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PubMed2004

Evolutionary tradeoff and equilibrium in an aquatic predator-prey system.

Due to the conventional distinction between ecological (rapid) and evolutionary (slow) timescales, ecological and population models have typically ignored the effects of evolution. Yet the potential for rapid evolutionary change has been recently established and may be critical to understanding how populations persist in changing environments. In this paper we examine the relationship between ecological and evolutionary dynamics, focusing on a well-studied experimental aquatic predator-prey system (Fussmann et al., 2000, Science, 290, 1358-1360; Shertzer et al., 2002, J. Anim. Ecol., 71, 802-815; Yoshida et al., 2003, Nature, 424, 303-306). Major properties of predator-prey cycles in this system are determined by ongoing evolutionary dynamics in the prey population. Under some conditions, however, the populations tend to apparently stable steady-state densities. These are the subject of the present paper. We examine a previously developed model for the system, to determine how evolution shapes properties of the equilibria, in particular the number and identity of coexisting prey genotypes. We then apply these results to explore how evolutionary dynamics can shape the responses of the system to 'management': externally imposed alterations in conditions. Specifically, we compare the behavior of the system including evolutionary dynamics, with predictions that would be made if the potential for rapid evolutionary change is neglected. Finally, we posit some simple experiments to verify our prediction that evolution can have significant qualitative effects on observed population-level responses to changing conditions.

Animals↗

PubMed2003

Conditioning and residual emotionality effects of predator stimuli: some reflections on stress and emotion.

The advantages of using predator-related odor stimuli to study emotional responses in laboratory tests depend on whether such stimuli do elicit a relatively complete pattern of emotionality. This has been confirmed for cat fur/skin odor stimuli, which elicit a range of defensive behaviors in rats that may be reduced by anxiolytic drugs, produce residual anxiety-like behavior in the elevated plus maze and support rapid aversive conditioning to the context in which they were encountered. Although the synthetic fox fecal odor, trimethylthiazoline (TMT), elicits avoidance similar to that seen in response to cat fur/skin odor, this avoidance does not respond to anxiolytic drugs. In addition, TMT does not produce residual anxiety-like behaviors in the elevated plus maze, nor does it support conditioning. As natural cat feces also elicit avoidance but fail to support conditioning, it is possible that the ability of a predator-related odor to serve as an effective unconditioned stimulus (US) relates to its predictive status with reference to the actual presence of the predator. Avoidance per se may reflect that a stimulus is aversive but not necessarily capable of eliciting an emotional response. This view is consonant with findings in a Mouse Defense Test Battery (MDTB) measuring a wide range of defensive responses to predator exposure. A contextual defense measure that may reflect either conditioned or residual but unconditioned emotional responses was almost never reduced by drug effects unless these also reduced risk assessment or defensive threat/attack measures. However, reductions in contextual defense without changes in flight/avoidance measures were much more common. These findings suggest that flight/avoidance, although it obviously may occur as one component of a full pattern of defensive and emotional behaviors, is also somewhat separable from the others. When-as appears to be the case with TMT-it is the major or perhaps only consistent defensive behavior elicited, this may reflect a stimulus that is aversive or noxious but with little ability to predict the presence of threat or danger. That such stimuli fail to support rapid aversive conditioning suggests the need for a reanalysis of the characteristics required for an effective aversive US.

Animals↗

PubMed2005

Sarcocystosis of chital-dhole: conditions for evolutionary stability of a predator parasite mutualism.

BACKGROUND: For parasites with a predator-prey life cycle, the completion of the life cycle often depends on consumption of parasitized prey by the predator. In the case of such parasite species the predator and the parasite have common interests and therefore a mutualistic relationship is possible. Some evidence of a predator-parasite mutualism was reported from spotted deer or chital (Axix axis) as a prey species, dhole or Indian wild-dog (Cuon alpinus) as the predator and a protozoan (Sarcocystis axicuonis) as the parasite. We examine here, with the help of a model, the ecological conditions necessary for the evolution and stability of such a mutualistic relationship. A two - level game theory model was designed in which the payoff of a parasite is decided not only by alternative parasite strategies but also by alternative host strategies and vice versa. Conditions for ESS were examined. RESULTS: A tolerant predator strategy and a low or moderately virulent parasite strategy which together constitute mutualism are stable only at a high frequency of recycling of parasite and a substantial prey - capture benefit to the predator. Unlike the preliminary expectation, parasite will not evolve towards reduced virulence, but reach an optimum moderate level of virulence. CONCLUSION: The available data on the behavioral ecology of dhole and chital suggest that they are likely to meet the stability criteria and therefore a predator-parasite mutualism can be stable in this system. The model also points out the gaps in the current data and could help directing further empirical work.

Animals↗

PubMed2003

A growth/mortality trade-off in larval salamanders and the coexistence of intraguild predators and prey.

Behavioral and morphological traits often influence a key trade-off between resource acquisition and vulnerability to predation, and understanding trait differences between species can provide critical insight into their interactions with other species and their distributions. Such an approach should enhance our understanding of the criteria for coexistence between species that can interact through both competition and predation (i.e. intraguild predators and prey). I conducted a common garden experiment that revealed strong differences between three guild members (larval salamanders Ambystoma laterale, A. maculatum, and A. tigrinum) in behavior, morphology, and growth in the presence and absence of a shared top predator (the larval dragonfly Anax longipes). All three species also reduced their activity and modified their tail fin depth, tail muscle length, and body length in response to non-lethal Anax. Species that act as intraguild predators were more active and could grow faster than their intraguild prey species, but they also suffered higher mortality in laboratory predation trials with Anax. I also used survey data from natural communities to compare the distribution of Ambystoma species between ponds differing in abiotic characteristics and predatory invertebrate assemblages. An intraguild prey species ( A. maculatum) was found more reliably, occurred at higher densities, and was more likely to persist late into the larval period in ponds with more diverse invertebrate predator assemblages. Taken together, these results indicate that top predators such as Anax may play an important role in influencing intraguild interactions among Ambystoma and ultimately their local distribution patterns.

Animals↗

PubMed1990

A model of predator-prey dynamics as modified by the action of a parasite.

A predator-prey population is described in which the prey population may be either a secondary host or a primary host to a parasite, but the predator is always a primary host. Those prey that have been invaded by the parasite have their behavior modified so as to make them more susceptible to predation. The model is described by a system of three autonomous ordinary differential equations. Conditions for persistence of all populations are given in the case that both populations are primary hosts. A brief discussion of the stability of the interior equilibrium is given.

Animals↗

PubMed2004

Protean behavior under barn-owl attack: voles alternate between freezing and fleeing and spiny mice flee in alternating patterns.

When attacking a spiny mouse in an experimental arena, a barn owl launched a few attacks from distant perches, made repetitive short-distance swoops in each attack and remained in the vicinity of the prey while chasing it. The spiny mouse fled in response, and typically oriented to face the owl whenever it stopped. When attacking a vole, the barn owl performed a greater number of attacks from distant perches, and left the vicinity of the prey after a few short-distance chases or capture attempts. Voles responded to these attacks in unspecific combinations of freezing and fleeing, and did not turn to face the owl when they stopped. Four conclusions are drawn from these encounters. First, two strategies characterized these predator-prey interactions; in one, both predator and prey continuously maintained awareness of each other's location; whereas in the other they continuously attempted to avoid the attention of the other. Second, responses of spiny mice and voles were a manifestation of protean behavior, with spiny mice fleeing in an alternating pattern and voles alternating between running and freezing. Third, locomotor response to owl attack comprised behavior that is an augmentation of normal behavior, with voles clinging to the walls and spiny mice running with frequent and irregular changes in direction. Fourth, the different defensive responses accord with the motor capacities and habitat of each rodent species. All in all, these results demonstrate the dynamic and multidimensional nature of predator-prey interactions.

Animals↗

PubMed2002

Non-human primate models for investigating fear and anxiety.

Non-human primates exhibit similar physiological and behavioral responses to anxiety-inducing situations as humans and have, in fact, been successfully employed in both conditioned (i.e. conflict paradigms) and ethologically based tests of fear/anxiety (i.e. involuntary isolation, social interaction, human threat, predator confrontation). In the last decade, a renewed and growing interest in non-human primate models has resulted from the use of the small callitrichid species in behavioral pharmacology and neuroscience. This review focuses on the available non-human primate models for investigating fear/anxiety, addressing their advantages, shortcomings, and conceptual framework on which they are based. Lastly, a new ethologically based model to study anxiety and fear-induced avoidance in callitrichids--the marmoset predator confrontation test--is discussed.

Animals↗

PubMed2003

Local selection and latitudinal variation in a marine predator-prey interaction.

Although pairs of species often interact over broad geographic ranges, few studies have explored how interactions vary across these large spatial scales. Surveys along 1500 kilometers of the Pacific coast of North America documented marked variation in the frequency of predation by the snail Nucella canaliculata on the intertidal mussel Mytilus californianus. Laboratory rearing experiments suggest that regional differences in drilling behavior have a genetic basis, and mitochondrial sequence variation confirms that gene flow is low among these snail populations. Marine communities separated by hundreds of kilometers may have intrinsically different dynamics, with interactions shaped by restricted gene flow and spatially varying selection.

Animals↗

PubMed1999

Prey capture in the weakly electric fish Apteronotus albifrons: sensory acquisition strategies and electrosensory consequences.

Sensory systems are faced with the task of extracting behaviorally relevant information from complex sensory environments. In general, sensory acquisition involves two aspects: the control of peripheral sensory surfaces to improve signal reception and the subsequent neural filtering of incoming sensory signals to extract and enhance signals of interest. The electrosensory system of weakly electric fish provides a good model system for studying both these aspects of sensory acquisition. On the basis of infrared video recordings of black ghost knifefish (Apteronotus albifrons) feeding on small prey (Daphnia magna) in the dark, we reconstruct three-dimensional movement trajectories of the fish and prey. We combine the reconstructed trajectory information with models of peripheral electric image formation and primary electrosensory afferent response dynamics to estimate the spatiotemporal patterns of transdermal potential change and afferent activation that occur during prey-capture behavior. We characterize the behavioral strategies used by the fish, with emphasis on the functional importance of the dorsal edge in prey capture behavior, and we analyze the electrosensory consequences. In particular, we find that the high-pass filter characteristics of P-type afferent response dynamics can serve as a predictive filter for estimating the future position of the prey as the electrosensory image moves across the receptor array.

Animals↗

PubMed2003

Helical Lévy walks: adjusting searching statistics to resource availability in microzooplankton.

The searching trajectories of different animals can be described with a broad class of flight length (lj) distributions with P(lj) = lj-mu. Theoretical studies have shown that changes in these distributions (i.e., different mu values) are key to optimizing the long-term encounter statistics under certain searcher-resource scenarios. In particular, they predict the advantage of Lévy searching (mu approximately 2) over Brownian motion (mu > or = 3) for low-prey-density scenarios. Here, we present experimental evidence of predicted optimal changes in the flight-time distribution of a predator's walk in response to gradual density changes of its moving prey. Flight times of the dinoflagellate Oxyrrhis marina switched from an exponential to an inverse square power-law distribution when the prey (Rhodomonas sp.) decreased in abundance. Concomitantly, amplitude and frequency of the short-term helical path increased. The specific biological mechanisms involved in these searching behavioral changes are discussed. We suggest that, in a three-dimensional environment, a stronger helical component combined with a Lévy walk searching strategy enhances predator's encounter rates. Our results support the idea of universality of the statistical laws in optimal searching processes despite variations in the biological details of the organisms.

Animals↗

PubMed2005

Predator-prey cycles from resonant amplification of demographic stochasticity.

We present the simplest individual level model of predator-prey dynamics and show, via direct calculation, that it exhibits cycling behavior. The deterministic analogue of our model, recovered when the number of individuals is infinitely large, is the Volterra system (with density-dependent prey reproduction) which is well known to fail to predict cycles. This difference in behavior can be traced to a resonant amplification of demographic fluctuations which disappears only when the number of individuals is strictly infinite. Our results indicate that additional biological mechanisms, such as predator satiation, may not be necessary to explain observed predator-prey cycles in real (finite) populations.

Animals↗

PubMed2002

Prey capture by larval zebrafish: evidence for fine axial motor control.

Swimming and turning behaviors of larval zebrafish have been described kinematically, but prey capture behaviors are less well characterized. High-speed digital imaging was used to record the axial kinematics of larval zebrafish as they preyed upon paramecia and also during other types of swimming. In all types of swim bouts, a series of traveling waves of bending is observed and these bends propagate along the trunk in the rostral to caudal direction. The prey capture swim bouts appeared to be more complex than other swim patterns examined. In the capture swim bouts, the initial bends were of low amplitude and were most prominent at far-caudal locations during each individual traveling wave of bending. Later bends in the bout (occurring just prior to prey capture) appeared to originate more rostrally and were of larger amplitude. These changes in bending pattern during capture swims were accompanied by a marked increase in tail-beat frequency. Associated with these axial kinematics were changes in heading and an abrupt increase in velocity close to the moment of prey capture. These changing patterns of bending suggest precise, bend-to-bend, neural control over both the timing and the rostral-caudal locus of bending. This degree of 'fine axial motor control' has not previously been described in the teleost behavioral literature and is notable because it occurs in larval zebrafish, where descending control signals are funneled through the roughly three-hundred neurons that project from brain into spinal cord. These findings will necessitate a significant increase in the complexity of current models of descending motor control in fishes.

Animals↗

PubMed2001

Global analysis of the Michaelis-Menten-type ratio-dependent predator-prey system.

The recent broad interest on ratio-dependent based predator functional response calls for detailed qualitative study on ratio-dependent predator-prey differential systems. A first such attempt is documented in the recent work of Kuang and Beretta(1998), where Michaelis-Menten-type ratio-dependent model is studied systematically. Their paper, while contains many new and significant results, is far from complete in answering the many subtle mathematical questions on the global qualitative behavior of solutions of the model. Indeed, many of such important open questions are mentioned in the discussion section of their paper. Through a simple change of variable, we transform the Michaelis-Menten-type ratio-dependent model to a better studied Gause-type predator-prey system. As a result, we can obtain a complete classification of the asymptotic behavior of the solutions of the Michaelis-Menten-type ratio-dependent model. In some cases we can determine how the outcomes depend on the initial conditions. In particular, open questions on the global stability of all equilibria in various cases and the uniqueness of limit cycles are resolved. Biological implications of our results are also presented.

Animals↗

PubMed2006

Population models with singular equilibrium.

A class of models of biological population and communities with a singular equilibrium at the origin is analyzed; it is shown that these models can possess a dynamical regime of deterministic extinction, which is crucially important from the biological standpoint. This regime corresponds to the presence of a family of homoclinics to the origin, so-called elliptic sector. The complete analysis of possible topological structures in a neighborhood of the origin, as well as asymptotics to orbits tending to this point, is given. An algorithmic approach to analyze system behavior with parameter changes is presented. The developed methods and algorithm are applied to existing mathematical models of biological systems. In particular, we analyze a model of anticancer treatment with oncolytic viruses, a parasite-host interaction model, and a model of Chagas' disease.

Algorithms↗

PubMed2001

Avoidance of conspecific injury-released chemical cues by free-ranging Gammarus lacustris (Crustacea:Amphipoda).

Behavioral responses to chemical cues have been demonstrated for a range of aquatic animals. Injury-released chemical alarm cues from conspecifics are released when a prey's predator is actively foraging. Detection of these cues elicits antipredator behaviors that reduce the probability of predation. Amphipod crustaceans in the genus Gammarus are widespread denizens of ponds and streams. Antipredator responses by Gammarus to conspecific alarm cues, and subsequent reduction of predation risk, are known from experiments in the laboratory. Here, we verify laboratory findings by demonstrating an avoidance response to alarm cues using a field population of G. lacustris. We used small traps baited with sponge blocks containing either water (control) or injury-released cues from Gammarus. We repeated the experiment twice. In both experiments, significantly fewer Gammarus were captured in traps with alarm cue sponges than in traps with water sponges. Predatory leeches Dina parva were attracted to Gammarus traps in the first experiment but not the second experiment. In the second experiment, we measured the individual weight of captured amphipods. Two size classes were present; small (1-5 mg) and large (35-108 mg). Both sizes contributed to the avoidance response. Within the large size class, small individuals were significantly less responsive to the alarm cue than large individuals, implying that small adult Gammarus may have different cost/benefit decision criteria for risk assessment than large Gammarus.

Animal Communication↗

PubMed2006

Use of urea as a chemosensory cloaking molecule by a bony fish.

Because urea is bioenergetically expensive to synthesize, few aquatic teleostean (bony) fish make or excrete much urea beyond early development and excrete the majority of nitrogenous waste as the readily diffusible ammonia. The gulf toadfish is one of a few adult teleostean fish that excretes predominantly urea. Most studies of chemosensing by fish predators have focused on amino acids as odorants, but we tested the chemo-attractiveness of both urea and ammonia. We report that characteristic "prey-attack" behaviors by a key toadfish predator, gray snapper, were elicited by low ammonia concentrations (<100 nmol N l(-1)) and similar urea concentrations blunted the ammonia-induced component of attacks. Thus, urea functions as a cloaking molecule, explaining why toadfish co-excrete urea with ammonia. Furthermore, ammonia waste is an important chemical attractant for piscine predators.

Amino Acids↗

PubMed2004

The rat exposure test: a model of mouse defensive behaviors.

In order to facilitate behavioral, and potentially pharmacological, analyses of risk assessment behaviors in mice, a rat exposure test (RET) was devised and evaluated. This test provides a home chamber connected via a tunnel to a rat (predator) exposure area. Familiar substrate is provided to permit burying, and mouse subjects are habituated to the apparatus prior to exposure to an amphetamine-activated rat. In comparison to toy-rat-exposed controls, rat-exposed BALB/c mice showed significantly more risk assessment [stretch attend posture (SAP) and stretch approach], freezing, and avoidance (time in the home chamber), and less time in contact with the wire mesh screen between itself and the threat stimulus. When BALB/c, C57BL/6, CD-1, and Swiss-Webster mice were compared in this test, the two inbred strains (BALB/c and C57BL/6) tended to show more extreme values of particular defensive behaviors, compared to the two outbred strains (Swiss-Webster and CD-1). C57BL/6 mice showed more avoidance and higher levels of SAP, freezing, and burying than BALB/c and more than one or both outbred strains as well. BALB/c mice showed little defensive burying, both in comparison to toy-exposed controls (Experiment 1), and in comparison to the three other strains in Experiment 2. These findings are somewhat at variance with characterizations of anxiety in C57BL/6 and BALB/c mice, based on tests utilizing novel areas and noxious stimuli, suggesting strain differences in defensiveness to such stimuli, compared to antipredator defense levels. Nonetheless, with the exception of burying in BALB/c mice, all strains showed all defensive behaviors measured to the rat stimulus. In particular, SAP levels were substantial in all strains tested, suggesting the usefulness of this test in assessment of the role of risk assessment in defense.

Aggression↗

PubMed1993

[Lemurs of Madagascar. Tests on evolution of primate communities].

The flora and fauna of Madagascar evolved rather independently from the African mainland. In contrast to other oceanic islands, Madagascar is large enough to house most major components of tropical ecosystems, allowing tests of evolutionary hypotheses on the level of complete communities. Taking lemurs, the primates of Madagascar, as an example, evolutionary hypotheses correctly predict the organization of their community structure with respect to ecological correlates. Lemur social systems and their morphological correlates, on the other hand, deviate largely from the typical mammalian pattern. Thus, the traditional hypotheses of behavioral ecology, based solely on resource distribution and predation pressure, are insufficient to explain the existing variability in lemur social systems. Other factors, such as activity patterns and avoidance of infanticide, may be equally important. Due to interspecific variation in these characters, lemurs offer the unique opportunity to determine the relative importance of these factors for the evolution of social systems.

Animals↗