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Recognition and avoidance of the odors of parasitized conspecifics and predators: differential genomic correlates.

In many species of animals chemical stimuli are an important source of information about the threats and dangers present in the social and non-social world. Olfactory cues play a fundamental role in modulating social recognition and interactions in a wide variety of mammals. Rodents, in particular, utilize chemical signals, to recognize and avoid conspecifics infected with parasites and other pathogens. Animals also respond to, and utilize, predator odor related information to assess and minimize their risk of predation. In this review, we briefly focus on the relations between odors, parasite recognition and avoidance and consider some of the associated hormonal, neural and genomic mechanisms. We describe how both male and female rodents distinguish between infected and uninfected males on the basis of odors, displaying aversive response to, and avoidance of, the urine odors of infected individuals. We further describe how the recognition and avoidance of the odors of infected individuals involves genes for the neuropeptide, oxytocin, (OT), and estrogenic mechanisms. We show that mice with deletions of the oxytocin gene (OT knockout mice (OTKO)) and mice whose genes for estrogen receptor (ER)-alpha or ER-beta [ER knockout mice (ERKO), alphaERKO and betaERKO] have been disrupted are specifically impaired in their recognition, avoidance, and memory of the odors of infected individuals. We contrast this with the recognition and display of aversive responses to predator (cat) odor that are insensitive to these genetic manipulations. These findings reveal some of the mechanisms associated with the olfactory mediated recognition of parasitized individuals and predators.

Animals↗

Lesions of structures showing FOS expression to cat presentation: effects on responsivity to a Cat, Cat odor, and nonpredator threat.

Exposure of rats to a cat elicits Fos activity in a number of brain areas or structures. Based on hodological relationships of these, Canteras has proposed a medial hypothalamic defense system, with input from several forebrain sites. Both electrolytic and neurotoxic lesions of the dorsal premammillary nucleus, which shows the strongest Fos response to cat exposure, produce striking decrements in a number of defensive behaviors to a cat or to cat odor stimuli, but do not have a major effect on either postshock freezing, or responsivity to the odor of a female in estrus. Neurotoxic lesions of the medial amygdala produce decrements in defensiveness to predator stimuli, particularly odor stimuli, that are consistent with a view of this structure as involved with allomonal cues. While dorsal hippocampal lesions had little effect on responsivity to predator stimuli, neurotoxic lesions of the ventral hippocampus reduced freezing and enhanced a variety of nondefensive behaviors to both cat odor and footshock, with similar reductions in defensiveness during context conditioning tests for cat odor, cat exposure and footshock. These results support the view that the dorsal premammillary nucleus is strongly and selectively involved in control of responsivity to predator stimuli. Structures with important input into the medial hypothalamic defense system appear also to be functionally involved with antipredator defensive behaviors, and these lesion studies may suggest specific hypotheses as to the particular defense functions of different areas.

Animals↗

Behavioral choices based on patch selection: a model using aggregation methods.

The aim of this work is to study the influence of patch selection on the dynamics of a system describing the interactions between two populations, generically called 'population N' and 'population P'. Our model may be applied to prey-predator systems as well as to certain host-parasite or parasitoid systems. A situation in which population P affects the spatial distribution of population N is considered. We deal with a heterogeneous environment composed of two spatial patches: population P lives only in patch 1, while individuals belonging to population N migrate between patch 1 and patch 2, which may be a refuge. Therefore they are divided into two patch sub-populations and can migrate according to different migration laws. We make the assumption that the patch change is fast, whereas the growth and interaction processes are slower. We take advantage of the two time scales to perform aggregation methods in order to obtain a global model describing the time evolution of the total populations, at a slow time scale. At first, a migration law which is independent on population P density is considered. In this case the global model is equivalent to the local one, and under certain conditions, population P always gets extinct. Then, the same model, but in which individuals belonging to population N leave patch 1 proportionally to population P density, is studied. This particular behavioral choice leads to a dynamically richer global system, which favors stability and population coexistence. Finally, we study a third example corresponding to the addition of an aggregative behavior of population N on patch 1. This leads to a more complicated situation in which, according to initial conditions, the global system is described by two different aggregated models. Under certain conditions on parameters a stable limit cycle occurs, leading to periodic variations of the total population densities, as well as of the local densities on the spatial patches.

Animals↗

Dorsal premammillary nucleus differentially modulates defensive behaviors induced by different threat stimuli in rats.

Lesions of the dorsal premammillary nucleus (PMd) have been reported to produce dramatic reductions in responsivity of rats to a live cat. Such lesions provide a means of analyzing the potentially differential neural systems involved in different defensive behaviors, and the relationship between these systems and concepts such as anxiety. Rats with bilateral electrolytic lesions of the PMd were run in an elevated plus maze (EPM), exposed first to cat odor and then to a live cat, and assessed for postshock freezing and locomotion. PMd lesions produced a dramatic reduction in freezing, avoidance, and stretch attend to the cat odor stimulus, and reduction in freezing, with greater activity, and enhanced stretch approach to cat exposure. However, PMd lesions had minimal effects in the EPM, and postshock freezing scores were unchanged. These results confirm earlier findings of reduced defensiveness of PMd-lesioned rats to a cat, extending the pattern of reduced defensiveness to cat odor stimuli as well, but also suggest that such lesions have few effects on nonolfactory threat stimuli.

Aggression↗

A predator's dilemma: prey choice and parasite susceptibility in three-spined sticklebacks.

The acanthocephalan parasite Pomphorhynchus laevis is transmitted by a suitable intermediate host such as the amphipod Gammarus pulex to its definitive fish host. Parasite-induced alterations in both appearance and behaviour concur to render infected gammarids more vulnerable to predation, thus promoting parasite transmission. Experimental infection of laboratory bred full- and half-sib groups of three-spined sticklebacks (Gasterosteus aculeatus) provided evidence that the parasite imposes a survival cost proportional to the severity of infection on its final host. Variation among sibships in the susceptibility to infection was consistent. When given a choice, fish consumed significantly more infected than uninfected prey. Overall, more resistant fish did not prey upon infected gammarids more often than did relatively susceptible fish. Only fish with a relatively high physical condition properly adjusted prey selection to the extent of their parasite susceptibility, thus exploiting the enhanced profitability of infected prey.

Acanthocephala↗

Predation of beetles (Tenebrio molitor) infected with tapeworms (Hymenolepis diminuta): a note of caution for the manipulation hypothesis.

Previous laboratory studies have indicated that tenebrionid beetles infected with Hymenolepis diminuta display behavioural alterations. These are assumed to increase the likelihood of this intermediate host being predated by the Rattus sp. definitive host, and hence provide support for the Manipulation Hypothesis. We tested this hypothesis by both assessing beetle behaviour and predation rates in semi-naturalistic environments. Behavioural assays showed that infected beetles were more often exposed (not concealed under boxes) than uninfected beetles. However, there were no differences in predation rates between infected and uninfected beetles. We discuss this in terms of the historical and evolutionary contexts under which such altered behaviours could evolve and their implications for the Manipulation Hypothesis.

Analysis of Variance↗

Electrophysiological evidence for transient topographic organization of retinotectal projections during optic nerve regeneration in the lizard, Ctenophorus ornatus.

In the lizard, Ctenophorus ornatus, anatomical studies have revealed that optic axons regenerate to visual centers within 2 months of nerve crush but that, from the outset, the regenerated projections lack topographic order (Beazley et al., 1997; Dunlop et al., 1997b). Here we assess the functional topography of the regenerated retinotectal projections by electrophysiological recording of extracellular multiunit responses to visual stimulation and by observing the lizards' ability to capture live prey. At the completion of the electrophysiology, DiI was applied locally to the retina and the topography of the tectal projection later assessed. Electrophysiology revealed that, at 2-4.2 months, responses were weak and habituated readily; no retinotopic order was detected. Between 4.5-6 months, responses were more reliable and the majority of lizards displayed a crude retinotopic order, especially in the ventro-temporal to dorso-nasal retinal axis. Although responses were variable between 6-9 months, they tended to be more reliable again thereafter. However, from 6-18 months, the projection consistently lacked topography with many retinal regions projecting to each tectal locus. Lizards, including those with electrophysiological evidence of crude retinotopy, were consistently unable to capture live prey using the experimental eye. Labelling with DiI confirmed the absence of anatomical retinotopy throughout. Taken together, the electrophysiological and anatomical data indicate that retinotopically appropriate axon terminals (or parts thereof) are transiently active whilst inappropriately located ones are silent. Presumably in lizard map-making cues fade with time and/or the mechanisms are lacking to stabilize and refine the ephemeral map. Moreover, the transient retinotopy is insufficient for useful visual function.

Animals↗

Tonic immobility in the blue crab (Callinectes sapidus, Rathbun) its relation to threat of predation.

The tonic immobility. response in the blue crab (Callinectes sapidus, Rathbun) was investigated in a series of seven experiments. Although reported to be a powerful variable in other species, preinduction electric shock produced inconsistent increases in the duration of tonic immobility with the blue crab; Manipulations that were more directly relevant to the fear of predation had considerably greater effects than shock. Physical damage to the chelipeds, mirror image stimulation, and immobilization beneath artificial glass eyes all produced significant prolongation of the immobility episode; Crabs immobilized on a bed of sand rather than on a hard surface showed shorter immobility durations, suggesting that opportunity for escape is an important variable affecting the immobility reactionmthe present results support the contention that threat of predation is the organizing principle behind tonic immobility

Animals↗

Stage-structured cycles promote genetic diversity in a predator-prey system of Daphnia and algae.

Competition theory predicts that population fluctuations can promote genetic diversity when combined with density-dependent selection. However, this stabilizing mechanism has rarely been tested, and was recently rejected as an explanation for maintaining diversity in natural populations of the freshwater herbivore Daphnia pulex. The primary limitation of competition theory is its failure to account for the alternative types of population cycles that are caused by size- or stage-dependent population vital rates--even though such structure both explains the fluctuating dynamics of many species and may alter the outcome of competition. Here we provide the first experimental test of whether alternative types of cycles affect natural selection in predator-prey systems. Using competing Daphnia genotypes, we show that internally generated, stage-structured cycles substantially reduce the magnitude of selection (thereby contributing to the maintenance of genetic diversity), whereas externally forced cycles show rapid competitive exclusion. The change in selection is ecologically significant, spanning the observed range in natural populations. We argue that structured cycles reduce selection through a combination of stalled juvenile development and stage-specific mortality. This potentially general fitness-equalizing mechanism may reduce the need for strong stabilizing mechanisms to explain the maintenance of genetic diversity in natural systems.

Animals↗

Training on a visual task improves the outcome of optic nerve regeneration.

Optic nerve regeneration in a lizard, Ctenophorus ornatus, is dysfunctional despite survival of most retinal ganglion cells and axon regeneration to the optic tectum. The regenerated retino-tectal projection at 6 months has crude topography but by 1 year is disordered; visually-elicited behavior is absent via the experimental eye. Here, we assess the influence of training on the outcome of optic nerve regeneration. Lizards were trained to catch prey presented within the monocular field of either eye. One optic nerve was then severed and visual stimulation resumed throughout regeneration. In the trained group, presentation was restricted to the eye undergoing optic nerve regeneration; for the untrained group, the unoperated eye was stimulated. Pupil responses returned in trained but not in untrained animals. At 1 year, trained animals oriented to and captured prey; untrained animals demonstrated minimal orienting and failed to capture prey. Regenerated retino-tectal projections were topographic in the trained but not in the untrained group as assessed by in vitro electrophysiological recording and by carbocyanine dye tracing. In vitro electrophysiological recording during application of neurotransmitter antagonists to the tectum revealed that the level of GABAergic inhibition was modest in trained animals but elevated in the untrained group; responses were mainly AMPA-mediated in both groups. We conclude that training improves the behavioral outcome of regeneration, presumably by stabilizing and refining the transient retino-tectal map and preventing a build-up of tectal inhibition. The results suggest that for successful central nerve regeneration to occur in mammals, it may be necessary to introduce training to complement procedures stimulating axon regeneration.

Animals↗

Ecological change, group territoriality, and population dynamics in Serengeti lions.

Territorial behavior is expected to buffer populations against short-term environmental perturbations, but we have found that group living in African lions causes a complex response to long-term ecological change. Despite numerous gradual changes in prey availability and vegetative cover, regional populations of Serengeti lions remained stable for 10- to 20-year periods and only shifted to new equilibria in sudden leaps. Although gradually improving environmental conditions provided sufficient resources to permit the subdivision of preexisting territories, regional lion populations did not expand until short-term conditions supplied enough prey to generate large cohorts of surviving young. The results of a simulation model show that the observed pattern of "saltatory equilibria" results from the lions' grouping behavior.

Algorithms↗

Long-term resource variation and group size: a large-sample field test of the Resource Dispersion Hypothesis.

BACKGROUND: The Resource Dispersion Hypothesis (RDH) proposes a mechanism for the passive formation of social groups where resources are dispersed, even in the absence of any benefits of group living per se. Despite supportive modelling, it lacks empirical testing. The RDH predicts that, rather than Territory Size (TS) increasing monotonically with Group Size (GS) to account for increasing metabolic needs, TS is constrained by the dispersion of resource patches, whereas GS is independently limited by their richness. We conducted multiple-year tests of these predictions using data from the long-term study of badgers Meles meles in Wytham Woods, England. The study has long failed to identify direct benefits from group living and, consequently, alternative explanations for their large group sizes have been sought. RESULTS: TS was not consistently related to resource dispersion, nor was GS consistently related to resource richness. Results differed according to data groupings and whether territories were mapped using minimum convex polygons or traditional methods. Habitats differed significantly in resource availability, but there was also evidence that food resources may be spatially aggregated within habitat types as well as between them. CONCLUSIONS: This is, we believe, the largest ever test of the RDH and builds on the long-term project that initiated part of the thinking behind the hypothesis. Support for predictions were mixed and depended on year and the method used to map territory borders. We suggest that within-habitat patchiness, as well as model assumptions, should be further investigated for improved tests of the RDH in the future.

Animals↗

[Behavioral pharmacology of amantadine with special references to the effect on abnormal behavior in mice and rats].

Behavioral effects of amantadine, especially on abnormal behavior in mice and rats, were reevaluated, in comparison with those of tricyclic antidepressants, methamphetamine and L-DOPA. 1) Amantadine at 10 approximately 50 mg/kg, i.p., tended to decrease ambulation and rearing in rats and mice. The drug at 50 mg/kg, i.p., caused piloerection and hyperirritability and at doses over 80 mg/kg, i.p., it impaired coordinated motor activity in rats. 2) Amantadine inhibited methamphetamine-induced hyperactivity, but apomorphine-induced stereotyped behavior was unaffected in rats. 3) Amantadine was equipotent to imipramine in suppressing haloperidol-induced catalepsy in rats, but L-DOPA was without effect. On the other hand, amantadine was 40 and 400 times as potent as imipramine and L-DOPA, respectively, in suppressing delta 9-tetrahydrocannabinol (THC)-induced catalepsy in rats. 4) Amantadine was as potent as imipramine in suppressing the muricide of olfactory bulbectomized rats, but was 3.5, 8.8 and 225.5 times as potent as methamphetamine, imipramine and L-DOPA, respectively, in inhibiting THC-induced reversible muricide in rats. 5) Amantadine at 50 mg/kg did not elicit circling behavior in the rat with unilateral nigral lesion induced by 6-hydroxydopamine. 6) Amantadine at high doses caused irritable aggression characterized by squealing in rats pretreated with intraventricular 6-hydroxydopamine. The most important characteristic of amantadine is its prominent effect suppressing the THC-induced catalepsy and muricide. This may be a reflection of the feature of amantadine activating the dopaminergic as well as the serotonergic systems.

Aggression↗

Species-specific danger signals, endogenous opioid analgesia, and defensive behavior.

The effects of handling stimuli and stress odors on species-specific defensive behavior and pain sensitivity were examined in rats. Animals not adapted to handling had longer jump latencies on the hot plate test of pain sensitivity than those with extensive handling experience. In a postshock freezing test, naltrexone enhanced defensive freezing relative to saline controls in nonadapted animals. However, naltrexone produced no such effect in rats that were adapted to handling. These two studies indicate that the handling procedure triggered an endogenous opioid analgesic response in rats not adapted to handling. Experiment 3 showed that a similar naltrexone-reversible opioid analgesia can be triggered by stress odors. Naltrexone, when compared to saline, enhanced postshock freezing in the presence of conspecific stress odors, but not in their absence. In Experiment 4, stress odors and nonadapted handling were able to activate defensive freezing directly, when tested in compound but not in isolation. The studies are consistent with the view that stress odors and handling stimuli are danger signals that activate endogenous opioid analgesia as well as defensive behavior, suggesting that analgesia is a component of the rat's defensive behavior system.

Adaptation, Psychological↗

Stoichiometry in producer-grazer systems: linking energy flow with element cycling.

All organisms are composed of multiple chemical elements such as carbon, nitrogen and phosphorus. While energy flow and element cycling are two fundamental and unifying principles in ecosystem theory, population models usually ignore the latter. Such models implicitly assume chemical homogeneity of all trophic levels by concentrating on a single constituent, generally an equivalent of energy. In this paper, we examine ramifications of an explicit assumption that both producer and grazer are composed of two essential elements: carbon and phosphorous. Using stoichiometric principles, we construct a two-dimensional Lotka-Volterra type model that incorporates chemical heterogeneity of the first two trophic levels of a food chain. The analysis shows that indirect competition between two populations for phosphorus can shift predator-prey interactions from a (+, -) type to an unusual (-, -) class. This leads to complex dynamics with multiple positive equilibria, where bistability and deterministic extinction of the grazer are possible. We derive simple graphical tests for the local stability of all equilibria and show that system dynamics are confined to a bounded region. Numerical simulations supported by qualitative analysis reveal that Rosenzweig's paradox of enrichment holds only in the part of the phase plane where the grazer is energy limited; a new phenomenon, the paradox of energy enrichment, arises in the other part, where the grazer is phosphorus limited. A bifurcation diagram shows that energy enrichment of producer-grazer systems differs radically from nutrient enrichment. Hence, expressing producer-grazer interactions in stoichiometrically realistic terms reveals qualitatively new dynamical behavior.

Animals↗