Comparative analysis of weight gain,hand/wrist maturation; and dental emergence rates in chimpanzees aged 0–24 months from varying captive environments

Variability during the first 2 years of growth and development is examined in captive chimpanzees. The mixed longitudinal study of 175 animals compares curves of weight, hand/wrist maturation, and dental emergence for groups within the sample which differ in sex, rearing circumstances (mother-reared versus hand-reared), and colony (Primate Foundation of Arizona, White Sands Research Center, and The University of Texas M.D. Anderson Cancer Center Department of Veterinary Sciences in Bastrop, Texas). Comparison of LOWESS fits of the curves, using a conservative jack-knife approach, reveals trends toward significant differences between colonies for weight (with 4 comparisons reaching significance) and between rearing groups for maturation (1 reaching significance). Results of a full versus reduced model approach show the same trends, for which significance is reached in a higher number of comparisons. The latter approach also indicates possible effects of sex and environmental differences on dental emergence rate. Difficulties with both approaches are discussed. It is concluded that the results are suggestive of significant sex and environmental effects on the variables monitored, justifying further analysis and continuation of the study. The study is significant in 1) providing norms specific to sex and rearing and colony environments with which individual colony animals may be compared in the assessment of their development and in 2) providing a standard, based upon data from a larger and more varied captive chimpanzee population than previously available, with which the dental emergence status and hand/wrist maturation of fossil apes and hominids may be compared. © 1996 Wiley-Liss, Inc.     

Growth and development of infant giant pandas (Ailuropoda melanoleuca) at the Wolong Reserve,China

From 1991 to 1993 inclusive, seven infant giant pandas (Ailuropoda melanoleuca) were born at the China Conservation and Research Center for the Giant Panda. Average daily weight gain in the first 6 months for mother-reared infants (n = 5) was 71.3 g/day; for one partially mother-reared and partially hand-reared infant, 41.5 g/day; and for one completely hand-reared infant, 50.3 g/day. There was a significant difference in growth rates across the first 6 months in all methods of rearing. In addition, a comparison of growth rates across the three rearing methods showed significant differences in the first, second, third, fifth, and sixth months. Average daily body length increase for mother-reared infants was 4.1 mm/day; for partially mother-reared/partially hand-reared infants, 4.0 mm/day; and for the completely hand-reared infant, 2.8 mm/day. In mother-reared infants, body length increase during the first month was significantly greater than during the following months, and was slowest during the sixth month. At birth, infants were all pink in color with a light white coat of lanugo. Black pigmentation was first noted at 7–10 days of age, which was also the time that initial hair coat growth was seen. Eyes opened at 35–48 days of age. Ears opened at 31–50 days of age. Deciduous dentition was first seen at 82–121 days of age, while permanent dentition began to erupt at 350 days of age. © 1996 Wiley-Liss, Inc.     

Correlates of sexual and maternal competence in captive gorillas

The captive gorilla population may not be self-sustaining, in part because the overall birth rate is low and because many potential founders have failed to reproduce. We used questionnaires to collect standardized biographies of every gorilla held in North America. These biographies were searched for factors associated with reproductive success or failure. Captive-born gorillas are reproducing at least as well as wild-borns, when reproductive success is expressed as number of infants per year of reproductive opportunity. Mother-reared females are reproductively more successful than hand-reared females, but there is no difference in the reproductive success of mother-reared and hand-reared males. Social access to conspecifics in the 1st year is associated with higher reproductive success, at least for females. Reproductively successful and unsuccessful gorillas of both sexes have had equal access to potential mates, but unsuccessful animals are less likely than successful ones to exhibit normal sexual behavior. We conclude that many cases of reproductive failure are due to deficits in sexual behavior, which in turn may result from lack of early social experience with conspecifics. Some reproductive failure involves medical problems; interventive diagnostic techniques continue to be useful, if only to identify healthy individuals that can be managed intensively. The probability that a female will be a competent mother is not affected by her being wild-born or captive-born, or mother-reared or hand-reared.     

Injury risks among chimpanzees in three housing conditions

Meeting the psychological needs of chimpanzees (Pan troglodytes) can be a challenge given their aggressiveness on the one hand and the complexity of their social lives on the other. It is unclear how to balance the need to provide opportunities for species-appropriate behavior against potential risks of injury chimpanzees may inflict on each other. This study evaluates the suggestion that simpler social environments protect chimpanzees from wounding. Over a two-year period all visible injuries to 46 adult males, 64 adult females, and 25 immature chimpanzees were recorded at the Yerkes Regional Primate Research Center. Approximately half of the subjects were mother-reared, and the rest were nursery-reared. Housing included compounds containing about 20 chimpanzees, interconnected indoor-outdoor runs for groups of up to 12 individuals, and smaller indoor-outdoor runs for pairs and trios. Annual wounding rates were calculated for serious wounds (extensive injuries and all those requiring veterinary intervention) as well as for minor wounds. Compound-housed chimpanzees incurred the highest level of minor wounding, but serious wounding levels were not affected by housing condition. Even with a period of dominance instability and elevated levels of wounding in one compound, compound chimpanzees were not injured more than those in smaller social groups over the long term. Nursery-reared females in moderate-sized groups were wounded more than mother-reared females. Also, nursery-reared males and females were wounded less often when paired with mother-reared companions. Overall, this study indicates that maintaining chimpanzees in pairs and trios would not be an effective means for reducing injuries. The management of wounding in chimpanzee colonies is influenced more by the sex and rearing composition of a colony.     

Formula-fed and breast-fed baboons: Weight growth from birth to adulthood

The objective of this study was to compare the immediate and long-term effects of type of infant diet (formula-fed vs. breast-fed) on the weight growth of baboons. Longitudinal (serial) weight data were collected from 44 savannah baboons (Papio cynocephalus anubis) for the total preadult period of growth from birth to approximately 8 years of age. Fifty percent of the sample (11 females and 11 males) were fed a standard infant formula by a nursery care technician for the first 18 weeks of postnatal life. The remaining 11 females and 11 males were breast-fed by their mothers for an equivalent period of time. After 18 weeks of age the animals were removed from their infant environments (either nursery or mother-reared) and assigned to one of four age-cohort social groups. Each age-cohort group was balanced for infant treatment and gender. The animals remained in these social groups for the duration of the project and were provided nutritionally identical postweaning diets. No significant weight differences were found between the two rearing conditions for either gender during the initial 15 weeks of growth. Following this period, however, females of breast-fed environments averaged greater weight at all ages than their bottle-fed peers. The intensity of the adolescent growth spurt was also slightly greater among the female breast-fed group. No postinfant average weight-per-age disparity was observed between the male treatment groups.     

Early predictors of self-biting in socially-housed rhesus macaques (Macaca mulatta)

The development of self-biting behavior in captive monkeys is little understood and poses a serious risk to their well-being. Although early rearing conditions may influence the expression of this behavior, not all animals reared under similar conditions self-bite. The purpose of this study was to examine the effects of three rearing conditions on biting behavior and to determine whether early infant behavior can predict later self-biting. The subjects were 370 rhesus macaques born at the National Institutes of Health (NIH) Animal Center between 1994 and 2004. They were reared under three conditions: mother-reared in social groups (n=183), peer-reared in groups of four (n=84), and surrogate-peer-reared (n=103). Significantly more surrogate-peer-reared animals self-bit compared to peer-only or mother-reared animals. There was no sex difference in self-biting, but this result may have been affected by a sex bias in the number of observations. The durations of behaviors exhibited by the surrogate-peer-reared subjects were recorded in 5-min sessions twice a week from 2 to 6 months of age while the animals were in their home cages and play groups. In the play-group situation, surrogate-peer-reared subjects who later self-bit were found to be less social and exhibited less social clinging than those that did not self-bite. Home-cage behavior did not predict later self-biting, but it did change with increasing age: surrogate clinging and self-mouthing decreased, while environmental exploration increased. Our findings suggest that surrogate rearing in combination with lower levels of social contact during play may be risk factors for the later development of self-biting behavior.     

Variables influencing the origins of diverse abnormal behaviors in a large sample of captive chimpanzees (Pan troglodytes)

The developmental origin of abnormal behaviors is generally associated with early rearing environments that lack sufficient physical and sensory stimulation. However, other factors should also be considered. A large sample of captive chimpanzees (128 males and 140 females) was surveyed for the presence or absence of 18 abnormal behaviors. Origin variables included the subject's source (zoo, pet, performer, or laboratory), rearing (mother‐ or hand‐reared), and sex. Animals were assessed while held at the Primate Foundation of Arizona, University of Texas M. D. Anderson Cancer Center, or White Sands Research Center. There was a confound among origin variables; more hand‐reared animals than expected were from laboratories. Logistic regression tested the relationship of rearing and source, with sex as a secondary predictor variable, to each of the abnormal behaviors. There was no clear association between any abnormal behavior and source. However, for coprophagy, relative to animals from the laboratory, zoo animals tended to show a higher prevalence, while performers tended to show a lower prevalence (when rearing and sex were controlled). Rocking and self‐sucking were significantly more likely in hand‐reared animals. Coprophagy and depilation of self were significantly more likely in mother‐reared animals. When rearing and source were statistically controlled, the only significant sex difference was a higher prevalence of coprophagy in females and a higher prevalence of rocking in males. In a second, smaller sample of 25 males and 33 females from Southwest Foundation for Biomedical Research, no significant sex association was found for coprophagy, urophagy, rocking, or self‐depilation. In this second sample, coprophagy was also significantly more likely in mother‐reared than hand‐reared subjects. The association of some abnormal behaviors with mother‐rearing suggests that some form of social learning may be involved in the origin of some of these behavior patterns. This indicates that some abnormal behaviors may not be always be indicative of reduced psychological well‐being in captive chimpanzees. Am. J. Primatol. 48:15–29, 1999. © 1999 Wiley‐Liss, Inc.     

Development,survival, body weight,longevity, and reproductive potential of Oemena hirta (Coleoptera: Cerambycidae) under different rearing conditions

Oemona hirta (F.) is a New Zealand native longicorn beetle, whose larvae bore into the wood of branches and stems of living trees and vines, causing serious damage. To explore effective methods for maintaining laboratory colonies and biology of immature stages of this species we evaluated four laboratory rearing methods with both natural and artificial diets and compared biological parameters of laboratory colonies with those of field-collected insects. On an artificial diet, approximately 40% of neonate and 70% of autumn- and 11% of winter-collected larvae reached adulthood. Neonate larvae could not complete their development in cut poplar (Populus nigra variety italica Koehne) twigs; however, when twigs were standing in water >46% of neonate larvae survived to adulthood. Mean larval development time ranged from approximately150 to almost 300 d, depending on rearing methods. Mean pupal stage ranged from 15 to 19 d. Adult females were significantly heavier than males. Although adult females from field-collected twigs and reared on the artificial diet had similar mean body weight, which was significantly greater than that of other rearing colonies, the potential fecundity was the highest in the former (83.1 +/- 29.4 eggs) and the lowest in the latter (33.5 +/- 9.1 eggs). In all laboratory-reared and field-collected insects, the total number of eggs produced (eggs laid + eggs that remained in dead female body) by females was positively correlated with their body weight. When larvae fed on the artificial diet, there was no positive correlation between the number of eggs successfully laid and female body weight. However, when larvae lived on natural food, a positive correlation was found. In laboratory colonies, mean longevity of females (36 to 52 d) was slightly greater than that of males (30 to 50 d) without significant difference between sexes but in the adults from field-collected twigs, males (52 d) lived significantly longer than females (33 d). In terms of time, labor, and the number of resulting adults, collecting larvae in the field in autumn and then transferring them onto artificial diet is the most effective method for maintaining a laboratory colony.     

Early social experience affects behavioral and physiological responsiveness to stressful conditions in infant rhesus macaques (Macaca mulatta)

Studies on early development have demonstrated the profound effects of early social experience on the behavioral development and physiology of young rhesus macaques. Given these relationships, we hypothesized that rhesus macaques exposed to different nursery-rearing conditions may develop unique biobehavioral profiles. If this is true, the assessment of temperament may allow us to pinpoint successful rearing environments, thus improving the overall health of nonhuman primates that are raised in captive environments. We conducted biobehavioral assessments in order to examine differences in the development of infants raised under four different peer-rearing conditions (continuous pairing (CP), intermittent pairing, CP with partner rotation, and intermittent rotational pairing) and compared these animals with data from a mother-reared control group. Overall, continuous rotationally paired animals were most similar to mother-reared controls on most behavioral and temperament measures, suggesting that more socially complex rearing environments (greater number of social partners) favor a more active behavioral style. Cortisol profiles of mother-reared controls were similar to both CP groups, and these three groups had higher cortisol concentrations than the intermittent rotational-pairing group. In addition, intermittently paired infants displayed a significantly higher frequency of self-stroke behavior during a human intruder challenge, an abnormal behavior also known as floating limb which has been shown to be a precursor of self-biting. Overall, the data are consistent with the idea that social complexity in the nursery, as operationalized in our continuous rotational pairing, leads to a biobehavioral profile that is most similar to that of infants raised by their mothers in large, socially complex, cages.     

What defines successful integration into a social group for hand-reared chimpanzee infants?

Hand-rearing of captive great ape infants is sometimes necessary but can have negative behavioral consequences. Modern hand-rearing protocols, including early integration into a diverse group of conspecifics, appear to reduce the negative consequences of hand-rearing, but the process of integration is not well studied. We investigated six potential metrics of success during the introduction of two hand-reared chimpanzee infants into a troop of nine other chimpanzees at the Maryland Zoo in Baltimore. Little aggression was observed and the infants continue to be maintained in the troop at publication. As we predicted, during the introduction the hand-reared infants showed consistent levels of stress-related behaviors, participated in affiliative interactions with all available partners, and acted, received, and mutually engaged in these interactions. Solitary behaviors by these infants were similar to a mother-reared infant in the same group. Each infant also formed a relationship with a specific female that involved nest-sharing, carrying, retrieval, and intervening to reduce risk to the infant; these relationships could be classified as allomothering because they involved maternal behavior but occupied significantly less of the infants’ time than a maternal relationship. Contrary to our prediction, the hand-reared infants therefore spent significantly less time in social behavior than a mother-reared infant of the same age. In addition, the hand-reared infants continued to show strong social preferences for each other as introductions progressed and to direct a low but consistent number of nonfeeding social behaviors to humans. The successful introduction of hand-reared infants appeared to involve adding conspecific social relationships to the infants’ social repertoire, but not eliminating social interactions directed at humans.     

Hand-rearing baboons for laboratory investigations

To conduct laboratory studies in unsedated animals that were similar anatomically and physiologically to man, five full term baboons (four Papio cynocephalus anubis, one Papio cynocephalus cynocephalus) were hand-reared. These infants were used as unsedated animal models in short-term lung clearance studies conducted from birth to 2 years of age. The hand-rearing techniques described here encouraged the formation of an infant-human rearer bond that permitted us to control the level of expressed aggressive behavior as the infant matured. These techniques resulted in baboons which displayed subordinate behavior, showed positive reception to human contact (without evidence of negative stereotypic behavioral anomalies) and remained cooperative subjects for our investigations of short-term pulmonary clearance. The baboons generally were above average in weight in comparison to conspecifically-reared baboons of similar age, sex and species. Representative lung retention curves presented on one baboon demonstrate the feasibility of lung clearance studies in these hand-reared animals. Due to its suitability for unsedated studies, this baboon model may be considered for other types of laboratory investigations.     

Early rearing interacts with temperament and housing to influence the risk for motor stereotypy in rhesus monkeys (Macaca mulatta)

Laboratory and zoo housed non-human primates sometimes exhibit abnormal behaviors that are thought to reflect reduced wellbeing. Previous research attempted to identify risk factors to aid in the prevention and treatment of these behaviors, and focused on demographic (e.g. sex or age) and experience-related (e.g. single housing or nursery rearing) factors. However, not all animals that display abnormal behavior possess these risk factors and some individuals that possess a risk factor do not show behavioral abnormalities. We hypothesized that other aspects of early experience and individual characteristics might identify animals that were more likely to display one specific abnormal behavior, motor stereotypy (MS). Using logistic regression we explored the influence of early rearing (involving four different types of rearing conditions), and variation in temperament, on likelihood of displaying MS while controlling for previously identified risk factors. Analyses indicated that having a greater proportion of life lived indoors, a greater proportion of life-indoors singly-housed, and a greater number of anesthesias and blood draws significantly increased the risk of displaying MS (P < 0.001). Rearing condition failed to independently predict the display of MS; however significant interactions indicated that single housing had a greater impact on risk for indoor-reared animals versus outdoor-reared animals, and for indoor mother-reared animals versus nursery-reared animals. There were no main effects of temperament, although interactions with rearing were evident: scoring high on Gentle or Nervous was a risk factor for indoor-reared animals but not outdoor-reared animals. The final model accounted for approximately 69.3 % of the variance in the display of MS, and correctly classified 90.6% of animals. These results indicate that previously identified risk factors may impact animals differently depending on the individual's early rearing condition. These results are also the first in non-human primates to demonstrate that individual difference factors, like temperament, could be additional tools to identify animals at highest risk for motor stereotypy.     

Body Weight and Rearing Conditions of Males, Female Choice and Paternities in a Small Mammal, Cavia aperea

Patterns of mate choice may be important determinants of a species' social organisation and mating system. At least two different aspects of female mate choice can be distinguished: choice of a social partner and choice of the genetic father of the offspring. Different characteristics of males can qualify them for these two roles. Although social and reproductive partners have been shown to differ in many species, social association times are often used in laboratory choice tests to infer reproductive preferences. The traits for which females may choose partners are diverse. Body size can correlate with the male's strength in defending resources or other abilities benefiting the female and her offspring. In species living in social groups, social skills learned from group members during infancy can be important for later reproductive success. In this laboratory study, we conducted choice tests with wild cavies, Cavia aperea , a harem-living species of South American rodents, to determine social preferences of females towards two simultaneously available males. For offspring sired during these tests, paternities were determined by microsatellite DNA profiling. Males used in the tests differed in body weight and in rearing conditions: Half of the males had been reared in the presence or absence of their father, respectively. Male rearing conditions had no effect on either female social preferences or paternities. Females significantly preferred heavier males as social partners. In five of six tests, the heavier male also sired the offspring. Sires were in most cases but not consistently socially preferred. Heavier males may be preferable as social partners because they are better able to provide females with resources or have more experience in paternal care or predator avoidance as weight correlates with age. When choosing reproductive partners, females may prefer other male traits and the distribution of paternities may also be influenced by sperm competition.     

Environmental and social influences on enclosure use and activity patterns of captive sloth bears (Ursus ursinus)

Four adult sloth bears (Ursus ursinus) at the Los Angeles Zoo were studied for 13 months to quantify the influence of environmental and social variables on enclosure use and activity. Observations were conducted in 1-hr samples, using 1-min scans, for a total of 150 hr distributed across two seasons and three time periods. Data were then subjected to chi-square and log-linear analyses. These showed that the bears' enclosure use and activity patterns were conditional on variations in the physical and social environments. Individual rearing history also influenced activity, as hand-reared bears showed significantly higher frequencies of self-directed and stereotyped behaviors than did mother-reared animals. Changes in group composition significantly affected the behavior patterns of the two females, which were most social when exhibited with a familiar male. Social behavior and nonsocial activity also decreased in the presence of an unfamiliar male. Further evaluations of exhibit design and management practices are encouraged to identify variables that increase the reproductive success, physical well-being, and educational value of captive sloth bears. © 1992 Wiley-Liss, Inc.     

Mortality, body weight, food and water consumption, and clinical signs in F344/DuCrj rats in studies of chronic toxicity and carcinogenicity.

In vivo historical control data, including mortality, body weight, food and water consumption, and clinical signs in F344/DuCrj rats were obtained from 11 long-term toxicity and carcinogenicity studies conducted at the Biosafety Research Center, Foods, Drugs and Pesticides, (An-Pyo Center) during the last five years. Survival at 109 weeks of age was 80.2% (min: 74%, max: 90%) in males and 80.5% (min: 72%, max: 92%) in females. The maximum mean body weights of males and females were 443.3 +/- 15.8 g (mean +/- S. D.) and 295.7 +/- 13.3 g respectively. Male rats attained their maximum body weight at 82.6 +/- 5.3 weeks of age, the females at 103.5 +/- 2.5 weeks of age. Clinical symptoms increased with age, particularly after 84 weeks of age, and included: wasting, piloerection, palpable subcutaneous and abdominal masses, and decreased spontaneous movement. Lowered body temperature and auricular pallor occurred commonly in moribund animals. The nature and grade of toxicity in the treated animals were generally disclosed by comparing with the behavior and signs in the control animals. The use of in-house, historical control data can be useful in subsequent evaluations of chronic toxicity and carcinogenicity studies.     

Stereotypic and self-injurious behavior in rhesus macaques: a survey and retrospective analysis of environment and early experience

Abnormal behavior in captive rhesus monkeys can range from active whole-body and self-directed stereotypies to self-injurious behavior (SIB). Although abnormal behaviors are common in singly-housed rhesus monkeys, the type and frequency of these behaviors are highly variable across individual animals, and the factors influencing them are equally varied. The purpose of this investigation was to survey abnormal behavior in a large population of rhesus macaques, to characterize the relationship between stereotypies and self-injury, and to identify potential risk factors for these aberrant behaviors. Behavioral assessments of 362 individually housed rhesus monkeys were collected at the New England Regional Primate Research Center (NERPRC) and combined with colony records. Of the 362 animals surveyed, 321 exhibited at least one abnormal behavior (mean: 2.3, range: 1-8). The most common behavior was pacing. Sex differences were apparent, with males showing more abnormal behavior than females. SIB was also associated with stereotypies. Animals with a veterinary record of self-injury exhibited a greater number of self-directed stereotypies than those that did not self-injure. Housing and protocol conditions, such as individual housing at an early age, longer time housed individually, greater number of blood draws, and nursery rearing, were shown to be risk factors for abnormal behavior. Thus, many factors may influence the development and maintenance of abnormal behavior in captive primates. Some of these factors are intrinsic to the individual (e.g., sex effects), whereas others are related to colony management practices, rearing conditions, and research protocols.     

Masturbation in captive Hylobates (gibbons)

Masturbation was observed in 9 of 52 captive adult Hylobates (gibbons) over a period of 17 years at the International Center for Gibbon Studies (ICGS). Four females and 5 males masturbated with varying degrees of regularity. The behavior may be related to a disruption in early rearing experiences of Hylobates. This is the first report on masturbation in these animals and documents that this behavior occurs infrequently in captive Hylobates. © 1994 Wiley-Liss, Inc.     

Breeding and rearing squirrel monkeys (Saimiri sciureus) in captivity.

A breeding colony of squirrel monkeys (Saimiri sciureus) was established to provide animals for behavioral research concerned with early development. The origin of the initial breeders was Peru, Colombia, and Bolivia. During the past 10 years, the colony has grown to 125 adult females, 20 adult males, and 120 immature animals of various ages. The annual conception rate for the last 5 years averaged 68%. This resulted in 84% viable births of which 82% survived past 6 months of age. The majority of the births (65%) occurred during June-August, and 87% during May-September. The most efficient and successful breeding strategy was to form mixed-sexed groups of 10--15 females and 2--3 males before the mating season began and to maintain the integrity of these groups with minimal interference. Progeny were reared apart from their natural mother without difficulty, and their growth and development were found to be similar to those of mother-reared monkeys.     

Grooming and resting of otters Lutra lutra in a marine habitat

The weight gain in lactating harbour seal pups and sex-specific growth curves are described. The relationship between body length, body weight and age were derived by regression analysis based on length and age data from 365 seals, and weight values from 136 seals. The asymptotic values of the curves describing body length were 148.0 cm and 147.2 cm in females and 153.9 cm and 155.5 cm in males using Gompertz and von Bertalanffy, respectively. The corresponding body weight values were 72.8 kg and 76.7 kg in females and 90.7 kg and 88.4 kg in males.     

Early behavioral development and temperamental traits in mother- vs peer-reared rhesus monkeys

Behavioral characteristics (“traits”) of 24 mother-reared and 24 nursery/peer-reared rhesus monkey infants were assessed via rating scales from the third through the seventh month of life while housed with their rearing partners, and from months 8–10 after all animals had been placed in novel peer groups. The animals were also tested at 8 months of age on the Bayley Scales of Infant Development adapted for nonhuman primates. During the early period effects noted were primarily developmental. Peer-reared animals were rated as increasingly less cautious and more attentive to the outside environment over time, while mother-reared infants declined in ratings of dependence on their mothers. All animals were rated as increasingly active and excitable, and less fearful, over time. For the period of months 8–10, peer-reared animals showed higher confidence ratings in month 10, and all animals showed a decline in attachment to cagemates. Mother-reared animals showed more hostility to the examiner during Bayley testing, whereas peer-reared animals showed more fear. Sex effects included greater ratings for independence from mothers, greater activity over both the early and later periods, greater excitability, greater attentiveness to the extra-cage environment, less cautiousness, and better performance on one Bayley problem for females.