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                    <text>Item ID Number

°1291

Author

Rose, Richard J.

Corporate Author
Report/Article Title Genetic Variance In Nonverbal Intelligence: Data From
the Kinships of Identical Twins

Journal/Book Title

Scienco

Year

1979

Month/Day

September 14

Color

n

Number of Images

3

DoSCrlpton NotOS

Alvin L Youn

9 fi'ecl this item under the category
"DDT/Human Toxicology and Environmental Fate"

Thursday, April 26, 2001

Page 1291 of 1328

�man, M. Kupcrsmith, E. Estcy, M. Goldstein,
Lancet 1976-11, 515 (1976); A. Licbcrman, T.
Miyamoto, A. F. Batlista, M. Goldstein, Neurology 25, 459 (1975); A. Liebcrman el al., N.
Engl J. Med. 295, 1400 (1976); H. L. Klawans,
W. J. Weiner, P. A. Nausieda, P. Volkman,
C. Goetz, M. D. Lupton, Neurology 27, 390
(1977).
3. M. O. Thorner, S. M. Ryan, J. A. H. Wass, A.
Jones, P. Bouloux, S. Williams, G. M. Besser,
J. Clin. Endocrinol. Melab. 47, 372 (1978);
L. Lemberger, R. Crabtree, J. Clemens, R.
W. Dyke, R. T. Woodburn, ibid. 39, 579
(1974).
4. After the administration of uC-labeled pcrgolide, total radiation in plasma reached its
peaked concentration at 1 to 2 hours, indicating
that the drug was absorbed during this period of
pharmacologic activity (A. Rubin el al,, unpublished observations).

5. Blood was drawn i» heparinized tubes daily at
8:00 a.m. and 9:30 a.m. for the determination of
prolactin, LH, FSH, growth hormone, cortisol,
and TSH.
6. V. N. Sinha, F. W. Selby, U. J. Lewis, W. P,
Vanderlaan,/. Clin. Endocrinol. Metab. 36, 509
(1973). The prolactin standard (V-L-S No. 1)
and specific antiserum were kindly provided by
the National Pituitary Agency, NIAMDD. Sensitivity of the assay is 0.5 ng/nil, and the coefficient of variation is 6 percent between 8 to 30 ng/
ml. A computer program was used for analysis
of the counting data. Control serum assayed
with each set of samples had an interassay variability of ± 5 percent.
7. R. W. Fuller, J. A. Clemens, E. C. Kornfeld, H.
D. Snoddy, E. B. Smalstig, N. J. Bach, Life
Sci., in press.
2 April 1979; revised 21 June 1979

Genetic Variance in Nonverbal Intelligence:
Data from the Kinships of Identical Twins
Abstract. The multiple relationships within kinships of adult monozygotic twins
permit incisive analyses to be made of genetic and environmental effects on behavioral traits. Data from families of 65 monozygotic twin pairs yield evidence of geneticvariance on the Block Design Test, a nonverbal measure of general intelligence.
A comparison of mental ability of foster children with that of their biological
and their adoptive parents was first reported in 1924 (/), and 1 year later H. J.
Mullet presented the first case report of
intellectual resemblance in monozygotic
(MZ) co-twins who had been separated
in infancy (2). In the succeeding halfcentury, studies of adopted children and
separated identical twins have had a central role in research on genetic and environmental determinants of cognitive
abilities. The resultant data have generated a continuing controversy (5), and its
resolution may require new research designs.
The families of adult identical twins
provide a new paradigm of particular
promise for behavior-genetic study (4).
In this report we describe the paradigm
and illustrate its application.
In Fig. 1 the multiple parent-offspring
relationships found within families of
adult identical twins arc diagramed. Children in each of the nuclear families derive half their genes from a twin parent,
those genes being identical with genes of
the parent's twin sister or brother (the
children's "twin aunt" or "twin uncle").
Since the children and the twin aunt or
uncle do not live in the same households,
their relationship is somewhat comparable to that between foster children
and their biological mother or father. In
this way, studies of children of MZ twins
provide a parallel to studies of adopted
children, with two important advantages: (i) There is no disruption of the nuclear family milieu; the children arc
reared by their biological parents in their
SCIENCE, VOL. 205, 14 SEPTEMBER 1979

own homes, (ii) In adoption studies biological fathers are rarely available for
study (5); the relationship of nephew or
niece to twin uncle parallels that of foster
children to their biological father.
The environmental covariance of a
nephew or niece and the twin aunt or
uncle will be markedly less than that of a
parent and child living in a common
household, it may not be zero, however,
because MZ twins may select or create
similar postmarital environments. An estimate of such effects can be obtained
from the resemblance of nephew or niece
to the spouse aunt in kinships of male
twins and to the spouse uncle in kinships
of female twins. In the absence of assorlative mating, the children share neither genes nor a household environment

with the spouse aunt or uncle, and any
behavioral resemblance would therefore
suggest an environmental covariance
common to all members of the kinship.
Because their twin parents have identical sets of nuclear genes, children of
MZ twins are genetically related to one
another as half-siblings; socially, they
are reared as cousins in separate homes.
Further, in contrast to conventional halfsiblings resulting from divorce, death of
a parent, or illegitimacy, MZ half-siblings are expected to be of the same age
and size. Accordingly, offspring of MZ
twins afford a unique human parallel to
methods employed in animal genetics,
where controlled matings of sires to multiple dams yield estimates of genetic, environmental, and maternal effects from a
nested analysis of variance of the fulland half-sib progeny. Better still, half-sib
progeny occur with equal frequency
among maternal and paternal MZ twins,
providing a balanced research paradigm
that cannot be achieved in lower animals
even with controlled matings. To illustrate the paradigm, we here present a
study in which it is applied to the Block
Design Subtest of the Wechsler Intelligence Scales.
Wechsler's Block Design Test is an
adaptation of one introduced in 1923 by
Kohs (6), who presented evidence that
results from his test correlated highly
with results from the Stanford Binet but,
unlike the Binet, had only a modest relation to level of education. Research with
Wechsler's adaptation supports Kohs's
assertion that the block test provides a
measure of general intelligence. Across a
wide age range, block design is highly related to the general factor common to all
Wechsler subtcsts (7); it is the most reliable of the nonverbal subtests, and no

Fig.
1.
Parent-offA
spring
relationships
in families of identical
twins. (A) The offspring of identical
twin mothers comprise a maternal halfsibship who genetically relate to their
twin aunt as closely as
Kinship of twin mothers
Kinship of twin fathers
they do to their own
mother. (B) In the abTwin mother or father—son/daughter
sence of assortative
Male
mating, offspring in a
illustrated for mother
Female
paternal half-sibship
Twin aunt or uncle—nephew/niece
Twin
share neither comillustrated for twin aunt
mon genes nor a comSpouse aunt or uncle—nephew/niece
mon
environment
with
their
spouse
illustrated for spouse aunt
aunt. In a parallel
manner, offspring of identical twin brothers genetically relate to their twin uncle as closely
as they do to their own father but offspring in a maternal half-sibship share neither common
gents nor a common environment with their spouse uncle.
0036-8075/79/0914-1153$00.50/0 Copyright © 1979

AAAS

1153

�Table 1. Regression and correlation analyses of Block Design Test scores and fingerprint ridge
counts.*
Block design scores
Relationship
Coefficient
Son or daughter on father or mother
Nephew or niece on twin uncle or aunt
Nephew or niece on spouse uncle or aunt
Offspring on midparent
Monozygotic twins
Full siblings
Half-siblings
Father-mother

Regressions
0.28 ± 0.04
0.23 ± 0.06
-0.01 ± 0.06
0.54 ± 0.07
Correlations
0.68 ± 0.06
0.24 ± 0.08
0.10 ± 0.12
0.06 ± 0.10

N

Ridge count

N

Coefficient

572
318
241
254

0.42
0.37
-0.06
0.82

±
±
±
±

0.05
0.05
0.07
0.07

564
310
247
254

65
297
318
102

0.96
0.36
0.17
0.05

±
±
±
±

0.03
0.08
0.12
0.10

60
296
310
98

The regressions and correlations for all genetic relationships differ significantly from zero; those for the two
nongenetic relationships, the spouse correlation and the regression of nephew or niece on spo'use uncle or
aunt, do not. Sample sizes for most of the estimates are modest, however, and confidence intervals are quite
large.

measure of cognitive ability except vocabulary has lower error variance (8).
Block design correlates highly with the
vocabulary and information subtests, yet
in contrast to these verbal tests is but
weakly correlated with education (9)
and, perhaps as a consequence, is not
influenced by assortative mating (10),
again in contrast to the verbal scales.
To serve as a genetic guide for the
analysis of block test scores, we have analyzed the fingerprint ridge counts of the
study population. Total ridge count is a
well-documented example of polygenic
inheritance (11) that is largely unaffected
by postnatal environmental influences,
the number of ridges being fixed about
the 12th week after conception.
The block test was administered to 550
members of 65 MZ twin kinships (12).
The regression and correlation results
for block design and total ridge count are
summarized in Table 1 (13). For both
traits, familial resemblance appears to be
a direct function of shared genes. The
parent-offspring regression is comparable in magnitude to that between twin
uncle or aunt and nephew or niece (14)
but in the absence of shared genes neither trait exhibits significant familial aggregation. There is no resemblance between spouse uncle or aunt and nephew
or niece nor between husband and wife.
Neither environmental covariance nor
assortative mating appears to influence
either trait.
The parallel pattern of results for
block design and for ridge count provides evidence of significant genetic Variation in nonverbal intelligence. We emphasize that the evidence is obtained
from normal children reared in their natural homes by their biological parents,
children who differ in no known way
from the larger population to which we
wish to draw inferences (75). Table 2
presents heritability estimates for both
1154

traits that can be derived from the data.
The estimates show remarkably close
agreement across the multiple genetic
relationships contained within each kinship, but the reader is cautioned that
they are not statistically independent.
Our analyses suggest that substantial
variance in Block Design Test scores is
genetic in origin (16). An equivalent conclusion, of course, is that substantial variance in those scores is attributable to
nongenetic influences, and the twin-family methods which establish genetic variance can also identify systematic sources
of environmental influence on nonverbal
IQ. 0ur methodology enables us to assess uniquely one potential source of environmental variation, that of maternal
effects. A nested analysis of variance of
the offspring data permits a direct comparison of the relative similarity of maternal and paternal half-siblings. In this
preliminary sample, we find no evidence
of maternal effects in block design data,
although such influences are present in
verbal IQ scores (17). Independently,
Table 2. Heritability estimates for Block Design Test scores and total ridge counts.*
Estimated h2
Relationship

Block
design
scores

Total
ridge
count

Midparent-offspring
Parent-offspring
Twin uncle or auntnephew or niece
Full siblings
Half-siblings

.54
.56

.82
.84

.46
.48
.40

.74
.72
.68

The ratio of additive genetic variance to total
phenotypic variation defines the heritability, li'2.
From the composition of phenotypic covariances,
the regression or correlation of relatives can be expressed as estimates of A 2 (19). Interpretation of the
estimates should take into account the imprecision
of the coefficients on which they are based, the restricted range of variation in our sample, and the fact
that possible effects of common environment and
dominance variation have been ignored.

however, a prenatal environmental influence has been demonstrated by contrasting MZ twins classified into mono- and
dichorionic placenta! types (18).
RICHARD J. ROSE
Department of Psychology,
Indiana University, Bloomington 47405
E. L. HARRISJ. C. CHRISTIAN
Department of Medical Genetics,
Indiana University School of Medicine,
Indianapolis 46202
W. E. NANCE
Department of Human Genetics,
Medical College of Virginia,
Richmond 23298
References and Notes

1. S. V. S. Theis, How Foster Children Turn Out
(State Charities Aid Association, New York,
1924).
2. H. ). Muller,/. Hercd. 16, 433 (1925).
3. L. J. Cronbach,/lm. Psychol. 30, 1 (1975); L. J.
Kamin, The Science and Politics of IQ (Erlbaum, Potomac, Md., 1974); H. Munsinger,
Psychol. Bull. 82, 623 (1975); L. J. Kamin, ibid.
85, 194 (1978); J. Shields, in Progress in Clinical
and Biological Research: Twin Research, W. E.
Nance, G. Allen, P. Parisi, Eds. (Liss, New
York, 1978), p. 79.
4. W. E. Nance and L. A. Corey, Genetics 83, 811
(1976); R. J. Rose, J. Z. Miller, M. DumontDriscoll, M. Evans,Behav. Genet. 9,71 (1979).
5. A serious problem with all published adoption
studies is that no data on biological fathers of the
adopted children are available. Consequently,
the extent of assortative mating by the biological
mothers is unknown. Significant assortative
mating is likely present [R. Plomin, J. C. DeFries, M. K. Roberts, Science 1%, 449 (1977)],
and estimates of genetic influence based on present adoption data may be inflated.
6. S. C. Kohs, Intelligence Measurement: A Psychological and Statistical Study Based upon the
Blocks-Design Tests (Macmillan, New York,
1923).
7. U. Gault, Aust. J. Psychol. 6, 85 (1954); J. Cohen, J. Consult. Psychol. 21, 451 (1957); ibid.
23, 285 (1959); A. J. Kaufman,/. Consult. Clin.
Psychol. 43, 135 (1973).
8. D. Wechsler, Manual for the Wechsler Adult
Intelligence Scale (Psychological Corp., New
York, 1955); Manual for the Wechsler. Intelligence Scale for Children—Revised (Psychological Corp., New York, 1974); National Center for Health Statistics, Intellectual Development of Youth as Measured by a Short Form of
the Wechsler Intelligence Scale (PHS Publ.
1000-Series 11-No. 129, Government Printing
Office, Washington, D.C., 1973).
9. J. E. Birren, K. F. Riegcl, D. E. Morrison, J.
Gerontol. 16,363 (1961); D. Wechsler, The Measurement and Appraisal of Adult Intelligence
(Williams &amp; Wilkins, Baltimore, cd. 4, 1958).
10. T. Williams, Behav. Genet. S, 405 (1975).
11. S. B. Holt, Br. Med. Bull. 17, 247 (1961).
12. There were 278 males and 272 females, of whom
130 were twin parents, 102 nontwin parents, and
318 offspring. The 232 parents ranged in age
from 31 to 64 years [mean = 43; standard deviation (S.D.) = 8.9], their 318 offspring from 6 to
38 years (mean = 15.8; S.D. = 7.1). This is a
volunteer, Caucasian sample. Some self-selection, expected in voluntary research, is evident
in socioeconomic and educational characteristics of the participants. The IQ data exhibit
elevated means and restricted variances: Wechsler scales are standardized on representative
national samples with mean = 10 and S.D. = 3;
Block Design Test scores of this sample have
mean = 11.98 and S.D. = 2.78. Means and variances are homogenous for the twin and nontwin
parents, for the children and adults, and for
males and females.
As part of a half-day protocol of medical and
behavioral research, the subjects were tested individually by trained examiners, the same team .
of examiners being assigned to all members of
each MZ kinship. Subjects aged 6 through 15
(189 offspring) were given the 1974 revision of
the Wechsler Intelligence Scale for Children
(WISC); subjects age 16 and over (129 offspring
and all parents) were tested with the WcchsSCIENCE. VOL. 205

�ler Adult Intelligence Scale (WAIS). The block
design subtests of the WISC and WAIS are quite
comparable. In samples of 16- to 17-year-old
youth, the intertest correlation of block design is
as high as its rctest reliability on either test alone
and exceeds that of any other performance subtest [R. T. Ross and J. Morledge, J. Consult.
Psychol. 31, 331 (1967); M. Y. Quereshi and J.
M. Miller,/. Educ. Meas. 7, 105 (1970)].
Raw scores from the block design subtest
were standardized according to norms in appendices in the Wechsler manuals (8) for 30 agebands from ages 6 to 16 and 7 age-bands from
ages 16 to 64.
Usable ridge count data were obtained from
522 members of 60 MZ kinships in the sample.
All family members were genotyped with multiple blood markers to confirm monozygosity,
for paternity exclusion, and for future linkage
analyses.
13. The analysis employed here is identical with that
previously reported for plasma cholesterol [J. C.
Christian and K. W. Rang, Am. J. Hum. Genet.
20, 462 (1977)]. Regression coefficients are used
for their predictive value between generations,
and correlation coefficients are used to measure
relationships within the same generation. Regression coefficients are the preferred measure
of parent-offspring resemblance, because they
are less affected by range restriction, assortativc
mating, and variable sibship size [J. C. DeFries,
R. C. Johnson, A. R. Kuse, G. E. McClearn, J.
Polovina, S.'G. Vandenberg, J. R. Wilson, Behav. Genet. 9, 23 (1979)]. Regression and intraclass correlation coefficients were estimated by
the pairwise method [B. Rosner, A. Donner, C.
H. Hennekens, Appl. Stat. 26, 179 (1977)]. The
regression of offspring on midparent value is included in Table 1, since in the absence of common environmental variance it is a direct estimate of heritability [A. Vetta, Soc. Biol. 24, 166
(1977); J. C. DeFries, G. C. Ashton, R. C. Johnson, A. R. Kuse, G. E. McClearn, M. P. Mi, M.
N. Rashad, S. G. Vandenberg, J. R. Wilson, Be. hav. Genet. 8, 281 (1978)].
14. Normative age-banding may not fully correct for
the substantial age differences intrinsic to our
measurement of adult-child resemblance. Inadequacies of age standardization and nonequivalence of the two tests would reduce adult-child
resemblance relative to that of sibs and half-sibs
who are more closely matched in age.
15. We assume that there exist no unique biases in
the parental behavior or family structure of MZ
twins or the experiences of their children. The
assumption is untested, but plausible: personality characteristics of MZ twins cannot be distinguished from those of singletons [J. C. Loehlin
and R. C. Nichols, Heredity, Environment and
Personality (Univ. of Texas Press, Austin,
(1976)], and the same is probably true of their
parental behavior. The frequency of MZ twinning is equivalent across social and ethnic
groups and is not known to be associated with
any systematic factors.
16. This analysis is consistent with evidence from a
recent adoption study in Minnesota [S. Scarr
and R. A. Weinberg, in Readings About Individual and Group Differences, L. Willerman and R.
G. Turner, Eds. (Freeman, San Francisco,
1979)].
17. R. J. Rose, paper presented at the Second International Congress on Twin Studies, Washington, D.C., 1977.
18. Chorion type is reported to influence similarity
of full-scale IQ's in MZ twin children [M. Melnick, N. C. Myrianthopoulos, J. C. Christian,
Am. J. Hum. Genet. 30, 425 (1978)], and we are
now studying block design performance in a
sample of adult MZ co-twins whose placentation
was documented at birth; preliminary results (R.
J. Rose et at., in preparation) suggest a significant effect of chorion type on within-pair variation.
19. The expected value of parent-offspring regression is fcz/2. To estimate hcritability, the regression of offspring on single parent is doubled.
Since the variance of midparent values is half
the variance of individual parent values, the regression of offspring on midparent provides a direct estimate of A 2 . If effects of dominance variation and common environments are ignored, z
the
correlation of full siblings also estimates /i /2,
that of half siblings A2/4. For details, see references cited in (13). The correlations of MZ twins
provide upper limit estimates of heritability confounded by their closely shared environments;
the estimates arc .68 for blocks and .96 for ridge
counts.
20. This is publication number 78-41 from the Indiana University Human Genetics Center. It
was supported by PHS grant GM 21054.
21 August 1978; revised 5 July 1979
SCIENCE, VOL. 205, 14 SEPTEMBER 1979

The Cheetah: Native American
Abstract. Two North American fossil species of large felids, hitherto regarded as
Late Cenozoic pumas (mountain lion), are in fact closely related to the living cheetah, Acinonyx, of Africa and Eurasia .Anew subgenus (Miracinonyx) is proposed for
the American species. Cheetahs and pumas may have had a common ancestor in the
Miocene of North America.
Fossils of Puma-like cats are relatively
common in the Late Cenozoic of North
America (/). One species of supposed
Puma, "Felis" studeri, from the Pliocene of the Texas panhandle, has long
been recognized as distinct from Puma
concolor because of morphological similarities with Old World cheetahs, but
previous work has attributed the similarities to parallel evolution (/, 2). Excavations at the Late Pleistocene deposits of
Natural Trap Cave, Wyoming (3), indicated that another species previously
referred to Puma, "Felis" trumani (4),
also possesses several characters of dentition, skull, and limb architecture that
are remarkably "cheetah-like." Again,
the similarities were attributed to parallelism, and "F." trumani was styled as
the "cheetah-like cat" (3). Continued
excavations at the Wyoming site have
yielded hundreds of bones of this felid
(5), and more recent work (6) has revealed numerous shared derived characters that link "F." studeri and "F." trumani to Old World Acinonyx. Other
work (7) has utilized multivariate comparison of upper and lower tooth rows to
group the two American species; evolutionary affinities with Old World cheetahs were also suggested.
Except for size differences and several
features which are interpreted as retained primitive characters, the fossils of
"F." studeri and "F." trumani are almost identical with Old World Acinonyx
species (6). The points of similarity are
so extensive and of such a complex nature that a hypothesis attributing their
origin to other than common genetic descent would require pushing the concept
of parallel evolution to an unprecedented
extreme.
The systematic paleontology follows:
Family: Felidae
Genus: Acinonyx
Miracinonyx subgen. nov.
Derivation of name. From Latin
"mirus": surprising, amazing; and
Acinonyx: Old World cheetah.
Diagnosis. Distinguished from Puma
and other medium-sized felids by elongation of distal limbs (radius-ulna, tibia-fibula, calcaneum, metapodials); braincase
short and expanded; postorbital constriction wide; frontals broad and flat; internal nares enlarged; orbital shelf (zygo-

matic process of maxilla) short; skull
highly arched; coronoid process of ramus weak, slopes noticeably posterior;
canines weak; short mandibular diastema; protocone reduced or absent; auditory bulla elongate and flattened anteriorly. The latter character distinguishes
Miracinonyx from Old World cheetahs
(8). Acinonyx studeri may be distinguished from A. trumani by greater overall skull size and elongate, widely spaced
occipital condyles (2). Old World cheetahs (subgenus Acinonyx) are distinguished from the subgenus Miracinonyx
by a prominent anterior or anterolingual
cusp on P3, inflation of frontal sinus and
auditory bullae, and greater development of the medial anteroposterior ridge
of the basioccipital.
Geographic distribution. Western
United States: Texas (2), Nevada (4),
Wyoming (3).
Temporal distribution. Middle Blancan to Late Pleistocene.
Type species. Acinonyx (Miracinonyx)
trumani (Orr, 1969) (4).
Included species. Type species and
Acinonyx (Miracinonyx) studeri (Savage,
1960) (2).
Description. The skull of Miracinonyx
is highly arched, with the facial and cranial regions sloping anteriorly and posteriorly from the interorbital area of the
frontals. Shortening of the facial region
and enlargement of the P3 have reduced
the upper diastema and crowded the P2
tightly between the P3 and canine. Both
upper and lower canines are reduced, the
lower prcmolars are relatively narrow,
and the protocone of the upper carnassial is greatly reduced or absent. The
zygomatic process of the maxilla, which
in Puma forms a distinct shelf in the orbit
floor, is reduced. Shortening of the cranial region of the skull gives a bulging appearance to the braincase; the anterior
portion of the zygomatic arch is shortened, and the postorbital constriction is
widened. The frontals are greatly widened and flat, with the orbits set far apart
and high on the face. The postorbital
process of the frontals is distinct and of a
sharp angular shape; because of frontal
sinus inflation the postorbital processes
of Old World Acinonyx appear more
rounded. Dorsally, the skull of Acinonyx
is readily distinguishable from Puma,
which has a prominent postorbital con-

0036-8075/79/0914-1155S00.50/0 Copyright © 1979 AAAS

1155

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                  <text>&lt;p style="margin-top: -1em; line-height: 1.2em;"&gt;The Alvin L. Young Collection on Agent Orange comprises 120 linear feet and spans the late 1800s to 2005; however, the bulk of the coverage is from the 1960s to the 1980s and there are many undated items. The collection was donated to Special Collections of the National Agricultural Library in 1985 by Dr. Alvin L. Young (1942- ). Dr. Young developed the collection as he conducted extensive research on the military defoliant Agent Orange. The collection is in good condition and includes letters, memoranda, books, reports, press releases, journal and newspaper clippings, field logs and notebooks, newsletters, maps, booklets and pamphlets, photographs, memorabilia, and audiotapes of an interview with Dr. Young.&lt;/p&gt;&#13;
&lt;p&gt;For more about this collection, &lt;a href="/exhibits/speccoll/exhibits/show/alvin-l--young-collection-on-a"&gt;view the Agent Orange Exhibit.&lt;/a&gt;&lt;/p&gt;</text>
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                <text>E. L. Harris</text>
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                <text>W. E. Nance</text>
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                <text>Genetic Variance in Nonverbal Intelligence: Data From the Kinships of Identical Twins</text>
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