AJP 363

American Journal of Primatology 44:43–56 (1998)

Relationship of Early Infant State Measures to Behavior

Over the First Year of Life in the Tufted Capuchin Monkey
(Cebus apella)
GAYLE BYRNE* AND STEPHEN J. SUOMI
Laboratory of Comparative Ethology, NICHD, NIH Animal Center, Poolesville, Maryland

Data on activity states were collected from 29 group-housed capuchin

monkey (Cebus apella) infants for 3 h each week from birth to 11 weeks
of age. The amounts of time spent in sleeping/drowsy, alert–quiet, and
alert–active states were measured in these subjects. Videotaped observa-
tions of these infants were recorded 3 times/week in the home cage over
the first year of life and were scored for a number of social and explor-
atory behaviors. The extent to which early infant activity state scores
predicted later behavior in the home cage was examined. Infant state
measures correlated significantly with home cage behavior during months
2–6 in that infants that had been more active in early infancy spent
more time alone, with other animals, and in exploration and play and
less time with mothers than did quieter infants. Early state measures
were less successful in predicting home cage scores beyond 8 months of
age, whereas differences in behavior attributable to housing variables
became more salient in the latter part of the first year. There was also a
negative correlation between mother and infant activity in months 2 and
3, in that more sedentary mothers tended to have more active infants.
Am. J. Primatol. 44:43–56, 1998. © 1998 Wiley-Liss, Inc.†

Key words: Cebus, capuchin; activity; infant; development; behavior

INTRODUCTION
Developmental psychologists have long looked for ways to predict childhood
behavioral tendencies from early infant measures. “Inhibited” children, at the
extreme of shyness and fearfulness, have been found to be more responsive to
sensory stimuli as neonates than others [Lagasse et al., 1989], suggesting that
their sensory arousal thresholds may be lower. Kagan and Snidman [1992] and
Calkins et al. [1996] linked differences in inhibition at 14 months to differences
in motor activity and negative affect during a standardized test at 4 months.
Infant temperament has also been related to neonatal stress reactivity. Gunnar
et al. [1995] found that greater neonatal reactivity was later associated with less
negative affect (distress to limitations) at 6 months of age.
Several studies have attempted to relate individual differences in infant ac-
tivity state profiles to later developmental outcomes. Anders et al [1985] found

*Correspondence to: Gayle Byrne, NIH Animal Center, P.O. Box 529, Poolesville, MD 20837. E-mail:
byrneg@lce.nichd.nih.gov

Received for publication 30 March 1997; revision accepted 24 September 1997.

†
© 1998 Wiley-Liss, Inc. This article is a US Government work and, as such, is in
the public domain in the United States of America.
44 / Byrne and Suomi

that temporal patterns and continuity of sleep were related to mental perfor-
mance in Bayley tests, with more mature patterns of sustained long sleep epi-
sodes associated with higher scores on the Bayley mental scale. They speculated
that regulation and sustainment of inhibition was largely determined by the
infant’s physical maturity at birth. Wakefulness, in contrast, seemed to depend
more on environmental factors than biological ones. Keener et al. [1988] linked
parental temperament reports at 6 months to objective sleep measures at that
age, especially those measures related to continuity and stability of sleep. Infants
judged as having easy temperaments had longer sleep periods and spent less
time out of their cribs at night. Fisher and Rinehart [1990] found that children
with sleep disturbances exhibited higher stress cortisol levels and were judged
as less regular in temperament. Halpern et al. [1994] also related sleep mea-
sures to later temperament: infants that spent more time awake, had less active
and more quiet sleep, and showed shorter sustained sleep periods at 3 weeks
were found to be more irritable, difficult to soothe, and less sociable and showed
fewer positive responses during a behavioral assessment at 3 months. Infants
that spent more time awake at night at 3 weeks were rated as being more irri-
table and more inhibited during the 3 month assessment. These authors suggested
that “infant sleep-wake characteristics and infant temperament reflect similar
aspects of infant biological organization” [Halpern et al. 1994:262].
Activity level is one of the most commonly used measures in assessment of
temperament; it is easy to observe and quantify, and it shows a fair amount of
reliability and continuity from infancy through childhood [Hubert et al. 1982;
Huttenen & Nyman, 1982; Worobey & Blajda, 1989]. Activity level has been shown
to have a heritable component [Saudino & Eaton, 1991], providing evidence for a
biological basis upon which an infant’s environment may act. Early differences
in activity level and state may reflect differences in response thresholds, in that
active infants may have lower thresholds of response to stimuli, whereas less
active infants respond only to more intense stimulation. The more inhibited infants
in Kagan and Snidman [1992] and Calkins et al. [1996] were theorized to have a
lower threshold of excitability of the amygdala, and they exhibited characteristi-
cally greater frontal EEG activation than other infants [Calkins et al. 1996].
The concept of temperament has been increasingly employed in studies of
individual differences in behavior and physiology in nonhuman primates (for re-
views see Higley & Suomi, 1989; Clarke & Boinski, 1995). Many of the findings
from human research have their parallels in the nonhuman primate world as
well. Early infant state organization and activity profiles would seem to offer
some promise to primatologists as outward manifestations of the biological dis-
positions with which an infant primate enters the world. These dispositions pre-
sumably affect the nature of the environment’s subsequent impact. The
environment exerts its greatest influence on an infant that is awake and atten-
tive; active infants encounter and interact with a wider range of stimuli in their
environment than do less active ones. The present study was developed to inves-
tigate whether early infant sleep patterns and activity states in capuchin mon-
keys were predictive of home cage behavior over the first year of life and whether
these measures could serve as reliable indicators of infant temperament. Although
home cage behavior in and of itself may not be a standardized measure of infant
temperament, individuals in our colony show considerable differences in levels of
activity, social, and exploratory behavior [Byrne & Suomi, 1995], the extremes of
which may serve as indicators of outgoing or inhibited temperaments such as
those seen in humans.
 Early Infant States and Behavior / 45

METHODS
Subjects
Twenty-nine tufted capuchin monkey infants (Cebus apella) from eight social
groups provided the data for this study (20 males, nine females). Seventeen (12
males, five females) lived in indoor cages (.9 × 1.8 × 1.7 m) and 12 (eight males,
four females) lived in indoor/outdoor runs during the fall and winter and in an
outdoor corncrib during the spring and summer.

Procedure
An infant activity state scoring system, first developed by Dorothy Fragaszy
[1989] for use in capuchin monkeys, was adapted for our research. Activity states
were recorded on the day of birth, day 4, week 1, and every week up until week
11. One these days, 3 separate hours of 1 min scan samples of infant and mother
activity were collected and summarized to create the measures listed in Table I.
Observations took place at 1100, 1300, and 1500 h. Interobserver reliability (per-
centage of agreements) of infant and mother activity states was assessed at 90%
at several points over the data collection period. Activity state and mother activ-
ity were expressed as a percentage of samples in which animals were scored in
each category, and weekly scores were combined to obtain monthly means (with
only the last 3 weeks included in month 3).
In addition, home cage behavior over the first year of life was videotaped in
three 10 min sessions each week. The videotapes were scored for the behaviors
listed in Table II by a single observer. Intraobserver reliability was assessed at
90% at the onset of the study. Weekly means for all these measures were ob-
tained, and monthly values were calculated from the weekly means for months
1–6 and months 8, 10, and 12.

Data analysis
Spearman rank correlations were calculated between infant state measures
in the first 3 months and monthly means of behaviors scored from videotapes over
the first year. An alpha level of .01 (two-tailed) was adopted for interpreting re-
sults, due to the large number of variables involved in the correlation analyses.

TABLE I. Infant and Mother Activity State Scoring Categories and Representative
Values Collapsed Over the First 3 Months*
Runs Cages Males Females
Mean SEM Mean SEM Mean SEM Mean SEM
Infant activity state
Sleep/drowsy 41.8 2.2 38.3 2.3 39.4 1.8 40.5 3.8
Alert–quiet 20.7 0.9 20.0 1.2 19.9 0.8 21.1 1.8
Alert–active 29.2 2.5 35.1 3.3 33.4 2.5 31.1 4.8
Mother activity
Sit 43.5 4.2 35.6 2.8 39.9 3.4 37.3 2.7
Stand 7.4 0.9 16.3 2.0 11.4 1.8 14.5 2.7
Lying down 13.1 2.5 6.1 1.8 10.5 2.2 6.2 1.2
Locomoting 11.6 1.4 21.7 2.7 15.9 2.3 20.4 3.2

*Behaviors are scored as % of samples observed in each category. Alert–quiet is analogous to Prechtl state 3
[Prechtl, 1974]; alert–active is analogous to Prechtl state 4.
46 / Byrne and Suomi
TABLE II. Behaviors Scored During Video Observations and Representative Values
Collapsed Over the First Year*
Runs Cages Males Females
Mean SEM Mean SEM Mean SEM Mean SEM
Interactions with mother
Approach/leave mother 3.0 0.3 7.3 0.9 5.1 0.7 6.4 1.4
Dorsal carry 50.7 2.4 42.1 3.0 44.5 2.7 48.1 3.3
Ventral carry 7.0 0.6 11.3 1.4 10.1 1.3 8.3 0.9
Proximity 13.3 1.0 22.8 2.0 18.4 1.9 19.7 2.6
Nonspecific contact 8.2 0.6 10.0 0.8 8.9 0.5 9.9 1.3
Grooming 3.2 0.5 4.9 0.7 4.3 0.7 3.9 0.6
Total time with mother 64.8 2.0 70.3 2.3 66.9 1.9 70.6 3.1
With other animals
Proximity 43.4 1.0 43.7 3.2 44.0 2.3 42.8 3.5
Contact 9.4 0.7 9.8 1.2 10.2 0.8 8.3 1.5
Total time with others 50.2 1.5 50.2 3.4 51.3 2.5 47.8 3.8
Alone 53.1 1.1 51.5 2.2 52.6 1.7 51.1 2.3
Explore environment 42.4 1.1 44.5 1.7 43.4 1.1 44.1 2.7
Social play 9.8 0.7 7.0 0.9 8.7 0.8 6.9 0.8
Self play 4.6 0.4 6.0 0.5 5.8 0.4 4.7 0.5
Object manipulation 2.9 0.4 3.1 0.6 2.7 0.4 3.7 0.8
(pounding/rubbing food
or objects)

*Most of these measures are arcsin percentages of observation time and are not mutually exclusive. Ap-
proaches/leaves to mothers and object manipulation were scored as frequency measures.

Byrne and Suomi [1995] described several effects of housing on home cage
behavior in these subjects. Mixed design ANOVAs were used to investigate the
effects of sex and housing on infant state and behavioral measures in the present
study. Percentage values were arcsin-transformed for use in ANOVAs [Lorenzen
& Anderson, 1993]. Sex and housing served as between subject factors and month
as a within subject factor in these analyses.
In addition, when correlations between infant state measures and home cage
behaviors were significant, multiple regressions were performed on the ranked
data to test the contributions of sex and housing to the variance in those home
cage measures. In these cases stepwise regressions were performed, which tested
the incremental variance added by sex and/or housing after the infant state mea-
sure had been entered.
Another possible influence on infant activity is the nature of mothers’ activ-
ity: infants’ ability to actively engage their environment could be influenced by
whether or not their mothers were sitting still or moving around. Thus, infant
activity scores were also correlated with maternal activity scores within each
month of observation.

RESULTS
Figures 1 and 2 present the significant rank correlations (P<.01) between
mean activity state scores for months 2 and 3 and behaviors scored from video
over the first year (none of the activity state variables in month 1 were predic-
tive of later behavior). These and the following results are presented as direc-
 Early Infant States and Behavior / 47

tions of correlations in order to illustrate the patterns that emerged in these

data; the absolute value of raw correlations that were significant ranged from
.475 to .732 and are available upon request. In Figures 1 and 2, positive cor-
relations are shown above the central axis, which represents months, and
negative ones are shown below. The number on the axis indicates in which
month the corresponding behavior was significantly correlated with the in-
fant state measure.
In general, scores indicative of lower activity levels during months 2 and 3
(sleep/drowsy, alert–quiet) were negatively correlated with measures of indepen-
dence and exploration. Conversely, more active scores were positively correlated
with those measures and negatively correlated with measures of maternal de-
pendence. The single exception to this pattern was a finding that infants who
slept/drowsed the most in month 2 played the most in month 12, a reversal of a
correlation shown in earlier months. This exception is most probably due to dif-
ferential effects of housing on behavior (discussed below).
Figure 1 illustrates the relationship between activity states and interactions
with mothers. Alert–quiet in month 2 (Fig. 1a) was negatively correlated with
nonspecific contact in month 2 and proximity and approach/leave in month 3, all
behaviors indicative of dismounting and separating from mothers. Alert–active,
in contrast, was positively correlated with contact, proximity, and approach/leave
in month 2 and negatively correlated with total time with mother in month 3
and dorsal contact in month 4. Sleep/drowsy in month 3 (Fig. 1b) was nega-
tively correlated with approach/leave in month 6. Alert–quiet in month 3 was
positively correlated with dorsal contact and total time with mothers in months
3–5. Alert–active scores in month 3 were positively correlated with proximity
to mothers in months 4 and 6 and approach/leave in month 6 and negatively
correlated with dorsal contact in months 4 and 5 and total time with mothers
in month 4.
Figure 2 describes significant rank correlations between infant state mea-
sures and other behaviors. Sleep/drowsy in month 2 (Fig. 2a) was negatively
correlated with time alone at 2 months, play in months 2 and 4 (although posi-
tively correlated with self-play at 12 months), and exploration in month 5. Alert–
quiet was negatively correlated with environmental exploration in months 3,4,
and 8 and with object manipulation in month 8. Alert–active during month 2
was positively correlated with exploration in months 2 and 5, time alone and in
self-play in month 3, and with object manipulation in month 4. Sleep/drowsy in
month 3 (Fig. 2b) was negatively correlated with exploration in months 6 and 12
and contact (and total time) with other animals in month 8. Alert–quiet in this
month was negatively correlated with time alone in months 4 and 5, self-play in
month 4, exploration in month 5, total time with others in month 6, and object
manipulation in month 8. Alert–active in month 3 was positively correlated with
contact with others in months 4 and 5, self-play in month 5, total time with
others in months 4 and 6, and exploration in months 4–8.

Sex and housing differences

ANOVAs revealed an interaction of sex × housing × month in levels of alert–
active over the first 3 months. Females in cages were less active in months 2 and
3 than any other animals (Fig. 3a) (F=3.943; 2, 42 df; P<.05). In home cage be-
havior, there was a sex × housing effect in environmental exploration (Fig. 3b):
males in runs explored slightly less than other animals (F=4.215; 1, 25 df; P=.051).
48 / Byrne and Suomi

Fig. 1. Significant (P<.01) rank correlations of infant activity state measures in month 2 (a) and month 3
(b) with interactions with mothers. Central axis indicates months; positive correlations are shown on top of
the axis, while negative ones are shown below. Numbers on the axis correspond to the month(s) in which the
corresponding behavior was correlated with the activity state measure indicated. Ap/lv, approach/leave; Con-
tact, contact with mother; Proximity, proximity to mother; Dorsal, dorsal contact with mother; Total, total
time spent mother.
 Early Infant States and Behavior / 49

Fig. 2. Significant (P<.01) rank correlations of infant activity state measures in month 2 (a) and month 3
(b) with other behaviors. Central axis indicates months; positive correlations are shown on top of the axis,
while negative ones are shown below. Numbers on the axis correspond to the month(s) in which the corre-
sponding behavior was correlated with the activity state measure indicated.
50 / Byrne and Suomi

Fig. 3. a: Scores for alert–active in months 1–3 for each sex/housing category. Vertical bars denote stan-
dard errors. b: Scores for environmental exploration collapsed over the first year for each sex/housing cat-
egory. Vertical bars denote standard errors.
 Early Infant States and Behavior / 51

Fig. 4. Housing differences in scores for approach/leave mothers (a), proximity to mothers (b), and ventral
contact with mothers (c) over the first year. Vertical bars denote standard errors.
52 / Byrne and Suomi

As in Byrne and Suomi [1995], several housing effects emerged for behavior
scores over the first year. Animals in cages approached/left mothers and were in
proximity to mothers more than those in runs in later months (Fig. 4). Percent of
time in ventral contact was higher in cages than in runs in the first 2 months.
Multiple regression analyses of those behaviors found to be correlated with
state measures revealed a significant (P<.05) contribution of housing to time alone
in month 2, proximity to mothers and dorsal contact in month 5, approach/leave
in month 6, and exploration in month 8. Sex contributed significantly to the
variance in month 4 values for dorsal contact, proximity and total time with
mothers, time alone, and self-play. Values for these variables in each of the hous-
ing conditions are shown in Table III. In all of the above cases, however, state
variables still contributed significantly to the variance in behavioral measures.

Influence of Maternal Activity

Correlation of mothers’ activity with infant states revealed that in month
1 there was no significant association between ranking of mother and infant
activity. In month 2 (Table IV), alert–active was positively associated with
mother sitting. In month 3, alert–quiet was positively correlated with mother
standing and locomoting, whereas alert–active was positively correlated with
mother sitting.

DISCUSSION
Tufted capuchin infants are typically in constant contact with mothers until
sometime in their second or third month, at which time they begin to separate
from mothers and explore their environments independently. By 6 months of
age, infants typically spend only about one-fourth of their time carried by moth-
ers, instead spending the majority of their time alone [Byrne & Suomi, 1995]. In
the present study, infants that spent more time on mothers and less time in

TABLE III. Significant Housing and Sex Differences (P<.05) Found in State–Behavior
Correlations as Revealed by Multiple Regression Analyses*

Runs Cages
Mean SEM Mean SEM

Alone month 2 6.2 2.4 1.4 0.8

Dorsal month 5 48.9 6.2 34.6 7.0
Prox month 5 17.0 3.0 28.0 3.0
Ap/lv month 6 3.5 1.4 8.4 1.2
Explore month 8 55.9 2.7 61.9 3.2
Runs Cages
Mean SEM Mean SEM
Dorsal month 4 52.1 5.6 72.4 6.0
Prox month 4 22.9 2.1 20.6 2.2
Total with mothers month 4 71.7 4.3 93.1 4.2
Time alone month 4 51.9 3.5 31.7 4.9
Self-play month 4 4.1 0.5 2.1 0.4

*Ap/lv, approach/leave mother (frequency/10 min); Prox, proximity to mother (this and following scores are
arcsin percentages of observation time, presented as means and standard errors).
 Early Infant States and Behavior / 53
TABLE IV. Correlations Between Maternal Activity and Infant State Scores in
Months 2 and 3*
Month 2 Month 3
Mother activity Sl/dr Al–qu Al–ac Sl/dr Al–qu Al–ac
Sit .087 –.096 .636a .427 –.130 .718a
Stand .108 .047 –.124 .448 .812a .144
Lie –.204 –.503 .164 –.463 –.476 .190
Locomotion .335 .516 –.429 .449 .775a .054
*Sl/dr, sleep/drowsy; Al–qu, alert–quiet; Al–ac, alert–active.
a
Spearman rank correlations significant at P<.01.

exploration during this transition period appeared to be sleepier or quieter dur-

ing activity state scoring in months 2 and 3. They slept more and spent more
time in alert–quiet and less time in an active state.
Those infants who spent more time separated from mothers, alone or with
other animals, in exploration, and play in home cage observations over the first
6 months appeared to be more awake or active during activity state scoring; for
example, they slept less and spent more time in an alert–active state.
The lack of correlation between infant state in month 1 and later behavior
may be due to the extreme immaturity of Cebus at birth. There is little variation
in the first month in activity state, with infants sleeping/drowsy about 50–65%
of the time and nursing or alert–quiet for the majority of the rest of the time
[Byrne & Suomi, 1995].
The relationship between early infant activity and behavior in the second
half of the first year was not as consistent as in the first 6 months. Examination
of housing differences in this study and in Byrne and Suomi [1995] suggests that
effects of housing became more salient in the latter half of the first year and may
obscure individual temperamental tendencies in most mundane situations. At
these later ages, when even the most inhibited of infants is becoming increas-
ingly independent, housing effects reflect differing constraints of the caging en-
vironments. Animals in small cages are less able to get away from their mothers
for extended periods. Group size in cages is usually smaller than in runs; thus,
infants tend to spend more time in self-play and environmental exploration and
less time in play with other animals. Nevertheless, the general pattern of corre-
lations established in the first 6 months continued to some extent in months 8–
12 for measures of exploration and interactions with other animals.
There were very few gender effects in levels of behavior in this study, and
those that were significant interacted with housing differences. Females in
cages tended to spend less time in alert–active but did not differ from other
animals in any other behavior. Sex differences in the present study should be
interpreted with caution, however, as male infants in our colony outnumber
females by two to one.
The link between infant activity state and later behavior is not necessarily a
causal one. Both early activity states and later behavior patterns may reflect
environmental effects (e.g., effects of prenatal variables, maternal activity, and
caretaking style, etc.). Indeed, Fragaszy [1995] reported differences in bout lengths
of states and periodicity of sleep/wake cycles under different rearing conditions
(mother-rearing and hand-rearing) in capuchins at 7–8 weeks of age. She also
reported a positive correlation between time spent in an alert–quiet state and
54 / Byrne and Suomi

maternal movement and a negative one between sleep and maternal movement.
Examination of mothers’ activity in this study suggests that, at least in older
infants, a more sedentary mother may actually promote greater activity in an
infant, whereas a more active mother, in constant motion, may discourage inde-
pendent infant activity. In our colony, the more sedentary mothers appear to be
the calmer, more equable ones, less likely to overreact to external stimuli, while
the ones in constant motion appear to be more excitable, nervous, hyperreactive
ones. If this characterization is accurate, calmer mothers would have infants
that appeared more independent, more exploratory, and less fearful, whereas
mothers that tended to be more skittish would have infants that appeared to
stay with them longer and explore less. The correspondence between mother and
infant activity is not absolute, however; in month 3, both sleep and alert–active
were correlated with mothers’ inactivity. In infants under 1 month of age, who
were only active 10–20% of the time [Byrne & Suomi, 1995], there was no signifi-
cant relationship between mother and infant activity.
Whatever infant tendencies may be, however, they must by necessity inter-
act with the existing environment. Sleepy infants end up staying closer to moth-
ers and may lose out on exploration, play experience, and contact with conspecifics.
More active infants engage in more exploration and therefore may be exposed to
more learning experiences in their social and physical environments, with all the
concomitant risks and benefits of those experiences. An interesting, varied envi-
ronment may encourage sustainment of an awake state beyond that seen in a
more monotonous one [Wolff, 1965]. Differences in reactivity affect behavior, which
in turn can affect how the environment exerts its impacts, which may then affect
future responsivity, etc.
Findings from the human literature cited earlier [e.g., Kagan & Snidman,
1992; Halpern et al. 1994] link extremely high levels of wakefulness and activity
to temperaments judged as difficult or inhibited, citing low arousal thresholds as
a possible mechanism underlying these aspects of development. In the present
study, however, increased waking activity appeared to be associated with infants
that were more exploratory and independent of mothers in early months, sug-
gesting a less fearful temperament. In the confines of this study, it is not pos-
sible to separate the effects of genetic vs. environmental contributions (i.e., calmer
mothers may have more independent infants because of a genetic disposition or
because of the differential ease of sustaining activity on a moving vs. sedentary
mother). Ongoing studies in our laboratory assessing infants’ adrenocortical re-
activity and subjective personality ratings are examining the relationship of re-
activity to early behavior and temperament in more detail.

CONCLUSIONS
1. Capuchin infants that appeared sleepier or quieter during early infant
activity state scoring (slept more, spent more time in a quiet and less time in an
active state) spent more time on mothers and less time in exploration during the
period from 2–6 months of age.
2. Conversely, those infants who appeared more awake or active during ac-
tivity state scoring (slept less, spent more time in an active state) spent more
time separated from mothers, alone or with other animals, and in exploration
and play over the first 6 months of life.
3. A relationship between early activity and later behavior was less apparent
in the second half of the first year, whereas effects of differing housing environ-
ments became more pronounced at this time.
 Early Infant States and Behavior / 55

4. At 2 and 3 months of age, infant activity was observed to vary inversely

with maternal activity to some extent. A calm, sedentary mother may facilitate
increased activity in an infant, whereas a more active mother may limit her
infant’s opportunities to explore independently.

ACKNOWLEDGMENTS
We thank Sarah Williams, Georgina Slavoff and Sarah Thrasher for assis-
tance in data collection during parts of this study and Evan Byrne for consulta-
tion on graphical presentation of data. This research was supported by the
Division of Intramural Research, National Institute of Child Health and Hu-
man Development.

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