differences in cell distribution, events possibly linked to
handedness. However, whether timing of zygotic division is associated
with handedness in MZ twins has recently been questioned (see Segal
and Montoya, 2018).
The majority of humans are classified as right-handed, while 10.6%
prefer using their left hand for most unimanual and bimanual tasks
(Papadatou-Pastou et al., 2020). In fact, a population-level minority of
left-handers has been observed consistently throughout history and
across cultures (Ocklenburg and Güntürkün, 2018). However, the
complexity of handedness is often underappreciated, and various as­
pects of this trait can be misunderstood. Although hand preference is
usually considered in terms of direction (left or right), it can also be
measured with reference to degree or strength (McManus and Bryden,
1992). For example, a given ‘right-hander’ may have a very strong
preference for using their right hand for a particular task or, alterna­
tively, this preference might be relatively weak. Additionally, rather
than simply assessing an individual’s hand preference, behavioural
measures can be employed to quantify the difference in skill between the
two hands (i.e., relative hand skill) (McManus and Bryden, 1992). As
with hand preference, relative hand skill can be considered in terms of
both direction (right > left or left > right) and strength (strong or weak
lateralisation).
It has been suggested that birth stress/trauma can alter handedness,
although empirical findings have been equivocal (Bakan et al., 1973;
Beaton, 2003; Coren and Porac, 1980; McManus, 1981; Nicholls et al.,
2012; Ooki, 2006; Satz, 1972; Searleman et al., 1989; Williams et al.,
1992). If such effects do occur, they would mostly be in the direction of
right to left, i.e., ‘pathological left-handedness’ (Satz, 1972), given the
higher frequency of left-handedness among individuals with brain
damage or marked cognitive impairment (Papadatou-Pastou and Tom­
prou, 2015). The birth circumstances of reared-apart twins are often
unknown—however, while birth trauma may precipitate shifts in
handedness from right to left, it is unlikely that MZ and DZ twins would
be affected equally since MZ twins experience more hazardous prenatal
environments. There is no extant theory as to how birth trauma in twins
would affect the strength of handedness, but it seems likely that MZ and
DZ twins would be equally affected.
Hand preference runs in families, with the children of two left-
handed parents exhibiting higher rates of left-handedness (~26%)
than the children of one left-handed and one right-handed parent
(~19%), and higher rates than the children of two right-handed parents
(~9%) (McManus and Bryden, 1992). Adoption studies indicate a ge­
netic basis, with the handedness of adoptees more closely resembling
that of their biological parents than adoptive parents; however, analyses
have often been underpowered, and findings have been somewhat
mixed (Carter-Saltzman, 1980; Saudino and McManus, 1998). Simple
Mendelian inheritance (Jordan, 1914) clearly does not conform to
empirical observation (McManus and Bryden, 1992), a fact exemplified
by the approximately 20% of MZ twin pairs that are discordant for di­
rection of hand preference (Pfeifer et al., 2022). Single gene models
(Annett, 2002; McManus, 2002) positing two alleles at a single
locus—one coding for right-handedness, the other for chance—have also
failed to account for this trait (McManus, 2022), especially as large-scale
molecular genetic studies have established that human handedness
shows polygenic effects (Cuellar-Partida et al., 2020).
More than a century ago, Danforth (1919, p. 400) commented that “a
surprising number of twin pairs seem to be composed of one right and
one left handed individual”. A recent meta-analysis (Pfeifer et al., 2022)
with N = 36,217 twin pairs (n = 15,506 MZ pairs; n = 20,711 DZ pairs)
2
Neuropsychologia 184 (2023) 108523
from k = 44 studies reported that the concordance rate for direction of
hand preference was slightly, but statistically significantly, higher in MZ
(80.49%) than DZ (79.27%) twin pairs. However, to effectively partition
variance between genetic factors and shared environmental factors,
studies of MZ (MZA) and DZ (DZA) reared-apart twin pairs are required.
2. Study 1: review of handedness in reared-apart twins
Handedness was reported for the very first pair of MZA twins (Bessie
and Jessie1) to appear in the academic literature (Muller, 1925; Pope­
noe, 1922; Saudek, 1933). Muller (1925, p. 438) commented that “Both
twins, moreover, had early shown lefthandedness, though B had been
broken of it.” A number of such case reports have since been published,
yet the only near-systematic compilation of their findings, that of Farber
(1981), predates much of the relevant literature. We, therefore, sum­
marise the available data here.
2.1. Literature search strategy
Handedness in reared apart twin pairs is represented by a chal­
lenging literature for several reasons. First, although in many cases
handedness has been reported, in no instance has it been the primary
focus of the study. This can make identifying such research difficult.
Second, as reared-apart twins embody such a rare and engaging phe­
nomenon, entire books are often written about individual cases. Some­
times these appear outside of the academic literature (e.g., Lindeman,
1969); in some cases, they are authored by the twins (e.g., Bordier and
Futerman, 2014; Schein and Bernstein, 2007), and books are sometimes
self-published (e.g., Glasby et al., 2013). Third, cases of reared-apart
twins have sometimes been identified from twin case series of genetic
and environmental studies of psychiatric conditions (Essen-Möller,
1941; Tienari, 1963a), e.g., schizophrenia (Gottesman and Shields,
1972; Kallmann, 1938; Kurihara, 1959; Mitsuda and Sakai, 1968), bi­
polar disorder (Stenstedt, 1952), psychosis (Kringlen, 1967a; Slater,
1953), as well as alcoholism (Kaij, 1960), lateral asymmetries (Bou­
terwek, 1943), criminal tendencies (Hayashi, 1963; Kranz, 1936; Lange,
1931; Yoshimasu, 1941), and the effects of head injury (Dencker, 1958).
In some instances, these data are complicated further by multiple
reporting (often including books and monographs) of the same twin
series (e.g., the case series reported on by Tienari [1963a; 1963b, 1966;
1968]). Additionally, there are many relevant articles, again including
books and monographs, published in languages other than English, e.g.,
German (Bouterwek, 1936, 1943; Essen-Möller, 1941; Kranz, 1936;
Prokop and Druml, 1973; Wagenseil, 1931), French (Franceschetti,
1948; Franceschetti et al., 1948), Japanese (Hayashi, 1963; Kurihara,
1959; Yoshimasu, 1941), Chinese (Gao et al., 2011).
Due to the above complications, search strings such as “Reared apart
twins” AND “Handedness” would likely be unreliable, and many rele­
vant articles would be missed. We, therefore, began by including all
cases from the three ‘classic’ studies of reared apart twins, that is, those
of Newman et al. (1937), Shields (1962), and Juel-Nielsen (1965, 1980),
and added to these data from the Minnesota Study of Twins Reared
Apart (MISTRA) (Segal, 2012), Swedish Adoption Twin Study of Aging
(SATSA) (Pedersen, 2015), Fullerton Study of Chinese Twins Adopted
Apart (Segal et al., 2011), the Finnish Twin Cohort Study (Kaprio et al.,
2019), and the Louise Wise Services/Child Development Center
(LWS/CDC) Twin Study (see Segal, 2021a). To identify other relevant
data, we examined all cases mentioned within reviews of reared apart
twin studies/cases, e.g., Farber (1981), Segal (2011a, 2012, 2021a),
Price (1968), Gottesman and Shields (1972), Tucker (1997), Slater
(1968), and Shields and Slater (1967). We returned to the original
literature for each case identified and translated those reports which
1
Please note that this pair is not the same as the Bessie and Jessie reported on
by Burlingham (1952).
G. Richards and N.L. Segal
were not published in English. We also checked reference lists for rele­
vant articles, and contacted authors of largescale studies for which we
knew reared-apart twin cases had been identified: that is, the SATSA
(Pedersen, 2015), Finnish Twin Cohort Study (Kaprio et al., 2019),
Osaka University Aged Twin Registry (Hayakawa et al., 2002)/Japanese
Study of Adult Twins Reared Apart and Growing Old Separately (Hay­
akawa et al., 2006), and the Chinese National Twin Registry (CNTR)
(Gao et al., 2011; Zhou et al., 2015). Therefore, although the current
paper departs from the Preferred Reporting Items for Systematic Re­
views and Meta-Analyses (PRISMA) (Moher et al., 2009) criteria, to the
best of our knowledge we have retrieved all available studies/cases.
3. Study 1 results
3.1. Number of reared apart twin pairs identified
We summarise case reports of same-sex reared-apart twins in
Table 1. It is not possible to determine with certainty exactly how many
reared-apart MZ male (MZAm), MZ female (MZAf), DZ male (DZAm),
and DZ female (DZAf) pairs have been identified in the scientific liter­
ature and/or mainstream media. However, we were able to identify n =
415 cases (n = 79 MZAm; n = 127 MZAf; n = 78 DZAm; n = 131 DZAf)
for which handedness data are available for both members of each twin
pair. This rises to n = 472 (n = 94 MZAm; n = 157 MZAf; n = 80 DZAm;
n = 141 DZAf) when also considering those cases of Shields (1962) for
which handedness data were missing or incomplete, and the n = 2 cases
of Tienari (1963a) and n = 1 case of Slater (1953) for which (left-)
handedness was specified for only one twin from each pair. This in­
creases further, to N = 560 (n = 141 MZAm; n = 184 MZAf; n = 89
DZAm; n = 146 DZAf), when also including other cases for which
handedness data are unavailable (see Table 1). Other possible cases of
same-sex twins reared apart were also identified from the literature but
not included in the current analysis; details of these and reasons for their
exclusion are provided below.
Sir Cyril Burt reported on a large sample of reared-apart twin pairs;
however, these twins are not considered here because the authenticity of
the data has been seriously questioned (for a discussion, see e.g., Tucker,
1997). The sample size increased across consecutive publications and
was reported on as follows: Burt, 1943, p. 91) (N = 15 pairs); Burt (1955,
p. 167–169) (N = 21 pairs); Burt (1958, p. 7) (“over 30 such cases”);
Conway (1958, p. 186) (N = 42); Burt (1966, p. 141) (N = 53 pairs).
Regardless, in the only mention of handedness, Burt (1966, p. 142)
stated that “Among our own cases we noted that mirror reversals
(opposite handedness, reversed patterns in finger-prints and
palm-prints, contrary directions in the crown whorl of the hair, etc.)
were nearly four times as frequent with monozygotic pairs as with
dizygotic”. From the available data it is therefore impossible to ascertain
concordance/discordance figures for this sample.
Although we do include in our review those cases in which zygosity
can be assumed with a reasonable degree of confidence but for which it
is less than certain, we did not consider the following cases further, as
zygosity information was missing completely. More specifically, there
are n = 3 pairs from Qingdao reported on by Gao et al. (2011) as being
present within the Chinese National Twin Registry, n = 2 male
twin-pairs from a triplet set in which one member was raised separately
to the other two (see Bordier and Futerman, 2014, p. 203), and n = 1
female twin-pair (Kathy and Betsy) for which no information on zygosity
was available (see Segal, 2021a, Chapter 5). Additionally, we are aware
of n = 6 cases of twins reared apart (separated at < 5 years of age) in the
Japanese Study of Adult Twins Reared Apart and Growing Old Sepa­
rately (Hayakawa et al., 2006), although we were unable to obtain
further details.
We do not consider the following cases further because details
regarding sex (and handedness) were not specified. Kallmann (1946)
reported on n = 59 MZA (Fig. 12., p. 317 in that paper) twin pairs who
were separated for more than five years (prior to onset of schizophrenia
3
Neuropsychologia 184 (2023) 108523
in the index case) – the number of DZA twin pairs included in the sample
is unclear – it is noted in the text that this “group of separated one-egg
pairs includes twins who developed schizophrenia at almost the same
time, although their separation took place soon after birth and led to
apparently very different life conditions.” (p. 316). Kallmann and Roth
(1956) also included data relating to four DZA twin pairs. Additionally,
based on a 1957 personal communication with Niels Juel-Nielsen,
Shields (1958, p. 116) stated that “a small series is also under investi­
gation by Professor Husén’s department in Stockholm.”; likewise, Con­
way (1958, p. 184) mentions, “In Stockholm Dr. Husen is carrying out an
analogous investigation.” However, further details of these twins appear
to be unavailable.
In addition to the n = 6 MZAm, n = 1 MZAf, and n = 2 DZAm “Twin
pairs who have been brought up in different homes” reported on by
Fukuoka (1937, p. 35), which are included in Table 1 of the current
paper, there is mention of n = 11 MZAm, n = 15 MZAf, n = 1 DZAm, and
n = 1 DZAf “Twin pairs who were brought up separately for some time,
then brought together and who have since been living in the same
home”. We do not include these latter twin pairs in our study because
degree of separation is unclear and appears likely to have been minimal.
We also note that Komai (1937, p. 4) mentioned, “HAGI’S material is the
same as FUKUOKA’S presented in this publication.” However, a Japanese
speaking colleague’s (Dr E. Self) reading of the paper by Hagi (1931)
revealed no mention of the study by Fukuoka (1937) or of reared-apart
twins. Additionally, Shields and Slater (1967, p. 581) noted that “In
Japan, Mitsuda (personal communication) has observed a series of MZ
twin pairs, in which one member had a schizophrenic or schizo-affective
illness; schizophrenia occurred in the co-twins in eight out of the 11
pairs brought up together, and in five out of the eight pairs brought up
apart.” It is unclear if these might be the same cases as reported on by
Inouye (1971, 1972), and so they are not considered separately within
our study. Likewise, Fisher (1958, p. 596) presented data, provided to
him by Eliot Slater, for n = 27 MZ twin pairs “separated at or shortly
after birth”, though additional details of this sample are unclear. Given
the chronicity, location, size of the sample, and the established collab­
oration between Eliot Slater and James Shields (Shields and Slater,
1967; Slater, 1953), it might be assumed that there is at least some
overlap between this sample and that of Shields (1962). Although un­
certain from the information available from these sources, given the
apparent likelihood of this being the case, we do not consider these data
separately here.
3.2. Hand preference of reared apart twins
Concordance/discordance rates for direction of hand preference in
MZA and DZA twin pairs identified from large studies, case series, and
case reports are presented in Table 2 and are summarized in Table 3. We
observed a slightly lower concordance rate in MZA (n = 206 pairs;
70.9% concordant, 29.1% discordant) than DZA (n = 209; 78.0%
concordant, 22.0% discordant) pairs, although this effect was not sta­
tistically significant χ2 (1) = 2.763, p = .096, φ = 0.082. However, the
direction of effect observed might relate to discordant handedness being
more striking in MZA than DZA pairs, an observation which could lead
to a greater likelihood of handedness being reported for the former than
the latter. Concordance was also slightly lower for both MZA and DZA
twins compared with the figures reported from the meta-analysis of
reared-together twins by Pfeifer et al. (2022) (MZT concordance =
80.49%; DZT concordance = 79.27%).
It is worth noting that handedness was not reported systematically
for the large cohort studied by Shields (1962), and it appears likely that
this was only noted when left-handedness was present in at least one
twin. Likewise, (left-) handedness for one twin, but no data for the other,
is reported for two MZAm pairs from Tienari (1963a) and one DZAm
pair from Slater (1953). We, therefore, imputed these missing values as
indicating right-handedness and present the aggregated data in Table 4.
As we had no relevant information from which to go by, we did not
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 1
Handedness data for MZA and DZA twin pairs from case reports.
Case Case no. in Original reference(s) Sex Zygosity Handedness Evidence Coding in
Farber (1981) reported in Table 2
primary source (current
study)

A and B N/A Townsend (circa 1874–1875, as cited M MZ Unknown

in Segal, 2012, p. 3–4)
a
Paul C. and Paul O. 118 Newman (1929a); Newman et al. M MZ RH/NRH “They are both right-handed” (Newman RH/NRH
(aka C and O) (1937, Case III) et al., 1937, p. 179); “Rl” (ibid, Table
43).
Ferdinand 418 Lange (1931, Concordant MZ pairs M MZ Unknown
Schweizer and Case 6)
Luitpold
Schweizer
a
Raymond and 144 Newman (1933); Newman et al. M MZ RH/RH “Both are right-handed” (Newman RH/RH
Richard (1937, Case VII) et al., 1937, p. 213); “RR” (ibid, Table
43).
a
Harold and Holden 320 Newman (1934a); Newman et al. M MZ LH/RH “Harold is left-handed; Holden, right- NRH/RH
(1937, Case IX) handed.” (Newman et al., 1937, p.
230); “LR” (ibid, Table 43).
g
Ronald and Dennis 232 Saudek (1934) M MZ RH/RH “Dextrality … Right-handed” [Ronald]; RH/RH
“Same” [Dennis] (Saudek, 1934, p. 18).
John and William 180 Rosanoff et al. (1935, Case 1); Farber, M MZ Unknown
Shields, and Craig (1979, as cited in
Farber, 1981)
A and B 412 Bouterwek (1936, Case E81); M MZ NRH/NRH “Sie geben an, die Linke nicht gerade zu NRH/NRH
g
Bouterwek (1943, Case E81) bevorzugen. Doch teilen beide die
Spielkarten mit der Linken aus; A
springt Stabhochsprung bei gegenüber
der normalen verkehrter Armstellung
besser und B spannt beim
Bogenschieẞen mit der Linken.” [They
state that they do not exactly favour the
left. But both deal the playing cards
with their left hands; A pole vaults
better than the normal inverted arm
position and B braces with his left hand
in archery.] (Bouterwek, 1936, p. 266);
“beide linkshändig” [both left handed]
(Bouterwek, 1943, p. 146); same
quotation as noted above from
Bouterwek (1936, p. 266) is reproduced
in Bouterwek (1943, p. 147).
A and B N/A Fukuoka (1937, MZm1) M MZ Unknown
A and B N/A Fukuoka (1937, MZm2) M MZ Unknown
A and B N/A Fukuoka (1937, MZm3) M MZ Unknown
A and B N/A Fukuoka (1937, MZm4) M MZ Unknown
A and B N/A Fukuoka (1937, MZm5) M MZ Unknown
A and B N/A Fukuoka (1937, MZm6) M MZ Unknown
a
Kenneth and Jerry 142 Newman et al. (1937, Case XIII) M MZ RH/RH “Both are right-handed in everything.” RH/RH
(Newman et al., 1937, p. 267); “RR”
(ibid, Table 43).
a
Edwin and Fred 182 Newman et al. (1937, Case XV) M MZ RH/RH “Both are fully right-handed.” ( RH/RH
Newman et al., 1937, p. 283); “RR”
(ibid, Table 43).
a
Gene and James 162 Newman et al. (1937, Case XVII) M MZ RH/LH “Gene is right-handed, while James is RH/NRH
partially left-handed.” (Newman et al.,
1937, p. 301); “Rl” (ibid, Table 43).
a
James and Reece 154 Newman et al. (1937, Case XVIII) M MZ RH/LH “Reece’s left arm is 2 inches shorter RH/NRH
than his right and is slightly deformed,
yet he is fully left-handed; while James
is definitely right-handed.” (Newman
et al., 1937, p. 309); “RL” (ibid, Table
43).
Alfred and August 414 Essen-Möller (1941, Case 3); Kaij M MZ Unknown
(1960, Case 19); Essen-Möller (1970,
Case N. 3)
g
J.J. and J.B. 106 Yates and Brash (1941) M MZ RH/RH “Reversed asymmetry or mirror- RH/RH
imagery of various features is
confirmatory but not necessary
evidence of monozygocity. J.J. and J.B.
did not appear to show this in any way;
certainly they did not in handedness
(both were right-handed)” (Yates and
Brash, 1941, p. 90).
Kazuo and Takau N/A Yoshimasu (1941, Case XVII) M MZ (?) Unknown
(continued on next page)

4
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 1 (continued )
Case Case no. in Original reference(s) Sex Zygosity Handedness Evidence Coding in
Farber (1981) reported in Table 2
primary source (current
study)

Millan and George 102 Stephens and Thompson (1943); M MZ RH/LH “George is left handed, while Millan is NRH/NRH
g
Stephens and Nunemaker (1950) right handed. Millan stated, however,
that he was originally left handed, but
when a youngster learned to use his
right hand because of an injury to his
left.” (Stephens and Thompson, 1943,
p. 111).
g
Philippe Joye and N/A Franceschetti (1948); Fraceschetti M MZ RH/RH “When going to sleep, both Ernstli and RH/RH
Charles Joye (né et al. (1948); McIndoe and Philippe suck their right thumbs” Joye
Ernstli Vatter) Franceschetti (1949); Joye (1954); (1954, p. 102); N. L. Segal (personal
(aka Victor J. and Segal (2009); Segal (2011a, p. communication with family members).
Eric V.) 40–53); Segal (2018, Case 1)
g
Earl and Frank 204 Burks and Roe (1949, Chapter 3) M MZ RH/RH “Both were righthanded.” (Burks and RH/RH
Roe, 1949, p. 17); Earl: “Right”; Frank:
“Right” (ibid, Table 5, p. 18).
James and Keith Group VI. Burks and Roe (1949, Chapter 5) M MZ (?) Not reported
(Doubtful
Zygosity)
c
Palle and Peter 108 Juel-Nielsen and Mogensen (1957); M MZ RH/RH “Both were right-handed.” ( RH/RH
Juel-Nielsen and Harvald (1958, Case Juel-Nielsen, 1980, Part 2, p. 25).
I); Mogensen and Juel-Nielsen
(1961a); Mogensen and Juel-Nielsen
(1961b); Mogensen and Juel-Nielsen
(1963); Juel-Nielsen (1965, Case I);
Juel-Nielsen (1980, Case I)
c
Kaj and Robert 206 Juel-Nielsen and Mogensen (1957); M MZ RH/RH “Both were right-handed.” ( RH/RH
Juel-Nielsen and Harvald (1958, Case Juel-Nielsen, 1980, Part 2, p. 139).
IV); Mogensen and Juel-Nielsen
(1961a); Mogensen and Juel-Nielsen
(1961b); Mogensen and Juel-Nielsen
(1963); Juel-Nielsen (1965, Case V);
Juel-Nielsen (1980, Case V)
g
A and B Group V Dencker (1958, Mz18) M MZ RH/RH “Left-handed”, “.” (Dencker, 1958, RH/RH
(Insufficient Table 35); note that although this only
Information) indicates the twins were alike for
handedness, it can be inferred from
elsewhere that they were both
right-handed as “The monozygotic
series contained 9 members who were
left-handed, and all of them had a
right-handed co-twin” (ibid, p. 100).
Peter and Bert 348 Kaij (1960, Case 17) M MZ Unknown
Philip and William Group V Kaij (1960, Case 18) M MZ Unknown
(Insufficient
Information)
Rudolph and Group VI Kaij (1960, Case 31) M MZ (?) Unknown
Edmund (Doubtful
Zygosity)
c
Viggo and Oluf 406 Mogensen and Juel-Nielsen (1961a); M MZ RH/RH “Both were right-handed.” ( RH/RH
Mogensen and Juel-Nielsen (1961b); Juel-Nielsen, 1980, Part 2, p. 285).
Mogensen and Juel-Nielsen (1963);
Juel-Nielsen (1965, Case XII);
Juel-Nielsen (1980, Case XII)
b
Alfred and Harry 174 Slater (1961, Case MZ 8); Shields M MZ RH/RH “Both right-handed.” (Slater, 1961, p. RH/RH
(1962, Case S m P 9) 375).
b
Herbert and 176 Slater (1961, Case MZ 11); Shields M MZ RH/RH “Both right-handed.” (Slater, 1961, p. RH/RH
Nicholas (1962, Case S m P 4); Gottesman and 376).
Shields (1972, Case MZ 17)
b
Richard and 236 Shields (1962, Case S m 1) M MZ Not reported RH/RH
Kenneth i
b
Bertram and 302 Shields (1962, Case S m 2) M MZ Not reported RH/RH
Christopher i
Russell and Tristram 126 Shields (1962, Case S m 3) M MZ LH (?)/LH “Both twins are left-handed.” (Shields, NRH/NRH
b
1962, p. 167). However, a hospital
report (from age 8) quoted (ibid, p. 166)
states Russell had “weakness of grip
and flexion of elbow on right side”; it is
stated elsewhere that an admission note
(age 14) mentioned he “had little use of
his right hand” (ibid, p. 166). He is also
noted to have had hemiplegia (this is
mentioned both in relation to the
admission note from age 14 [ibid, p.
(continued on next page)

5
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 1 (continued )
Case Case no. in Original reference(s) Sex Zygosity Handedness Evidence Coding in
Farber (1981) reported in Table 2
primary source (current
study)

166], as well as in a general comment

by the author [ibid, p. 167]).
b
Frederick and Peter 322 Shields (1962, Case S m 5) M MZ Not reported RH/RH
i

Foster and Francis i 324 Shields (1962, Case S m 6) M MZ Not reported RH/RH b

Rodney and Barry i 220 Shields (1962, Case S m 7) M MZ Not reported RH/RH b

Edward and Keith i 190 Shields (1962, Case S m 8) M MZ Not reported RH/RH b
b
William and Stanley 246 Shields (1962, Case S m 10) M MZ Not reported RH/RH
i

Timothy and Kevin i 308 Shields (1962, Case S m 11) M MZ Not reported RH/RH b

James and Robert i 228 Shields (1962, Case S m 12) M MZ Not reported RH/RH b

Patrick and Victor i 334 Shields (1962, Case S m 13) M MZ Not reported RH/RH b

Hubert and Brian i 336 Shields (1962, Case S m 14) M MZ Not reported RH/RH b
b
Benjamin and 328 Shields (1962, Case S m 15) M MZ Not reported RH/RH
Ronald i
A and B i N/A Shields (1962, Suppl. Case 2) M MZ Not reported RH/RH b
g
A and B Group V Hayashi (1963, Case X); Hayashi M MZ LH/RH “Bは右利きであるが,Aは左利き。” [“B NRH/RH
(Insufficient (1967, Case 10) is right-handed, while A is left-
Information) handed.”] (Hayashi, 1963, p. 163).
A and B Group V Hayashi (1963, Case XIII); Hayashi M MZ Unknown
(Insufficient (1967, Case 13)
Information)
A and B Group V Hayashi (1963, Case XIV); Hayashi M MZ Unknown
(Insufficient (1967, Case 14)
Information)
A and B Group V Hayashi (1963, Case XV); Hayashi M MZ Unknown
(Insufficient (1967, Case 15)
Information)
A and B 222 Tienari (1963a, Case 173); M MZ Not reported
Tienari (1963b); Tienari (1966, Case
173)
A and B h 354 Tienari (1963a, Case 466); M MZ RH (?)/LH “HISTORY OF B. Left-handed.” ( RH/NRH g

Tienari (1963b); Tienari (1966, Case Tienari, 1963a, p. 133).

466)
A and B N/A Tienari (1963a, Case 703); M MZ Not reported
Tienari (1963b); Tienari (1966, Case
703)
A and B h N/A Tienari (1963a, Case 989); M MZ RH (?)/LH “HISTORY OF B. Left-handed.” ( RH/NRH g

Tienari (1963b); Tienari (1966, Case Tienari, 1963a, p. 86).

989)
A and B 166 Tienari (1963a, Case 1080); M MZ Not reported
Tienari (1963b); Tienari (1966, Case
1080)
A and B 240 Kringlen (1964, Case V); Kringlen M MZ Unknown
(1967a, Case A); Kringlen (1967b,
Case 22)
A and B Group V Kringlen (1967a, Case C); Kringlen M MZ Unknown
(Insufficient (1967b, Case 66)
Information)
A and B N/A Kringlen (1967b, Case 50) M MZ Unknown
A and B m N/A Mitsuda and Sakai (1968, Case 16) M MZ Unknown
e
Tony Milsai and 202 Lindeman (1969) (also mentioned M MZ RH/RH ““Are you right-handed all the way? RH/RH
Roger Brooks k studies of these twins by Shields, and asked Tony.
by Machover [ibid, p. 201–202]) “I bat and throw righty,” Roger
answered, “but I kick lefty.”
“I do everything righty,” Tony said” (
Lindeman, 1969, p. 144).
A and B N/A Inouye (1971); Inouye (1972, Case M MZ Unknown
490)
A and B N/A Inouye (1971); Inouye (1972, Case M MZ Unknown
4143)
A and B N/A Inouye (1971); Inouye (1972, Case M MZ Unknown
4145)
A and B N/A Inouye (1971); Inouye (1972, Case M MZ Unknown
4155)
A and B N/A Inouye (1971); Inouye (1972, Case M MZ Unknown
4218)
A and B N/A Inouye (1971); Inouye (1972, no case M MZ Unknown
number, see p. 184–185)
Alois H. and Oskar 332 Prokop and Druml (1973) M MZ Unknown
H.
Twin 1 and Twin 2 N/A Sudarsky et al. (1983) M MZ Unknown
A and B N/A Gao et al. (2011, MZm1) M MZ Unknown
(continued on next page)

6
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 1 (continued )
Case Case no. in Original reference(s) Sex Zygosity Handedness Evidence Coding in
Farber (1981) reported in Table 2
primary source (current
study)

A and B N/A Gao et al. (2011, MZm2) M MZ Unknown

A and B N/A Gao et al. (2011, MZm3) M MZ Unknown
A and B N/A Gao et al. (2011, MZm4) M MZ Unknown
A and B N/A Gao et al. (2011, MZm5) M MZ Unknown
A and B N/A Gao et al. (2011, MZm6) M MZ Unknown
A and B N/A Gao et al. (2011, MZm7) M MZ Unknown
A and B N/A Gao et al. (2011, MZm8) M MZ Unknown
A and B N/A Gao et al. (2011, MZm9) M MZ Unknown
A and B N/A Gao et al. (2011, MZm10) M MZ Unknown
A and B N/A Gao et al. (2011, MZm11) M MZ Unknown
Matías and Marcos N/A Segal (2011a, p. 79–80); Segal (2018, M MZ Unknown
Case 6)
Dan Matthews and N/A Bordier and Futerman (2014, p. M MZ Unknown
Twin 194–196)
AT and LT j N/A Segal and Diamond (2014) M MZ RH/A N. L. Segal (personal communication). NRH/NRH
g
g
William and Jorge N/A Segal et al. (2017); Segal (2018, Case M MZ RH/RH “All four twins are right-handed, but RH/RH
7a); Segal and Montoya (2018); Segal not to the same degree. William is the
et al. (2018); Segal et al. (2019) strongest right-hander, followed by
Wilber, Jorge, and Carlos in that order,
with Carlos on the brink of
ambidexterity (Segal and Montoya,
2018, p. 187–188).
g
Wilber and Carlos N/A Segal et al. (2017); Segal (2018, Case M MZ RH/RH See quotation in cell above. RH/NRH
7b); Segal and Montoya (2018); Segal
et al. (2018); Segal et al. (2019)
d
Howard and Doug N/A Segal (2021a) M MZ RH/LH “The twins appear to be opposite- RH/NRH
handed.” (Segal, 2021a, p. 430); “The
twins in the one male set also appear to
be opposite-handed.” (ibid, p. 208
[Footnote 38 from p. 167]).
e
Robert Shafran and N/A Segal (2021a) M MZ LH/RH “The triplets’ hand preference scores NRH/RH
Edward Gelland varied somewhat–Bob is left-handed,
(triplets) k and Dave and Eddy are both right-
handed” (Segal, 2021a, p. 184);
“Unseen films of Bob as a youngster,
made by researchers during home visits
and retrieved from the Yale archive,
show him drawing designs, sorting
pictures, and building a block tower,
mostly using his left hand.” (ibid, p.
184); “Bob’s score indicated
left-handedness, whereas Eddy’s and
David’s scores indicated
right-handedness. Attempts were made
to shift Bob’s handedness from left to
right when he learned to write in
school, but they were unsuccessful
(ibid, p. 435 [Footnote 60 from p. 184]);
“All three identical females sets and the
male triplets in the twin study had a
left-handed member.” (ibid, p. 208).
e
Robert Shafran and N/A Segal (2021a) M MZ LH/RH See quotations in cell above. NRH/RH
David Kellman
(triplets) k
e
Edward Gelland and N/A Segal (2021a) M MZ RH/RH See quotations in cell two above. RH/RH
David Kellman
(triplets) k
Sir Mohamed Farah N/A Burley (2022) M MZ RH/? Handedness for writing determined
(né Hussein from pictures of Sir Mohamed Farah
Kahin) and signing autographs.
Hassan Kahin
g
A and B N/A Fernandes et al. (in preparation) M MZ RH/RH RH/RH
Christian Immerlich N/A Kranz (1936) M DZ Unknown
and Florian
Immerlich
Ottokar Immel and N/A Kranz (1936) M DZ Unknown
Roland Immel
A and B N/A Fukuoka (1937, DZm1) M DZ Unknown
A and B N/A Fukuoka (1937, DZm2) M DZ Unknown
John and Terry h N/A Slater (1953, Case 236) M DZ RH (?)/LH “Terry is left-handed.” (Slater, 1953, p. RH/NRH g

46).
(continued on next page)

7
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 1 (continued )
Case Case no. in Original reference(s) Sex Zygosity Handedness Evidence Coding in
Farber (1981) reported in Table 2
primary source (current
study)

A and B h N/A Shields (1962, Case Ds 1) M DZ LH/RH (?) “A, asthenic build, left-handed” ( NRH/RH b

Shields, 1962, p. 247).

A and B N/A Tienari (1966, Case 1011 [p. 22]) M DZ RH/? “In 1975, he replied that was right-
handed as a child and now writes with
his right-hand.” (J. Kaprio, personal
communication).
A and B N/A Gao et al. (2011, DZm1) M DZ Unknown
A and B N/A Gao et al. (2011, DZm2) M DZ Unknown
A and B N/A Gao et al. (2011, DZm3) M DZ Unknown
George Skrzynecky N/A Segal (2015) M DZ (?) Unknown
and Lucian
Poznanski
g
Greek twins N/A N. L. Segal (personal communication, M DZ (?) RH/RH (?) N. L. Segal (personal communication RH/RH
2022) with one twin).
Bessie and Jessie 116 Popenoe (1922); Muller (1925); F MZ RH/LH “Both twins, moreover, had early NRH/NRH
g
Saudek (1933) shown lefthandedness, though B had
been broken of it” (Muller, 1925, p.
438).
a
Alice and Olive (aka 210 Newman (1929b); Newman et al. F MZ LH/RH “Olive is right-handed and Alice is left- NRH/RH
A and O) (1937, Case I) handed.” (Newman et al., 1937, p.
165); “LR” (ibid, Table 43).
a
Eleanore and 208 Newman (1929c); Newman et al. F MZ RH/RH “Both are fully right-handed.” Newman RH/RH
Georgiana (aka E (1937, Case II) et al. (1937, p. 169); “RR” (ibid, Table
and G) 43).
Miriam and Mary Group V Wagenseil (1931) F MZ Unknown
(Insufficient
Information)
a
Mabel and Mary 318 Newman (1932a); Newman et al. F MZ RH/RH “Both are right-handed” (Newman RH/RH
(1937, Case IV) et al., 1937, p. 189); “RR” (ibid, Table
43).
a
Edith and Fay (aka B 158 Newman (1932b); Newman et al. F MZ RH/RH “both are fully right-handed” (Newman RH/RH
and D) (1937, Case V) et al., 1937, p. 198); “RR” (ibid, Table
43).
a
Ada and Ida 244 Newman (1932c); Newman et al. F MZ RH/RH “Both are right-handed” (Newman RH/RH
(1937, Case VI) et al., 1937, p. 206); “RR” (ibid, Table
43).
g
Anna and Corinne N/A Seeman and Saudek (1933, Case 1) F MZ RH/RH “Dextrality … Right-handed” [Anna]; RH/RH
“Same” [Corinne] (Seeman and Saudek,
1933, p. 271).
g
Leta and Lota N/A Seeman and Saudek (1933, Case 2) F MZ RH/RH “Dextrality … Right” [Leta]; “Same” RH/RH
[Lota] (Seeman and Saudek, 1933, p.
283).
a
Mildred and Ruth 150 Newman (1934b); Newman et al. F MZ LH/RH “Mildred is left-handed; Ruth, right- NRH/RH
(aka M and R) (1937, Case VIII) handed.” (Newman et al., 1937, p.
222); “LR” (ibid, Table 43).
A and B 234 Bouterwek (1936, Case E115); F MZ Not reported
Bouterwek (1943, Case E115)
A and B N/A Fukuoka (1937, MZf1) F MZ Unknown
a
Betty and Ruth 152 Newman et al. (1937, Case X) F MZ RH/A “Betty is fully right-handed, Ruth RH/NRH
distinctly ambidextrous.” (Newman
et al., 1937, p. 238); “AR” (ibid, Table
43).
a
Gladys and Helen 124 Newman et al. (1937, Case XI) F MZ RH/RH “Both are strongly right-handed.” ( RH/RH
Newman et al., 1937, p. 249); “RR”
(ibid, Table 43).
a
Thelma and Zelma 160 Newman et al. (1937, Case XII) F MZ A (LH?)/RH “Zelma is fully right-handed, and NRH/RH
Thelma partly left-handed or
ambidextrous.” (Newman et al., 1937,
p. 258); “AR” (ibid, Table 43).
a
Esther and Ethel 120 Newman et al. (1937, Case XIV) F MZ LH/RH “Ethel is fully right-handed, while NRH/RH
Esther is reported to be naturally left-
handed though trained to use the right
hand. Ethel has a short defective little
finger on her right hand, and Esther the
same deformity on her left hand.” (
Newman et al., 1937, p. 276); “Ethel is
much stronger in hand grip, registering
70 pounds with right hand and 68
pounds with left hand, while Esther
registered 62 pounds with right and 40
pounds with left hand.” (ibid, p. 275);
“It will be remembered that Esther is
reported to be naturally left-handed but
(continued on next page)

8
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 1 (continued )
Case Case no. in Original reference(s) Sex Zygosity Handedness Evidence Coding in
Farber (1981) reported in Table 2
primary source (current
study)

trained to use her right hand.” (ibid, p.

279); “LR” (ibid, Table 43).
Maxine and Virginia 164 Newman et al. (1937, Case XVI) F MZ RH/RH “RR” (Newman et al., 1937, Table 43). RH/RH a
a
Augusta and Helen 408 Newman et al. (1937, Case XIX) F MZ LH/RH “Helen is fully right-handed; Augusta NRH/RH
was originally left-handed but has
learned to use her right hand for most
monomanual operations.” (Newman
et al., 1937, p. 318); “RL” (ibid, Table
43).
Mary and Gertrude 122 Rosanoff et al. (1937, Case 91) F MZ Unknown
Agnes Emmaline N/A Rosanoff et al. (1937, Case 119) F MZ Unknown
and Madeline
Jennie
Kaete St. and Lisa St. 226 Kallmann (1938, p. 207–211) F MZ Unknown
g
Lois and Louise 138 Gardner and Newman (1940) F MZ RH/RH “Both are strongly right-handed” ( RH/RH
Gardner & Newman, 1940, p. 122).
Adelaide and 242 Burks (1942) F MZ A/A Twin A: “Writes from copy … Right … NRH/NRH
g
Beatrice Tosses ball … RRR”; Twin B: “Writes
from copy … Right … Tosses ball …
LLL” (Burks, 1942, Table I, p. 38); “It is
interesting to note that both twins
showed early left-handed tendencies
but that A was changed to the right
hand for all unimanual activities,
whereas B’s parents, on physician’s
advice, permitted her to use the left
hand, the change to the right hand for
writing being made voluntarily.” (ibid,
p. 40–41); Twin A: “Handedness … Left
tendency as baby. Mother tied left
hand; did not punish. A uses right
hand”; Twin B: “Handedness … Left
tendency as baby. Mother tied left hand
but stopped on doctor’s order. Later B
learned to write with right hand of own
accord; uses left otherwise” (ibid,
Table II p. 47).
Florence and Edith 134 Craike and Slater (1945) F MZ Unknown
g
Clara and Doris 212 Burks and Roe (1949, Chapter 2) F MZ RH/A Clara: “Right”; Doris: “Right and left; RH/NRH
can use left to sew” (Burks and Roe,
1949, Table 1, p. 5).
g
Gertrude and Helen 186 Burks and Roe (1949, Chapter 4) F MZ RH/RH Gertrude: “Right”; Helen: “Right” ( RH/RH
Burks and Roe, 1949, Table 8, p. 31).
Esther and Elvira 330 Schwesinger (1952) F MZ Unknown
A and B 156 Stenstedt (1952, Case 106 [p. 74–75]) F MZ Unknown
A and B l 404 Kallmann and Roth (1956) F MZ Unknown
c
Olga and Ingrid 104 Juel-Nielsen and Mogensen (1957); F MZ RH/RH “Both were right-handed.” ( RH/RH
Juel-Nielsen and Harvald (1958, Case Juel-Nielsen, 1980, Part 2, p. 57).
II); Mogensen and Juel-Nielsen
(1961a) (?); Mogensen and
Juel-Nielsen (1961b); Mogensen and
Juel-Nielsen (1963); Juel-Nielsen
(1965, Case II); Juel-Nielsen (1980,
Case II)
c
Maren and Jensine 146 Juel-Nielsen and Mogensen (1957); F MZ RH/RH Jensine: “When she commenced school, RH/NRH
Mogensen and Juel-Nielsen (1961a) she had a pronounced tendency to use
(?); Mogensen and Juel-Nielsen her left hand for writing and drawing
(1961b); Mogensen and Juel-Nielsen but this was quickly corrected by her
(1963); Juel-Nielsen (1965, Case III); teacher, and since then she had felt
Juel-Nielsen (1980, Case III) naturally right-handed.” (Juel-Nielsen,
1980, Part 2, p. 74); “Both were
right-handed.” (ibid, Part 2, p. 77).
c
Ingegerd and 188 Juel-Nielsen and Mogensen (1957); F MZ RH/RH “Both were right-handed.” ( RH/RH
Monika Juel-Nielsen and Harvald (1958, Case Juel-Nielsen, 1980, Part 2, p. 106).
III); Mogensen and Juel-Nielsen
(1961a) (?); Mogensen and
Juel-Nielsen (1961b); Mogensen and
Juel-Nielsen (1963); Juel-Nielsen
(1965, Case IV); Juel-Nielsen (1980,
Case IV)
c
Martha and Marie 168 Juel-Nielsen and Mogensen (1957); F MZ RH/RH “Both were right-handed.” ( RH/RH
Mogensen and Juel-Nielsen (1961a) Juel-Nielsen, 1980, Part 2, p. 158).
(?); Mogensen and Juel-Nielsen
(continued on next page)

9
G. Richards and N.L. Segal
Table 1 (continued )
Case Case no. in Original reference(s)
Farber (1981)
(1961b); Mogensen and Juel-Nielsen
(1963); Juel-Nielsen (1965, Case VI);
Juel-Nielsen (1980, Case VI)
Kamma and Ella 114 Juel-Nielsen and Mogensen (1957);
Juel-Nielsen and Harvald (1958, Case
V); Mogensen and Juel-Nielsen
(1961a) (?); Mogensen and
Juel-Nielsen (1961b); Mogensen and
Juel-Nielsen (1963); Juel-Nielsen
(1965, Case VII); Juel-Nielsen (1980,
Case VII)
Signe and Hanne 130 Juel-Nielsen and Mogensen (1957);
Juel-Nielsen and Harvald (1958, Case
VI); Mogensen and Juel-Nielsen
(1961a) (?); Mogensen and
Juel-Nielsen (1961b); Mogensen and
Juel-Nielsen (1963); Juel-Nielsen
(1965, Case VIII); Juel-Nielsen (1980,
Case VIII)
Karin and Kristine 178 Juel-Nielsen and Mogensen (1957);
Mogensen and Juel-Nielsen (1961a)
(?); Mogensen and Juel-Nielsen
(1961b); Mogensen and Juel-Nielsen
(1963); Juel-Nielsen (1965, Case IX);
Juel-Nielsen (1980, Case IX)
Petrine and Dorthe 184 Juel-Nielsen and Harvald (1958, Case
VII); Mogensen and Juel-Nielsen
(1961a) (?); Mogensen and
Juel-Nielsen (1961b); Mogensen and
Juel-Nielsen (1963); Juel-Nielsen
(1965, Case X); Juel-Nielsen (1980,
Case X)
Astrid and Edith 170 Juel-Nielsen and Harvald (1958, Case
VIII); Mogensen and Juel-Nielsen
(1961a) (?); Mogensen and
Juel-Nielsen (1961b); Mogensen and
Juel-Nielsen (1963); Juel-Nielsen
(1965, Case XI); Juel-Nielsen (1980,
Case XI)
A and B 340 Kurihara (1959, 17-A and 17-B [also
referred to as MZ-4192]);
Inouye (1971); Inouye (1972, Case
4192)
Jessie and Winifred 238 Shields (1962, Case S f 1)
i
Neuropsychologia 184 (2023) 108523
Sex Zygosity Handedness Evidence Coding in
reported in Table 2
primary source (current
study)
F MZ A/A “V female 49 … 2) hemicrania left- NRH/NRH
c
handed” (Juel-Nielsen and Harvald,
1958, Table 1, p. 58); “In two of the
pairs of twins, one of the twins (V2 and
VII1) was ambidextrous and probably
originally left-handed.” (ibid, p. 63);
Kamma: “She stressed that she was
trained not to prefer writing with her
left hand and considered that she
originally had been left-handed. Since
leaving school she had used her right
hand in a natural manner.” (
Juel-Nielsen, 1980, Part 2, p. 169); Ella:
“She was left-handed, was taught to
write with her right hand but continued
to prefer her left when for example she
used an ax or saw, was rather clumsy in
some types of work, and was quite
certain that she was left-handed.” (ibid,
Part 2, p. 173); “Both were
ambidextrous, Ella probably
predominantly lefthanded.” (ibid, Part
2, p. 177).
c
F MZ RH/RH “Both were right-handed.” ( RH/RH
Juel-Nielsen, 1980, Part 2, p. 207).
c
F MZ RH/RH “Both twins were right-handed.” ( RH/RH
Juel-Nielsen, 1980, Part 2, p. 224).
F MZ RH/A “VII female 71 1) left-handed” ( RH/NRH c
Juel-Nielsen and Harvald, 1958,
Table 1, p. 58); “In two of the pairs of
twins, one of the twins (V2 and VII1)
was ambidextrous and probably
originally left-handed.” (ibid, p. 63);
Dorthe: “She believed that she had been
left-handed as a child and preferred
using her left hand, but at school she
was forced to use her right hand. She
still wrote with her right hand but used
her left hand when working with a
hammer or throwing, and she felt that if
she lost her right hand, she “would
quickly be able to use the left”.” (
Juel-Nielsen, 1980, Part 2, p. 244);
“Petrine was right-handed, Dorthe
ambidextrous, probably originally
left-handed.” (ibid, Part 2, p. 247).
c
F MZ RH/RH “Both were right-handed.” ( RH/RH
Juel-Nielsen, 1980, Part 2, p. 268).
F MZ Unknown
b
F MZ Not reported RH/RH
(continued on next page)
10
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 1 (continued )
Case Case no. in Original reference(s) Sex Zygosity Handedness Evidence Coding in
Farber (1981) reported in Table 2
primary source (current
study)

A and B h 326 Shields (1962, Case S f 2) F MZ RH (?)/LH “B is taller but lighter in weight and has RH/NRH b

had more illness; she is left-handed.” (

Shields, 1962, p. 192).
Valerie and Joyce h 338 Shields (1962, Case S f 3) F MZ LH/RH (?) “Valerie’s periods started a month or NRH/RH b

two before Joyce’s and she has usually

been bigger; she is left-handed.” (
Shields, 1962, p. 192).
A and B i 402 Shields (1962, Case S f 4) F MZ Not reported RH/RH b

Megan and Polly i 172 Shields (1962, Case S f 5) F MZ Not reported RH/RH b

A and B i 140 Shields (1962, Case S f 6) F MZ Not reported RH/RH b

b
Jenny and Kathleen 304 Shields (1962, Case S f 7) F MZ Not reported RH/RH
i

Olive and Madge i 112 Shields (1962, Case S f 8) F MZ Not reported RH/RH b
b
Madeline and Lilian 110 Shields (1962, Case S f 9) F MZ RH (?)/LH “Lilian is taller and heavier and is left- RH/NRH
h
handed.” (Shields, 1962, p. 201).
b
Marjorie and Norah 128 Shields (1962, Case S f 10) F MZ LH/RH “Marjorie went to a convent school NRH/RH
h
(family not R.C.) where she was a
boarder from 10 to 17½. Though
naturally left-handed she was taught to
write with her right hand.” (Shields,
1962, p. 203). However, in the
following section (without prior
mention of Norah’s handedness) it is
stated “As already mentioned Norah is
left-handed, except for writing.” (ibid,
p. 203). This is interpreted as being a
typo, and so this pair is coded as LH/RH
rather than RH/LH.
Molly and Dorothy i 416 Shields (1962, Case S f 11) F MZ Not reported RH/RH b
Pauline and Sally i 410 Shields (1962, Case S f 12) F MZ Not reported RH/RH b
Viola and Olga i 344 Shields (1962, Case S f 13) F MZ Not reported RH/RH b
b
Millicent and Edith 148 Shields (1962, Case S f 14) F MZ RH (?)/LH “Edith is left-handed.” (Shields, 1962, RH/NRH
h
p. 213).
Joan and Dinah i 352 Shields (1962, Case S f 15) F MZ Not reported RH/RH b

June and Clara i 224 Shields (1962, Case S f 16) F MZ Not reported RH/RH b
b
Jacqueline and 230 Shields (1962, Case S f 17) F MZ Not reported RH/RH
Beryl i
Christine and Nina i 312 Shields (1962, Case S f 18) F MZ Not reported RH/RH b

Herta and Berta i 136 Shields (1962, Case S f 19) F MZ Not reported RH/RH b
b
Charlotte and Laura 306 Shields (1962, Case S f 20) F MZ Not reported RH/RH
i
h b
Mary and Nancy 218 Shields (1962, Case S f 21) F MZ LH/RH (?) “Mary is left-handed.” (Shields, 1962, NRH/RH
p. 227).
b
Olwen and Gwladys 214 Shields (1962, Case S f 22) F MZ Not reported RH/RH
i

Annie and Trixie i 314 Shields (1962, Case S f 23) F MZ Not reported RH/RH b

Joanna and Isobel i 216 Shields (1962, Case S f 24) F MZ Not reported RH/RH b

Odette and Fanny i 342 Shields (1962, Case S f 25) F MZ Not reported RH/RH b

Amy and Teresa i 132 Shields (1962, Case S f 26) F MZ Not reported RH/RH b

Dora and Brenda i 350 Shields (1962, Case S f 27) F MZ Not reported RH/RH b

Maisie and Vera i 346 Shields (1962, Case S f 28) F MZ Not reported RH/RH b
b
Adeline and 422 Shields (1962, Case S f 29) F MZ Not reported RH/RH
Gwendolen i
A and B i N/A Shields (1962, Suppl. Case 1) F MZ Not reported RH/RH b

A and B 316 Kringlen (1967a, Case B); Kringlen F MZ (?) Unknown

(1967b, Case 48)
A and B 420 Kringlen (1967a, Case D); Kringlen F MZ Unknown
(1967b, Case 55)
A and B 310 Inouye (1971); Inouye (1972, Case F MZ Unknown
4138)
A and B N/A Inouye (1971); Inouye (1972, Case F MZ Unknown
4223)
A and B N/A Inouye (1971); Inouye (1972, Case F MZ Unknown
4317)
A and B N/A Chess, Ladimer, and Thompson F MZ Unknown
(unpublished, as cited in Thomas and
Chess, 1977)
d
Paula and Elyse N/A Schein and Bernstein (2007); Segal F MZ RH/RH “Elyse and I are both right-handed” ( RH/RH
(2021a) Schein and Bernstein, 2007, p. 63);
“They are both right-handed.” (Segal,
2021a, p. 271).
g
US and SK N/A Segal and Hur (2008) F MZ RH/RH Crovitz-Zener Self-Report Handedness RH/RH
Inventory: “Both twins were right-
(continued on next page)

11
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 1 (continued )
Case Case no. in Original reference(s) Sex Zygosity Handedness Evidence Coding in
Farber (1981) reported in Table 2
primary source (current
study)

handed, although SK scored slightly

higher (27) than her twin sister (23),
indicating that she was less strongly
right-handed. Their handedness scores
place them both within the range
including 98% of right-handers” (Segal
and Hur, 2008, p. 544).
g
Begoña and Delia N/A Segal and Blandón-Gitlin (2010); F MZ RH/RH N. L. Segal (personal communication). RH/RH
Segal (2011a); Segal (2018, Case 5)
A and B N/A Gao et al. (2011, MZf1) F MZ Unknown
A and B N/A Gao et al. (2011, MZf2) F MZ Unknown
A and B N/A Gao et al. (2011, MZf3) F MZ Unknown
A and B N/A Gao et al. (2011, MZf4) F MZ Unknown
A and B N/A Gao et al. (2011, MZf5) F MZ Unknown
A and B N/A Gao et al. (2011, MZf6) F MZ Unknown
A and B N/A Gao et al. (2011, MZf7) F MZ Unknown
g
Nora Laster and N/A Segal (2011a, p. 26–27) F MZ RH/RH N. L. Segal (personal communication). RH/RH
Chun Hei
g
Kasia and Edyta N/A Segal (2011a, p. 59–66); Segal (2018, F MZ RH/LH “RL—MZAf (Poland, left twin was RH/NRH
Case 3) originally left, then some switch by
teachers” (N. L. Segal, personal
communication with parent).
Mari Tairí and Tairí N/A Segal (2011a, p. 67–78); Segal (2018, F MZ Unknown
Mari Case 4a)
g
Dorothy Loader and N/A Segal (2011b); Glasby et al. (2013) F MZ LH/LH “Dorothy is left-handed and I am too. I RH/NRH
Rosabelle (Belle) do recall however my foster Mum
Glasby telling me that when I was young that
my foster Dad held down my right hand
purposely so that I learned to write with
my left hand. I have no idea why he did
this and I didn’t think to ask. Out of
interest, I use scissors in my right hand
but I think that Dorothy still uses her
left hand for that purpose too.” (N. L.
Segal, personal communication with
Rosabelle Glasby).
AH and BH N/A Segal (2012, p. 266–267) F MZ Unknown
g
Samantha Futerman N/A Bordier and Futerman (2014); Segal F MZ RH/RH Anaïs: “Although I was ambidextrous, I RH/NRH
and Anaïs Bordier and Cortez (2014); Segal (2016) tended to use my left hand, but the
(aka US and FR) French educators were trained to
encourage us to be right-handed.” (
Bordier and Futerman, 2014, p. 61);
Samantha: “We were both
right-handed, although Anaïs could use
both hands.” (ibid, p. 282); Samantha:
“I am right-handed, and Anaïs is
ambidextrous. She only uses her right
hand about a quarter of the time. I don’t
know if Dr. Segal’s testing considered
that Anaïs had been encouraged to use
her right hand in school, part of French
educational aversion to
left-handedness. When she broke her
right arm riding her horse at age fifteen,
she had no choice but to write with her
left hand. Anyway, she is considered
ambidextrous, and I am right-handed.”
(ibid, p. 285); “Both twins were
right-handed, although FR uses both
hands. She had broken her right arm at
age fifteen, and used her left during
recovery” (Segal and Cortez, 2014, p.
102).
Siam and Fabienne N/A Segal (2014, Case 2) F MZ Unknown
g
US and SK N/A Segal et al. (2015a) F MZ RH/RH “Both SK and US use their right hand RH/RH
exclusively for unimanual tasks such as
writing and cutting” (Segal et al.,
2015a, p. 298).
f
Audrey and Gracie N/A Segal (2017) F MZ LH/RH “Audrey is left-handed and Gracie is NRH/RH
right-handed.” (Segal, 2017, p. 181).
d
Anne and Susan N/A Segal (2021a) F MZ LH/RH “Susan is right-handed and Anne is left- NRH/RH
handed.” (Segal, 2021a, p. 127).
(continued on next page)

12
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 1 (continued )
Case Case no. in Original reference(s) Sex Zygosity Handedness Evidence Coding in
Farber (1981) reported in Table 2
primary source (current
study)
d
Melanie and Ellen N/A Segal (2021a) F MZ LH/RH “Melanie is left-handed and Ellen is NRH/RH
right-handed.” (Segal, 2021a, p. 139);
“they knew that they were
opposite-handed. Melanie, the
left-handed twin, had a longer left arm
and Ellen, the right-handed twin, had a
longer right arm.” (ibid, p. 147); “All
three identical females sets and the
male triplets in the twin study had a
left-handed member.” (ibid, p. 208).
d
Sharon and Lisa N/A Segal (2021a) F MZ LH/RH “Sharon is left-handed and Lisa is right- NRH/RH
handed.” (Segal, 2021a, p. 208); “All
three identical females sets and the
male triplets in the twin study had a
left-handed member.” (ibid, p. 208).
g
Dolores and Mirta N/A Segal (2021b, Case 1) F MZ RH/RH “Both twins are right-handed.” (Segal, RH/RH
2021b, p. 245).
g
Linh (Nugyen Thi N/A Segal (2021b, Case 2) F MZ RH/RH “Both twins are right-handed.” (Segal, RH/RH
Hoai Linh) and 2021b, p. 246).
Alice Cecilia Linh
Svensson
g
Elpis and Maria N/A Segal (2022) F MZ RH/RH “physical similarities, including right- RH/RH
hand preference” (Segal, 2022, p. 167)
g
SK and US N/A Segal and Hur (2022) F MZ RH/RH N. L. Segal (personal communication). RH/RH
g
A and B (born in N/A Segal (unpublished Case 1) F MZ RH/RH “RH/RH.” (N. L. Segal, personal RH/RH
Malaysia, raised communication).
in Singapore)
A and B N/A Segal (unpublished Case 2) F MZ LH/LH “some intentional switching by others NRH/NRH
g
later in life” (N. L. Segal, personal
communication).
g
A and B (born in N/A N. L. Segal (personal communication) F MZ RH/RH N. L. Segal (personal communication). RH/RH
South Korea,
raised in the US)
g
A and B N/A N. L. Segal (personal communication) F MZ RH/LH RH/NRH
Martha and Lydia N/A Slater (1953, Case 52) F DZ Unknown
Esther and Rachel N/A Slater (1953, Case 225) F DZ (?) LH/LH “This is the only pair where it is NRH/NRH
g
reported that both twins are left-
handed.” (Slater, 1953, p. 46); “Both
are left-handed.” (ibid, p. 334).
A and B N/A Juel-Nielsen and Harvald (1958, p. F DZ Unknown
63)
A and B h N/A Shields (1962, Case Ds 2) F DZ LH/RH (?) “A (like mother and aunt) has had NRH/RH b

migrainous headaches from age of 3,

liable to occur when excited; also a dry
skin conditions (familial), left-handed.”
(Shields, 1962, p. 247).
A and B i N/A Shields (1962, Case Ds 3) F DZ Not reported RH/RH b

A and B i N/A Shields (1962, Case Ds 4) F DZ Not reported RH/RH b

A and B i N/A Shields (1962, Case Ds 5) F DZ Not reported RH/RH b

A and B i N/A Shields (1962, Case Ds 6) F DZ Not reported RH/RH b

A and B i N/A Shields (1962, Case Ds 7) F DZ Not reported RH/RH b

A and B i N/A Shields (1962, Case Ds 8) F DZ Not reported RH/RH b

A and B i N/A Shields (1962, Case Ds 9) F DZ Not reported RH/RH b

A and B i N/A Shields (1962, Case Ds 10) F DZ Not reported RH/RH b

A and B i N/A Shields (1962, Case Ds 11) F DZ Not reported RH/RH b

A and B N/A Gao et al. (2011, DZf1) F DZ Unknown

A and B N/A Gao et al. (2011, DZf2) F DZ Unknown
g
Jennifer and N/A Segal (2011a, p. 67–78); Segal (2018, F DZ (?) Not reported N. L. Segal (personal communication). RH/RH
Samantha Case 4b)
g
US and UK N/A Segal et al. (2015b) F DZ RH/RH Crovitz and Zener Group Test for RH/RH
Assessing Hand and Eye Dominance:
“US was right-handed (score = 15),
while UK showed mixed handedness
(score = 34). UK, originally right-
handed, had broken her right wrist in
2010 and used her left hand during
recovery. However, she used her right
hand to place pegs in the dominant
hand condition.” (Segal et al., 2015b, p.
113).
d
Michele and Allison N/A Segal (2021a) F DZ RH/RH N. L. Segal (personal communication). RH/RH
Paula and Marjorie N/A Segal (2021a) F DZ Unknown
BL and Twin N/A N. L. Segal (personal communication) F DZ (?) Unknown “RH-/RH; BUT BOTH PARTLY AMBI” NRH/NRH
g
(N. L. Segal, personal communication).

13
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Note. Other than the MZAm triplets, Robert Shafran, David Kellman, and Edward Gelland (who were also included in the Louise Wise Services [LWS]/Child
Development Center [CDC] Twin Study) and Tony and Roger (who are described in the book by Lindeman [1969]), case reports relating to twins in the MISTRA are not
included here (see Segal [2012] e.g., for Jim Lewis and Jim Springer, Oskar Stöhr and Jack Yufe, Dorothy and Bridget, Barbara and Judy, Mark Newman and Jerry
Levey, Jennifer Mitchell and Margaret Williams; Segal [2011a, p. 53–59, 2018, Case 2] for George and Brent; Bordier and Futerman [2014, p. 132–133] for DZAf pair
Kerrie and Amy).
a
Newman et al. (1937).
b
Shields (1962).
c
Juel-Nielsen (1965, 1980).
d
LWS/CDC Twin Study.
e
Minnesota Study of Twins Reared Apart (MISTRA).
f
Fullerton Study of Chinese Twins Adopted Apart.
g
Case reports.
h
Left-handedness was noted for one twin, but data were not reported for the other twin (assumed right-handed). Note that a preliminary report from Tienari’s
(1963b, p. 395) study stated, “In cases where one of the twins was left-handed and the other right-handed, dexterity tests were used as a check on this circumstance.”
i
Handedness not reported for this case, but left-handedness was noted in other cases in the same series (both twins assumed right-handed).
j
This pair is included in Table 2 as an MZAm pair; although assigned male at birth, both twins later assumed female gender identities.
k
Twin pairs included in Table 2 as aggregated data for the MISTRA rather than as case reports.
l
The sex of this pair is reported as female by Farber (1981, Table 3.1, p. 58–59).
m
It is unclear whether this pair is also included as one of the cases reported on by Inouye (1972), as it is stated in that publication that “Some pairs were also included
in a study by Mitsuda and Sakai (1968), who kindly permitted us to look through the case records. The present author also provided Dr. Mitsuda case records to avail in
his study.” (Inouye, 1972, p. 183).

impute values for the two cases (one MZAm [Burley, 2022]; one DZAm
[Tienari, 1966, Case 1011]) in which right-handedness was established
for one twin, but no details were available for the other. Concordance
rates were fairly similar to our original estimates: MZA (n = 251 pairs;
72.9% concordant, 27.1% discordant), DZA (n = 221; 77.8% concor­
dant, 22.2% discordant), χ2 (1) = 1.526, p = .217, φ = 0.057.
4. Study 2: Strength of hand preference and relative hand skill
in the Minnesota Study of Twins Reared Apart (MISTRA)
Strength of handedness, while an important manifestation of hand
preference, has received relatively little attention in the scientific liter­
ature, especially in studies of twins reared apart. Study 2, therefore,
examined this variable as part of the relevant data gathered in the
Minnesota Study of Twins Reared Apart (MISTRA), conducted by Bou­
chard et al. (1990). The rearing environments of the MISTRA twins are
described briefly below, with more detailed description of participants,
zygosity diagnosis, tests, inventories, findings, and conclusions provided
by Segal (2012).
5. Materials and methods
5.1. Participants
Reared-apart twins in the MISTRA had to have been separated by
four years of age and to have spent their formative childhood years
apart. The mean age at separation was 218.21 days (SD = 343.59) and
ranged between 0 and 1644 days. None were raised in seriously
impoverished homes, coming mostly from lower-middle class to middle
class families. The majority of twins (70.2%) were reared by unrelated
individuals, whereas the remainder were variously reared by biological
relatives or multiple biological relatives, with some raised in “complex”
situations. More mothers and fathers of MZA than DZA twins pursued
professional than non-professional occupations. Most family environ­
mental variables showed little correlation with twins’ measures of in­
telligence and other behaviours.
As part of the standard test battery, twins completed the Crovitz-
Zener Handedness Inventory (CZHI) (Crovitz and Zener, 1962), a
self-report assessment of hand preference, and the Purdue Pegboard
(PPB) task (Tiffin, 1968), a behavioural measure of hand skill. Hand­
edness data (scores from at least one of the following: CZHI, PPB
right-hand speed, or PPB left-hand speed) were available for the mem­
bers of 81 MZA twin pairs (35 male pairs [43.2%]; 46 female pairs
[56.8%]) and 55 DZA twin pairs (12 male pairs [21.8%]; 26 female pairs
[47.3%]; 17 [30.9%] male-female pairs). Within these data are three
triplet sets: two sets consist of three MZA males (i.e., three MZA twin
pairs each), whereas the other set consists of a pair of MZA female twins
and their DZA male co-triplet (organised into one MZA female pair, and
two DZA male-female pairs). The age range of the N = 136 twin pairs
included in the current study was 12–77 years (M = 41.87, SD = 13.93).
5.2. Zygosity diagnosis
When the MISTRA began in 1979, twins provided blood samples that
were analyzed across multiple serological systems by the Minneapolis
War Memorial Blood Bank. These data in combination with dermato­
glyphic measures, ponderal index and cephalic index yielded a proba­
bility of less than .001 (Lykken, 1978). As genetic technology advanced
in later years, zygosity diagnosis was accomplished by DNA testing.
Additional details about this process are reported by Segal (2012).
5.3. Measures
5.3.1. Strength of hand preference
The Crovitz-Zener Handedness Inventory (CZHI; Crovitz and Zener,
1962) is a self-report measure of hand preference. The current study
employed a modified version of the original measure for which scores
were reported from 1 (left) to 9 (right), with those further from 0 indi­
cating stronger hand preference. Analyses presented here are based on
14 items: preferred hand used for writing, holding a nail while
hammering, throwing a ball, holding a bottle when removing the top,
drawing, peeling a potato, cutting with scissors, using a knife, holding a
needle when threading, holding a glass when drinking, holding a
toothbrush, holding a dish when wiping, and using a tennis racquet.
Note that five items (nail, bottle, peeling potato, needle, dish) are
reverse scored, as it is the non-dominant hand that is specified in their
response. Items from the CZHI are known to correlate strongly with
similar items from the more widely used Edinburgh Handedness In­
ventory (Oldfield, 1971), r = .80–.90 (Bryden, 1977), and internal
consistency in the current study was high, Cronbach’s α = 0.942.
To facilitate calculation of a laterality index (LI), scores for each of
the 14 CZHI items were recoded as − 4 (left) to +4 (right), with 0 indi­
cating no preference. LI was then calculated as: (R - L)/(R + L) × 100. In
this formula, R represents the sum of positive values, and L represents
the sum of (absolute) negative values. Scores therefore range from
− 100, which indicates left preference (including items with no prefer­
ence) for all items, to +100, which indicates right preference (including
items with no preference) for all items. To examine strength of hand
preference (strong or weak), we removed the sign for the LI so that all
values were positive. Scores closer to 100 therefore indicate stronger

14
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 2
Concordance/discordance for direction of hand preference in monozygotic (MZA) and dizygotic (DZA) twins reared apart.
Male Female

Study/cohort Measure(s) MZA DZA MZA DZA

RH/ NRH/ RH/ RH/ NRH/ RH/ RH/ NRH/ RH/ RH/ NRH/ RH/
RH NRH NRH RH NRH NRH RH NRH NRH RH NRH NRH

Newman et al. (1937) a Unspecified 3 0 4 0 0 0 6 0 6 0 0 0

Shields (1962) b Unspecified 15 1 0 0 0 1 24 0 6 9 0 1
Juel-Nielsen (1965, 1980) Unspecified 3 0 0 0 0 0 6 1 2 0 0 0
Louise Wise Services/Child Unspecified 0 0 1 0 0 0 1 0 3 2 0 0
Development Center (LWS/
CDC) Twin Study (Segal,
2021a) c
Minnesota Study of Twins Writing hand preference 20 2 8 7 0 4 28 2 9 18 0 4
Reared Apart (MISTRA) (from Crovitz-Zener
(current study) d Handedness Inventory)
Fullerton Study of Chinese Unspecified 0 0 1 1 0 0 7 2 5 0 1 3
Twins Adopted Apart (Segal
et al., 2011)
Finnish Twin Cohort Study ( Child handedness (right, 7 0 2 32 1 4 7 0 5 31 0 10
Kaprio et al., 2019) left, both)
Swedish Adoption Twin Study Dominant hand (right, 9 0 4 21 0 7 12 0 3 42 2 14
of Aging (Pedersen, 2015) left, both)
Case studies (see Table 1) Varied 7 3 4 1 0 1 14 3 5 2 2 0
Total (all data) 64 6 24 62 1 17 105 8 44 104 5 32
Total (excluding missing/ 51 6 22 62 1 15 81 8 38 95 5 31
incomplete data)

Note. More detailed handedness data for case reports are provided in Table 1. ‘Total (all data)’ aggregates all available data, including cases from Shields (1962) for
which handedness was not reported, as well as two cases from Tienari (1963a) and one case from Slater (1953) that reported (left-) handedness for only one twin from
each pair (all missing values from those cases have been inferred to indicate right-handedness); ‘Total (excluding missing/incomplete data)’ includes all available data
other than these cases.
a
Handedness data for all twin-pairs studied by Newman et al. (1937) are reported in Table 43 of that publication. However, these data are not entirely consistent
with those presented in the individual case reports (see Table 1). In particular, Case III is reported as “Rl” in Table 43 of Newman et al. (1937), but in the text of the case
report it is specified that both twins are right-handed (RH). Where non-right-handedness (NRH) is indicated (in Table 43 and/or the text of the case report of Newman
et al., [1937]), we have coded such cases as NRH.
b
Handedness was not systematically reported by Shields (1962), although is mentioned in case reports for 9 of the 57 twin-pairs. Within these case reports, it is
specified that 10 (8.8%), or possibly 11 (9.6%; see Case S f 10), of the 114 twins were left-handed. As this is very close to the overall 9.13% prevalence estimate for
left-handedness in reared-together twins reported by a recent meta-analysis (Pfeifer et al., 2022), it is conceivable that handedness had only been noted in the text in
cases of non-right-handedness. Further support for this idea is provided in that Slater (1961) reported on handedness for two of the pairs later studied by Shields (1962)
(but on which Shields did not specify handedness), and in each case both twins were right-handed.
c
Handedness data from the MZAm triplet set included as part of the LWS/CDC study are not included here because they are also included within the data from the
MISTRA. Additionally, Abrams (1986) reported on an MZA female pair referred to pseudonymously as ‘Amy and Beth’ (for Amy “it was reported that she was definitely
right-handed.” [p. 44]; Beth “became left-handed.” [p. 49]); it is unclear to which pair this corresponds, and these data are not included here. Likewise, Abrams and
Neubauer (1994) referred pseudonymously to one MZAf pair (‘Abby and Amy’) and one MZAm pair (‘Barry and Ben’), though did not comment on their handedness.
d
Handedness data for the MISTRA have also been presented by Segal (2012, p. 294–295). Table 13–1 of that publication (p. 294) reports the following: MZA twins
(n = 74 pairs): Right-Left, 15%; Right-Mixed, 8%; Left-Left, 0%; Left-Mixed, 1%; Mixed-Mixed, 3%; Right-Right, 73% (MZA hand-concordant = 76%; MZA hand-­
discordant = 24%); DZA twins (n = 55 pairs): Right-Left, 18%; Right-Mixed, 15%; Left-Left, 2%; Left-Mixed, 2%; Mixed-Mixed, 5%; Right-Right, 58% (DZA hand
concordant = 65%; DZA hand discordant = 35%). Data presented in Study 1 of the current paper differ for several reasons: (1) we included only same-sex twin-pairs,
(2) those who reported attempts at handedness switching (whether successful or not) are coded as NRH, and (3) these data relate specifically to writing hand, whereas
the data presented by Segal (2012) relate to Crovitz-Zener Handedness Inventory (Crovitz and Zener, 1962) scores.

hand preference, regardless of direction. 5.4. Statistics

5.3.2. Strength of relative hand skill
The Purdue Pegboard (PPB; Tiffin, 1968) task is a test of manual
dexterity in which participants are timed while placing metal pins, one
at a time, into a row of holes in a pegboard, first with the dominant hand
and then with the non-dominant hand. The measure is known to exhibit
high test-retest reliability (e.g., r = 0.76; Doyen and Carlier, 2002). As
with the CZHI, we computed an absolute (i.e., unsigned) LI from this
task as a measure of the strength of relative hand skill. In this case we
used absolute values of (L - R)/(L + R) × 100. That is because higher
scores on the PPB indicate slower hand speed, i.e., low values for the
right hand relative to the left hand signify right-handedness (opposite to
the scoring system for the CZHI). A score of 0, therefore, indicates no
difference in completion time for the right and left hands, whereas
scores closer to 100 indicate greater disparity between the hands. In
addition to the absolute LI, we examined right-hand speed and left-hand
speed as separate outcome variables.
Males are more likely than females to indicate being left-, mixed-, or
non-right-handed (Papadatou-Pastou et al., 2008). Cohort effects have
also been reported, with studies published earlier than 1976 finding a
lower prevalence of left-handedness than later studies (Papadatou-Pas­
tou et al., 2020). Additionally, females are, on average, faster than males
on the PPB, and performance on this task reduces with age for both
males and females (Agnew et al., 1988). For these reasons, we regressed
sex, age, sex × age, age2, and sex × age2 on each of our outcome vari­
ables (unsigned CZHI score, unsigned PPB LI score, right-hand PPB
speed, and left-hand PPB speed), and used the residual scores in sub­
sequent analyses to control for age and sex effects according to the
methods of McGue and Bouchard (1984).
To determine MZA and DZA within-pair similarity for handedness
outcomes we computed intraclass correlation coefficients (ICC), using
two-way mixed effects models with single measures and absolute
agreement definition. Given that shared environmental effects are ab­
sent for MZA twins, the ICC for this group can be considered a direct

15
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 3 Table 4
Summary concordance/discordance (excluding missing/incomplete data) for Summary concordance/discordance (all data) for direction of hand preference in
direction of hand preference in twins reared apart. twins reared apart.
RH/RH NRH/NRH RH/NRH RH/RH NRH/NRH RH/NRH

Males (n ¼ 157 MZA (n = 51 (64.6%) 6 (7.6%) 22 Males (n ¼ 174 MZA (n = 64 (68.1%) 6 (6.4%) 24
pairs) 79) (27.8%) pairs) 94) (25.5%)
DZA (n = 62 (79.5%) 1 (1.3%) 15 DZA (n = 62 (77.5%) 1 (1.3%) 17
78) (19.2%) 80) (21.3%)
Females (n ¼ 258 MZA (n = 81 (63.8%) 8 (6.3%) 38 Females (n ¼ 298 MZA (n = 105 (66.9%) 8 (5.1%) 44
pairs) 127) (29.9%) pairs) 157) (28.0%)
DZA (n = 95 (72.5%) 5 (3.8%) 31 DZA (n = 104 (73.8%) 5 (3.5%) 32
131) (23.7%) 141) (22.7%)
All twins (n ¼ 415 MZA (n = 132 (64.1%) 14 (6.8%) 60 All twins (N ¼ 472 MZA (n = 169 (67.3%) 14 (5.6%) 68
pairs) 206) (29.1%) pairs) 251) (27.1%)
DZA (n = 157 (75.1%) 6 (2.9%) 46 DZA (n = 166 (75.1%) 6 (2.7%) 49
209) (22.0%) 221) (22.2%)
Concordant Discordant Concordant Discordant
Males (n ¼ 157 MZA (n = 57 (72.2%) 22 (27.8%) Males (n ¼ 174 MZA (n = 70 (74.5%) 24 (25.5%)
pairs) 79) pairs) 94)
DZA (n = 63 (80.8%) 15 (19.2%) DZA (n = 63 (78.8%) 17 (21.3%)
78) 80)
Females (n ¼ 258 MZA (n = 89 (70.1%) 38 (29.9%) Females (n ¼ 298 MZA (n = 113 (72.0%) 44 (28.0%)
pairs) 127) pairs) 157)
DZA (n = 100 (76.3%) 31 (23.7%) DZA (n = 109 (77.3%) 32 (22.7%)
131) 141)
All twins (n ¼ 415 MZA (n = 146 (70.9%) 60 (29.1%) All twins (N ¼ 472 MZA (n = 183 (72.9%) 68 (27.1%)
pairs) 206) pairs) 251)
DZA (n = 163 (78.0%) 46 (22.0%) DZA (n = 172 (77.8%) 49 (22.2%)
209) 221)

Note. Data included here are derived from the samples studied by Newman et al.
(1937) (n = 7 MZAm pairs; n = 12 MZAf pairs), Shields (1962) (n = 3 MZAm
pairs), and Juel-Nielsen (1965, 1980) (n = 3 MZAm pairs; n = 9 MZAf pairs); the
Louise Wise Services (LWS)/Child Development Center (CDC) Twin Study (see
Segal, 2021a) (n = 1 MZAm pair; n = 4 MZAf pairs; n = 2 DZAf pairs); Minnesota
Study of Twins Reared Apart (MISTRA) (Segal, 2012) (n = 30 MZAm pairs; n =
39 MZAf pairs; n = 11 DZAm pairs; n = 22 DZAf pairs); Fullerton Study of
Chinese Twins Adopted Apart (Segal et al., 2011) (n = 1 MZAm pair; n = 14
MZAf pairs; n = 1 DZAm pair; n = 4 DZAf pairs); Finnish Twin Cohort Study
(Kaprio et al., 2019) (n = 9 MZAm pairs; n = 12 MZAf pairs; n = 37 DZAm pairs;
n = 41 DZAf pairs); Swedish Adoption Twin Study of Aging (SATSA) (Pedersen,
2015) (n = 13 MZAm pairs; n = 15 MZAf pairs; n = 28 DZAm pairs; n = 58 DZAf
pairs); as well as all miscellaneous case reports (n = 12 MZAm pairs; n = 22
MZAf pairs; n = 1 DZAm pair; n = 4 DZAf pairs). Cases with missing or
incomplete handedness data from Shields (1962) (n = 13 MZAm pairs; n = 30
MZAf pairs; n = 1 DZAm pair; n = 10 DZAf pairs), Tienari (1963a) (n = 2 MZAm
pairs), and Slater (1953) (n = 1 DZAm pair) are not included here.
Note. Data included here are derived from the cohorts studied by Newman et al.
(1937) (n = 7 MZAm pairs; n = 12 MZAf pairs); Shields (1962) (n = 16 MZAm
pairs, n = 30 MZAf pairs; n = 1 DZAm pair; n = 10 DZAf pairs [note that an
earlier paper by Shields, 1958, reports on n = 13 MZAm and n = 25 MZAf of
these twin pairs); and Juel-Nielsen (1965, 1980) (n = 3 MZAm pairs; n = 9 MZAf
pairs); the Louise Wise Services (LWS)/Child Development Center (CDC) Twin
Study (see Segal, 2021a) (n = 1 MZAm pair; n = 4 MZAf pairs; n = 2 DZAf pairs);
Minnesota Study of Twins Reared Apart (MISTRA) (Segal, 2012) (n = 30 MZAm
pairs; n = 39 MZAf pairs; n = 11 DZAm pairs; n = 22 DZAf pairs); Fullerton Study
of Chinese Twins Adopted Apart (Segal et al., 2011) (n = 1 MZAm pair; n = 14
MZAf pairs; n = 1 DZAm pair; n = 4 DZAf pairs); Finnish Twin Cohort Study
(Kaprio et al., 2019) (n = 9 MZAm pairs; n = 12 MZAf pairs; n = 37 DZAm pairs;
n = 41 DZAf pairs); Swedish Adoption Twin Study of Aging (SATSA) (Pedersen,
2015) (n = 13 MZAm pairs; n = 15 MZAf pairs; n = 28 DZAm pairs; n = 58 DZAf
pairs); as well as all available miscellaneous case reports (n = 14 MZAm pairs; n
= 22 MZAf pairs; n = 2 DZAm pairs; n = 4 DZAf pairs).

estimate of heritability (Segal, 2012). We also compared the ICCs be­ Table 5
Reported writing hand stratified by sex and twin type in the MISTRA.
tween MZA and DZA twins using Fisher’s r-to-z tests. We did this because
a significantly higher ICC for MZAs than DZAs would provide further MZAm DZAm MZAf DZAf
evidence of genetic influence should such effects emerge. Statistical n % n % n % n %
analyses were conducted using IBM SPSS version 27 (IBM Corp, 2020), RH total 51 85.0% 17 77.3% 73 93.6% 43 95.6%
and data visualisation was accomplished with the ggplot2 package LH total 9 15.0% 5 22.7% 5 6.4% 2 4.4%
(Wickham, 2009) in RStudio version 1.3.1073 (R Core Team, 2020).
Note. 26 twins reported that they had been encouraged to switch from LH to RH,
although in 7 cases this proved unsuccessful. Handedness reported here relates
6. Results to preference for the writing hand at the time of measurement, regardless of
whether switching of handedness had been attempted in childhood. DZAf, fe­
Most participants, regardless of twin type, reported writing with male dizygotic reared-apart twins; DZAm, male dizygotic reared-apart twins;
their right hand (Table 5). Scores on the CZHI and PPB were uncorre­ MZAf, female monozygotic reared-apart twins; MZAm, male monozygotic
lated, whether signed, rs (214) = − 0.096, p = .161) or unsigned, rs (214) reared-apart twins.
= 0.102, p = .136.
Intraclass correlation coefficients (ICCs) were calculated for each excluding twin pairs for which one or both members reported attempts
handedness measure (Table 6) and scatterplots are displayed in Fig. 1. to switch their handedness during childhood and when restricting the
Strength of hand preference was significantly positively correlated analysis to only right-handed participants. The only notable differences
within DZA pairs, but not within MZA pairs. Strength of relative hand were that (1) the DZA ICC for strength of hand preference was not sig­
skill showed no evidence of correlation within MZA or DZA twin pairs, nificant when considering only those participants that did not report
but significant positive correlations for right-hand and left-hand speed attempted handedness switching during childhood, and (2) the DZA
on the PPB task were observed in MZA twins. No such effects were correlation was significantly stronger than the MZA correlation only
observed in DZA twins, although the difference in slopes was not sta­ when the analysis was restricted to right-handed participants (Table 6).
tistically significant. This pattern of findings remained similar when

16
G. Richards and N.L. Segal Neuropsychologia 184 (2023) 108523

Table 6
Intraclass correlation coefficients for handedness measures in MZA and DZA twins in the MISTRA.
MZA DZA Comparison

Inclusion criteria n ICC p n ICC p z p

Hand preference strength All participants 80 0.019 .433 54 0.333 .007 − 1.81 .070
Non-switched 62 0.043 .370 45 0.230 .064 − 0.95 .342
Non-switched RH 51 − 0.112 .781 37 0.490 < .001 − 2.89 .004
Relative hand skill strength All participants 74 0.043 .357 45 − 0.143 .842 0.96 .337
Non-switched 59 0.126 .171 38 − 0.138 .813 1.23 .219
Non-switched RH 49 0.131 .185 31 − 0.199 .862 1.39 .165
Right-hand speed All participants 76 0.332 .002 47 0.053 .362 1.53 .126
Non-switched 64 0.339 .003 41 0.065 .343 1.39 .165
Non-switched RH 55 0.384 .002 34 0.231 .089 0.75 .453
Left-hand speed All participants 76 0.318 .002 47 0.150 .157 0.93 .352
Non-switched 64 0.289 .009 41 0.199 .108 0.46 .646
Non-switched RH 55 0.380 .002 34 0.086 .316 1.38 .168

Note. Hand preference strength is the absolute (unsigned) laterality index (LI) for the Crovitz-Zener Handedness Inventory (CZHI; Crovitz and Zener, 1962); relative
hand skill strength is the absolute (unsigned) LI for the Purdue Pegboard (PPB; Tiffin, 1968) task; right- and left-hand speed are the times taken to complete the PPB
task with each hand, respectively (each outcome variable has been residualised to control for effects of age and sex). Statistically significant effects (p < .05, two-tailed)
are presented in bold. DZA, dizygotic reared-apart twins; MZA, monozygotic reared-apart twins.

Fig. 1. Scatterplots for similarity in (a) strength of hand preference (b) strength of relative hand skill, (c) right hand speed, and (d) left hand speed, stratified by
zygosity.
Note. CZHI, Crovitz-Zener Handedness Inventory; DZA, dizygotic reared-apart twins; LI, laterality index; MZA, monozygotic reared-apart twins; PPB, Purdue
Pegboard Task.

17
G. Richards and N.L. Segal
7. Discussion
The current research provides a synthesis of the reared-apart twin
literature as regards hand preference (Study 1) and examines several
features of handedness obtained from participants in the Minnesota
Study of Twins Reared Apart (MISTRA) (Study 2). Study 1 identified that
at least N = 560 MZA and same-sex DZA twin pairs have come to the
attention of researchers. Direction of hand preference data were avail­
able for both members of n = 415 cases, and concordance was slightly
higher in DZA (78.0%) than MZA (70.9%) twin pairs. Although the focus
here is on handedness, it is hoped that our compilation of case reports
can serve as a useful resource for researchers interested in studying
reared-apart twins. The findings from Study 2 offer evidence of mod­
erate heritability for right-hand and left-hand completion speed for the
Purdue Pegboard (PPB) task, implying genetic influence on motor skill.
However, MZA twin pairs were no more alike than chance for strength of
hand preference or relative hand skill. This lack of MZA twin pair
resemblance could possibly be linked to unusual prenatal developmental
influences unique to MZ twins (e.g., differential cell distribution asso­
ciated with zygotic division) that may affect cotwins differently
(McNamara et al., 2016; Segal, 2021a). We also observed statistically
significant similarity for strength of hand preference in DZA twin pairs.
Further research will be needed to resolve these unexpected results.
The finding that MZA twin pairs were similar for right-hand and left-
hand task completion speed for the PPB is consistent with both reared-
together and reared-apart twin studies reporting that MZ twins are
more alike than chance for motor performance (Fox et al., 1996; Jäncke
and Steinmetz, 1995) and dominant hand psychomotor speed (Simonen
et al., 1998). These findings are also consistent with the observation that
studies using the classic twin design have provided evidence of genetic
influence for most measured human individual difference traits studied
thus far (Polderman et al., 2015). However, it should be noted that
heritability estimates for hand speed derived from MZA intraclass cor­
relation coefficients (ICC) in the current study were moderate, implying
that considerable variance for this characteristic is explained by
non-shared environmental factors.
Strength of relative hand skill, as measured via the PPB task, did not
show evidence of heritability. This finding is consistent with an adoption
study (Saudino and McManus, 1998) that observed no resemblance
between adopted children and their biological parents for strength of
handedness. It is also consistent with research showing no statistically
significant resemblance in strength of relative hand skill within MZ twin
pairs reared together (Jäncke and Steinmetz, 1995). Although these
findings suggest no genetic basis for strength of relative hand skill,
McManus and Bryden (1992) estimated that only ~10% of the variance
in this factor is familial. Large sample sizes would, therefore, be required
to detect statistically significant effects, implying that the current study
may lack adequate statistical power.
We observed an unexpected effect in that strength of hand preference
on the CZHI was more similar than chance for DZA twin pairs, but not for
MZA twin pairs. Considering the absence of shared environmental fac­
tors present for DZA twins, this effect could be attributed to genetic
influence, with the lack of significant effect in MZA twins representing a
Type 2 error. Alternatively, the significant DZA correlation could be a
Type 1 error. The effect in DZA pairs was statistically significant when
examining the whole cohort, or when the analysis was limited to right-
handed participants who had not been subject to attempts at handedness
switching during childhood, but not when examining the subset of
participants that included left-handers who had not experienced at­
tempts at handedness switching. Likewise, the difference in strength
between the MZA and DZA correlations was notably only statistically
significant when the analysis was restricted to right-handers who had
not experienced attempts at handedness switching. It is unclear why the
findings differ in this manner. Regardless, considering the paucity of
literature relating to genetic and environmental influences on strength
of hand preference (regardless of direction) and relative hand skill,
18
Neuropsychologia 184 (2023) 108523
further large-scale research that includes reared-together and reared-
apart twin pairs, combined with molecular genetic techniques, could
prove informative with respect to this question.
A strength of the current research is that we used a reared-apart twin
study, representing one of the most powerful designs for examining the
relative importance of genetic and environmental effects on behavioural
traits. However, while this design allows researchers to separate genetic
from shared environmental influences, reared-apart twin pairs are rare;
thus, obtaining sufficient data to achieve adequate levels of statistical
power is challenging. Nevertheless, Lykken et al. (1981) showed how
statistical power increased by 20 percent when studying twins reared
apart versus twins reared together.
Another possible concern is that our survey of the literature of
reared-apart twins identified data for more MZA (n = 325) than DZA (n
= 235) pairs, even though same-sex DZ twins are approximately as
common as MZ twins in the general population. Interestingly, this
pattern was not observed when considering only those cases for which
handedness data were available (MZA, n = 206; DZA, n = 209). In some
research, e.g., Newman et al. (1937), DZA pairs were not studied, and in
other instances it is clear that DZA pairs were studied in less detail than
MZA pairs (e.g., Shields, 1962). Notably, although Tienari (1963a,
1963b) specified that all MZA pairs in his case series were investigated,
elsewhere it is mentioned that a DZAm pair, raised apart from age 2–15
years, was not followed-up (Tienari, 1966). Likewise, although Juel-­
Nielsen (1965, 1980) dedicated (two editions of) an entire book to the
study of 12 pairs of MZA pairs, a DZAf pair, separated until age 28, was
only mentioned briefly in a separate publication (Juel-Nielsen and
Harvald, 1958). Additionally, as Segal (2012) has pointed out, MZA
twins are more likely to find each another than DZA twins because
confusion by others (due to physical resemblance) may facilitate their
reunion. Consistent with this view, 65 of 89 (73.0%) DZAm and 99 of
146 (67.8%) DZAf cases identified in our review came from either the
SATSA or the Finnish Twin Cohort Study, both of which are
population-based. Notably, we also identified only two case reports that
were dedicated to the study of a specific DZA twin-pair (one DZAm
[Segal, 2015]; one DZAf [Segal et al., 2015b]). Although the study of
MZA twins is particularly compelling, we emphasise the need to also
study DZA pairs to obtain adequate comparison. However, despite the
foregoing, the body of work produced by the MISTRA yielded findings
consistent with extant reared-together twin studies (Segal, 2012). The
applicability of most twin study findings to the non-twin population has
also been demonstrated for the majority of measured traits (Christensen
et al., 1995).
We administered the Crovitz-Zener Handedness Inventory (CZHI;
Crovitz and Zener, 1962) rather than the more widely used Edinburgh
Handedness Inventory (EHI; Oldfield, 1971). However, although dif­
ferences are noted, considerable consistency is generally observed
across handedness questionnaires as regards their content and response
formats (Papadatou-Pastou et al., 2013). Notably, items from the CZHI
and EHI that assess the same manual activities tend to show a high de­
gree of agreement (Bryden, 1977). Likewise, although we utilised the
PPB task (Tiffin, 1968) as a measure of hand speed and strength of
relative hand skill, alternative paradigms (e.g., Annett Pegboard task
[Annett, 1970]; Tapley and Bryden circle-marking task [Tapley and
Bryden, 1985]; finger-tapping [Peters and Durding, 1978]) are some­
times used in laterality research. Simultaneous consideration of multiple
measures of relative hand skill suggests that although direction (i.e.,
right or left) is consistent across individuals, the different tasks relate to
separable processes (McManus et al., 2016). Further research may
therefore consider administering a range of relative hand skill measures,
as these could exhibit different levels of heritability. It would also be of
interest to examine genetic correlations between these distinct aspects of
handedness.
G. Richards and N.L. Segal
8. Conclusion
We present a comprehensive summary of hand preference in reared-
apart twins (Study 1) and the first study of strength of hand preference
and relative hand skill in twin pairs from the MISTRA (Study 2).
Although we observed evidence consistent with moderate genetic effects
on right-hand and left-hand speed, we did not find evidence of herita­
bility for strength of relative hand skill. We also found greater than
chance similarity for strength of hand preference in DZA, but not MZA
twins. Although this last effect was unexpected, it may reflect the rela­
tively small sample of DZA pairs present within the dataset. Further
largescale molecular genetics and reared-together twin studies may offer
a better understanding of the nature and underpinnings of strength of
handedness.
Data sharing
The datasets generated and/or analyzed during the current study are
not publicly available due to limited ethics approval for the wider
Minnesota Study of Twins Reared Apart (MISTRA) and to the specific
consent provided by the participants. They may be available from the
second author on reasonable request and pending approval of any future
analyses by the MISTRA. SPSS syntax and R code used to run analyses for
Study 2 are publicly available on the Open Science Framework (htt
ps://osf.io/e4ysv/).
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgments and funding sources
Preliminary findings were presented on June 23rd, 2022, at the 52nd
Annual Meeting of the Behavior Genetics Association (Richards and Segal,
2022) and on August 25th, 2022, at the 8th North Sea Meeting on Lat­
erality. The authors thank Prof. Nancy L. Pedersen for providing and
interpreting the SATSA data and Prof. Jaakko Kaprio for sharing the
handedness data for reared apart twins from the Finnish Twin Cohort
Study. For their assistance in interpreting research published in German
and Japanese, we thank, respectively, Dr Patrick Rosenkranz and Dr
Eriko Self. We also thank the team based at the Walton Library, New­
castle University, for their tireless efforts in searching for and locating
many obscure academic articles that we could not otherwise have ob­
tained. The SATSA was supported by the National Institutes of Health
(grant numbers R01 AG04563, AG10175, AG028555), the MacArthur
Foundation Research Network on Successful Aging; the Swedish Council
for Working Life and Social Research (FAS/FORTE) (grant numbers
97:0147:1B, 2009-0795), and the Swedish Research Council (grant
numbers: 825-2007-7460, 825-2009-6141). Support from the Academy
of Finland (grant #336823 to Jaakko Kaprio) enabled provision of the
Finnish Twin Cohort Study data.
References
Abrams, S., 1986. Disposition and the environment. Psychoanal. Stud. Child 41 (1),
41–60. https://doi.org/10.1080/00797308.1986.11823450.
Abrams, S., Neubauer, P.B., 1994. Hartmann’s vision: identical twins and developmental
organizations. Psychoanal. Stud. Child 49 (1), 49–59. https://doi.org/10.1080/
00797308.1994.11823050.
Agnew, J., Bolla-Wilson, K., Kawas, C.H., Bleecker, M.L., 1988. Purdue pegboard age and
sex norms for people 40 years old and older. Dev. Neuropsychol. 4 (1), 29–35.
https://doi.org/10.1080/87565648809540388.
Annett, M., 1970. The growth of manual preference and speed. Br. J. Psychol. 61 (4),
545–558. https://doi.org/10.1111/j.2044-8295.1970.tb01274.x.
Annett, M., 2002. Handedness and Brain Asymmetry: the Right Shift Theory. Psychology
Press.
Bakan, P., Dibb, G., Reed, P., 1973. Handedness and birth stress 11, 363–366.
19
Neuropsychologia 184 (2023) 108523
Beaton, A.A., 2003. The nature and determinants of handedness. In: Hugdahl, K.,
Davidson, R.J. (Eds.), The Asymmetrical Brain. MIT Press, pp. 105–158. https://doi.
org/10.7551/mitpress/1463.003.0006.
Bordier, A., Futerman, S., 2014. Separated @ Birth: a True Love Story of Twin Sisters
Reuinited. Berkley Books.
Bouchard Jr., T.J., Lykken, D.T., McGue, M., Segal, N.L., Tellegen, A., 1990. Sources of
human psychological differences: the Minnesota study of twins reared apart. Science
250 (4978), 223–228. https://doi.org/10.1126/science.2218526.
Bouterwek, H., 1936. Erhebungen an eineiigen Zwillingspaaren über Erbanlage und
Umwelt als Charakterbildner. Z. Menschl. Vererb-u. Konstit-Lehre 20, 265–275.
Bouterwek, H., 1943. Erbe und Persönlichkeit: Charakterologische Ergebnisse der
Zwillingsforschung. Franz Deuticke.
Bryden, M.P., 1977. Measuring handedness with questionnaires. Neuropsychologia 15
(4–5), 617–624. https://doi.org/10.1016/0028-3932(77)90067-7.
Burks, B.S., 1942. A study of identical twins reared apart under differing types of family
relationships. In: Terman, L.M. (Ed.), Studies in Personality. McGraw-Hill,
pp. 35–69. https://doi.org/10.1111/j.1536-7150.1978.tb02790.x.
Burks, B.S., Roe, A., 1949. Studies of identical twins reared apart. Psychological
Monographs: General and Applied, 63(5), i–62. https://doi.org/10.1037/h0093608.
Burley, L., 2022. The Real Mo Farah. British Broadcasting Company.
Burlingham, D., 1952. Twins: a study of three pairs of identical twins with 30 charts.
International Universities Press.
Burt, C., 1955. The evidence for the concept of intelligence. Br. J. Educ. Psychol. 25 (3),
158–177. https://doi.org/10.1111/j.2044-8279.1955.tb03305.x.
Burt, C., 1958. The inheritance of mental ability. Am. Psychol. 13 (1), 1–15. https://doi.
org/10.1037/h0049002.
Burt, C., 1966. The genetic determination of differences in intelligence: a study of
monozygotic twins reared together and apart. Br. J. Psychol. 57 (1–2), 137–153.
https://doi.org/10.1111/j.2044-8295.1966.tb01014.x.
Burt, C., 1943. Ability and income. Br. J. Educ. Psychol. 13 (2), 83–98. https://doi.org/
10.1111/j.2044-8279.1943.tb02725.x.
Carter-Saltzman, L., 1980. Biological and sociocultural effects on handedness:
comparison between biological and adoptive families. Science 209 (4462),
1263–1265. https://doi.org/10.1126/science.7403887.
Christensen, K., Vaupel, J.W., Holm, N.V., Yashin, A.I., 1995. Mortality among twins
after age 6: fetal origins hypothesis versus twin method. BMJ 310 (6977), 432–436.
https://doi.org/10.1136/bmj.310.6977.432.
Conway, J., 1958. The inheritance of intelligence and its social implications. British
Journal of Statistical Psychology 11 (2), 171–190. https://doi.org/10.1111/j.2044-
8317.1958.tb00012.x.
Coren, S., Porac, C., 1980. Birth factors and laterality: effects of birth order, parental age,
and birth stress on four indices of lateral preference. Behav. Genet. 10 (2), 123–138.
https://doi.org/10.1007/BF01066263.
Craike, W.H., Slater, E., 1945. Folie à deux in uniovular twins reared apart. Brain: J.
Neurol. 68, 213–221. https://doi.org/10.1093/brain/68.3.213.
Crovitz, H.F., Zener, K., 1962. A group-test for assessing hand- and eye-dominance. Am.
J. Psychol. 75 (2), 271–276. https://doi.org/10.2307/1419611.
Cuellar-Partida, G., Tung, J.Y., Eriksson, N., Albrecht, E., Aliev, F., Andreassen, O.A.,
Barroso, I., Beckmann, J.S., Boks, M.P., Boomsma, D.I., Boyd, H.A., Breteler, M.M.B.,
Campbell, H., Chasman, D.I., Cherkas, L.F., Davies, G., de Geus, E.J.C., Deary, I.J.,
Deloukas, P., Medland, S.E., 2020. Genome-wide association study identifies 48
common genetic variants associated with handedness. Nat. Human Behav. https://
doi.org/10.1038/s41562-020-00956-y.
Danforth, C.H., 1919. Resemblance and difference in twins: twins that look and act alike
attract attention first, while dissimilar ones are apt to be overlooked. J. Hered. 10
(9), 399–409. https://doi.org/10.1093/oxfordjournals.jhered.a101956.
Dencker, S.J., 1958. A follow-up study of 128 closed head injuries in twins using co-twins
as controls. Acta Psychiatrica et Neurologica 33 (Suppl. 123), 1–125.
Doyen, A.-L., Carlier, M., 2002. Measuring handedness: a validation study of Bishop’s
reaching card test. Laterality: Asymmetries Body, Brain Cognit. 7 (2), 115–130.
https://doi.org/10.1080/13576500143000140.
Essen-Möller, E., 1941. Psychiatrische Untersuchungen an einer Serie von zwillingen.
Acta Psychiatrica et Neurologica, Suppl. 23, 1–200.
Essen-Möller, E., 1970. Twenty-one psychiatric cases and their MZ cotwins: a thirty
years’ follow-up. Acta Genet. Med. Gemellol.: Twin Res. 19 (1–2), 315–317. https://
doi.org/10.1017/s1120962300025798.
Farber, S.L., 1981. Identical Twins Reared Apart: a Reanalysis. Basic Books, Inc.,
Publishers.
Fernandes, E. de S., Correia, J. L., Hellen, H. V. V., Silva, M. Jr., Segal, N. L., & Otta, E. (in
preparation). Brazilian twins reared apart since newborns: A Case Study.
Fisher, R.A., 1958. Cancer and smoking. Nature 182 (4635). https://doi.org/10.1038/
182596a0, 596.
Fox, P.W., Hershberger, S.L., Bouchard Jr., T.J., 1996. Genetic and environmental
contributions to the acquisition of a motor skill. Nature 384 (6607), 356–358.
https://doi.org/10.1038/384356a0.
Fraceschetti, A., Bamatter, F., Klein, D., 1948. Valeur des tests cliniques et sérologiques
en vue de l’identification de deux jumeaux univitellins, dont l’un a été échangé par
erreur. Bull. Schweiz. Akad. Med. Wiss. 4 (5–6), 433–444. https://doi.org/10.5169/
seals-306949.
Franceschetti, A., 1948. L’importance déterminante des facteurs rhésus dans un cas
médico-légal. Arch. Julius Klaus-Stiftung für Vererbungsforsch. Sozialanthropol.
Rassenhyg. 23 (3–4), 460–464.
Franceschetti, A., Bamatter, F., Klein, D., 1948. Valeur des tests cliniques et sérologiques
en vue de l’identification de deux jumeaux univitellins, dont l’un a été échangé par
erreur. Bull. Schweiz. Akad. Med. Wiss. 4 (5–6), 433–444. https://doi.org/10.5169/
seals-306949.
G. Richards and N.L. Segal
Fukuoka, G., 1937. Anthropometric and psychometric studies on Japanese twins. In:
Komai, T. (Ed.), Contributions to the Genetics of the Japanese Race II: Studies on
Japanese Twins, pp. 9–47.
Gao, W.-J., Li, L.-M., Cao, W.-H., Zhan, S.-Y., Lv, J., Pang, Z.-C., Chen, W.-J., Wang, S.-J.,
Chen, R.-F., Hu, Y.-H., 2011. A comparative study on blood pressure, obesity,
smoking and alcohol drinking behavior in Chinese twins reared apart and together.
J. Peking Univ. (Heal. Sci.) 43 (3), 329–332. V43/I3/329. http://xuebao.bjmu.edu.
cn/EN/Y2011/.
Gardner, I.C., Newman, H.H., 1940. Mental and physical traits of identical twins reared
apart: case XX. Twins Lois and Louise. J. Hered. 31 (3), 119–126. https://doi.org/
10.1093/oxfordjournals.jhered.a104855.
Glasby, M., Loader, D., Glasby, R., 2013. Buy One Get One Free: A Book about Fact Being
Stranger than Fiction. www.wanowandthen.com.
Gottesman, I.I., Shields, J., 1972. Schizophrenia and Genetics: a Twin Study Vantage
Point. Academic Press.
Hagi, A., 1931. The blood types of twins as studied with saliva. Tokio Iji Shinshi 2731,
1455–1462.
Hayakawa, K., Shimizu, T., Kato, K., Onoi, M., Kobayashi, Y., 2002. A gerontological
cohort study of aged twins: the Osaka University Aged Twin Registry. Twin Res. 5
(5), 387–388. https://doi.org/10.1375/136905202320906147.
Hayakawa, K., Kato, K., Onoi, M., Hayashi, C., Yang-Ping, C., Kanamori, M., Doi, S.,
Kikuchi, H., Nishihara, R., Kadota, K., 2006. The Japanese study of adult twins
reared apart and growing old separately. Twin Res. Hum. Genet. 9 (6), 806–807.
https://doi.org/10.1375/twin.9.6.806.
Hayashi, S., 1963. A study of juvenile delinquency by twin method. Hanzaigaku Zasshi
29, 153–172.
Hayashi, S., 1967. A study of juvenile delinquency in twins. In: Mitsuda, H. (Ed.), Clinical
Genetics in Psychiatry: Problems in Nosological Classification. Igaku Shoin Ltd.,
pp. 373–378
IBM Corp, 2020. IBM SPSS Statistics for MacOS. Version 27.0. IBM Corp.
Inouye, E., 1971. Monozygotic twins with schizophrenia reared apart in infancy.
Excerpta Med. Int. Congr. Ser. 233, 93.
Inouye, E., 1972. Monozygotic twins with schizophrenia reared apart in infancy. Jpn. J.
Hum. Genet. 16 (3), 182–190.
Jäncke, L., Steinmetz, H., 1995. Hand motor performance and degree of asymmetry in
monozygotic twins. Cortex 31 (4), 779–785. https://doi.org/10.1016/S0010-9452
(13)80028-7.
Jordan, H.E., 1914. Hereditary lefthandedness, with a note on twinning. (Study III).
J. Genet. 4 (1), 67–81. https://doi.org/10.1007/bf02981820.
Joye, M., 1954. He Was Not My Son. Rinehart & Company, Inc.
Juel-Nielsen, N., 1965. Individual and Environment: Monozygotic Twins Reared Apart.
International Universities Press.
Juel-Nielsen, N., 1980. Individual and Environment: Monozygotic Twins Reared Apart.
International Universities Press.
Juel-Nielsen, N., Harvald, B., 1958. The electroencephalogram in uniovular twins
brought up apart. Acta Genet. Stat. Med. 8 (1), 57–64. https://doi.org/10.1159/
000151054.
Juel-Nielsen, N., Mogensen, A., 1957. Uniovular twins brought up apart: (preliminary
report of a psychiatric-psychological study). Acta Genet. Stat. Med. 7 (2), 430–433.
Kaij, L., 1960. Alcoholism in twins: studies on the etiology and sequels of abuse of
alcohol. Almqvist & Wiksell.
Kallmann, F.J., 1938. The genetics of schizophrenia. J. J. Augustin.
Kallmann, F.J., 1946. The genetic theory of schizophrenia: an analysis of 691
schizophrenic twin index families. Am. J. Psychiatr. 103 (3), 309–322. https://doi.
org/10.1176/ajp.103.3.309.
Kallmann, F.J., Roth, B., 1956. Genetics aspects of preadolescent schizophrenia. Am. J.
Psychiatr. 112 (8), 599–606. https://doi.org/10.1176/ajp.112.8.599.
Kaprio, J., Bollepalli, S., Buchwald, J., Iso-Markku, P., Korhonen, T., Kovanen, V.,
Kujala, U., Laakkonen, E.K., Latvala, A., Leskinen, T., Lindgren, N., Ollikainen, M.,
Piirtola, M., Rantanen, T., Rinne, J., Rose, R.J., Sillanpää, E., Silventoinen, K.,
Sipilä, S., Waller, K., The older Finnish Twin Cohort — 45 years of follow-up, 2019.
Twin Res. Hum. Genet. 22 (4), 240–254. https://doi.org/10.1017/thg.2019.54.
Komai, T., 1937. Review of literature on twin studies in Japan. In: Komai, T. (Ed.),
Contributions to the Genetics of the Japanese Race II: Studies on Japanese Twins,
pp. 1–8.
Kranz, H., 1936. Lebensschicksale krimineller Zwillinge. Verlag von Julius Springer.
Kringlen, E., 1964. Schizophrenia in male monozygotic twins. Acta Psychiatr. Scand. 40
(Suppl. 178), 1–76.
Kringlen, E., 1967a. Heredity and Environment in the Functional Psychoses. Heinemann.
Kringlen, E., 1967b. Case Histories Supplement. Norwegian Monographs on Medical
Science.
Kurihara, M., 1959. A study of schizophrenia by twin method. Psychiatr. Neurol. Jpn. 61
(13), 1721–1741.
Lange, J., 1931. Crime as Destiny: a Study of Criminal Twins. George Allen & Unwin Ltd.
Lindeman, B., 1969. The twins who found each other. Pocket Books second ed.
Lykken, D.T., 1978. The diagnosis of zygosity in twins. Behav. Genet. 8 (5), 437–473.
https://doi.org/10.1007/BF01067939.
Lykken, D.T., Geisser, S., Tellegen, A., 1981. Heritability Estimates from Twin Studies:
the Efficiency of the MZA Design. Unpublished Manuscript. University of Minnesota.
McGue, M., Bouchard Jr., T.J., 1984. Adjustment of twin data for the effects of age and
sex. Behav. Genet. 14 (4), 325–343. https://doi.org/10.1007/BF01080045.
McIndoe, A., Franceschetti, A., 1949. Reciprocal skin homographs in a medico-legal case
of familial identification of exchanged identical twins. Br. J. Plast. Surg. 2, 283–289.
McManus, I.C., 1981. Handedness and birth stress. Psychol. Med. 11 (3), 485–496.
https://doi.org/10.1017/S0033291700052806.
20
Neuropsychologia 184 (2023) 108523
McManus, I.C., 2002. Right Hand, Left Hand: the Origins of Asymmetries in Brains,
Bodies, Atoms and Cultures. Weidenfeld & Nicolson.
McManus, I.C., 2022. Cerebral polymorphisms for lateralisation: modelling the genetic
and phenotypic architectures of multiple functional modules. Symmetry, 14(4) 814.
https://doi.org/10.3390/sym14040814.
McManus, I.C., Bryden, M.P., 1992. The genetics of handedness, cerebral dominance and
lateralization. In: Rapin, I., Segalowitz, S.J. (Eds.), Handbook of Neuropsychology,
Vol. 6: Child Neuropsychology. Elsevier Science Publishers B.V., pp. 115–144
McManus, I.C., Van Horn, J.D., Bryden, P.J., 2016. The Tapley and Bryden test of
performance differences between the hands: the original data, newer data, and the
relation to pegboard and other tasks. Laterality: Asymmetries of Body, Brain and
Cognit. 21 (4–6), 371–396. https://doi.org/10.1080/1357650X.2016.1141916.
McNamara, H.C., Kane, S.C., Craig, J.M., Short, R.V., Umstad, M.P., 2016. A review of
the mechanisms and evidence for typical and atypical twinning. Am. J. Obstet.
Gynecol. 214 (2), 172–191. https://doi.org/10.1016/j.ajog.2015.10.930.
Mitsuda, H., Sakai, T., 1968. Comparative studies with monozygotic twins discordant for
typical and atypical schizophrenia. Jpn. J. Hum. Genet. 13 (3), 183–188.
Mogensen, A., Juel-Nielsen, N., 1961a. Factors influencing the selection and rejection of
Szondi’s pictures: a study of uniovular twins brought up apart. Acta Psychiatr.
Scand. 37 (1), 32–36. https://doi.org/10.1111/j.1600-0447.1961.tb06157.x.
Mogensen, A., Juel-Nielsen, N., 1961b. Monozygotic twins reared apart. Part 1.
Intelligence. Acta Psychologica 91, 426–427. https://doi.org/10.1016/S0001-6918
(61)80190-X.
Mogensen, A., Juel-Nielsen, N., 1963. Proceedings of the Sixteenth International
Congress of Psychology, Bonn 1960: Monozygotic Twins Reared Apart, vol. 3,
pp. 22–23. I. Intelligence. Excerpa Criminologica.
Moher, D., Liberati, A., Tetzlaff, J., Altman, D.G., Antes, G., Atkins, D., Barbour, V.,
Barrowman, N., Berlin, J.A., Clark, J., Clarke, M., Cook, D., D’Amico, R., Deeks, J.J.,
Devereaux, P.J., Dickersin, K., Egger, M., Ernst, E., Gøtzsche, P.C., Tugwell, P., 2009.
Preferred reporting items for systematic reviews and meta-analyses: the PRISMA
statement. PLoS Med. 6 (7), e1000097. https://doi.org/10.1371/journal.
pmed.1000097.
Muller, H.J., 1925. Mental traits and heredity: the extent to which mental traits are
independent of heredity, as tested in a case of identical twins reared apart. J. Hered.
16 (12), 433–448. https://doi.org/10.1093/oxfordjournals.jhered.a102537.
Newman, H.H., 1929a. Mental traits of identical twins reared apart: case III. Twins ‘C’
and ‘O’. J. Hered. 20 (4), 153–166. https://doi.org/10.1093/oxfordjournals.jhered.
a103166.
Newman, H.H., 1929b. Mental and physical traits of identical twins reared apart: case I.
Twins ‘A’ and ‘O’. J. Hered. 20 (2), 49–64. https://doi.org/10.1093/oxfordjournals.
jhered.a103142.
Newman, H.H., 1929c. Mental and physical traits of identical twins reared apart: case II.
Twins ‘E’ and ‘G’. J. Hered. 20 (3), 97–104. https://doi.org/10.1093/oxfordjournals.
jhered.a103163.
Newman, H.H., 1932a. Mental and physical traits of identical twins reared apart: case IV.
Twins Mary and Mabel, and review of the first four cases studied. J. Hered. 23 (1),
3–18. https://doi.org/10.1093/oxfordjournals.jhered.a103509.
Newman, H.H., 1932b. Mental and physical traits of identical twins reared apart: case V.
Twins ‘B’ and ‘D’. J. Hered. 23 (8), 297–303. https://doi.org/10.1093/
oxfordjournals.jhered.a103639.
Newman, H.H., 1932c. Mental and physical traits of identical twins reared apart: case VI.
Twins Ada and Ida. J. Hered. 23 (9), 369–377. https://doi.org/10.1093/
oxfordjournals.jhered.a103664.
Newman, H.H., 1933. Mental and physical traits of identical twins reared apart: case VII.
Twins Richard and Raymond. J. Hered. 24 (5), 209–214. https://doi.org/10.1093/
oxfordjournals.jhered.a103778.
Newman, H.H., 1934a. Mental and physical traits of identical twins reared apart: case IX.
Twins Harold and Holden. J. Hered. 25 (4), 137–143. https://doi.org/10.1093/
oxfordjournals.jhered.a103900.
Newman, H.H., 1934b. Mental and physical traits of identical twins reared apart: case
VIII. Twins ‘M’ and ‘R’. J. Hered. 25 (2), 55–60. https://doi.org/10.1093/
oxfordjournals.jhered.a103878.
Newman, H.H., Freeman, F.N., Holzinger, K.J., 1937. Twins: A Study of Heredity and
Environment. University of Chicago Press.
Nicholls, M.E.R., Johnston, D.W., Shields, M.A., 2012. Adverse birth factors predict
cognitive ability, but not hand preference. Neuropsychology 26 (5), 578–587.
https://doi.org/10.1037/a0029151.
Ocklenburg, S., Güntürkün, O., 2018. The Lateralized Brain: the Neuroscience and
Evolution of Hemispheric Asymmetries. Academic Press.
Oldfield, R.C., 1971. The assessment and analysis of handedness: the Edinburgh
Inventory. Neuropsychologia 9 (1), 97–113. https://doi.org/10.1016/0028-3932
(71)90067-4.
Ooki, S., 2006. Nongenetic factors associated with human handedness and footedness in
Japanese twin children. Environ. Health Prev. Med. 11 (6), 304–312. https://doi.
org/10.1007/bf02898021.
Papadatou-Pastou, M., Tomprou, D.-M., 2015. Intelligence and handedness: meta-
analyses of studies on intellectually disabled, typically developing, and gifted
individuals. Neurosci. Biobehav. Rev. 56, 151–165. https://doi.org/10.1016/j.
neubiorev.2015.06.017.
Papadatou-Pastou, M., Martin, M., Munafò, M.R., Jones, G.V., 2008. Sex differences in
left-handedness: a meta-analysis of 144 studies. Psychol. Bull. 134 (5), 677–699.
https://doi.org/10.1037/a0012814.
Papadatou-Pastou, M., Martin, M., Munafò, M.R., 2013. Measuring hand preference: a
comparison among different response formats using a selected sample. Laterality:
Asymmetries of Body, Brain and Cognit. 18 (1), 68–107. https://doi.org/10.1080/
1357650X.2011.628794.
G. Richards and N.L. Segal
Papadatou-Pastou, M., Ntolka, E., Schmitz, J., Martin, M., Munafò, M.R., Ocklenburg, S.,
Paracchini, S., 2020. Human handedness: a meta-analysis. Psychol. Bull. 146 (6),
481–524. https://doi.org/10.1037/bul0000229.
Pedersen, N.L., 2015. Swedish adoption/twin study on aging (SATSA), 1984, 1987, 1990,
1993, 2004, 2007, and 2010. Inter-university Consortium for Political and Social
Research [distributor], 10.3886/ICPSR03843.2015-05-13, v2.
Peters, M., Durding, B.M., 1978. Handedness measured by finger tapping: a continuous
variable. Can. J. Psychol. 32 (4), 257–261. https://doi.org/10.1037/h0081694.
Pfeifer, L.S., Schmitz, J., Papadatou-Pastou, M., Peterburs, J., Paracchini, S.,
Ocklenburg, S., 2022. Handedness in twins: meta-analyses. BMC Psychology 10 (1),
11. https://doi.org/10.1186/s40359-021-00695-3.
Polderman, T.J.C., Benyamin, B., de Leeuw, C.A., Sullivan, P.F., van Bochoven, A.,
Visscher, P.M., Posthuma, D., 2015. Meta-analysis of the heritability of human traits
based on fifty years of twin studies. Nat. Genet. 47 (7), 702–709. https://doi.org/
10.1038/ng.3285.
Popenoe, P., 1922. Twins reared apart. J. Hered. 13 (3), 142–144. https://doi.org/
10.1093/oxfordjournals.jhered.a102184.
Price, J., 1968. The genetics of depressive behavior. In: Coppen, A., Walk, A. (Eds.),
Recent Developments in Affective Disorders, British Journal of Psychiatry. Special
Publication No. 2, pp. 37–54 (Headley Bros).
Prokop, H., Druml, W., 1973. Diskordantes Auftreten einer depressiv gefärbten Katatonie
bei eineiigen Zwillingen im Jugendalter. Wien. Z. Nervenheilkd. Deren Grenzgeb. 31
(2), 156–166.
R Core Team., 2020. R: a language and environment for statistical computing.
R Foundation for Statistical Computing. https://www.r-project.org/.
Richards, G., Segal, N.L., 2022. Strength of hand preference and relative hand skill in
twins reared apart. Behav. Genet. 52, 388. https://doi.org/10.1007/s10519-022-
10119-6.
Rosanoff, A.J., Handy, L.M., Rosanoff Plesset, I., 1935. The etiology of manic-depressive
syndromes with special reference to their occurrence in twins. Am. J. Psychiatr. 91
(4), 725–762. https://doi.org/10.1176/ajp.91.4.725.
Rosanoff, A.J., Handy, L.M., Rosanoff Plesset, I., 1937. The etiology of mental deficiency
with special reference to its occurrence in twins: a chapter in the genetic history of
human intelligence. Psychological Monographs, 48(4), i–137. https://doi.org/10.1037
/h0093544.
Satz, P., 1972. Pathological left-handedness: an explanatory model. Cortex 8 (2),
121–135. https://doi.org/10.1016/S0010-9452(72)80013-3.
Saudek, R., 1933. Identical twins reared apart: a comparative study of various tests of
their intellectual, emotional, and social attitudes. Character and Personality 2 (1),
22–40. https://doi.org/10.1111/j.1467-6494.1933.tb02079.x.
Saudek, R., 1934. A British pair of identical twins reared apart. Character and Personality
3 (1), 17–39. https://doi.org/10.1111/j.1467-6494.1934.tb01980.x.
Saudino, K., McManus, I.C., 1998. Handedness, footedness, eyedness and earedness in
the Colorado Adoption Project. Br. J. Dev. Psychol. 16 (2), 167–174. https://doi.org/
10.1111/j.2044-835X.1998.tb00916.x.
Schein, E., Bernstein, P., 2007. Identical Strangers: A Memoir of Twins Separated and
Reunited. Hachette Australia.
Schwesinger, G.C., 1952. The effect of differential parent-child relation on identical twin
resemblance in personality. Acta Genet. Med. Gemellol. 1 (1), 40–47. https://doi.
org/10.1017/s1120962300042360.
Searleman, A., Porac, C., Coren, S., 1989. Relationship between birth order, birth stress,
and lateral preferences: a critical review. Psychol. Bull. 105 (3), 397–408. https://
doi.org/10.1037/0033-2909.105.3.397.
Seeman, E., Saudek, R., 1933. The handwriting of identical twins reared apart. Character
and Personality 1 (4), 268–285.
Segal, N.L., 2009. Twins reared apart: a forgotten case. Twin Res. Hum. Genet. 12 (1),
123–126. https://doi.org/10.1375/twin.12.1.123.
Segal, N.L., 2011a. Someone Else’s Twin: the True Story of Babies Switched at Birth.
Prometheus Books.
Segal, N.L., 2011b. Reunited twins: spouse relations. Twin Res. Hum. Genet. 14 (3),
290–294. https://doi.org/10.1375/twin.14.3.290.
Segal, N.L., 2012. Born Together—Reared Apart: the Landmark Minnesota Twin Study.
Harvard University Press. https://doi.org/10.4159/harvard.9780674065154.
Segal, N.L., 2014. Art for twins: Yorùbá artists and their statues. Twin Res. Hum. Genet.
17 (3), 215–221. https://doi.org/10.1017/thg.2014.18.
Segal, N.L., 2015. The fourth International Network of Twin Registries: overview from
Osaka. Twin Res. Hum. Genet. 18 (6), 812–818. https://doi.org/10.1017/
thg.2015.81.
Segal, N.L., 2016. Twinsters the movie: reared apart twins in real time. Twin Res. Hum.
Genet. 19 (1), 80–85. https://doi.org/10.1017/thg.2015.98.
Segal, N.L., 2017. Reared-apart Chinese twins: chance discovery. Twin Res. Hum. Genet.
20 (2), 180–185. https://doi.org/10.1017/thg.2017.9.
Segal, N.L., 2018. Twins switched at birth: frequency, life histories, twin relationships,
and critical issues. Twin Res. Hum. Genet. 21 (5), 477–483. https://doi.org/
10.1017/thg.2018.48.
Segal, N.L., 2021a. Deliberately Divided: Inside the Controversial Study of Twins and
Triplets Adopted Apart. Rowman & Littlefield.
Segal, N.L., 2021b. Twins separated at birth: across a country and around the world.
Twin Res. Hum. Genet. 24 (4), 244–250. https://doi.org/10.1017/thg.2021.30.
Segal, N.L., 2022. Reared-apart and reunited: extraordinary dynamics. Twin Res. Hum.
Genet. 25 (3), 165–170. https://doi.org/10.1017/thg.2022.21.
Segal, N.L., Blandón-Gitlin, I., 2010. Twins switched at birth: a case from the Canary
Islands. Twin Res. Hum. Genet. 13 (1), 115–119. https://doi.org/10.1375/
twin.13.1.115.
21
Neuropsychologia 184 (2023) 108523
Segal, N.L., Cortez, F.A., 2014. Born in Korea-adopted apart: behavioral development of
monozygotic twins raised in the United States and France. Pers. Indiv. Differ. 70,
97–104. https://doi.org/10.1016/j.paid.2014.06.029.
Segal, N.L., Diamond, M., 2014. Identical reared apart twins concordant for
transsexuality. J. Exp. Clin. Med. 6 (2), 74. https://doi.org/10.1016/j.
jecm.2014.02.007.
Segal, N.L., Hur, Y.-M., 2008. Reared apart Korean female twins: genetic and cultural
influences on life histories, physical and health-related measures, and behavioral
traits. Int. J. Behav. Dev. 32 (6), 542–548. https://doi.org/10.1177/
0165025408097133.
Segal, N.L., Hur, Y.-M., 2022. Personality traits, mental abilities and other individual
differences: monozygotic female twins raised apart in South Korea and the United
States. Pers. Indiv. Differ. 194 (111643) https://doi.org/10.1016/j.
paid.2022.111643.
Segal, N.L., Montoya, Y.S., 2018. Accidental Brothers: the Story of Twins Exchanged at
Birth and the Power of Nature and Nurture. St. Martin’s Press.
Segal, N.L., Stohs, J.H., Evans, K., 2011. Chinese twin children reared apart and reunited:
first prospective study of co-twin reunions. Adopt. Q. 14 (1), 61–78. https://doi.org/
10.1080/10926755.2011.557953.
Segal, N.L., Hur, Y.-M., Graham, J.L., 2015a. Korean twins reared apart: genetic and
cultural influences on behavior and health. Adopt. Q. 18 (4), 291–310. https://doi.
org/10.1080/10926755.2015.1088110.
Segal, N.L., Cortez, F.A., Zettel-Watson, L., Cherry, B.J., Mechanic, M., Munson, J.E.,
Velázquez, J.M.A., Reed, B., 2015b. Genetic and experiential influences on behavior:
twins reunited at seventy-eight years. Pers. Indiv. Differ. 73, 110–117. https://doi.
org/10.1016/j.paid.2014.09.017.
Segal, N.L., Montoya, Y.S., Loke, Y.J., Craig, J.M., 2017. Identical twins doubly
exchanged at birth: a case report of genetic and environmental influences on the
adult epigenome. Epigenomics 9 (1), 5–12. https://doi.org/10.2217/epi-2016-0104.
Segal, N.L., Montoya, Y.S., Becker, E.N., 2018. Twins reared apart and twins in families:
the findings behind the fascination. Twin Res. Hum. Genet. 21 (4), 295–301. https://
doi.org/10.1017/thg.2018.34.
Segal, N.L., Montoya, Y.S., Peña, F.Y., Burgos, S., Katz, X., 2019. Eye refraction in doubly
exchanged monozygotic twins. Twin Res. Hum. Genet. 22 (3), 177–182. https://doi.
org/10.1017/thg.2019.26.
Shields, J., 1958. Twins brought up apart. Eugen. Rev. 50 (2), 115–123.
Shields, J., 1962. Monozygotic Twins Brought up Apart and Brought up Together: an
Investigation into the Genetic and Environmental Causes of Variation in Personality.
Oxford University Press.
Shields, J., Slater, E., 1967. Genetic aspects of schizophrenia. Hosp. Med. 1, 579–584.
Simonen, R.L., Videman, T., Battié, M.C., Gibbons, L.E., 1998. Determinants of
psychomotor speed among 61 pairs of adult male monozygotic twins. J. Gerontol.
Series A: Biol. Sci. Med. Sci. 53A (3), M228–M234. https://doi.org/10.1093/gerona/
53A.3.M228.
Slater, E., 1953. Psychotic and Neurotic Illnesses in Twins (Medical Research Council
Special Report Series, No. 278). Her Majesty’s Stationery Office.
Slater, E., 1961. The thirty-fifth Maudsley lecture: ‘Hysteria 311’. J. Ment. Sci. 107 (448),
359–381. https://doi.org/10.1192/bjp.107.448.359.
Slater, E., 1968. A review of earlier evidence on genetic factors in schizophrenia.
J. Psychiatr. Res. 6 (Suppl. 1), 15–26. https://doi.org/10.1016/0022-3956(68)
90005-8.
Stenstedt, Å., 1952. A study in manic-depressive psychosis: clinical, social and genetic
investigations. Acta Psychiatr. Scand. 28 (Suppl. 79), 1–111.
Stephens, F.E., Nunemaker, J.C., 1950. Spondylitis in identical twins reared apart.
J. Hered. 41 (11), 283–286. https://doi.org/10.1093/oxfordjournals.jhered.
a106058.
Stephens, F.E., Thompson, R.B., 1943. The case of Millan and George: identical twins,
reared apart. J. Hered. 34 (4), 109–114. https://doi.org/10.1093/oxfordjournals.
jhered.a105259.
Sudarsky, L., Myers, R.H., Walshe, T.M., 1983. Huntington’s disease in monozygotic
twins reared apart. J. Med. Genet. 20 (6), 408–411. https://doi.org/10.1136/
jmg.20.6.408.
Tapley, S.M., Bryden, M.P., 1985. A group test for the assessment of performance
between the hands. Neuropsychologia 23 (2), 215–221. https://doi.org/10.1016/
0028-3932(85)90105-8.
Thomas, A., Chess, S., 1977. Temperament and Development. Brunner/Mazel.
Tienari, P., 1963a. Psychiatric illnesses in identical twins. Acta Psychiatr. Scand. 39
(Suppl. 171), 9–195.
Tienari, P., 1963b. A psychiatric twin study (preliminary report). Acta Psychiatr. Scand.
38 (Suppl. 169), 393–397.
Tienari, P., 1966. On intrapair differences in male twins with special reference to
dominance-submissiveness. Acta Psychiatr. Scand. 42 (Suppl. 188), 1–166. https://
doi.org/10.1111/j.1600-0447.1966.tb01912.x.
Tienari, P., 1968. Schizophrenia in monozygotic male twins. In: Rosenthal, D., Kety, S.S.
(Eds.), The Transmission of Schizophrenia. Pergamon, pp. 27–36.
Tiffin, J., 1968. Purdue pegboard examiner manual. Science Research Associates.
Tucker, W.H., 1997. Re-reconsidering Burt: beyond a reasonable doubt. J. Hist. Behav.
Sci. 33 (2), 145–162. https://doi.org/10.1002/(sici)1520-6696(199721)33:2<145::
aid-jhbs6>3.0.co;2-s.
Wagenseil, F., 1931. Zwei Mitteilungen über die erbbiologische Bedeutung der eineiigen
Mehrlinge. Z. Für Konstitutionslehere 15, 632–645.
Wickham, H., 2009. ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag.
https://doi.org/10.1007/978-0-387-98141-3.
Williams, C.S., Buss, K.A., Eskenazi, B., 1992. Infant resuscitation is associated with an
increased risk of left-handedness. Am. J. Epidemiol. 136 (3), 277–286. https://doi.
org/10.1093/oxfordjournals.aje.a116493.
G. Richards and N.L. Segal
Yates, N., Brash, H., 1941. An investigation of the physical and mental characteristics of
a pair of like twins reared apart from infancy. Ann. Eug. 11 (1), 89–101. https://doi.
org/10.1111/j.1469-1809.1941.tb02276.x.
Yoshimasu, S., 1941. Psychopathie und Kriminalität. Die Bedeutung der Erbanlage und
Umwelt für die Entstehung von Verbrechen im Lichte der Zwillingsforschung.
Psychiatr. Neurol. Jpn. 45 (9), 455–531.
22
Neuropsychologia 184 (2023) 108523
Zhou, B., Gao, W., Lv, J., Yu, C., Wang, S., Liao, C., Pang, Z., Cong, L., Dong, Z., Wu, F.,
Wang, H., Wu, X., Jiang, G., Wang, X., Wang, B., Cao, W., Li, L., 2015. Genetic and
environmental influences on obesity-related phenotypes in Chinese twins reared
apart and together. Behav. Genet. 45 (4), 427–437. https://doi.org/10.1007/
s10519-015-9711-0.