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Population variation in asymmetry and diversity from finger to finger for digital ridge-counts.

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Population Variation in Asymmetry and Diversity from
Finger to Finger for Digital Ridge-counts
RICHARD L. JANTZ
Department of Anthropology, University of Tennessee,
Knoxville, Tennessee 37916
KEY WORDS
tion
Dermatoglyphics
. Asymmetry.
. Ridge-counts . Racial varia-
ABSTRACT
Population variation in ridge-count asymmetry and diversity
from finger to finger has received scant attention in dermatoglyphic studies.
Asymmetry, in particular, has generally been attributed to environmental effects
operating during the formation of dermal ridges. Examination of samples from
several groups of diverse racial background revealed the existence of considerable population variation with respect to finger ridge-count asymmetry and diversity from finger to finger. Patterning along population lines suggests a genetic
rather than environmental basis for such variation. The genetic mechanisms responsible for ridge-counts may also mediate asymmetry and diversity, or the
degree of developmental stability in different populations may itself be under
genetic control.
It has long been recognized that homologous digits of the same individual often
display different dermatoglyphic patterns
(Cummins and Midlo, '61). The consistency
in levels of asymmetry among diverse
groups led these authors to conclude that
there was little in the way of racial variation with respect to pattern type asymmetry. More recently a number of investigators have examined bilateral asymmetry
with respect to ridge-counts. Asymmetrical
ridge-counts have been attributed to "developmental indeterminancy" (Rothammer
et al., ' 7 3 ) , and increases i n asymmetry
have been observed in chromosomal mosaics (Polani and Polani, '69) and cleft
palate patients (Adams and Niswander,
'67). Family studies have indicated that
asymmetry lacks a genetic basis entirely
(Holt, '54) or has only a small component
(Singh, '70).
Likewise, it has long been recognized
that ridge-counts on non-homologous digits
are correlated but may show considerable
variation in some individuals. Racial variation has been shown to exist with respect to ridge-count diversity from finger
to finger (Jantz, '74), and Holt ('60)
demonstrated a substantial genetic comAM. J. PHYS. ANTHROP., 42: 215-224.
ponent in her family material. Neither
asymmetry nor diversity from finger to
finger has ever been systematically considered from the standpoint of population
variation despite the information that
might result from such a n approach concerning the genetic and developmental
nature of dermatoglyphics. The aim of this
study is to investigate ridge-count asymmetry and diversity among non-homologous digits in a number of diverse groups
of mankind. The extent of racial and population variation is explored along with relationships among asymmetry, diversity
from finger to finger and ridge-counts.
MATERIALS AND METHODS
A number of different measures have
been used to quantitatively evaluate asymmetry in finger ridge-counts. Holt ('54)
used summed ridge-counts of right and
left hands, Parsons ('64) the summed
absolute differences between homologous
digits and Singh ('68) the mean of the
squared differences between homologous
digits. I n any measure of asymmetry it
would seem desirable to square differences
between homologous digits to emphasize
the larger and presumably more important
215
216
RICHARD L. JANTZ
asymmetries. For present purposes, the
following measure of asymmetry seems
appropriate :
i = l
Taking the square root of squared differences will result in a more symmetrical
distribution than simply using A'.
Holt ('58) evaluated diversity from
finger to finger by computing the quantity :
i=l
where q, is the ridge-count for the ith digit
and Q is Lq, or the total ridge count.
Later Holt ('60) used the measure
S/ \/lOor the standard deviation of counts
over 10 fingers to study familial correlations. It is apparent that ridge-count
asymmetry a s well as diversity among
non-homologous digits will contribute to
S/\/lo. The effect of asymmetry may be
easily eliminated by summing the ridgecounts of homologous digits so that only
diversity among non-homologous digits
contributes to the value. Holt's measure
then becomes :
s
/
d
5
=
j
F
Computation of these two values for
each individual provides convenient, independent, numerical expressions of that
individual's degree of asymmetry and diversity among non-homologous digits. Population characteristics can then be determined and comparisons made in the
usual manner.
The samples available for study consist
of two groups of European extraction
(English and American White), four of
African extraction (Dogon, Efe Pygmy,
Bedik-Bassari and American Negro) and
one Polynesian group (Easter Island).
Ideally, one would want to have samples
representing the major groups of mankind, but the present samples comprehend
virtually the total known range of ridgecount variation in man from the lowest
(Pygmy) to the highest (Easter Island).
The American White sample (133 males
and 132 females) was drawn from the
student body of the University of Tennessee. Only students who gave their birthplace as Tennessee or adjacent states were
included in the sample so that it may be
considered broadly representative of the
White population of the Southeastern
United States. No information as to the
ethnic composition was obtained, but the
majority of the subjects are probably of
British Isles extraction, since the Southeast was settled mainly by people of Scottish, Irish and English descent. Ridgecounts and S f i data for this sample appear in Jantz ('74).
The English sample (151 males and 151
females) consists of the parents of Dr. S.
B. Holt's large family sample. Only parents were included to eliminate related individuals and to keep the sample size on
more or less the same order as the other
groups in the study. This sample is broadly
representative of the population of Southeastern England and in particular, the
London area. Ridge-count data have been
reported in Holt ('58, '68) and S/\/10data
appear in Holt ('58) and Jantz ('74).
American Negro (102 males, 122 females) dermatoglyphic prints were obtained principally from students in elementary and junior high schools in Knoxville, Tennessee, but a few came from the
University of Tennessee as well. Most of
the subjects were born in Knoxville, but
many had parents who came to Knoxville
from many other areas in the Southern
United States. This sample can be taken
as representative of Southern Blacks, generally, and not of the Black population of
Knoxville in particular. Ridge-counts and
S / f i data have been presented in Jantz
('74).
Efe Pygmy (152 males and 53 females)
and Dogon (169 males and 100 females)
prints are those located in the Institute of
Human Biology, Utrecht, Netherlands. The
Pygmy prints were collected in the 1930's
by Dr. P. Julien, and ridge-counts of this
sample are presented by Glanville ('69).
Dogon dermatoglyphics were collected during the Tellem Expeditions of 1964-1965,
conducted by the Institute of Human
Biology, Utrecht. Finger dermatoglyphics
of this sample have been presented by
Huizinga ('65) and Glanville and Huizinga
('66).
217
RIDGE-COUNT ASYMMETRY AND DIVERSITY
Bedik-Bassari (103 males and 55 females) are two related tribal groups located
in Eastern Senegal. Dermatoglyphic prints
for both groups are located in the MusCe
de l'Homme, Paris. The dermatoglyphics
of the Bedik have been presented by
Gomila et al. ('65); those of the Bassari
are as yet unpublished. Data for these two
groups have been pooled for purposes of
the p r e s e n t analysis. Ridge-count and
S / d l O data for this sample appear in
Jantz ('74).
Easter Island (146 males and 141 females) dermatoglyphic prints were collected by Dr. R. J. Meier during the course
of his human biological investigations.
Meier ('69) identified approximately 14%
of his sample as containing no recent admixture, 39% were mixed with other Polynesians, and 47% possessed some European admixture. An analysis of the Easter
Island dermatoglyphics appears in Meier
('69).
All ridges were counted according to
methods set forth in Holt ('68), the larger
count being used in the case of whorls.
Ridge-counts of American Whites and
Negroes and Bedik-Bassari were made by
the author; for the remainder the counts
made by previous investigators were made
available to me. Intergroup differences
were evaluated by calculating the standard
error of the difference between means for
each pairwise comparison as described by
Fisher ('58). Results may then be presented as normal deviates, with probabilities obtained from a normal curve table.
This procedure is based on the assumption
that variables being tested are normally
distributed, a requirement not met by the
measures of diversity and asymmetry used
in this study. However, the distribution of
differences between means will frequently
approximate the normal distribution more
closely than the variables themselves, so
results obtained in this manner should be
acceptable for present purposes.
RESULTS
The means and standard deviations for
total ridge-count (TRC), S / 6 a n d m
are presented in table 1. TRC values show
no remarkable features and are in accord
with previous concepts of population variation. It is apparent that neither S/V% nor
*means
bear a systematic relationship
to the pattern of variation established by
TRC. In general the Ne ro populations
have the lowest S 6 a n d A' values, indicating greater homogeneity among fingers,
and less asymmetry, as well as the lowest
ridge-counts. However, the Easter Islanders, with the highest ridge-counts, are characterized by s/= and
values only
slightly above those of the Negroes, while
English and American Whites show the
greatest inter-finger heterogeneity and
asymmetry.
The magnitude of population differences
for all three measures is indicated by the
normal deviates for each painvise comparison in table 2. The two groups on the extremes of the ridge-count range, Easter
Islanders and Pygmies, differ significantly
from all other groups, and of course from
each other, with respect to TRC. English
males differ from the various Negro groups,
but beyond that, significant differences are
few. For samples of the size used here, it
is difficult to demonstrate significant differences for groups falling in the intermediate ridge-count range. The same is
F
*
TABLE 1
Means and standard deviations f o r TRC, S/\/rand 4Z
Males
Means
TRC
Sldd
Females
Standard deviations
dm
Means
TRC
SId3
dxi
TRC
46.93
55.74
45.71
47.25
45.32
50.11
49.40
2.79
3.29
3.20
2.79
2.78
2.83
2.59
4.07
4.07
3.98
3.14
3.91
3.37
3.62
162.30
127.87
126.08
129.21
118.44
122.95
97.66
S / d s fl
Standard deviations
TRC
SldS
di@
52.22
51.37
50.02
47.05
47.46
28.50
49.92
3.09
3.35
3.22
2.97
3.68
3.21
2.73
3.84
4.27
4.22
3.54
3.33
3.28
4.00
~~
Easter Island
English
Am. White
Am.Negro
Dogon
Bedik-Bassari
Pygmy
175.53
145.76
134.74
130.81
126.46
121.55
96.65
6.83 8.17
7.88 9.20
7.35 8.98
5.96 7.14
6.51 8.06
5.72 7.02
5.55 7.41
6.54
7.18
6.94
6.27
6.82
6.22
5.49
7.93
9.10
8.00
7.97
8.10
7.42
7.69
218
RICHARD L. JAN’IZ
a,
true for S / 6 and
with only the
groups on the extremes of the ranges,
English and American Whites on the
one hand, and the Negro samples on the
other showing consistently significant differences.
It is of considerable interest to note the
close correspondence between the means
of S / 6 and
among the different
samples. As can be seen from figure 1,
groups with high S / d 5 values are also
the most asymmetrical, and conversely,
groups with greatest symmetry show the
greatest amount of homogeneity among
non-homologous fingers. This is the more
surprising since it is not apparent in individuals on the intrasample level. Correlation coefficients between
and
S/ G r a n g e from 0.02 in Dogon females,
to
0.40 in American White females,
with most of the values falling between
0.20 and
0.35. This might suggest
that common factors are responsible for
asymmetry and inter-finger variation, but
that they tend to take one form or the other
in the individual, culminating in similar
values at the population level.
Sex differences for TRC, S e and
are shown in table 3. None of the differ-
+
+
t
Ap
I
0 Females
t
m
+
+
t
I
6.0
1
I
7.0
8.0
S/E
Fig. 1 Bivariate plot showing the relationship
between mean values of S/&and fi for each
sample (abbreviations: E.I., Easter Island; E,
English; AW, American White; AN, American
Negro; D, Dogon; BB, Bedik-Bassari; P, Pygmy).
ences is statistically significant except
those for TRC in Easter Island and English
samples. In most human populations males
have the larger ridge-counts, and that bias
exists in the present samples, except for
a
TABLE 2
Normal deviates
of
all pairwise comparisons f o r TRC, S/*
TRC
Comparison
E. Island X English
E. Island x Am. White
E. Island x Am. Negro
E. Island X Dogon
E. Island x Bedik & Bassari
E. Island x Pygmy
English X Am. White
English x Am. Negro
English x Dogon
English x Bedik & Bassari
English x Pygmy
Am. White x Am. Negro
Am. White X Dogon
Am. White x Bedik & Bassari
Am. White x Pygmy
Am. Negro x Dogon
Am. Negro x Bedik & Bassari
Am. Negro x Pygmy
Dogon x Bedik & Bassari
Dogon x Pygmy
Bedik & Bassari X Pvnmv
1
2
P < 0.05.
P < 0.01.
3P
< 0.001.
Males
4.9g3
7.35
7.35
9.403
8.59
14.123
1.83
2.29
3.37
3.61
8.11
0.64
1.57
2.08
6.76
0.74
1.36
5.55
0.81
5.61
3.91
S N 5
Females
Males
Females
5.67
5.85
5.40
6.78
6.73
7.93
0.30
0.22
1.49
0.87
3.763
0.51
1.19
0.58
3.503
1.68
1.09
3.91
0.74
2.49
3.21
2.96
1.45
2.42
1.01
3.08
4.11
1.37
4.98 3
3.99
5.60
6.85
3.55 3
2.40
4.16
5.18
1.58
0.62
1.19
2.27
3.22
0.41
1.70
1.03
0.73
0.60
0.65
2.32
0.63
2.38
0.81
.
1.89
3.67
1.72
0.26
1.40
3.10 2
1.19
0.11
1.70
1.05
2.53 2
1.27
’
~~
and
fiz
Males
2.19
1.68
2.25
0.26
2.44‘
1.69
0.46
4.55 3
2.57
4.67
1.69
3.973
2.02’
4.103
3.463
2.12’
0.26
0.64
1.13
1.53
0.89
dD
Females
2.47 ’
1.76
0.07
0.53
0.93
0.37
0.61
2.41
1.92
2.99
0.37
1.70
1.24
2.39
1.65
0.47
0.99
0.42
1.37
0.75
0.38
219
RIDGECOUNT ASYMMETRY AND DIVERSITY
the Pygmies and Bedik-Bassari. S/
and
sex differences do not parallel those
of TRC, and i t is noteworthy that in all
the Black samples, except for S / a in
Pygmies, the females exceed the males,
while the reverse is true among Easter
Islanders, English, and American Whites.
These last three are characterized by the
greatest sex differences in TRC. While no
sex difference in ~ / G o d
r F is significant, the consistent patterning suggests
that this reflects a biological reality rather
than sampling fluctuations,
and kurtosis (gz) for the
S/ s a n d A2 distributions are presented
in table 4. Both variables are characterized
by a strong positive skew and a mild leptokurtosis. Most gl values are significantly
different from zero, and all are positive.
Fewer of the g2 values attain statistical significance, but those which do are all positive. The shapes, of these distributions
contrast with those usually encountered
for TRC, which are negatively skewed and
flattened (Holt, '55; Jantz, '74).
Finally, it is of interest to examine the
relationship between TRC, S/ 6and
The correlation coefficients of TRC with
S/ G a n d
are shown in table 5. About
half attain statistical significance, and all
but those for Pygmies, Easter Island males,
and American Negro males for TRC and
are negative. The generally negative
nature of these correlations agrees with
that obtained by Holt ('58) between TRC
and S2,and between TRC and S/mfor
American Whites, American Negroes, and
Bedik and Bassari (Jantz, '74).
The contribution of each of the digits
to S/ 6and
was ascertained by computing the correlation coefficient of each
digit with the asymmetry and diversity
values. These results are set forth in figure
2. Most striking is the strong contribution
made by certain fingers, and the apparent
absence of relationship on the part of
others. There is in fact, remarkable conformity both by sex and group concerning
the attern of correlations, particularly for
S/ 5. For this variable, most groups display negative correlations on the order of
0.35-0.45 for digits I1 and 111, and correlations of approximately zero for the remainder of the digits. The Pygmies and
Easter Islanders do not show negative correlations i n digits I1 and 111, but instead
display elevations in the other digits, thus
preserving the characteristic shape of the
curve, with dips corresponding to digits
I1 and 111. It would appear then, that the
level of inter-finger diversity in a n individual results from interaction between two
groups of digits, with digits I1 and I11 behaving as one unit, and I , IV, and V as
m,
SkewnessP)
T
TABLE 3
Sex differences ( m a l e - f e m a l e ) f o r T R C , S / v 3
and diF
Easter Island
English
Am. White
Am. Negro
Dogon
Bedik-Bassari
PY gmy
1 P
ZP
+
+
+
+ 0.29
+0.70
+0.41
-0.31
- 0.30
- 0.50
+0.06
13.23 1
+17.89
8.66
1.60
8.02
- 1.40
- 1.01
+
+0.24
+0.10
4-0.18
- 0.83
-0.12
- 0.40
-0.28
< 0.05.
< 0.01.
TABLE 4
Skewness (gl) and kurtosis
(92)
for S / &
and dAZ
Skewness (gl)
s/fi
Group
E. Island
English
Am. White
A m . Negro
Dogon
Bedik-Bassari
Pygmy
1
2
3
P
P
P
< 0.05.
< 0.01.
< 0.001.
Kurtosis (g2)
vm
v=
S / G
Males
Females
Males
Females
Males
Females
0.731
0.338
0.723
0.838
0.533
0.951
0.429
0.868
0.726
0.423
1.268 3
1.226
1.106
0.409
1.1003
0.467
0.720
0.729
1.000
1.243 3
0.746
1.648
0.947
0.876
0.634 2
0.627
0.435
0.774
0.413
-0.359
0.678
0.327
0.245
0.473
0.214
1.1462
0.453
-0.415
2.621 3
1.807
0.926
0.225
Males
Females
1.619 3
6.555 3
-0.479
1.587 3
0.215
0.395
0.690 -0.158
1.278
0.226
1.662 - 0.870
0.982
1.718
220
RICHARD L. JANTZ
TABLE 5
Correlation coeficients of TRC w i t h S/dg
and d z
Group
Easter Island
English
Am. White
Am. Negro
Dogon
Bedik-Bassari
Pygmy
1P
2
TRC and S / d 5
Males
TRC and VAT
Females
Males
-0.02
-0.21' -0.03
-0.25 -0.20
-0.34 -0.10
-0.2S3 -0.05
- 0.22
- 0.13
+0.18' +0.25
+0.14
-0.12
-0.17
+0.01
+0.05
Females
-0.19
-0.06
-0.14
-0.18
- 0 . 1 6 ' -0.06
- 0.31 3 - 0.331
4-0.19 +0.02
< 0.05.
< 0.001.
P < 0.01.
3
P
the other. Disparity in ridge-counts between these groups tends to produce high
values of S/ 6
Correlation of the individual digits with
is less clearly patterned than is the
case in S / a . Here too, however, digit 111,
and to a lesser degree, digit I1 are negatively correlated with asymmetry, so that
the pattern is similar to that of S\/Js although marked by less fluctuation.
DISCUSSION
The data presented in this paper suggest
that diversity from finger to finger and
asymmetry between homologous fingers,
have considerably greater biological significance than has previously been accorded
them. In these samples the magnitude of
inte opulation differences revealed by
S / 5 and
approaches that for TRC,
but the patterning of interpopulation differences is quite different. These results
also raise some interesting questions concerning the underlying causes of asymmetry and inter-finger diversity. First, it
is useful to consider the present results in
the light of the limited number of studies
dealing with the same topic.
So far as I am aware, a study by Salzano
and Benevides ('74) constitutes the only
other instance where racial differences in
ridge-count asymmetry were sought. Their
samples, however, consisting of Brazilian
Whites, and Brazilian Blacks with various
degrees of white admixture, failed to show
any significant differences in asymmetry,
and no systematic influence of white admixture. Their samples did show generally
lower asymmetry in Negroes, and i t is per-
\;p
haps important that mean TRC values
were very similar in all of their groups.
Studies concerned with diversity among
non-homologous fingers are likewise scarce.
In a previous study using the measure of
diversity S/
I demonstrated the existence of interpopulation variation among
American Whites, American Blacks, English, and Bedik-Bassari. Sunderland and
Coope ('69) and Basu ('72) calculated
Holt's S' for Jivaro and Mysore Brahman
samples respectively, but the results are
not presented in such a way a s to facilitate
comparison with present data. With such
limited comparable information, it is sufficient simply to point to the existence of
considerable intergroup heterogeneity, and
a more general picture of the extent of
variation within and among the major
groups of man must await further investigations.
For purposes of assessing the racial significance of inter-finger diversity and asymmetry it is desirable to have information
concerning their genetic and environmental components. Such information can
be obtained from family studies, but here
too we are faced with a paucity of information. Holt's ('60) familial correlations for
S / m reveal a substantial genetic component, and interpopulation variation apparent among the present samples is
explicable in those terms. Asymmetry, however, while showing some hereditary influence, apparently has a high environmental component (Singh, '70). This is the
case as well for dental asymmetry (Bailit
et al., '70) but unlike dental asymmetry it
is not possible to suggest health, nutritional, or other environmental circumstances as responsible for dermatoglyphic
asymmetry. For example, American Negroes, whose environment is no doubt very
different from that of African Negroes, are
most similar to Africans with respect to
both
and S / G . Levels of inter-finger
diversity and asymmetry appear to be patterned along population lines rather than
related to any identifiable environmental
factor.
Is it possible to relate the observed interpopulation variation to existing hypotheses
concerning possible underlying genetic
mechanisms? Two possible explanations
suggest themselves, neither of which can
m,
22 1
RIDGE-COUNT ASYMMETRY AND DIVERSITY
+4
-4
+4
-
- Male
S/6
r
---- Female
r
English
-2
-4
-6 (
1
1
1
1
1
1
1
1
1
1
L
I
I
I
I
I
I
I
I
I
+2 r
American
White
-2
-4
c
Q)
.-
-6
,
I
/
.
,
c
.2
-2
r'\
\---
-4
~
0
,
\
-2
-4
+4
,
'\
'
, \I/
\---'
II
'I
'
,-.
.
,
\
Bedik
+6 r
Digit
Fig. 2 Correlation coefficients of individual digits with S/&
ordinate is X 10.
Digit
and
m. Scale
of the
222
RICHARD L. JANTZ
be accepted or rejected on present evidence. Holt (’58) suggested, in her study
of the relationship of S’ to TRC, that the
TRC values showing the highest Sz might
be expected to consist of individuals most
heterozygous in the polygenic system responsible for TRC. If one assumes that the
genetic mechanism responsible for ridgecounts in different populations is the same,
apart from differences in frequency of the
various alleles, then the proportion of
heterozygotes, or more accurately, the ratio
of heterozygotes to homozygotes, will vary
accordingly. Although we know little about
gene action concerning ridge-counts, it is
at least reasonable to suggest that individuals heterozygous at the most loci will display the highest levels of asymmetry or
inter-finger diversity.
Looking at the data from this standpoint,
the Pygmies, who presumably have the
highest frequency of genes resulting in low
ridge-counts, rank lowest with respect to
S / f i and
As ridge-counts increase
there is generally a concommitant rise in
these values, until they reach a maximum
in the English, followed by a decline again
in Easter Islanders, where the highest
ridge-counts are found. Easter Islanders
presumably have high frequencies of those
alleles resulting in high ridge-counts, and
thus fewer heterozygotes. Groups such a s
English or American Whites, under this
assumption, would have intermediate allelic frequencies, and as TRC increases or
decreases, so also does the fraction of
homozygotes, reflected in greater homogeneity among the ten fingers. The low
heritability just noted for asymmetry might
also be explained in these terms, since
genotypes homozygous for contrasting
pairs of alleles would be grouped into the
same phenotypic class for purposes of
computing familial correlations.
Another possibility relates to genetic control of developmental stability, or canalization. Waddington (’60) has shown that
some Drosophila genotypes possess greater
abilities to withstand environmental insults
than others, so that canalization itself is
under genetic control. Mather’s (’53) experiments with asymmetry in sternoplueral
chaetae number in Drosophila revealed
evidence for a genetic component controlling developmental asymmetry, and dif-
ferent Drosophila lines showed different
degrees of asymmetry which were independent of environment. It is quite likely
that similar mechanism exist which would
result in different degrees of developmental
asymmetry in man. As far as the present
samples are concerned, it is perhaps significant that groups showing greatest
homogeneity among digits are, or recently
were, tribal populations, where environmental stresses might be expected to be
greater than those experienced by groups
with a long history of civilization. Under
such circumstances genotypes better able
to resist environmental insults would be at
a selective advantage, resulting in greater
developmental stability for such groups.
It is not possible on present evidence, to
say which, if either, of these two hypotheses is the most tenable, and for that
matter they are not necessarily mutually
exclusive. Appropriate family material,
and additional population data are necessary before attempting to carry these ideas
any further. It is clear from present evidence however, that relationships among
digits are just as important as the number
of ridges as far as population variation is
concerned.
ACKNOWLEDGMENTS
I am grateful to the following people for
making dermatoglyphic prints or data
available to me: Dr. S. B. Holt (English),
Professor R. Gessain and Madame M.
Gessain (Bedik-Bassari), Professor J. Huizinga (Dogon and Pygmy) and Dr. R. J.
Meier (Easter Island). I am also grateful
to Dr. Roy Wallace, Superintendent of Instruction, Knoxville Public Schools; to the
principals and students of the several elementary and junior high schools, and to
the students at the University of Tennessee
for their cooperation in this study. My
appreciation for much helpful discussion
goes to Dr. Sarah Holt and Professor
C. A. B. Smith of the Galton Laboratory.
This investigation was supported by a
National Institutes of Health Fellowship
No. 1 FO GM55508-01 from the National
Institute of General Medical Sciences.
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