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Individuals with major depressive disorder (MDD) can experience reduced motivation and cognitive function, leading to challenges with goal-directed behavior. When selecting goals, people maximize ‘expected value’ by selecting actions that maximize potential reward while minimizing associated costs, including effort ‘costs’ and the opportunity cost of time. In MDD, differential weighing of costs and benefits are theorized mechanisms underlying changes in goal-directed cognition and may contribute to symptom heterogeneity.
Methods
We used the Effort Foraging Task to quantify cognitive and physical effort costs, and patch leaving thresholds in low effort conditions (reflecting perceived opportunity cost of time) and investigated their shared versus distinct relationships to clinical features in participants with MDD (N = 52, 43 in-episode) and comparisons (N = 27).
Results
Contrary to our predictions, none of the decision-making measures differed with MDD diagnosis. However, each of the measures was related to symptom severity, over and above effects of ability (i.e. performance). Greater anxiety symptoms were selectively associated with lower cognitive effort cost (i.e. greater willingness to exert effort). Anhedonia and behavioral apathy were associated with increased physical effort costs. Finally, greater overall depression was related to decreased patch leaving thresholds.
Conclusions
Markers of effort-based decision-making may inform understanding of MDD heterogeneity. Increased willingness to exert cognitive effort may contribute to anxiety symptoms such as worry. Decreased leaving threshold associations with symptom severity are consistent with reward rate-based accounts of reduced vigor in MDD. Future research should address subtypes of depression with or without anxiety, which may relate differentially to cognitive effort decisions.
Edited by
Jeremy Koster, Max Planck Institute for Evolutionary Anthropology, Leipzig,Brooke Scelza, University of California, Los Angeles,Mary K. Shenk, Pennsylvania State University
Evolutionary psychology and human behavioral ecology are two complementary approaches to understanding human behavior within the evolutionary social sciences. This chapter reviews the shared histories and current states of these approaches, with an eye toward the futures of both. The chapter explore the many ways in which the theoretical and empirical approaches of evolutionary psychology and human behavioral ecology overlap and complement each other, highlighting the synergies that lie at the intersection of these approaches. The chapter also addresses the ways in which these approaches diverge in their theoretical assumptions and the phenomena that they have been used to study. Understanding these divergences can help to identify gaps in the understanding of human behavior, and it can point to ways to bridge those gaps through broadening the visions of each approach. The chapter reviews directions in which evolutionary psychology and human behavioral ecology might evolve in the coming years, including expanding the range of phenomena under study, looking to new sources of theory, and increasing diversity and representation of researchers and communities involved in research.
Edited by
Alexandre Caron, Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD), France,Daniel Cornélis, Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD) and Foundation François Sommer, France,Philippe Chardonnet, International Union for Conservation of Nature (IUCN) SSC Antelope Specialist Group,Herbert H. T. Prins, Wageningen Universiteit, The Netherlands
African buffalo herd size varies across their distribution range from as few as 5–10 in the rainforests of West and Central Africa to as many as 2000 individuals in the floodplains of eastern and southern Africa. The home range size of African buffalo also varies greatly, with those of savanna buffalo herds generally ranging between 50 and 350 km2. The larger home ranges are generally observed in areas where resources are spatially segregated, and where herds are forced to undertake seasonal movements. In contrast, forest buffalo exhibit smaller home ranges (<10 km2) due to a less pronounced seasonality of the environment, and a more homogeneous spatial arrangement of resources. African buffalo are ruminants, essentially feeding on grass and roughage. This species is capable of subsisting on pastures too coarse and too tall for most other herbivores. The African buffalo occupies an important niche, opening up habitats that are preferred by short-grass grazers. Although the African buffalo primarily is a grazer, savanna buffalo can partially switch their diet to browse when grasses become tall and lignified. The ability of the African buffalo to cope with contrasting environmental conditions throughout most sub-Saharan ecosystems, by modulating a large array of biological traits, highlights a high degree of behavioural plasticity.
Common marmosets, Callithrix jacchus, are widely used by research laboratories and are commonly provided with food in bowls. These centralised, unchallenging sources of food result in high foraging success for low foraging effort. Foraging devices, which require more skill and effort for foraging success, may broaden the behavioural profiles of marmosets by including more elements of their natural ethogram, reflecting improved welfare. The behaviour of eight female common marmosets was examined as a function of four different food distributions: food centrally located in a stationary bowl; food in a bowl that changed location each day; food centrally located in a stationary bowl, in addition to hidden food in a clustered food source (cluster feeder) or hidden food in dispersed food sources (dispersed feeders). Both the cluster and dispersed feeder distributions increased foraging, and there was a trend for reduced scratching and grooming in the presence of the feeders compared with the bowl-only treatments. The cluster feeder increased the amount of time a marmoset spent in a large room annexed to the home rooms more than the dispersed feeders, and this effect was sustained throughout the day after the feeders had been removed. Both feeders increased activity and movements within all areas of the annexed room compared with the bowl-only treatments; therefore, both feeder types improved the welfare of the captive marmosets more than the provision of food bowls.
There is a growing awareness that non-human primates kept in zoos and laboratories deserve more species-appropriate stimulation because of their biological adaptation to a challenging environment.
Numerous attempts have been made to effectively emulate the gathering and processing aspects of natural feeding. Whole natural food-items, woodchips mixed with seeds, the puzzle ceiling and the puzzle feeder stocked with ordinary biscuits, cost little or nothing but induce sustained food gathering and/or food processing. Turf and fleece substrates sprinkled with particles of flavoured food, foraging trays, probe feeders and puzzles baited with food treats also promote more foraging behaviour, but they are relatively expensive and require added labour time to load and clean them.
In the wild, chimpanzees spend most of their time foraging, so any device that stimulates this behaviour in captivity could potentially be effective enrichment. A simple grass foraging device constructed of a polyvinyl chloride (PVC) pipe cut in half lengthwise and planted with rye grass seed was designed to allow captive chimpanzees living in non-grassy enclosures to exhibit foraging similar to that of their wild counterparts. The grass containers were attached to the outside of six different enclosures. Observational data were collected on 14 adult chimpanzees (eight females, six males) within groups of either two or four members. A total of 54 hours of behavioural observations were conducted and comparisons were made across three conditions: baseline; grass container; grass container with extra foraging material (one half cup of sunflower seeds). Subjects used the grass container for 4.0 per cent of their time, but for 19.8 per cent of their time when the grass container with extra foraging material There was no statistical evidence of habituation to the device. Overall, the grass container only increased time spent foraging when it contained additional food items. Since behavioral benefits associated with this device are few, its potential application is limited.
A captive group of white-faced capuchins, Cebus capucinus, was presented with four deep litters in simultaneous choice (or preference) tests. A floor covering of ground corn cob, woodchips, wood wool or peat was presented once in each quarter of the group ‘s indoor floor-area for 14 consecutive days, and the layout of the litters was rotated after each such period. The monkeys were observed on 10 days in each period to determine the occurrence of locomotion, foraging, play, and social contact on each of the litters. The ground corn cob was clearly the least attractive floor covering for the monkeys, while peat and wood wool proved to be the most attractive. Most instances of social contact occurred on the peat, due to the occurrence of communal peat-bathing, while wood wool afforded the most play. The provision of different litter types in different areas of the indoor enclosure is a simple means of promoting a greater range of natural activities in captive primates, and probably also in other animals.
The purpose of this study was to enrich parrot enclosures by creating foraging opportunities appropriate for the species and to investigate the possible preference for a variable versus a constant food supply. The foraging device comprised of a length of wood (2×0.08×0.08m) with 50 holes (0.02m diameter x 0,02m depth) drilled into one face. Food was placed in the holes of the foraging device in one of two distributions: ‘constant’, one food item in every hole (total = 50 food items) or ‘variable’, 5 food items in 10 of the holes (total = 50 food items). The holes were then covered with starch paper. During the enrichment period the parrots spent significantly more time allopreening than in the baseline or post-enrichment periods. The results also provide some evidence of contrafreeloading in parrots, but no preference for a variable over a constant food source. The study shows that providing extra foraging opportunities for parrots is a useful form of enrichment.
The influence of an environmental enrichment programme on the searching behaviour of separate groups of male and female bush dogs at Edinburgh Zoo was evaluated. The enrichment programme involved hiding food in specially constructed wood-piles and other appropriate places in the bush dogs’ enclosures. Behavioural data were recorded morning and afternoon for 20 consecutive experimental days and were compared to pre- and post-enrichment programme data, representing basal conditions. Data were collected over a 10-period for both pre and post enrichment phases. Activities were recorded under seven behavioural categories. There was no significant difference between sexes in the proportion of time spent performing searching behaviour so data were pooled. The enrichment programme appeared to cause an increase (P < 0.01) in searching behaviour from initial basal conditions of 2.7per cent to 6.1 per cent of total recorded data points. There was a subsequent decrease in searching behaviour (P < 0.01) to 2.5 per cent when basal conditions were reinstated. All dogs showed increases in searching behaviour when enrichment and basal data were compared. The effectiveness of the enrichment programme in terms of increasing the proportion of time spent in searching behaviour showed a significant decline (P <0.005) over time, probably relating to the dogs increasing proficiency at finding food. It is suggested that the enhanced levels of searching behaviour represent an improvement in welfare.
The nose-ringing of outdoor pigs (Sus scrofa), although commonly practiced as a means to inhibit rooting behaviour and therefore reduce pasture damage and soil erosion, has been questioned on ethical grounds and alternatives are being sought. In this experiment, the effect of overground environmental enrichment was assessed as a possible alternative. 12 multiparous sows were housed in groups of four and randomly allocated to one of three treatments in a 3 × 3 Latin square design. Treatments were: 1) no environmental enrichment, 2) edible overground enrichment in the form of grass silage, and 3) inedible overground enrichment in the form of branches and tyres. Sows that received silage as overground enrichment spent significantly less time rooting the paddock (P < 0.01) than did sows on the other two treatments. The absence of a significant difference between treatments in overall foraging time budgets suggests that the manipulation of edible substrates may substitute for rooting behaviour in outdoor sows.
The welfare of caged laying hens could be improved by placing objects in the feed trough. Such objects might (a) simulate general ground-litter thus promoting more normal foraging activity and (b) give hens the opportunity to work ‘for feed - a behaviour usually thwarted in conventional cages. Spherical objects with various characteristics were placed in the feed trough of a tier of caged laying hens (n = 16). The hens pecked frequently at the objects, moving them to the trough space of adjacent cages. The mean proportion of hen heads over the trough containing these objects was significantly greater than before the objects were present (35.3 cf 32.9%) and significantly greater than the proportion of heads over a similar trough containing no objects (33.6%). Thirty days later, the mean proportions were still significantly different (33.5 cf 31.0%) showing that there was little habituation. Daily manual scattering of the objects increased the distance they were subsequently moved by the hens (23.0 cf 19.3 cm/day) indicating increased pecking activity. In a second study 12 hens were given a choice of feeding from troughs containing 0, 12 or 36 spherical objects. There was no overall preference to feed from any of the troughs. All the hens fed from troughs containing the objects, possibly indicating that the opportunity to move the objects and forage or work for feed was desired on occasions. Brightly coloured spherical objects are considered to be a promising method of successful environmental enrichment for caged laying hens. Their use to improve the welfare of caged laying hens appears to be practical and reasonably inexpensive.
Environmental enrichment by increasing foraging behaviour and providing food item choice are widely practised and generally accepted as effective methods for reducing stereotypic behaviour in captive animals. In this study, the effectiveness of increasing foraging patch choice and food item choice on reducing motor stereotypy in two captive vicugna were examined. For the purposes of the study, first, browse was added to the vicugna's enclosure as an additional forage item and, second, the vicugna's normal feed was divided: half being provided in the indoor quarters and half in the outdoor yard. The results revealed that providing browse as an additional forage item increased the observed stereotypic behaviour; however, dividing the vicugna's feed, and therefore increasing forage patch choice, decreased stereotypy. This study was limited because of the small sample size and because the area in which the vicugna were performing stereotypic behaviour was partially visually obscured. However, this study has implications for animal welfare because it highlights the need to evaluate the suitability of foraging enrichment items, and suggests that more research into accommodating the adaptive foraging behaviour of this species in captivity may be necessary.
This study is an assessment of the use of mealworm dispensers as environmental enrichment devices for Rodrigues fruit bats (Pteropus rodncensis). Captive animals frequently receive easily consumed food at set times and locations, which often minimizes the time they spend searching for and processing food. The mealworm dispensers used in this study provide an unpredictable food source, which allows the link between foraging and feeding to be reinstated. Mealworm dispensers were placed into the Rodrigues fruit bat enclosure at the Jersey Wildlife Preservation Trust and the behaviour of the bats recorded over 14 days. For 7 days the dispensers were empty but, for the remaining 7, 20 mealworms were placed in each dispenser. The number of bats feeding declined with increasing time from initial food presentation in all cases, but the presence of mealworms in the dispensers decreased the rate of decline. In addition, the number of bats active within 20cm of the food in the dishes and on the heater tops increased significantly when mealworms were present. Although the presence of mealworms had no effect on the number of flights made by the group of bats as a whole, both the number of bats on the enclosure floor and the amount of aggression observed in the enclosure decreased when mealworms were present.
Installation of mealworm dispensers meant that the bats found food items as a consequence of their natural exploratory and foraging behaviour, and as such they provided important ingredients for approximating a natural habitat and improving welfare.
The provision of foraging opportunities may be a simple way of improving an animal's welfare, but this approach has been neglected for laboratory rats (Rattus norvegicus). Standard housing contains little enrichment, and food is often provided ad libitum, which may result in inactivity and obesity, especially in mature males. Foraging enrichments may offer a way to correct these deficiencies. This study compared three potential enrichments — a limited-access hopper, gnawing sticks and a foraging device — to standard housing and feeding conditions, in order to examine their effects on rat body weight, food consumption, behaviour and preferences. The subjects were 12 mature male Wistar rats. Effects were assessed from daily weighing and from video records of the rats' behaviour over 24 h periods. The rats' preferences were determined using a four-way test system in which they could choose between a standard cage and cages offering the three potential enrichments. Compared to the standard housing and feeding, the limited-access hopper had a tendency to reduce food consumption, but the time spent feeding increased. The gnawing sticks provided the rats with the opportunity to gnaw, but did not affect other behaviours or body weight. The foraging device had the benefits of reducing aggression and allowing the rats to search for and manipulate food, but resulted in significant gains in body weight. Additionally, the foraging device was the preferred feeding source. Of the four possible feeding locations, the rats spent the least amount of time in the standard cage. The foraging device provided the most benefits but requires further modification to address problems of obesity.
An attempt was made to encourage more foraging behaviour in eight pair-housed adult rhesus macaques (Macaca mulatta). No special device and no special food were used Daily commercial dry food rations (238g per animal) consisting of 33 bar-shaped or 16 star-shaped biscuits per animal were placed on the mesh ceiling of the cages instead of in the feed-boxes. This induced an 80-fold increase (17.0 vs 1362.9s) and 289-fold increase (12.3 vs 3551.4s), respectively, in foraging time. The animals spent on average 9.6 per cent and 24.7 per cent respectively, of four-hour observation sessions foraging for biscuits from the mesh ceiling. Working for their food did not discourage them from eating all left-overs in the course of a day regardless of the shape of the biscuits.
It was concluded that the new feeding procedure enhanced the animals’ behavioural well-being by encouraging foraging activities thereby helping to counteract understimulation.
How do mushroom foragers make safe and efficient decisions under uncertainty, or deal with the genuine risks of misidentification and poisoning? This article is an inquiry into ecological rationality, heuristics, perception, and decision-making in mushroom foraging. By surveying 894 Finnish mushroom foragers, this article illustrates how socially learned rules of thumb and heuristics are used in mushroom foraging, and how simple heuristics are often complemented by more complex and intuitive decision-making. The results illustrate how traditional foraging cultures have evolved precautionary heuristics to deal with uncertainties and poisonous species, and how foragers develop selective attention through experience. The study invites us to consider whether other human foraging cultures might use heuristics similarly, how and why such traditions have culturally evolved, and whether early hunter-gatherers might have used simple heuristics to deal with uncertainty.
The welfare of captive animals is influenced by their ability to express natural behaviours. Foraging is one behaviour that may be particularly important in this respect; many species will continue to work for food even when it is freely available. The role of substrate, and in particular particle size, on the foraging behaviour of goldfish (Carassius auratus) was examined through three repeated measures experiments. In the first, tanks were set up with five uniform substrates: plastic grid, coarse sand, fine gravel, pebbles, and cobbles. In the second, fish were provided with a choice between coarse sand and fine gravel, fine gravel and pebbles, and pebbles and cobbles. In the third, they were provided with two choices between coarse sand and cobbles, one where the sand contained more food and one where the cobbles did. Our results show that particle size significantly affected the amount of time goldfish spent foraging, and that goldfish exhibited foraging behaviour even in the absence of a substrate they can manipulate. Goldfish foraged longest when provided with coarse sand. Fish foraged significantly longer over smaller particle size substrates when given a choice, although they did not distinguish between the two finest substrates, coarse sand and gravel. Increases in total time spent foraging were achieved through more, rather than longer, bouts. Food density did not significantly alter preference for smaller particle substrates. In general, coarse sand (1.5 mm) was found to be the most appropriate substrate in terms of facilitating natural foraging behaviours. These findings are discussed with respect to the welfare and husbandry of goldfish and aquarium fish in general.
While mixed-species flocks of birds (hereafter ‘flocks’) have been widely studied, few studies have looked at the effect of habitat structure on flock presence and flocking propensity within a site. Here, we employ a use-availability approach in locations with flocks and random locations to ask whether habitat characteristics influence the presence of flocks, and whether structurally similar microhabitats support compositionally similar flocks. We also examine the effect of habitat on flock size and species richness, and the effect of intraspecifically gregarious flock participants on habitat selection. We find that flocks use a narrow subset of available tree density and canopy cover variation and prefer relatively less-dense areas with large trees and a complex foliage structure. Similar microhabitats do not result in compositionally similar flocks, and while foliage complexity was associated with flock size, no habitat characteristics influenced species richness. Flocks led by the intraspecifically gregarious western crowned warbler (Phylloscopus occipitalis), a potential nuclear species, showed preference for high foliage complexity and tree density. Thus, habitat preferences of intraspecifically gregarious species, which are followed by other species, could play a strong role in habitat selection in flocks. This suggests that degraded forests that cannot provide a suitable range of tree density, canopy cover, and/or complex vegetation structure may not support some core flock species around which flocks form, which may lead to decreased flocking in those patches.
There are surprisingly few experimental studies directly comparing the cognition of primate species representing distinct phylogenetic groupings, specialized foraging ecologies, or unique social structures. Although researchers have focused on the role of foraging and social ecology in predicting cognition, they have examined social and foraging strategies in a nuanced fashion that would permit an understanding of how specific aspects of a species’ natural environment might sculpt the evolution of specific forms of cognition. In the absence of such studies, and a clear consensus as to whether cognition should best be viewed as domain-general or domain-specific suites of abilities, it is challenging to draw conclusions as to (1) cognitive differences between primate families or (2) selection pressures responsible for shaping differences. We conclude, based on paltry but accumulating evidence, that there is little utility in postulating separate physical and social domains. In addition, we see little evidence that group-living species are cognitively advantaged compared to primates that exhibit other social structures. Lastly, we advocate for greater attention to reproductive and parental strategies and individual differences in ontogenetic experiences that may color species-level comparisons.
Edited by
Irene Cogliati Dezza, University College London,Eric Schulz, Max-Planck-Institut für biologische Kybernetik, Tübingen,Charley M. Wu, Eberhard-Karls-Universität Tübingen, Germany
Information-seeking is usually conceived of as gathering information to make better decisions by observing and sampling from the external world. But for humans and many other intelligent agents, much of that information, once gathered, is also stored to guide future decisions, necessitating mechanisms for seeking information in some form of inner space. Here we survey various types of evidence suggesting that strategies adapted for search in external spatial environments are also used to seek information internally from memory. These include foraging strategies such as area-restricted search, which adaptively balances exploitation of locally clustered resources with exploration for resources more widely dispersed. We also describe how internal search satisfies the predictions of external foraging theory via the Marginal Value Theorem and show how these predictions can be used to investigate individual differences in memory search such as those caused by aging and cognitive impairment. Finally, we consider evidence that the structure of inner space may be a result of the very processes we use to search it.