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Spatial and reversal learning in congeneric lizards with different foraging strategies.

Environmental demands that require intensive search for mates, food and nest sites are correlated with efficient spatial memory in many mammalian and avian species. This convergence of evidence has led to the view that spatial memory, and the neurological structures associated with it, have been selected in niches that require memory for the location of goal objects. Whether such evolutionary demands are also correlated with nonspatial abilities that require flexible use of associations similar to those required for spatial memory has not been well studied. In addition, correlations between niche types and the use of spatial or nonspatial memory have not been investigated in nonmammalian, nonavian taxa. In this study, we investigated the relationship between foraging strategies and performance on two tasks, one spatial and the other nonspatial, in congeneric lizard species: Acanthodactylus boskianus, an active forager that collects clumped sedentary prey, Acanthodactylus scutellatus, a sit-and-wait predator that collects distributed mobile prey. The two species did not differ in their performance of a spatial memory task, but A. boskianus, the active forager, performed better on the reversal of a visual discrimination, a nonspatial task. These findings question the generality of the spatial adaptation model for vertebrates. We present the pliancy hypothesis, which we developed to account for these results. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Patch time allocation and patch sampling by foraging great and blue tits.

The rate at which parents deliver energy to their brood is an important factor in avian reproduction because poor condition caused by malnutrition may reduce the offspring's survival to breeding. Models of central place foraging predict that nesting parents should optimize their prey delivery rate by minimizing travelling distances and by selecting patches where the gain per unit cost is high. I investigated the allocation of searching time amongst food patches in the home ranges of breeding great tits, Parus major, and blue tits P. caeruleus, by radiotracking. The density of locations in individual trees was positively correlated with prey biomass within trees and negatively with the distance of the trees from the nest. These two factors explained 52% of the variance in the allocation of the birds' search time. In rich patches, food was reduced considerably within 20 m of the nests, and the birds' travelling distances increased significantly during the nestling period. In parallel to foraging selectively in rich resources near the nest, the birds continually sampled the trees in their territory. The average surplus search time due to resource exploration was 1.52 times (range 1.25-1.99) the expected search time if the birds had exclusively used the most profitable patch. Despite considerable effort in patch sampling, the overall search time per unit prey was 30% better than expected by an equal use of trees. The results suggest that foraging tit parents come close to the maximum rate of prey delivery possible in a given patch distribution. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Social learning of a novel foraging patch in families of free-living Florida scrub-jays.

Free-living juvenile Florida scrub-jays, Aphelocoma coerulescens, learned to forage in a novel patch (the centre of a ring) when in proximity to other family members that foraged successfully. We were able to distinguish the contributions of social learning and of individual learning, and to show that social learning occurred. The foraging task required individual jays to dig for peanut bits (chopped fragments) buried in sand in the centre of a 33-cm plastic ring. Jays were trained in their family groups to perform the task during a summer season, and were allowed to perform the task in the presence of juveniles (aged 40-85 days) in later years. Jays living in 18 control families received partial exposure to the training situation, but received no exposure to the ring before being presented with the task in the presence of their young. Juveniles in 16 families with trained jays were able to witness demonstrations and to scrounge peanut pieces from the models as they completed the task. These 41 juveniles learned more of the task than the 33 juveniles in control families. Seven juvenile jays and two older, nonbreeding jays in the trained families completed the task at least once, whereas no jays in control families completed the task. A modified task that prevented snatching also prevented transmission of the complete task, although the 22 juveniles that observed the modified task learned more of the task than the 33 control juveniles. Further analysis indicated that demonstrations had their greatest effect in increasing the probability that juveniles would enter the ring. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Diffusion of foraging innovations in the guppy.

The way in which novel learned behaviour patterns spread through animal populations remains poorly understood, despite extensive field research and the recognition that such processes play an important role in the behavioural development, social interactions and evolution of many animal species. We conducted a series of controlled diffusions of foraging information in replicate experimental populations of the guppy, Poecilia reticulata. We presented novel foraging tasks over 15 trials to mixed-sex groups, made up of food-deprived and nonfood-deprived adults (experiment 1) or small, young fish and old, large adults (experiment 2). In these diffusions, knowledge of a route to a feeder could spread through the group by subjects learning from others, discovering the route for themselves, or, most likely, by some combination of these social and asocial learning processes. We found a striking sex difference, with novel foraging information spreading at a significantly faster rate through subgroups of females than of males. Females both discovered the goal and learned the route more quickly than males. Food-deprived individuals were faster at completing the tasks over the 15 trials than nonfood-deprived guppies, and there was a significant interaction between sex and size, with a sex difference in adults but not young individuals. There was also an interaction between sex and hunger level, with food deprivation having a stronger effect on male than female performance. We suggest that information may diffuse in a similar nonrandom or 'directed' manner through many natural populations of animals. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Experimental evidence that group foragers can converge on predicted producer-scrounger equilibria.

When foraging together, animals are often observed to feed from food discoveries of others. The producer-scrounger (PS) game predicts how frequently this phenomenon of food parasitism should occur. The game assumes: (1) at any moment all individuals can unambiguously be categorized as either playing producer (searching for undiscovered food resources) or scrounger (searching for exploitation opportunities), and (2) the payoffs received from the scrounger tactic are negatively frequency dependent; a scrounger does better than a producer when the scrounger tactic is rare, but worse when it is common. No study to date has shown that the payoffs of producer and scrounger conform to the game's assumptions or that groups of foragers reach the predicted stable equilibrium frequency (SEF) of scrounger, whereby both tactics obtain the same payoff. The current study of three captive flocks of spice finches, Lonchura punctulata, provides the first test of the PS game using an apparatus in which both assumptions of the PS game are met. The payoffs to the scrounger, measured as feeding rate (seeds/s), were highly negatively frequency dependent on the frequency of scrounger. The feeding rate for scrounger declined linearly while the rate for producer either declined only slightly or not at all with increasing scrounger frequency. When given the opportunity to alternate between tactics, the birds changed their use of each, such that the group converged on the predicted SEF of scrounger after 5-8 days of testing. Individuals in this study, therefore, demonstrated sufficient plasticity in tactic use such that the flock foraged at the SEF of scrounger. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Social influences on foraging in vertebrates: causal mechanisms and adaptive functions.

We summarize 20 years of empirical and theoretical research on causes and functions of social influences on foraging by animals. We consider separately studies of social influence on when, where, what and how to eat. Implicit in discussion of the majority of studies is our assumption that social influences on foraging reflect a biasing of individual learning processes by social stimuli rather than action of independent social-learning mechanisms. Our review of theoretical approaches suggests that the majority of formally derived hypotheses concerning functions of social influence on foraging have not yet been tested adequately and many models are in need of further refinement. We also consider the importance to the future of the field of integrating 'top-down' and 'bottom-up' approaches to the study of social learning. Copyright 2001 The Association for the Study of Animal Behaviour.

Journal Article↗

Implications of predator foraging on aphid pathogen dynamics.

The foraging behavior of starved and nonstarved second and fourth instar Coccinella septempunctata larvae on dead Acyrthosiphon pisum aphids, either infected with the entomopathogenic fungus Erynia neoaphidis (sporulating) or uninfected, was examined. Larvae searched for longer and fed less when presented with infected rather than uninfected A. pisum. Although no sporulating infected aphids were completely consumed, both adult and larval ladybirds can still be considered as intraguild predators. In a further study, fourth instar larvae fed on dying infected, dead infected (not sporulating), and dead uninfected aphids for similar periods of time but again the infected aphids were seldom entirely consumed. Live uninfected aphids were fed upon for significantly longer than any other prey. Infected aphids which were damaged at an early stage of infection (0, 1, or 2 days after inoculation) did not sporulate, whereas damaged moribund aphids (3 days after inoculation) did subsequently sporulate. Damaged sporulating cadavers continued to sporulate. However, damage to moribund and sporulating infected aphids, both mechanical or due to C. septempunctata feeding, reduced the number of conidia subsequently produced. Larval feeding caused the most significant reduction. Under laboratory conditions, C. septempunctata foraging on infected aphids did, therefore, reduce the pathogen density. However, conidia produced from a damaged cadaver resulted in levels of transmission to healthy aphids comparable to that resulting from an intact cadaver. Furthermore, the presence of a foraging adult ladybird resulted in a significant increase in transmission of the fungus to healthy aphids. Preliminary studies to assess the potential of other aphid natural enemies as intraguild predators illustrated that adults of the generalist carabid, Pterostichus madidus, entirely consumed sporulating cadavers. Third instar lacewing, Chrysoperla carnea, and hoverfly, Episyrphus balteatus, larvae never fed on sporulating cadavers. The ecological implications of these results are discussed.

Animals↗

Optimal Foraging and Predator-Prey Dynamics

A system consisting of a population of predators and two types of prey is considered. The dynamics of the system is described by differential equations with controls. The controls model how predators forage on each of the two types of prey. The choice of these controls is based on the standard assumption in the theory of optimal foraging which requires that each predator maximizes the net rate of energy intake during foraging. Since this choice depends on the densities of populations involved, this allows us to link the optimal behavior of an individual with the dynamics of the whole system. Simple qualitative analysis and some simulations show the qualitative behavior of such a system. The effect of the optimal diet choice on the stability of the system is discussed.

Journal Article↗

Optimal foraging and predator-prey dynamics, II.

In this paper we consider one-predator-two-prey population dynamics described by a control system. We study and compare conditions for permanence of the system for three types of predator feeding behaviors: (i) specialized feeding on the more profitable prey type, (ii) generalized feeding on both prey types, and (iii) optimal foraging behavior. We show that the region of parameter space leading to permanence for optimal foraging behavior is smaller than that for specialized behavior, but larger than that for generalized behavior. This suggests that optimal foraging behavior of predators may promote coexistence in predator-prey systems. We also study the effects of the above three feeding behaviors on apparent competition between the two prey types.

Animals↗

Metabolic rate during foraging in the honeybee.

The metabolic rate of free-flying honeybees, Apis mellifera ligustica, was determined by means of a novel respirometric device that allowed measurement of CO2 produced by bees foraging under controlled reward at an artificial food source. Metabolic rate increased with reward (sugar flow rate) at the food source. In addition, there was no clear-cut dependence of metabolic rate on load carried during the visit, neither as crop load nor as supplementary weights attached to the thorax. The hypothesis that metabolic rate, as well as foraging and recruiting activities, depend on the motivational state of the foraging bee determined by the reward at the food source is discussed.

Animals↗

Postnatal growth, age estimation and development of foraging behaviour in the fulvous fruit bat Rousettus leschenaulti.

This study documents the postnatal growth, age estimation and development of the foraging behaviour of the fulvous fruit bat Rousettus leschenaulti under captive conditions. At birth, the young were naked and pink with closed eyes and folded pinnae. By day four of age, their eyes had opened and the pups began to move. The mean length of forearm in 5-day-old pups was 24.9 mm and body mass was 10.8 g, equivalent to 32.3% and 14.2% of the values from postpartum females. The length of forearm and body mass increased linearly until 45 and 50 days, respectively, and thereafter maintained an apparent stability. The epiphyseal gap of the fourth metacarpal-phalangeal joint increased until 15 days, then decreased linearly until 75 days and thereafter closed. Age was estimated quantitatively, based on linear changes observed in the length of the forearm and epiphyseal gap. Pups began to roost separately, but adjacent to their mothers when 30 days old and flew clumsily when they were about 40 days old. After attaining clumsy flight, the young bats made independent foraging attempts feebly by biting and licking small fruit pieces. Young bats were engaged in suckling as well as ingesting fruits when they were about 50 days old. Between 55 and 65 days, they flew well and fed on fruits. At the age of 75 days, the young bats were completely weaned and at two months, their foraging behaviour was similar to that of their mothers. There was no significant difference in the growth pattern of the young maintained in captivity compared with those under natural conditions.

Animals↗

Behavioral genomics of honeybee foraging and nest defense.

The honeybee has been the most important insect species for study of social behavior. The recently released draft genomic sequence for the bee will accelerate honeybee behavioral genetics. Although we lack sufficient tools to manipulate this genome easily, quantitative trait loci (QTLs) that influence natural variation in behavior have been identified and tested for their effects on correlated behavioral traits. We review what is known about the genetics and physiology of two behavioral traits in honeybees, foraging specialization (pollen versus nectar), and defensive behavior, and present evidence that map-based cloning of genes is more feasible in the bee than in other metazoans. We also present bioinformatic analyses of candidate genes within QTL confidence intervals (CIs). The high recombination rate of the bee made it possible to narrow the search to regions containing only 17-61 predicted peptides for each QTL, although CIs covered large genetic distances. Knowledge of correlated behavioral traits, comparative bioinformatics, and expression assays facilitated evaluation of candidate genes. An overrepresentation of genes involved in ovarian development and insulin-like signaling components within pollen foraging QTL regions suggests that an ancestral reproductive gene network was co-opted during the evolution of foraging specialization. The major QTL influencing defensive/aggressive behavior contains orthologs of genes involved in central nervous system activity and neurogenesis. Candidates at the other two defensive-behavior QTLs include modulators of sensory signaling (Am5HT(7) serotonin receptor, AmArr4 arrestin, and GABA-B-R1 receptor). These studies are the first step in linking natural variation in honeybee social behavior to the identification of underlying genes.

Animals↗

The influence of the physical environment on the self-organised foraging patterns of ants.

Among social insects such as ants, scouts that modulate their recruiting behaviour, following simple rules based on local information, generate collective patterns of foraging. Here we demonstrate that features of the abiotic environment, specifically the foraging substrate, may also be influential in the emergence of group-level decisions such as the choice of one foraging path. Experimental data and theoretical analyses show that the collective patterns can arise independently of behavioural changes of individual scouts and can result, through self-organising processes, from the physico-chemical properties of the environment that alter the dynamics of information transfer by chemical trails.

Animals↗

Genetic variations of cell wall digestibility related traits in floral stems of Arabidopsis thaliana accessions as a basis for the improvement of the feeding value in maize and forage plants.

Floral stems of Arabidopsis thaliana accessions were used as a model system relative to forage plant stems in genetic variation studies of lignin content and cell wall digestibility related traits. Successive investigations were developed in a core collection of 24 Arabidopsis accessions and in a larger collection of 280 accessions. Significant genetic variation for lignin content in the cell wall, and for the two in vitro cell wall digestibility investigated traits, were found both in the core collection and in the large collection. Genotype x environment interactions, investigated in the core collection, were significant with a few genotypes contributing greatly to interactions, based on ecovalence value estimates. In the core collection, genotypes 42AV, 224AV, and 8AV had low cell wall digestibility values, whatever be the environmental conditions. Genotype 157AV, observed only in one environment, also appeared to have a low cell wall digestibility. Conversely, genotypes 236AV, 162AV, 70AV, 101AV, 83AV had high cell wall digestibility values, genotype 83AV having a slightly greater instability across differing environments than others. The well-known accession Col-0 (186AV) appeared with a medium level of cell wall digestibility and a weak to medium level of interaction between environments. The ranges of variation in cell wall digestibility traits were higher in the large collection than in the core collection of 24 accessions, these results needing confirmation due to the lower number of replicates. Accessions 295AV, 148AV, and 309AV could be models for low stem cell wall digestibility values, with variable lignin content. Similarly, accessions 83AV and 162AV, already identified from the study of the core collection, and five accessions (6AV, 20AV, 91AV, 114AV, and 223AV) could be models for high stem cell wall digestibility values. The large variations observed between Arabidopsis accessions for both lignin content and cell wall digestibility in floral stems have strengthened the use this species as a powerful tool for discovering genes involved in cell wall biosynthesis and lignification of dicotyledons forage plants. Investigations of this kind might also be applicable to monocotyledons forage plants due to the basic similarity of the genes involved in the lignin pathway of Angiosperms and the partial homology of the cell wall composition and organization of the mature vascular system in grasses and Arabidopsis.

Animal Feed↗

Cyclodiene insecticide, DDE, DDT, arsenic, and mercury contamination of big brown bats (Eptesicus fuscus) foraging at a Colorado Superfund site.

Rocky Mountain Arsenal (RMA) National Wildlife Area, near Denver, Colorado, is a Superfund site contaminated by past military and industrial uses, including pesticide manufacturing. From an ecosystem standpoint, the most critical contaminants at RMA are certain cyclodiene insecticides and metabolites, p,p'-DDE, p,p'-DDT, arsenic, and mercury. Bats are important ecosystem components that can be impacted by persistent contaminants because of their position in the food chain and their potential longevity and thus duration of exposure. Big brown bats (Eptesicus fuscus) were captured (n = 51) while foraging at RMA in the summers of 1997 and 1998 for determination of concentrations of contaminants of concern in carcasses, brains, and stomach contents. Adult females (n = 15) were also tracked by radiotelemetry to determine locations of nearest maternity roosts for sampling of guano for contaminant analysis and inspection for potential contaminant-induced mortality. Bats captured while foraging at RMA had measurable quantities of dieldrin and DDE in masticated insect samples from stomach contents and significantly higher concentrations of dieldrin, DDE, DDT, and mercury (juveniles) in carcasses than big brown bats (n = 26) sampled at a reference area 80 km to the north. Concentrations of dieldrin and DDE in brains of bats captured while foraging at RMA were also greater than in bats from the reference area, but not high enough to suggest mortality. Maximum concentrations of DDE, DDT, and cyclodienes in brains of big brown bats were found in adult males from RMA. Guano from the two closest known roosts had significantly higher concentrations of dieldrin, DDE, and mercury than guano from two roosts at the reference area. Dieldrin concentrations in carcasses of bats from RMA were highest in juveniles, followed by adult males and adult females. DDE concentrations in carcasses were lowest in adult females at both sites and highest in adult males at RMA. No contaminant-related mortality was obvious at the small maternity colonies near RMA. Big brown bats show higher contamination than most other mammals previously sampled at this site. Concentrations and proportions of samples with detectable residues of dieldrin in carcasses of big brown bats from RMA were similar to or exceeded reports for this species from elsewhere in the United States some 25 years ago, prior to or just following restrictions on use of this compound.

Animals↗

Octopamine modulates responsiveness to foraging-related stimuli in honey bees (Apis mellifera).

The biogenic amine neurochemical octopamine is involved in the onset of foraging behaviour in honey bees. We tested the hypothesis that octopamine influences honey bee behavioural development by modulating responsiveness to task-related stimuli. We examined the effect of octopamine treatment on responsiveness to brood pheromone (an activator of foraging) and to the presence of older bees in the colony (an inhibitor of foraging in young bees). Octopamine treatment increased responsiveness to brood pheromone and decreased responsiveness to social inhibition. These results identify octopamine both as an important source of variation in response thresholds and as a modulator of pheromonal communication in insect societies. We speculate that octopamine plays more than one role in the organisation of behavioural development indicating a very high level of integration between the neurochemical system and the generation of complex behaviour.

Aging↗

Increase in dance imprecision with decreasing foraging distance in the honey bee Apis mellifera L. is partly explained by physical constraints.

Honey bee foragers communicate the direction and distance of both food sources and new nest sites to nest mates by means of a symbolic dance language. Interestingly, the precision by which dancers transfer directional information is negatively correlated with the distance to the advertised food source. The 'tuned-error' hypothesis suggests that colonies benefit from this imprecision as it spreads recruits out over a patch of constant size irrespective of the distance to the advertised site. An alternative to the tuned-error hypothesis is that dancers are physically incapable of dancing with great precision for nearby sources. Here we revisit the tuned-error hypothesis by studying the change in dance precision with increasing foraging distance over relatively short distances while controlling for environmental influences. We show that bees indeed increase their dance precision with the increase in foraging distance. However, we also show that dance performed by swarm-scouts for a nearby (30 m) nest site, where there could be no benefit to imprecision, are either without or with only limited directional information. This result suggests that imprecision in dance communication is caused primarily by physical constraints in the ability of dancers to turn around quickly enough when the advertised site is nearby.

Animal Communication↗

Modulation of sucrose response thresholds in honey bees (Apis mellifera L.): influence of genotype, feeding, and foraging experience.

The perception of sugar is important to honey bees for making foraging decisions. We measured bees' perception by determining what concentration of sucrose touched to the antennae elicited the proboscis extension response (response threshold). A low response threshold (extension at low concentration) suggests a high perceptual value of sucrose. and vice versa. Perception of sucrose solutions differed between two artificially selected genotypic strains and was modulated by the bees' recent feeding experiences. Bees offered 10%, 30%, or 50% sucrose solutions in small cages overnight, and in large flight-cages or free-flying in the field for several days, had subsequent response thresholds positively correlated to the concentration offered. Empty bees, whether they were nectar, water or pollen foragers, dancers or non-dancers, had a significantly lower threshold than loaded bees. Crop volume affected response thresholds directly and independently of sucrose concentration. We interpret these findings as multiple mechanisms that operate in different time scales, modulating perception of sucrose. Changes occurred in the time scale of evolutionary processes as demonstrated by genotypic differences. Changes with foraging experience occur in hours or minutes while effects of crop filling are instantaneous.

Adaptation, Physiological↗