Melinda A. Novak

A comparison of paternal behaviour in the meadow vole Microtus pennsylvanicus, the pine vole M. pinetorum and the prairie vole M. cchrogaster

Paternal care in microtines has been studied infrequently and few studies have compared patterns of direct and indirect paternal investment. The paternal behaviour of three vole species, the meadow vole (Microtus pennsylvanicus), the pine vole (M. pinetorum) and the prairie vole (M. ochrogaster) was examined in a semi-natural setting. Prairie and pine voles were found to exhibit high levels of paternal care. Prairie vole males contributed the most direct care by remaining in the natal nest for long periods of time in contact with the pups. Pine voles contributed less direct care than prairie voles as they spent less time in the natal nest with their offspring. In addition, both prairie and pine vole males were observed to groom their pups and retrieve them back to the nest area. Prairie vole males also engaged in such indirect forms of care as nest construction and maintenance, while pine voles provided indirect care in the form of tunnel construction and food caching. Meadow vole males were the least paternal of the three species and rarely engaged in either direct or indirect care. These findings support predictions that M. pennsylvanicus is promiscuous and that male and female meadow voles occupy separate territories. They are also consistent with studies which indicate that prairie and pine voles are monogamous and have a structured social organization with members interacting closely with one another.

Sex and species differences in the vasopressin innervation of sexually naive and parental prairie voles, Microtus ochrogaster and meadow voles, Microtus pennsylvanicus

To study whether central systems that are implicated in functions associated with reproduction show different changes in males and females that become parental, the central vasopressin (AVP) innervation was compared in two species of voles: prairie voles, in which males and females provide parental care, and meadow voles, in which only females provide parental care. For both species, the densities of AVP‐immunoreactive (AVP‐ir) fibers in the lateral septum, lateral habenular nucleus, medial preoptic area and paraventricular nucleus of the thalamus were compared in males and females that were sexually inexperienced or had become parents 6 days before sacrifice. The lateral septum and lateral habenular nucleus presumably receive their projections from the bed nucleus of the stria terminalis and medial amygdaloid nucleus, while the other two areas presumably receive their projections from the suprachiasmatic nucleus. Differences between sexually naive and parental animals were found only in the presumed projections of the bed nucleus of the stria terminalis and medial amygdaloid nucleus. In both species, AVP‐ir fiber densities in the lateral habenular nucleus and the lateral septum were much greater in males than in females regardless of parental state. In prairie voles, AVP‐ir fiber density in the lateral septum and lateral habenular nucleus was reduced in parental males, while no differences were found in females. In parental meadow voles, the AVP‐ir fiber density in the lateral septum did not show changes, while the fiber density in the lateral habenular nucleus was increased. The reduction in AVP‐ir fiber density in parental prairie vole males and the absence of such a reduction in meadow vole males may be related to differences in their contribution to parental care.

A COMPARISON OF MATERNAL BEHAVIOUR IN THE MEADOW VOLE (MICROTUS PENNSYLVANICUS), PRAIRIE VOLE (M. OCHROGASTER) AND PINE VOLE (M. PINETORUM)

Abstract The maternal behaviour of three species of voles was compared using a semi-naturalistic laboratory system. Meadow vole females spent more time out of the nest, and exhibited less maternal behaviour in terms of nursing and brooding, and more non-social behaviour such as locomoting, eating, and drinking, than the females of the other two species. Across all species, maternal behaviour decreased from parturition onward, while non-social behaviour increased. Paternal care was well developed in the prairie vole and pine vole, but was never observed in the meadow vole. During the first postnatal week, male prairie voles showed a tendency to enter the nest when the female left the young unattended. This trend was not apparent in the male pine vole. The physical parameters of pup development, including eye opening and the development of fur, were similar in all three species. Behavioural development, however, was most rapid in the meadow vole, intermediate in the prairie vole, and slowest in the pine vole. These results are compared with those of previous field and laboratory studies, and are discussed with reference to the life-history strategy of each species.

Minireview: Hair Cortisol: A Novel Biomarker of Hypothalamic-Pituitary-Adrenocortical Activity

Activity of the hypothalamic-pituitary-adrenocortical (HPA) axis is commonly assessed by measuring glucocorticoids such as cortisol (CORT). For many years, CORT was obtained primarily from blood plasma or urine, whereas later approaches added saliva and feces for noninvasive monitoring of HPA functioning. Despite the value of all these sample matrices for answering many research questions, they remain limited in the temporal range of assessment. Plasma and saliva are point samples that vary as a function of circadian rhythmicity and are susceptible to confounding by environmental disturbances. Even urine and feces generally assess HPA activity over a period of only 24 h or less. We and others have recently developed and validated methods for measuring the concentration of CORT in the body hair of animals (e.g. rhesus monkeys) and scalp hair of humans. CORT is constantly deposited in the growing hair shaft, as a consequence of which such deposition can serve as a biomarker of integrated HPA activity over weeks and months instead of minutes or hours. Since the advent of this methodological advance, hair CORT has already been used as an index of chronic HPA activity and stress in human clinical and nonclinical populations, in a variety of laboratory-housed and wild-living animal species, and in archival specimens that are many decades or even centuries old. Moreover, because human hair is known to grow at an average rate of about 1 cm/month, several studies suggest that CORT levels in hair segments that differ in proximity to the scalp can, under certain conditions, be used as a retrospective calendar of HPA activity during specific time periods preceding sample collection.

Cohabitation alters vasopressin innervation and paternal behavior in prairie voles (Microtus ochrogaster)

The density of vasopressin-immunoreactive (AVP-ir) fibers in the lateral septum and lateral habenular nucleus is lower in prairie vole fathers--which display paternal behavior under natural conditions-than in sexually naive males. To see if these changes occur before or after the birth of pups, and whether they are related to changes in paternal behavior, we tested paternal responsiveness and measured AVP-ir fiber density in the lateral septum, lateral habenular nucleus, medial preoptic area, and paraventricular nucleus of the thalamus of sexually naive males and females (0P) and breeding pairs that were sacrificed shortly after mating (3P); during early (13P); or late gestation (21P); or after the birth of pups (6PP). Paternal responsiveness was increased in 3P males and reached a plateau in 13P males. AVP-ir fiber density did not change in the medial preoptic area and the paraventricular nucleus of the thalamus. The fiber density in the lateral septum and lateral habenular nucleus was affected differently in males and females. Among males, 3P animals had the lowest fiber density, while 13P and 6PP animals had an intermediate, and 0P and 21P animals the highest fiber density, whereas among females, no differences in fiber density were found. A second experiment showed that the decrease in fiber density in 3P males could be induced by cohabitation with an unfamiliar female but not by an unfamiliar male nor by relocation to a novel cage. The changes in AVP-ir fiber density shortly after mating suggest that these fibers may be involved in paternal responsiveness as well as in various other social behaviors that change after mating.

Where's the ball? Two- and three-year-olds reason about unseen events.

Children 2, 2 1/2, and 3 years of age engaged in a search task in which they opened 1 of 4 doors in an occluder to retrieve a ball that had been rolled behind the occluder. The correct door was determined by a partially visible wall placed behind the occluder that stopped the motion of the unseen ball. Only the oldest group of children was able to reliably choose the correct door. All children were able to retrieve a toy that had been hidden in the same apparatus if the toy was hidden from the front by opening a door. Analysis of the younger childrens errors indicated that they did not search randomly but instead used a variety of strategies. The results are consistent with the Piagetian view that the ability to use representations to guide action develops slowly over the first years of life.

Do rhesus monkeys (Macaca mulatta) attribute knowledge and ignorance to others

The ability of 4 rhesus monkeys (Macaca mulatta) to understand the causal connection between seeing and knowing was investigated. The subjects were tested to determine if they could discriminate between information provided by experimenters who randomly alternated between roles of guesser and knower. In a series of tests, the knower either hid food under 1 of 3 cups or watched as someone else hid the food. The guesser waited outside the room or covered her or his head until the food was hidden. The subjects watched this procedure occur but could not see which cup the food was hidden under. The knower pointed to the correct cup while the guesser pointed to an incorrect one. None of the macaques provided any evidence that they realized the different states of knowledge possessed by the guesser and knower. The results are consistent with the hypothesis that rhesus macaques are incapable of making inferences about the mental states of others.

Influence of the social environment on parental behavior and pup development of meadow voles (Microtus pennsylvanicus) and prairie voles (M. ochrogaster)

Parental behavior and pup development in meadow and prairie voles (Microtus pennsylvanicus and M. ochrogaster) were examined. Social units were manipulated for the presence of fathers and of juveniles. Meadow vole mothers alone with pups showed less maternal care and spent more time resting than did mothers with juveniles or with fathers and juveniles. Pups reared only with mothers developed faster than did pups under other conditions. Number of animals in the social unit was negatively comelated with the rate of pup development

Role reversal by rhesus monkeys, but no evidence of empathy

Abstract Four adult rhesus monkeys, Macaca mulatta , were individually trained to interact with a human partner on a social task. In each macaque-human dyad, one of the participants (the informant) could see where a food reward was hidden on an apparatus, but was unable to retrieve it. The other participant (the operator) could operate the apparatus by pulling one of three handles, but could not see where the food was hidden. Two of the macaques were initially trained to be informants, and their behaviour was shaped until they adopted an unambiguous ‘pointing’ gesture in front of the correct cup. The other two were trained as operators and responded to the pointing gestures of their human partner. The subjects were trained until their performances were almost errorless. Finally, the operator-informant roles in each dyad were reversed. None of the subjects showed an immediate comprehension of their new role. This result contrasts with previous research with chimpanzees, Pan troglodytes , and indicates potentially important species differences in social intelligence.

Object permanence in orangutans (Pongo pygmaeus) and squirrel monkeys (Saimiri sciureus).

The authors tested orangutans (Pongo pygmaeus) and squirrel monkeys (Saimiri sciureus) on object permanence tasks. In Experiment 1, orangutans solved all visible displacements and most invisible displacements except those involving movements into 2 boxes successively. In Experiment 2, performance of orangutans on double invisible displacements and control displacements (assessing simple strategies) was compared. Orangutans did not use the simple strategy of selecting the box visited last by the experimenter. Instead, poorer performance on double invisible displacements may have been related to increased memory requirements. In Experiment 3, squirrel monkeys were tested using the procedure of Experiment 1. Squirrel monkeys solved visible but did not comprehend invisible displacements. Results suggest that orangutans but not squirrel monkeys possess Stage 6 object permanence capabilities.