AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 52:169-173 (1980)

Fluctuating Dental Asymmetry: A Measure of

Developmental Instability in Down Syndrome
HOWARD S. BARDEN
Uniuersity of Illinois at Chicago Circle, Chicago, Illinois 60680

KEY WORDS Down Syndrome, Dental asymmetry,

Mentally retarded, Development, Aneuploidy
ABSTRACT Subjects with Down syndrome provide a useful model for inves-
tigating the effect of chromosomal aneuploidy on developmental pathways. Studies
suggest that a major effect of trisomy is a decrease in developmental stability. The
present study examines fluctuating dental asymmetry in Down syndrome.
Mesiodistal crown diameters were measured from dental casts of 114 Down
syndrome subjects. Correlation coefficients for antimeric permanent teeth served
as an index of dental asymmetry. These values were compared with normal values
obtained from the literature.
Fluctuating dental asymmetry is thought to reflect the relative success of devel-
opmental homeostasis in countering developmental disturbances. Down syndrome
subjects have significantly increased dental asymmetry. In addition, they show a
disproportionate increase in dental asymmetry for those teeth reported to have the
least developmental stability. These results support the contention that the
chromosomal imbalance in Down syndrome results in amplified developmental
instability.

Structures of the body are influenced by both
genetic and environmental factors during
growth and development. Studies of subjects
with abnormal growth and development have
contributed substantially to our understanding
of growth processes. Recent studies (Shapiro,
'70, '75) indicate that subjects with Down syn-
drome provide a model for investigating the
effects of chromosomal aneuploidy on devel-
opmental pathways.
Amplified developmental instability is a term
used to describe the effect that the condition of
trisomy i n Down syndrome has on devel-
opmental pathways (Shapiro,'70).Morphologic
structures develop along specific devel-
opmental pathways or tracks (Waddington,
'42). Developmental tracts represent the out-
come of coadapted genetic and chromosomal
systems which have evolved because they pro-
duce phenotypes selected over time (Shapiro,
'70). Normal phenotypes are believed to be sta-
bilized within certain limits by processes
termed canalization (Waddington, '42) or de-
velopmental homeostasis (Lerner, '54; Mayr,
'63). Theoretically, developmental pathways
are buffered (canalized) so that they are rela-
tively unaffected by genetic and environmental
disturbances. Developmental pathways are
thought to differ in their stability according to
their degree of canalization (Bader, '65; Sha-
piro, '70).The extra chromosome in Down syn-
drome (trisomy 21) may upset the evolved genic
and chromosomal systems and r e s u l t i n
amplified developmental instability in Down
syndrome subjects (Shapiro, '75).
Shapiro has supported his contention of
amplified developmental instability in Down
syndrome with studies of dental morphology
(Shapiro, '701, palate (Shapiro et al., '67; Sha-
piro, '75) and dermatoglyphics (Shapiro, '75).
These studies demonstrate an increased pheno-
typic variability in Down syndrome subjects.
Importantly, those traits most affected by envi-
ronmental influences in normal subjects are
disproportionately more deviant in Down syn-
drome subjects than are more stable traits.
Recent studies have examined the relative
roles of genetic and environmental factors in
the aetiology of dental asymmetry. Because the
genetic contribution to both sides of bilaterally
symmetric organisms is assumed to be identi-
cal, the magnitude of asymmetry is thought to
Received December 14, 1978; accepted June 3, 1979.

0002-9483/80/5202-0169$01.400 1980 ALAN R. LISS, Inc. 169

170 HOWARD S. BARDEN

reflect the relative success that developmental Bp-30 perforator which punches onto paper
homeostasis has in countering developmental tape for conversion to computer tape or cards
disturbances. Some traits are thought to be bet- (Tally Corporation, Kent, Washington). This
ter buffered and thus more developmentally system eliminates errors involved in reading
stable than others (Bader, '65). Dental asym- and recording caliper measurements and in
metry is randomly distributed with respect to converting the data to a form acceptable by a
side in humans (Garnet al., '66; '67; Lundstrom computer. Measurements were carried out by
'60; Moorrees and Reed, '64). Random side dif- a n experienced technician trained by Dr. Lassi
ferences is termed fluctuating asymmetry (Van Alvesalo. Measurement errors were below
Valen, '62). 5 0.15 mm on repeated measurements.
Studies of fluctuating dental asymmetry of Discrepancies between the number of dental
laboratory animals exposed to various stressful casts measured and the sample sizes in Table 1
stimuli (Siegel and Doyle, '75a, b; Siegel et al., reflect t h e fact t h a t tooth loss among in-
'77; Siegel and Smookler, '73) and among stitutionalized subjects is frequently high and
human populations known to be exposed to dif- that many subjects had mixed dentitions. Sam-
ferences in environmental stresses (Bailit et ple sizes in Table 1refer to the number of indi-
al., '70; Doyle and Johnston, '77; Perzigian, '77) viduals studied with the bilateral presence of a
conclude that fluctuating dental asymmetry particular tooth, rather than the number of
may reflect the significant impact that envi- teeth measured.
ronmental factors have on growth and devel- Although the sample size of each tooth type
opment. Potter and Nance ('76) found little evi- was not published in the study of normal sub-
dence for a genetic basis ofdental asymmetry in jects (Moorrees and Reed, '641, the number of
humans. What constitutes an environmentally individuals ranged from 138to 193for all tooth
induced developmental disturbance, however, types except for the second molars (range:
is unclear. A recent study (DiBennardo and 8S96). The lower limits of these ranges were
Bailit, '78) indicated that dental asymmetry of used for statistical tests comparing normal
permanent teeth among Japanese children did subjects with Down syndrome subjects.
not relate to five factors that were assumed to Intra-class correlation coefficients (r) were
be relevant parameters of prenatal stress. computed between measurements of mesio-
The present study compares fluctuating den- distal diameters of antimeric teeth. The propor-
tal asymmetry in Down syndrome with that of tion of intra-individual variation due to fluctu-
normals derived from the literature (Moorrees ating asymmetry is equal to 1-r (Bailit e t al.,
and Reed, '64). Fluctuating dental asymmetry '70). Thus the lower the correlation, the greater
is used to support the contention that aneu- the asymmetry. The values of r were trans-
ploidy produces amplified developmental in- formed into Fisher's z values. Group differences
stability in Down syndrome subjects. were assessed using Student's t test. Because
correlation coefficients for the entire sample
MATERIALS AND METHODS frequently exceeded the correlation coefficients
Data were collected from dental casts ob- for either sex, they are presented for the com-
tained from two sources: a) 35 Down syndrome bined sexes after averaging the findings for the
subjects (16males, 19 females) collected by the males and females following the procedure of
author and Dr. Gerald Larson from residents a t Moorrees and Reed ('64).
Central Wisconsin Center for t h e Devel-
RESULTS
opmentally Disabled, Madison (Barden, '74);
b) 79 Down syndrome subjects (45 males, 34 Table 1 lists the correlation coefficients for
females) collected by Dr. Verner Alexandersen mesiodistal crown diameters of Down syn-
from residents of the Central Wisconsin Center drome subjects and normal subjects (Moorrees
and the Southern Wisconsin Center for the De- and Reed, '64). In comparison with normal val-
velopmentally Disabled, Union Grove, and ues, dental asymmetry in Down syndrome is
from subjects examined in private homes in the significantly greater in 10 of the 14 teeth
Madison area (Alexandersen, '69). measured. In all but one case, the distal tooth
Conventional maximum mesiodistal crown within each morphologic tooth group h a s
diameters were taken on all available perma- greater asymmetry than the mesial tooth. The
nent teeth using a n automated caliper designed one exception occurs with the lower incisor
at the University of Washington. This caliper group which shows identical asymmetry for
employs a Helios needle-point caliper in the both I2 and 11. According to Dahlberg ('511, the
RUR series with 0.05 mm readout and a Tally distal tooth within a morphologic tooth group is
 DENTAL ASYMMETRY IN DOWN SYNDROME 171

TABLE 1. Correlation Coefficientsfor Mesiodistal Crown Diameters: An In&x of Asymmetry

Tooth Down Normal' Tooth Down Normal'

Maxilla Mandible
M2 n 10 82 M2 n 15 82
r 0.50 0.95 r 0.70 0.92
2 0.55 1.83 z 0.87 1.59
tz 3.25* tz 2.33*
M1 n 56 138 M1 n 66 138
r 0.67 0.92 r 0.79 0.92
z 0.81 1.59 z 1.07 1.59
tz 4.8W tz 3.39
P2 n 25 138 P2 n 29 138
r 0.70 0.89 r 0.79 0.87
z 0.87 1.42 z 1.07 1.33
tz 2.41* tz 1.22
P1 n 36 138 P1 n 42 138
r 0.88 0.91 r 0.91 0.87
z 1.38 1.53 z 1.53 1.33
tz 0.78 tz 1.07
C n 22 138 C n 39 138
r 0.93 0.93 r 0.83 0.93
z 1.66 1.66 z 1.19 1.66
tz 0.00 tz 2.51*
I2 n 29 138 I2 n 54 138
r 0.61 0.90 r 0.85 0.93
z 0.71 1.47 z 1.26 1.66
tz 3.56* tz 2.44*
I1 n 46 138 I1 n 53 138
r 0.91 0.96 r 0.85 0.94
2 1.53 1.95 z 1.26 1.74
tz 2.39 tz 2.91*
'Indicates significance a t P<O.O5
'(from Moorrees and Reed, '64)

the least developmentally stable, with the one
exception of the mandibular incisors where I2
is more stable than 11. The present study re-
sults suggest that in Down syndrome, fluctuat-
ing dental asymmetry is disproportionately in-
creased in the least stable teeth.
DISCUSSION
The interaction of genetic information and
the local tissue environment in which a tooth is
growing is of critical importance to normal
tooth development. Physical, chemical, nutri-
tional, or other environmental disturbances
can result in abnormalities in local tissue con-
ditions which affect the stable, continuous ex-
pression of genetic information relating to
normal tooth development.
Fluctuating asymmetry, characteristically
found to some degree in all bilateral traits, is
believed to be a consequence of the interaction
of various environmental influences with poly-
genic inheritance during development. Rela-
tive developmental stability may be assessed
according to the degree of fluctuating asym-
metry present. Increased fluctuating asym-
metry indicates t h a t genetic information is rel-
atively less able than is normal to effectively
control development in the presence of disturb-
ing factors (Adams and Niswander, '67).
Previous studies (Shapiro, '71; '75) suggest
that a major effect of the trisomic condition in
Down syndrome is a decrease in developmental
stability. This contention is indirectly sup-
ported by studies which show high frequencies
of dental abnormalities in Down syndrome
subjects (Cohen, '71; Cohen and Diner, '70;
Cohen et al., '70; Kraus et al., '67; Kraus et al.,
'68; Shapiro, '70).
In the present study, fluctuating dental
asymmetry is consistently greater in Down
syndrome than in normal subjects, a finding in
agreement with a previous study (Garn et al.,
'70). Assuming that fluctuating dental asym-
metry is induced by environmental factors in-
terfering with normal dental morphogenesis
(Bailit et al., '70; DiBennardo and Bailit, '78;
Perzigian, '77; Siegel and Doyle, '75a; Siegel
and Smookler, '731, the finding of increased
fluctuating asymmetry in Down syndrome
suggests that this aneuploid condition signifi-
cantly increases t h e influence of envi-
ronmental disturbances. Similar results should
172 HOWARD S. BARDEN
be found for subjects with other chromosomal
imbalances.
Shapiro (’75) contends that Down syndrome
subjects, because of their chromosomal imbal-
ance, should show amplified developmental in-
stability in those traits which show reduced
developmental stability in normal subjects.
The principles of Butler’s morphogenetic field
concept (’39)applied to human dentition (Dahl-
berg, ’51)hold that, in general, the first tooth
(mesial)in each morphologic tooth group is the
most developmentally stable. The one excep-
tion is the mandibular incisor group.
The present study supports the contention of
amplified developmental instability in Down
syndrome. Results show that fluctuating den-
tal asymmetry is not only increased, but the
degree of increase over normal asymmetry val-
ues is greater in the least stable tooth within a
morphologic tooth group. This result was not
consistently found in a previous study (Garn et
al., ’70).Maxillary I2 is found to have one of the
greatest deviations from normal asymmetry
values, a finding which corresponds to the re-
port that the maxillary I2 is the most variable
of all the permanent teeth (Lundstrom, ’60).
Conversely, the maxillary canine, the most
stable tooth in the upper dentition (Osborne et
al., ’581, is shown to have no deviation from
normal asymmetry.
The results of this study on Down syndrome
subjects favors t h e contention t h a t chro-
mosomal aneuploidy results i n a reduction
in the stability of normal developmental path-
ways. Fluctuating dental asymmetry may
serve as a useful measure of this developmental
instability.
ACKNOWLEDGMENTS
I wish to express my appreciation to Dr.
Verner Alexandersen for the use of his dental
casts of Down syndrome subjects. Dr. Gerald
Larson gave generously of his time and exper-
tise in obtaining the remainder of the dental
casts used in this study. Without the coopera-
tion of Dr. B. Harper, Dr. John Toussaint, and
other staff members and the residents of Cen-
tral and Southern Wisconsin Centers for the
Developmentally Disabled, this study could not
have been accomplished. Special thanks go to
Ms. Helen Bennett for performing the tooth
measurements. Critical comments on t h e
manuscript were graciously provided by Dr. A.
Dahlberg, Dr. R. Osborne, and Dr. B. Shapiro.
Partial support for this work came from the
University of Illinois Research Foundation and
the University of Wisconsin Graduate Re-
search Committee. Computer services used in
the research were provided in part by the Corn-
puting Center of the University of Illinois a t
Chicago Circle.
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