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Variation in predation risk and vole feeding behaviour: a field test of the risk allocation hypothesis.

Many prey animals experience temporal variation in the risk of predation and therefore face the problem of allocating their time between antipredator efforts and other activities like feeding and breeding. We investigated time allocation of prey animals that balanced predation risk and feeding opportunities. The predation risk allocation hypothesis predicts that animals should forage more in low- than in high-risk situations and that this difference should increase with an increasing attack ratio (i.e. difference between low- and high-risk situations) and proportion of time spent at high risk. To test these predictions we conducted a field test using bank voles ( Clethrionomys glareolus) as a prey and the least weasel ( Mustela nivalis nivalis) as a predator. The temporal pattern and intensity of predation risk were manipulated in large outdoor enclosures and the foraging effort and patch use of voles were measured by recording giving-up densities. We did not observe any variation in feeding effort due to changes in the level of risk or the proportion of time spent under high-risk conditions. The only significant effect was found when the attack ratio was altered: the foraging effort of voles was higher in the treatment with a low attack ratio than in the treatment with a high attack ratio. Thus the results did not support the predation risk allocation hypothesis and we question the applicability of the hypothesis to our study system. We argue that the deviation between the observed pattern of feeding behaviour of bank voles and that predicted by the predation risk allocation hypothesis was mostly due to the inability of voles to accurately assess the changes in the level of risk. However, we also emphasise the difficulties of testing hypotheses under outdoor conditions and with mammals capable of flexible behavioural patterns.

Animals↗

Intraguild predation and interference competition on the endangered dragonfly Aeshna viridis.

We examined the effects of intraguild predation (IGP) and interference competition on an endangered dragonfly, Aeshna viridis Eversm. (Odonata: Anisoptera). A. viridis is rare in Europe due to the decrease in suitable habitats harboring the macrophyte Stratiotes aloides L. Stratiotes plants are the principal oviposition substrate for A. viridis females and protect the larvae of A. viridis from fish predation. In our study lakes A. viridis larvae are sympatric with larvae of Aeshna grandis and Aeshna juncea. The susceptibility of A. viridis larvae to IGP by similar-sized larvae of A. grandis and A. juncea was tested in a laboratory predation experiment. Microhabitat use of A. viridis and A. grandis was studied in the laboratory to determine the possible effects of interference competition on the spatial distribution of A. viridis larvae. Our results show that at least in laboratory conditions, A. viridis is susceptible to IGP and interference competition. In competition, A. grandis larvae dominated the middle and outer portion of S. aloides rosettes whereas A. viridis stayed in the inner parts. When A. grandis larvae were absent, A. viridis colonized the middle and outer parts of the rosettes. We conclude that asymmetric predation between odonate larvae of equal size can be intense, and that both IGP and interference competition affect A. viridis. Although natural habitat complexity diminishes their impact, these interactions may nevertheless influence the distribution of A. viridis in S. aloides waters and restrict its microhabitat use in S. aloides rosettes.

Animals↗

Experimental confirmation of aggressive mimicry by a coral reef fish.

A number of potential mimetic relationships between coral reef fishes have been described, but the underlying mechanisms are poorly understood. Similarities in colour between species have often been attributed to aggressive mimicry (where predators resemble models in order to deceive prey), however this has not been tested. The fang blenny, Plagiotremus rhinorhynchos is a specialized predator that feeds on tissues of other fishes. Some individuals appear to mimic the harmless cleaner wrasse Labroides dimidiatus in order to deceive fish visiting cleaning stations, thereby increasing access to food. In this study, the ecological relationship between the mimic and model was examined at Kimbe Bay (Papua New Guinea) and the hypothesis that colour similarities represent facultative aggressive mimicry was experimentally evaluated. Some juveniles exhibited a striking resemblance to the juvenile colouration of the cleaner wrasse, but only when in close proximity to the wrasse and only when similar in size. As predicted for mimics, P. rhinorhynchos co-occurred with L. dimidiatus, but was rare relative to the model. Among site comparisons showed that the abundance of mimetic type blennies was positively correlated with the abundance of juvenile cleaner wrasses. Approximately 50% of all P. rhinorhynchos were found </=1 m from the nearest L. dimidiatus, a distance significantly shorter than expected if they were not associated. A cleaner wrasse removal experiment was carried out to test whether the colour displayed by the blenny and its foraging success were contingent upon the presence of a model. In all cases, removal of the model prompted a rapid colour change to a general non-mimetic colouration in P. rhinorhynchos. Removal of L. dimidiatus also resulted in a approximately 20% reduction in the average foraging success of the blenny compared to controls, supporting the hypothesis that the blenny is a facultative aggressive mimic of the cleaner wrasse.

Animals↗

Dynamics of demographically open mutualists: immigration, intraspecific competition, and predation impact goby populations.

Although it is now recognized that mutualistic species are common and can have stable populations, the forces controlling their persistence are poorly understood. To better understand the mechanisms that impact the stability of obligate mutualists, I conducted several field experiments within a sandy coral reef lagoon in Moorea, French Polynesia that manipulated densities of fish (gobies) that interact mutualistically with shrimp. Obligate, mutualistic partnerships of gobies and shrimp are common on Indo-Pacific coral reefs and have been shown previously to interact as follows: shrimp construct burrows in which both species reside, and gobies warn shrimp of predators through tactile communication. Augmentation of gobies by up to 100% above ambient densities within 9 m2 plots produced no change in overall density of gobies or shrimp because gobies competed intraspecifically for a limited number of shrimp burrows and smaller gobies were outcompeted by larger individuals. I used predators to assess the impact of goby removal on the stability of goby and shrimp populations. First, although surveys taken throughout the lagoon revealed no relationship between goby and predator densities, predators correlated negatively with the proportion of adult gobies and positively with the proportion of small gobies paired with large shrimp. Second, experimental augmentation of predators resulted in a dramatic reduction of adult gobies within predator-addition plots, but had no impact on overall densities as immigrants rapidly replaced the missing adult gobies. Furthermore, goby turnover resulted in an increase in the proportion of small gobies paired with large shrimp because body sizes of gobies and shrimp in a burrow were similar prior to predator introduction, and predators apparently had a greater impact on gobies than shrimp. The mechanisms that prevent expansion (intraspecific competition) and collapse (immigration) of goby-shrimp populations likely contribute to local-scale stability of mutualistic populations in other terrestrial and aquatic environments.

Animals↗

Manipulations of dietary tryptophan: effects on mouse killing and brain serotonin in the rat.

Maintaining rats on a tryptophan-free diet for 4--6 days induced mouse killing in non-killer rats, and significantly facilitated killing in killer rats, as indicated by shorter latencies to kill the mice. The killing responses were similar in topography to the natural killing responses. These changes in killing behavior did not appear to be due to generalized changes in irritability. The increased killing after maintenance on a tryptophan-free diet was accompanied by a 26% reduction in brain serotonin (5-HT) and a 29% reduction in brain 5-hydroxyindoleacetic acid (5-HIAA). When the tryptophan-free diet was supplemented with L-tryptophan (0.5 or 2%), brain 5-HT and 5-HIAA were increased above control levels, and the rat's killing response appeared normal both in terms of latency and topography, similar to that seen in control chow fed animals. While rats consumed less of the tryptophan-free and tryptophan supplemented diets, control subjects deprived of chow such that they lost as much weight as rats fed the tryptophan-free diet, did not show changes in killing behavior. These results are consistent with the hypothesis that central serotonergic systems exert inhibitory control over mouse killing behavior in rats.

Aggression↗

Afferent projections to quiet attack sites in cat hypothalamus.

Quiet biting attack by a cat on a rat was elicited by electrical stimulation of sites in the cat's lateral hypothalamus. Horseradish peroxidase was deposited at the attack sites. Cells containing reaction products were found in gyrus proreus, anterior and central medial amygdaloid nuclei, lateral and medial preoptic areas, substantia innominata, the bed nuclei of stria terminalis, and anterior commissure. The dorsomedial area of the hypothalamus, paraventricular nucleus, supramammillary region, and posterior hypothalamic area also contained reactive cells. In the midbrain the ventral tegmental area of Tsai, the dorsal and superior central nuclei of the raphe, central gray matter and interpeduncular nucleus were regions with reactive cells. In the pontine region, the locus coeruleus, parabrachial nuclei, nucleus of the lateral lemniscus, and the dorsal tegmental nucleus of Gudden all had reactive cells. There are many structures which send afferent projections to quiet attack sites located in the hypothalamus and the pontine tegmentum. The commonality of afferents to attack sites lends credence to the notion that a complex, distributed, interactive network underlies the neural basis of attack behavior.

Afferent Pathways↗

Temporal effects of castration on emotionality and shock-induced aggression in adult male rats.

Alterations in shock-induced aggressive behavior, general emotional reactivity and flinch thresholds in 61 and 82 day old male rats were analyzed at either 3 or 6 weeks following castration. Castrated animals were significantly less aggressive than sham-operated controls at the 6 week but not the 3 week interval. These findings could not be attributed to changes in flinch threshold as neither treatment nor time affected this measure. With regard to emotionality, castrated animals were significantly more reactive than the control animals at the 3 week but not 6 week interval. The data are interpreted as indicating that castration altered shock-induced aggression by preventing a maturational increase in such behavior. These behavioral alterations are compared to previously reported time-dependent changes in brain monoaminergic dynamics following gonadectomy. The conclusions support the concept of a necessary balance between catecholaminergic and serotonergic systems with regard to affective behavior.

Age Factors↗

Components of predation defense behavior in chickens: evidence for endogenous rhythmicity.

The manifestation of diurnal periodicity and the extent of its control by the photoperiod was assessed in three predation defense reactions which constitute either components or outcomes of a predator-prey interaction sequence. Two-hundred White Leghorn chicks were reared from hatching for one week in either 24L or 12L and then tested at one of two clock hours previously demonstrated to define peak and trough response for one of the components. Putative evidence was found for an endogenous source of the periodicity manifested in all reactions. Maintenance schedule did not entrain the periodicity, but simple room entry and handling elicited anti-predator reactions, the extent of which varied as a function of clock hour. A general model of predation defense behavior was proposed.

Animals↗

Mouse killing in rats: a comparison of spontaneous killers and rats with lesions of the medial hypothalamus or the medial accumbens nucleus.

Mouse killing was observed and videotaped at forty-eight hr following surgery in rats with lesions of the medial hypothalamus or the medial accumbens. The initial attacks and killing bites of the lesioned rats were directed at the anterior dorsal surface, predominantly to the regions of the neck, shoulders, and thorax and did not differ from those of spontaneous mouse killing rats. The latency to attack was significantly shorter for the lesioned animals but the time required to kill following the attack tended to be longer. Lesioned animals spent significantly more time biting the prey following the kill and left significantly more bite marks on the prey than did the spontaneous killers. When the dead prey was moved about the cage following the kill, the lesioned animals showed a significantly greater tendency to attack it than did the spontaneous killers. Following the test of mouse killing, each rat was successively exposed to a freshly killed mouse, a cotton wad, and a wood block. The lesioned animals attacked the dead mouse and the cotton wad as though they were live mice whereas the spontaneous killers did not. These results suggest that while the killing is similar for lesioned and spontaneous killers, the lesioned animals show a heightened response to the killing experience. This is manifested in an exaggeration of attack behaviors toward prey and prey-like stimulus objects following an initial killing experience.

Aggression↗

Twilight activity and antipredator behavior of young fowl housed in artificial or natural light.

Twenty-four-hr patterns of running wheel activity (Experiment 1) and death feigning, an antipredator behavior (Experiment 2), were studied in domestic chicks housed outside, in natural lighting, or indoors, with light onset and offset timed to coincide with the upper limbs of local sunrise and sunset, respectively. Although chicks housed outside were more active and displayed stronger death feigning reactions, the daily patterns of each activity were highly similar in the two groups. Activity peaked during the period corresponding to evening twilight and was negligible during the morning twilight period; in contrast, death feigning peaked during the morning twilight period. Activity measures indicated that chicks on the artificial light schedule had learned to anticipate dark by day 5, and subsequent ontogenetic changes in activity occurred exclusively in the evening twilight phase.

Aging↗

The effects of refuges on predator-prey interactions: a reconsideration.

Prey refuges are widely believed to prevent prey extinction and damp predator-prey oscillations. A review of the empirical evidence suggests that refuges are indeed capable of playing the former role. But the conditions under which they do so are not understood, nor is there any solid evidence for an effect on population fluctuations. The intuitive view that refuges act to stabilize equilibria and damp predator-prey oscillations is based in several theoretical studies of extremely simple models. Using a more realistic model, I show that several kinds of refuges can exert a locally destabilizing effect and create stable, large-amplitude oscillations which would damp out if no refuge was present. This finding contrasts sharply with the usual view. I argue that current evidence is tol weak, and the range of theoretically possible effects is too broad, to justify any simple characterization of refuge effects in nature. Manipulative empirical studies are an important first step toward correcting this situation, and I discuss some important factors to consider in their design.

Animals↗

Problem solving ability of Octopus vulgaris Lamarck (Mollusca, Cephalopoda).

Experiments presented in this study show that Octopus vulgaris Lamarck is able to open transparent glass jars closed with a plastic plug and containing a live crab (Carcinus mediterraneus). The animals remove the plus (Operandum: O) and seize the crab (Predation :P) in one single attack. The number of unsuccessful attacks appears to decrease over a series of trials (p less than .01); during the same period exploration time remains unchanged. There is a statistically significant increase in performance over trials for O (p less than .01) and P (p less than .05) mean times analyzed by single factor ANOVA, suggesting that the learning process is accomplished either by stimulus-response association or by trial and error. We propose that Octopus vulgaris is capable of learning the solutions of both problems, Operandum and Predation, thus showing a highly developed ability of "integration" of the behavioral program.

Animals↗

A neuroethological approach to hamster vision.

The contributions of the midbrain optic tectum to visuomotor behaviors likely to be important to hamsters in the wild were studied, including aperture detection, insect catching, and barrier avoidance. Following tectal undercuts, hamsters ceased to make direct approaches to apertures in the posterior 180 degrees of the visual field; this appeared to be mediated by a loss of exploratory or scanning head movements. Reorientation to and pursuit of crickets jumping out of grasp into the visual periphery was impaired, though initial approach to them was not. Barrier avoidance was unaffected by tectal undercuts. This pattern is similar to the contribution of the frog and toad optic tectum to analogous visuomotor tasks. The contribution of the tectum to searching and scanning in the hamster is an extension of the basic orienting capabilities dependent on optic tectum in anurans.

Animals↗

Behavioural responses to electrical and visual stimulation of the toad tectum.

Slow potential shifts in brain structures have been recorded and correlated with motivational state in several species. Previous studies have also found that application of an electrical current to the surface of brain tissue generates such slow potential shifts. The present study was conducted to examine if imposed dc shifts to the brain influenced motivation in the toad (Bufo bufo). Toads (B. bufo) had stimulating electrodes implanted on the surface of each optic tectum. After 1 day of recovery combined dc stimuli and a prey-like visual stimulus were presented to the animal. A current-dependent increase in prey-catching activities occurred with dc currents from 0.1 to 500 micro A and in avoidance behaviours from 50 to 500 micro A. There is also evidence of additivity of dc and visually induced negativity increasing some behaviours. The dc current was applied in order to start a movement of ions through the brain structure but more specifically through radial glia. The resulting flux of ions is thought to be responsible for the recorded slow potential shift associated with motivation and these experiments hopefully shed further light on the possible neuromodulatory role played by radial glia through the spatial buffering of potassium and the associated slow potential shifts.

Animals↗