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The effect of early poststressor intervention with sertraline on behavioral responses in an animal model of post-traumatic stress disorder.

Whereas several well-controlled studies have established the selective serotonin reuptake inhibitors (SSRIs) as the recommended first-line pharmacotherapeutic agents for acute and chronic post-traumatic stress disorder (PTSD), drug interventions in the acute postexposure phase have not been studied to the same extent and tend to be largely speculative. This study employed an animal model which assesses prevalence of individual stress-response behavior patterns in order to assess the short-term effects of a brief treatment regimen with an SSRI (sertraline) administered immediately after stress-exposure, with those of an identical delayed regimen and of saline. Prevalence rates of rats displaying extreme anxiety-like behavioral responses to predator stress, compared to partial and minimal responses, were assessed in the elevated plus maze and startle response paradigms, with and without intraperitoneal administration of sertraline for 7 days immediately postexposure, or 7 days after exposure. Immediate postexposure administration of sertraline reduced anxiety-like and avoidant behavior, decreased hyperarousal responses and diminished the overall incidence of extreme (PTSD-like) behavioral responses, compared to the delayed treatment regimen and to saline controls. Brief immediate poststress exposure treatment with sertraline reduced prevalence rates of extreme behavioral disruption in the short-term. SSRI drugs are thus worthy of further investigation as agents of secondary prevention in the acute aftermath of stress-exposure.

Animals↗

Size-dependent interactions inhibit coexistence in intraguild predation systems with life-history omnivory.

Growth in body size during ontogeny often results in changes in diet, leading to life-history omnivory. In addition, growth is often dependent on food density. Using a physiologically structured population model, we investigated the effects of these two aspects of individual growth in a system consisting of two size-structured populations, an omnivorous top predator and an intermediate consumer. With a single shared resource for both populations, we found that life-history omnivory decreases the likelihood of coexistence between top predator and intermediate consumer in this intraguild predation (IGP) system. This result contrasts with previous unstructured models and stage-structured models without food-dependent development. Food-dependent development and size-dependent foraging abilities of the predator resulted in a positive feedback between foraging success on the shared resource at an early life stage and foraging success on the intermediate consumer later in life. By phenomenologically incorporating this feedback in an unstructured IGP model, we show that it also demotes coexistence in this simple setting, demonstrating the robustness of the negative effect of this feedback.

Animals↗

Nitric oxide synthase activity in the dorsal periaqueductal gray of rats expressing innate fear responses.

Immunohistochemical studies have shown nitric oxide synthase (NOS)-positive neurons in the dorsolateral sector of periaqueductal gray (PAG), a neural site known to be critical for the expression of defensive responses. In the present study, we first characterized the dorsal PAG (dPAG) NOS, and then examined NOS activity and cyclic GMP (cGMP) accumulation in the dPAG of rats exposed to a predator (cat) for 15 min. NOS activity evaluated by enzymatic conversion of [3H]arginine to [3H]citrulline in dPAG of exposed rats increased 14.6% and the cGMP radioimmunoassay showed an increase of 30.6% in relation to non-exposed rats. These results suggest an involvement of the NO/cGMP pathway in the dPAG during defensive responses.

Analysis of Variance↗

Arms races between and within species.

An adaptation in one lineage (e.g. predators) may change the selection pressure on another lineage (e.g. prey), giving rise to a counter-adaptation. If this occurs reciprocally, an unstable runaway escalation or 'arms race' may result. We discuss various factors which might give one side an advantage in an arms race. For example, a lineage under strong selection may out-evolve a weakly selected one (' the life-dinner principle'). We then classify arms races in two independent ways. They may be symmetric or asymmetric, and they may be interspecific or intraspecific. Our example of an asymmetric interspecific arms race is that between brood parasites and their hosts. The arms race concept may help to reduce the mystery of why cuckoo hosts are so good at detecting cuckoo eggs, but so bad at detecting cuckoo nestlings. The evolutionary contest between queen and worker ants over relative parental investment is a good example of an intraspecific asymmetric arms race. Such cases raise special problems because the participants share the same gene pool. Interspecific symmetric arms races are unlikely to be important, because competitors tend to diverge rather than escalate competitive adaptations. Intraspecific symmetric arms races, exemplified by adaptations for male-male competition, may underlie Cope's Rule and even the extinction of lineages. Finally we consider ways in which arms races can end. One lineage may drive the other to extinction; one may reach an optimum, thereby preventing the other from doing so; a particularly interesting possibility, exemplified by flower-bee coevolution, is that both sides may reach a mutual local optimum; lastly, arms races may have no stable and but may cycle continuously. We do not wish necessarily to suggest that all, or even most, evolutionary change results from arms races, but we do suggest that the arms race concept may help to resolve three long-standing questions in evolutionary theory.

Adaptation, Biological↗

A plant pathogen reduces the enemy-free space of an insect herbivore on a shared host plant.

An important mechanism in stabilizing tightly linked host-parasitoid and prey-predator interactions is the presence of refuges that protect organisms from their natural enemies. However, the presence and quality of refuges can be strongly affected by the environment. We show that infection of the host plant Silene latifolia by its specialist fungal plant pathogen Microbotryum violaceum dramatically alters the enemy-free space of a herbivore, the specialist noctuid seed predator Hadena bicruris, on their shared host plant. The pathogen arrests the development of seed capsules that serve as refuges for the herbivore's offspring against the specialist parasitoid Microplitis tristis, a major source of mortality of H. bicruris in the field. Pathogen infection resulted both in lower host-plant food quality, causing reduced adult emergence, and in twofold higher rates of parasitism of the herbivore. We interpret the strong oviposition preference of H. bicruris for uninfected plants in the field as an adaptive response, positioning offspring on refuge-rich, high-quality hosts. To our knowledge, this is the first demonstration that plant-inhabiting micro-organisms can affect higher trophic interactions through alteration of host refuge quality. We speculate that such interference can potentially destabilize tightly linked multitrophic interactions.

Adaptation, Physiological↗

Aposematism: what should our starting point be?

The evolution of aposematism is considered to be a major evolutionary problem because if new aposematic forms emerged in defended cryptic populations, they would face the dual problems of rarity and conspicuousness. We argue that this commonly assumed starting point might not have wide validity. We describe a novel evolutionary computer model in which prey evolve secondary defences and become conspicuous by moving widely over a visually heterogeneous habitat. Unless crypsis imposes high opportunity costs (for instance, preventing prey from efficient foraging, thermoregulation and communication), costly secondary defences are not predicted to evolve at all. However, when crypsis imposes opportunity costs, prey evolve secondary defences that facilitate raised behavioural conspicuousness as prey exploit opportunities within their environment. Optimal levels of secondary defence and of behavioural conspicuousness increase with population sizes and the costs imposed by crypsis. When prey are already conspicuous by virtue of their behaviours, the evolution of aposematic appearances (bright coloration, etc.) is much easier to explain because aposematic traits add little further costs of conspicuousness, but can bring large benefits.

Adaptation, Physiological↗

Frequency-dependent predation, crypsis and aposematic coloration.

Frequency-dependent predation may maintain or prevent colour pattern polymorphisms in prey, and can be caused by a variety of biological phenomena, including perceptual processes (search images), optimal foraging and learning. Most species are preyed upon by more than one predator species, which are likely to differ in foraging styles, perceptual and learning abilities. Depending upon the interaction between predator vision, background and colour pattern parameters, certain morphs may be actively maintained in some conditions and not in others, even with the same predators. More than one kind of predator will also affect stability, and only slight changes in conditions can cause a transition between polymorphism and monomorphism. Frequency-dependent selection is not a panacea for the explanation of variation in animal colour patterns, although it may be important in some systems.

Animals↗

The evolutionary response of predators to dangerous prey: hotspots and coldspots in the geographic mosaic of coevolution between garter snakes and newts.

The "geographic mosaic" approach to understanding coevolution is predicated on the existence of variable selection across the landscape of an interaction between species. A range of ecological factors, from differences in resource availability to differences in community composition, can generate such a mosaic of selection among populations, and thereby differences in the strength of coevolution. The result is a mixture of hotspots, where reciprocal selection is strong, and coldspots, where reciprocal selection is weak or absent, throughout the ranges of species. Population subdivision further provides the opportunity for nonadaptive forces, including gene flow, drift, and metapopulation dynamics, to influence the coevolutionary interaction between species. Some predicted results of this geographic mosaic of coevolution include maladapted or mismatched phenotypes, maintenance of high levels of polymorphism, and prevention of stable equilibrium trait combinations. To evaluate the potential for the geographic mosaic to influence predator-prey coevolution, we investigated the geographic pattern of genetically determined TTX resistance in the garter snake Thamnophis sirtalis over much of the range of its ecological interaction with toxic newts of genus Taricha. We assayed TTX resistance in over 2900 garter snakes representing 333 families from 40 populations throughout western North America. Our results provide dramatic evidence that geographic structure is an important component in coevolutionary interactions between predators and prey. Resistance levels vary substantially (over three orders of magnitude) among populations and over short distances. The spatial array of variation is consistent with two areas of intense evolutionary response by predators ("hotspots") surrounded by clines of decreasing resistance. Some general predictions of the geographic mosaic process are supported, including clinal variation in phenotypes, polymorphism in some populations, and divergent outcomes of the interaction between predator and prey. Conversely, our data provide little support for one of the major predictions, mismatched values of interacting traits. Two lines of evidence suggest selection is paramount in determining population variation in resistance. First, phylogenetic information indicates that two hotspots of TTX resistance have evolved independently. Second, in the one region that TTX levels in prey have been quantified, resistance and toxicity levels match almost perfectly over a wide phenotypic and geographic range. However, these results do not preclude the role the nonadaptive forces in generating the overall geographic mosaic of TTX resistance. Much work remains to fill in the geographic pattern of variation among prey populations and, just as importantly, to explore the variation in the ecology of the interaction that occurs within populations.

Adaptation, Physiological↗

Habitat specialization and adaptive phenotypic divergence of anuran populations.

We tested for adaptive population structure in the frog Rana temporaria by rearing tadpoles from 23 populations in a common garden experiment, with and without larval dragonfly predators. The goal was to compare tadpole phenotypes with the habitats of their source ponds. The choice of traits and habitat variables was guided by prior information about phenotypic function. There were large differences among populations in life history, behaviour, morphological shape, and the predator-induced plasticities in most of these. Body size and behaviour were correlated with predation risk in the source pond, in agreement with adaptive population divergence. Tadpoles from large sunny ponds were morphologically distinct from those inhabiting small woodland ponds, although here an adaptive explanation was unclear. There was no evidence that plasticity evolves in populations exposed to more variable environments. Much among-population variation in phenotype and plasticity was not associated with habitat, perhaps reflecting rapid changes in wetland habitats.

Adaptation, Physiological↗

Evolution of motor patterns in tetraodontiform fishes: does muscle duplication lead to functional diversification?

Several times within the teleost fish order Tetraodontiformes singular jaw adducting muscles have been effectively 'duplicated' by physical subdivision to produce new muscles. This morphological system provides an opportunity to investigate how the functional complexity of muscular systems changes with evolutionary increases in the number of component muscles. In this study we asked if muscle duplication has lead to functional diversification by comparing the motor patterns of muscles that result from subdivision events. The activity patterns of five different sets of duplicated muscles were quantified with electromyographic recordings (EMG) from four individuals in each of three species during processing of three prey types. Prey varied in durability and elusiveness (live fiddler crabs, pieces of squid tentacle and live paeneid shrimps). For each cycle of prey processing, measurements were made of the relative onset time of each adductor muscle, the duration of each burst of activity, and the relative intensity of each activity burst. Two types of functional divergence of muscles were observed in analyses of variance conducted on the EMG variables. In two of the 15 variables examined, the timing of activity of the descendant set of muscles differed. In another three of the 15 variables, there were significant interactions between muscle and prey type, indicating a prey effect which differed in the descendant muscles. Overall, evidence of motor divergence was found in three of five cases of muscle duplication. This indicates that muscle subdivision has led to increased functional complexity of the jaw-adductor muscle system in tetraodontiform fishes.

Animals↗

Configurational prey selection by Bufo, Alytes, Bombina and Hyla.

The aim of the present study is to answer the following questions; (1) Is the ability of toads to distinguish between different configurational prey stimuli based on learning during ontogeny? (2) Is the configurational meaning of a stimulus influenced by its distance from the animal? (3) Are there differences in configurational prey selection among different anuran species? (1) Experiments with postmetamorphic juveniles of Alytes obstetricans indicate that preferences for worm-like objects can be obtained immediately after metamorphosis. This preference undergoes some specialization during ontogeny. (2) The configurational meaning of 'worm-like' and 'antiworm-like' objects is -- within broad ranges -- independent of the distance from the animal so far as has been investigated in A. obstetricans and Bufo bufo. (3) The configurational discrimination ability (between worm-like and antiworm-like objects) has common basic components in different anuran species, so far investigated in the following species: B. bufo, Hyla arborea, Hyla cinerea, A. obstetricans. Bombina variegata. There are remarkable species differences for preferred absolute prey sizes, which may depend on different body sizes and in particular upon the size of the mouth.

Age Factors↗

Vertebrate predation in Cebus capucinus: meat eating in a neotropical monkey.

A long-term study of two groups of white-faced capuchins (Cebus capucinus) in Santa Rosa National Park in Costa Rica provides evidence of unusually high levels of vertebrate predation compared to those reported in other field studies of Cebus. The hunting techniques for different prey types are described, and several questions concerning vertebrate predation in primates are addressed. Why is there variation between individuals and between groups in the rate of predation? Why do males hunt more than females? Previous hypotheses to explain hunting in Old World primates are applied to this Neotropical example. Finally, I argue that successful vertebrate predation can readily arise in species like Cebus, which are characterized by opportunistic foraging patterns, manipulative and cognitive skills and well-developed techniques for locating and subduing invertebrate prey.

Animals↗

The orienting response of Lake Michigan mottled sculpin is mediated by canal neuromasts.

Lake Michigan mottled sculpin, Cottus bairdi, exhibit a naturally occurring and unconditioned orienting response that can be triggered by both live prey and chemically inert vibrating spheres, even in blinded animals. CoCl(2)-induced reductions of the orienting response demonstrate that the lateral line is required for this behavior in the absence of non-mechanosensory cues (such as vision), but shed no light on the relative contributions of superficial and canal neuromasts to this behavior. To determine the relative roles of these two subsystems, we measured the frequency with which mottled sculpin oriented towards a small vibrating sphere before and after two treatments: (i) immersion of fish in a solution of gentamicin, an aminoglycoside antibiotic that damages hair cells in canal, but not superficial, neuromasts; and (ii) scraping the skin of the fish, which damages the superficial, but not the canal, neuromasts. To ensure that both superficial and canal neuromasts were adequately stimulated, we tested at different vibration frequencies (10 and 50 Hz) near or at the best frequency for each type of neuromast. At both test frequencies, response rates before treatment were greater than 70 % and were significantly greater than 'spontaneous' response frequencies measured in the absence of sphere vibration. Response rates fell to spontaneous levels after 1 day of gentamicin treatment and did not return to pre-treatment levels for 10-15 days. In contrast, response rates stayed approximately the same after superficial neuromasts had been damaged by skin abrasion. Scanning electron microscopy confirmed hair cell damage (loss of apical cilia) in canal, but not superficial, neuromasts of gentamicin-treated animals after as little as 24 h of treatment. The sensory epithelium of canal neuromasts gradually returned to normal, following a time course similar to behavioral loss and recovery of the orienting response, whereas that of superficial neuromasts appeared normal throughout the entire period. This study shows that the orienting response of the mottled sculpin is mediated by canal neuromasts.

Animals↗

Aggregation of helminths: the role of feeding behavior of fish hosts.

Individual Arctic charr (Salvelinus alpinus) from Fjellfrøsvatn, northern Norway, could be categorized by their stomach contents as zooplanktivores or benthivores. Feeding specialization among these fish was evident from negative correlations between helminths transmitted by pelagic copepods (Diphyllobothrium dendriticum and D. ditremum) and those transmitted by the benthic amphipod Gammarus lacustris (Cystidicola farionis and Cyathocephalus truncatus). Occurrences of parasite species acquired from the same types of invertebrate were positively correlated in the fish. Strong relationships among habitat use, diet, and helminth infections among the Arctic charr indicated persistent foraging patterns involving long-term habitat use and feeding specialization. The distribution of all parasite species was highly aggregated in the fish samples, measured by the exponent k of the fitted negative binomial distributions (range: 0.5-7.5) and the variance-to-mean ratios (s2/mean, range: 5-85). Charr specializing on either copepods or Gammarus predominantly contributed to high-intensity class intervals within the overall frequency distributions of the corresponding parasite species. Such fish had low infection intensities of helminths transmitted by other prey organisms. The detailed analyses of the parasite frequency distributions for fish with different habitat or feeding preferences evidently show how heterogeneity in trophic behavior contributes strongly to the commonly observed aggregation of helminths among hosts under natural conditions.

Amphipoda↗

Assessment of prey vulnerability through analysis of wolf movements and kill sites.

Within predator-prey systems behavior can heavily influence spatial dynamics, and accordingly, the theoretical study of how spatial dynamics relate to stability within these systems has a rich history. However, our understanding of these behaviors in large mammalian systems is poorly developed. To address the relationship between predator selection patterns, prey density, and prey vulnerability, we quantified selection patterns for two fine-scale behaviors of a recovering wolf (Canis lupus) population in Yellowstone National Park, Wyoming, USA. Wolf spatial data were collected between November and May from 1998-1999 until 2001-2002. Over four winters, 244 aerial locations, 522 ground-based telemetry locations, 1287 km of movement data from snow tracking, and the locations of 279 wolf kill sites were recorded. There was evidence that elk (Cervus elaphus) and bison (Bison bison) densities had a weak effect on the sites where wolves traveled and made kills. Wolf movements showed a strong selection for geothermal areas, meadows, and areas near various types of habitat edges. Proximity to edge and habitat class also had a strong influence on the locations where elk were most vulnerable to predation. There was little evidence that wolf kill sites differed from the places where wolves traveled, indicating that elk vulnerability influenced where wolves selected to travel. Our results indicate that elk are more vulnerable to wolves under certain conditions and that wolves are capable of selecting for these conditions. As such, vulnerability plays a central role in predator-prey behavioral games and can potentially impact the systems to which they relate.

Animals↗