Deanne F. Johnson

The time window of feeding.

This paper contrasts the traditional depletion/repletion model of food intake with a longer-term perspective which focuses on function rather than mechanism. We review naturalistic observations as well as the economic relationships we have discovered in the laboratory by a cost/benefit analysis of feeding in a closed economy. We have manipulated feeding costs by means of operant methodology and have explored two classes of cost, the cost of initiating meals and the cost of consuming food. We conclude that when an animal can detect the cost/benefit structure of its habitat, its feeding behavior will tend to maximize benefits relative to cost in apparent anticipation of its nutritional requirements and environmental conditions. The time window over which these feeding decisions are integrated may be minutes, hours, months, or seasons depending upon the animals niche and current habitat. Feeding mechanisms based on momentary deficits and repletions are inadequate to explain these strategies, leaving the field of feeding mechanisms open for major discovery and revision.

Activity patterns in rats (Rattus norvegicus) as a function of the cost of access to four resources.

Patterns of eating, drinking, wheel running, and nesting were recorded in 2 experiments in which rats (Rattus norvegicus) lived in a laboratory environment that provided food, water, a running wheel, and a nest box. Access to each resource was contingent on the completion of a fixed ratio of bar presses and once earned remained available until the resource was not used for 10 consecutive min. In all cases an increase in the access price of a resource produced a decrease in the frequency with which the resource was accessed. This reduction in bout frequency was countered by an increase in bout size, which was compensatory for eating and nearly so for drinking, but which was only partially compensatory for wheel running. Nest bout size did not change significantly as nest price increased. The bout patterns of these 4 activities changed independently of one another, and the probabilities of behavioral transitions did not indicate strong links between any pairs of activities.

Development in Japanese macaques (Macaca fuscata) : Sexually dimorphic behavior during the first year of life

We have documented several sexually dimorphic patterns of behavior that develop during the first year of life in infant Japanese macaques and their mothers. Mothers treated their infants differently by sex—mothers of males broke contact with them and retrieved them more frequently than did mothers of females. And mothers of male infants moved more frequently than did mothers of female infants. Male infants played more, played in larger groups, and mounted more frequently; female infants groomed and spent more time close to other monkeys in larger social groups than did males. Female infants were also punished by other group members more frequently than were male infants. We conclude that male and female Japanese macaque infants receive differential treatment early in life by both their own mothers and other animals, and males and females in turn treat their mothers and other animals differently. There appears to be a reciprocal relationship between the behavior of infants, mothers and other social partners that contributes to the development of sexually dimorphic patterns of behavior.

Changes in rat's meal patterns as a function of the caloric density of the diet

Rats in a laboratory foraging paradigm were offered each of four diets which differed in caloric density, and intakes, meal frequencies, meal sizes, and eating rates were monitored. The rats maintained a constant daily caloric intake by eating more frequent, larger meals of the lower density foods. However, caloric meal size was not regulated, and significant correlations between meal size and the length of the post-meal interval were rarely found. The 24-hour pattern of calorie intake was the same regardless of diet. Higher-calorie foods were consumed at a faster rate within meals than were lower-calorie foods. The feeding patterns observed suggest that caloric intake may be regulated over a time frame of several meals rather than on a meal-to-meal basis.

Aggressive behavior in a confined troop of Japanese macaques: Effects of density, season, and gender

A troop of Japanese macaques (Macaca fuscata) confined in a 2-acre outdoor corral increased from 107 to 192 individuals during the 5-year tenure of a project that assessed the effects of density, season, and gender on the expression of adult aggressive behavior. Two statistical subgroups of 16 males and 28 females that were adults at the start of the project and that survived until its completion were studied intensively. There were significant season and sex differences in all groups: males were much more aggressive than females, and males were most aggressive during the fall and and winter mating season; females were most aggressive during the spring and summer birth season. Only the 16 adult males increased their frequency of aggressive behavior as the population density increased. This increase was due to the greater number of potential antagonists available each year.

Who is in Charge? Animal vs Experimenter Control

In the 1920s Curt Richter (1927) stated that the central problem for psychology was to discover the determinants of the initiation and termination of bouts of behavior. Ignoring this challenge, experimentation in animal psychology has been dominated by the session paradigm in which animals work in brief sessions for a resource of which they have been deprived. In this open economy, no behavioral strategy of the animal can meet its demand, and the beginnings and ends of bouts are controlled by the experimenter; thus, Richters problem cannot be addressed. In contrast, in a free-feeding, closed economy, the animal controls the initiation and termination of feeding and can regulate its intake, and bout patterns can be observed. If the paradigm is modified to simulate a habitat where resources are distributed discontinuously and the animal must work to discover and procure access to a commodity before it can be used, behavioral strategies allowing the animal to regulate its intake while tending to maximize the ratio of benefits to costs are revealed. We offer an answer to Richters question based on a cost/benefit analysis of feeding behavior in this foraging paradigm. We show that the time and energy costs of resource acquisition and resource consumption are powerful determinants of the pattern of resource use, and that they have different and independent effects. The former costs are reduced by reducing the frequency of initiating bouts, and the latter costs, by altering the rate and amount of consumption. Further, the time window of these relations is much longer than expected from analyses in the session paradigm. We conclude that the recurrent nature of behavior is due to the discontinuous distribution of resources rather than to cycles of physiological depletion and repletion, and that the determinants of bout initiation and termination lie in the economics of the allocation of time and effort to different resources and activities.

Development of partner preferences in Japanese Macaques (Macaca fuscata): Effects of gender and kinship during the second year of life

Quantitative data are presented on the effects of subject sex, partner sex,and kinship on the social interactions of 18 juveniles of the Oregon troop of Japanese macaques (Macaca fuscata).Data on these subjects as infants were also used to detail maturational changes in partner sex preferences. Nine males and nine females, whose multiparous mothers represented a cross section of dominance ranks, were observed using a focal-animal technique. Juveniles of both sexes engaged in more proximity, contact, grooming, mounting, aggression, and social play with kin than with nonkin partners. They initiated less contact with females and more contact with males during their second year. They initiated more grooming and aggression during their second year than their first year, with females displaying a strong preference for grooming females and males specifically aggressing males more during the second year. Aggression was higher between same-sexed partners than between opposite-sexed partners. Males engaged in more social interactions with males during the second year than the first year of life. Males played more than females during both years. Males played more with males during the second year than the first year, and males played with males more than did females during the second year. We conclude that sex differences in behavioral frequencies become evident during the first year of life, and sex differences in partner preferences emerge during the second year of life.

Social behavior of infant and mother Japanese Macaques (Macaca fuscata): Effects of kinship, partner sex, and infant sex

The behavioral interactions of 22 infant and mother Japanese macaques with other group members were studied. Focal-animal observations were made from the time of each infant’s birth until 1 year of age. Infants and mothers both displayed exceedingly strong preferences for associating with matrilineal kin and, specifically, for female kin. The degree of genetic relatedness was positively correlated with levels of spatial proximity, contact, grooming, aggression, and play. Overall frequencies of interactions with nonkin were very low, and partner sex was not an important factor in interactions with nonkin. There were no significant differences between male and female infants in interactions with kin versus nonkin. There was only one significant difference between male and female infants in interactions with males versus females: female infants showed stronger preferences for initiating proximity with females over males than did male infants. Because mothers provide the focal point for infant interactions during the first year of life, we compared the behavior of infants and mothers. Mothers were the recipients of more social interactions than were infants, mothers engaged in more grooming than did infants, and infants engaged in more social play than did mothers. These findings are only partially consistent with kin-selection theory, and the inadequacies of studying matrilineal kin discrimination to test kin selection are reviewed. The near-absence of infant sex differences in associations with social partners suggests that although maternal kin other than the mother are important to infant socialization, they probably do not contribute to the development of behavioral sex differences until after the first year of life.

Caloric regulation and patterns of food choice in a patchy environment: The value and cost of alternative foods

Rats in a laboratory foraging paradigm had 24-hr-per-day access to a feeder where they could search, by completing a fixed number of bar presses, for an opportunity to eat one of a pair of foods differing in caloric density (2.5, 3, 3.5, or 4 kcal/g) and, in Experiment 2, the price of food pellets (10 to 50 bar presses per pellet). The rats could either accept the opportunity, and eat a meal, or reject it in favor of further search. Daily caloric intake was relatively constant. The rats always included both foods in their diet, but, for any particular food, the degree of inclusion in the diet and of acceptance of meal opportunities, the meal size, and the rate of eating were all functions not only of the price and caloric value of that food but also of the price and value of the alternately-available food. The patterns of intake for one food relative to those for the other available food were strongly correlated with the relative rate of calorie intake during consumption of that food compared to the other. Although the rats appeared to be sensitive to the local rates of calorie flow, they did not maximize daily calories consumed per time spent feeding.

Patch choice and meal size of foraging rats as a function of the profitability of food

Abstract Rats lived in a laboratory environment that simulated foraging costs with bar-pressing requirements. The rats encountered sequential opportunities to eat meals in two food patches that differed in the size and/or price of food pellets. The rats accepted more opportunities and ate larger meals in the patch offering the more-profitable food, whether the larger or the lower-priced pellets. Despite widely varying patterns of intake between the patches, total daily intake was constant across most conditions. The extent to which the less-profitable food was included in the diet was a function of the size of the profitability difference. Profitability at each patch was calculated both as g earned per bar press and as g earned per min during meals. These two measures were correlated, but relative g/min was a better predictor of the relative feeding patterns. The results were not in accord with a simple, effort-minimization model of optimal foraging, although they were compatible with the notion that the cost of suboptimal behaviour will influence its occurrence.