The adaptive dynamics of Lotka-Volterra systems with trade-offs

We analyse the adaptive dynamics of a generalised type of Lotka-Volterra model subject to an explicit trade-off between two parameters. A simple expression for the fitness of a mutant strategy in an environment determined by the established, resident strategy is obtained leading to general results for the position of the evolutionary singular strategy and the associated second-order partial derivatives of the mutant fitness with respect to the mutant and resident strategies. Combinations of these results can be used to determine the evolutionary behaviour of the system. The theory is motivated by an example of prey evolution in a predator-prey system in which results show that only (non-EUS) evolutionary repellor dynamics, where evolution is directed away from a singular strategy, or dynamics where the singular strategy is an evolutionary attractor, are possible. Moreover, the general theory can be used to show that these results are the only possibility for all Lotka-Volterra systems in which aside from the trade-offs all parameters are independent and in which the interaction terms are of quadratic order or less. The applicability of the theory is highlighted by examining the evolution of an intermediate predator in a tri-trophic model.     

Permanence of some ecological systems with several predator and one prey species

The stability criterion used in the following is permanence. Permanence means that all trajectories starting in the interior are ultimately bounded away from the boundary and that this bound is independent of the initial values. Hence sufficiently small fluctuations cannot lead to extinction of any species. In the following we deal with one-prey, two-predator resp. one-prey, three-predator systems and a one-prey, two-predator, one-top-predator system with three trophic levels. It turns out that the characterization of permanence for such models described by Lotka-Volterra dynamics is rather simple and elegant.     

Adaptive dynamics of Lotka-Volterra systems with trade-offs: the role of interspecific parameter dependence in branching

We determine the adaptive dynamics of a general Lotka-Volterra system containing an intraspecific parameter dependency--in the form of an explicit functional trade-off between evolving parameters--and interspecific parameter dependencies--arising from modelling species interactions. We develop expressions for the fitness of a mutant strategy in a multi-species resident environment, the position of the singular strategy in such systems and the non-mixed second-order partial derivatives of the mutant fitness. These expressions can be used to determine the evolutionary behaviour of the system. The type of behaviour expected depends on the curvature of the trade-off function and can be interpreted in a biologically intuitive manner using the rate of acceleration/deceleration of the costs implicit in the trade-off function. We show that for evolutionary branching to occur we require that one (or both) of the traded-off parameters includes an interspecific parameter dependency and that the trade-off function has weakly accelerating costs. This could have important implications for understanding the type of mechanisms that cause speciation. The general theory is motivated by using adaptive dynamics to examine evolution in a predator-prey system. The applicability of the general theory as a tool for examining specific systems is highlighted by calculating the evolutionary behaviour in a three species (prey-predator-predator) system.     

The roles of ecological and behavioral observation in species recognition among primates

Closely related living mammal species present numerous difficulties of recognition and delineation, not least because populations as well as individual organisms fulfill both genetic and economic functions. However, since species are often distributed geographically across multiple ecosystems or environments, it is evident that species as wholes, except where coterminous with the local populations of which they are composed, cannot be said to play unitary ecological roles. Aspects of the economic activity of organisms thus may not be admitted as elements in species recognition. Those who study the behavior of mammals in their natural habitats must therefore focus upon behaviors which preserve the genetic integrity of species if they wish to contribute to the systematic question of species identification. «Isolation» concepts of species are not served by data of this kind, since they emphasize interspecies discontinuities; the «recognition» concept of species, however, specifically focuses upon behaviors of this kind as they contribute to the «Specific-Mate Recognition System». Among primates such behaviors are likely to involve signalling of various kinds; where this is visual it is reasonable to expect to find morphological or chromatic correlates; where such cues are auditory or, to a lesser extent, olfactory, it is less probable that such correlates will be found to exist.     

Life history and mating systems select for male biased parasitism mediated through natural selection and ecological feedbacks

tlsb-1%Males are often the ‘sicker’ sex with male biased parasitism found in a taxonomically diverse range of species. There is considerable interest in the processes that could underlie the evolution of sex-biased parasitism. Mating system differences along with differences in lifespan may play a key role. We examine whether these factors are likely to lead to male-biased parasitism through natural selection taking into account the critical role that ecological feedbacks play in the evolution of defence. We use a host-parasite model with two-sexes and the techniques of adaptive dynamics to investigate how mating system and sexual differences in competitive ability and longevity can select for a bias in the rates of parasitism. Male-biased parasitism is selected for when males have a shorter average lifespan or when males are subject to greater competition for resources. Male-biased parasitism evolves as a consequence of sexual differences in life-history that produce a greater proportion of susceptible females than males and therefore reduce the cost of avoiding parasitism in males. Different mating systems such as monogamy, polygyny or polyandry did not produce a bias in parasitism through these ecological feedbacks but may accentuate an existing bias.     

Effects of Hebbian learning on the dynamics and structure of random networks with inhibitory and excitatory neurons

The aim of the present paper is to study the effects of Hebbian learning in random recurrent neural networks with biological connectivity, i.e. sparse connections and separate populations of excitatory and inhibitory neurons. We furthermore consider that the neuron dynamics may occur at a (shorter) time scale than synaptic plasticity and consider the possibility of learning rules with passive forgetting. We show that the application of such Hebbian learning leads to drastic changes in the network dynamics and structure. In particular, the learning rule contracts the norm of the weight matrix and yields a rapid decay of the dynamics complexity and entropy. In other words, the network is rewired by Hebbian learning into a new synaptic structure that emerges with learning on the basis of the correlations that progressively build up between neurons. We also observe that, within this emerging structure, the strongest synapses organize as a small-world network. The second effect of the decay of the weight matrix spectral radius consists in a rapid contraction of the spectral radius of the Jacobian matrix. This drives the system through the "edge of chaos" where sensitivity to the input pattern is maximal. Taken together, this scenario is remarkably predicted by theoretical arguments derived from dynamical systems and graph theory.     

Relationships among recent models for insect population dynamics with variable rates of development

A classification scheme for those population models which allow variation in development rates is proposed, based on two ways of modifying standard age-structured models. The resulting classes of models are termed development index models and sojourn time models. General formulations for the two classes of models are developed from two basic balance equations, and numerous specific models from the literature are shown to fit into the scheme. Concepts from competing risks theory are shown to be important in understanding the interplay between mortality and maturation. Relationships among the classes are investigated both for the most general forms of the models and for the simpler forms often used. The scheme can provide guidance in developing appropriate insect population models for specific modelling situations.Contribution 3878871     

A test of hybrid growth disadvantage in wild, free-ranging species pairs of threespine stickleback (Gasterosteus aculeatus) and its implications for ecological speciation

Ecological speciation is the evolution of reproductive isolation as a direct or indirect consequence of divergent natural selection. Reduced performance of hybrids in nature is thought to be an important process by which natural selection can favor the evolution of assortative mating and drive speciation. Benthic and limnetic sympatric species of threespine stickleback (Gasterosteus aculeatus) are adapted to alternative trophic niches (bottom browsing vs. open water planktivory, respectively) and reduced feeding performance of hybrids is thought to have contributed to the evolution of reproductive isolation. We tested this “hybrid‐disadvantage hypothesis” by inferring growth rates from otoliths sampled from wild, free‐ranging benthic, limnetic, and hybrid sticklebacks in two lakes. There were significant differences in growth rate between lakes, life‐history stages, and among years (maximum P = 0.02), as well as interactions between most factors, but not between hybrid and parental species sticklebacks in most comparisons. Our results provide little evidence of a growth disadvantage in hybrid sticklebacks when free‐ranging in nature. Although trophic ecology per se may contribute less to ecological speciation than envisioned, it may act in concert with other aspects of stickleback biology, such as interactions with parasites, predators, competitors, and/or sexual selection, to present strong multifarious selection against hybrids.     

CSS, NIS and dynamic stability for two-species behavioral models with continuous trait spaces

Static continuously stable strategy (CSS) and neighborhood invader strategy (NIS) conditions are developed for two-species models of frequency-dependent behavioral evolution when individuals have traits in continuous strategy spaces. These are intuitive stability conditions that predict the eventual outcome of evolution from a dynamic perspective. It is shown how the CSS is related to convergence stability for the canonical equation of adaptive dynamics and the NIS to convergence to a monomorphism for the replicator equation of evolutionary game theory. The CSS and NIS are also shown to be special cases of neighborhood p^*- superiority for p^* equal to one half and zero, respectively. The theory is illustrated when each species has a one-dimensional trait space.     

Effects of variation in individual growth on plant species coexistence

Abstract. Both size structure and variability (spatial heterogeneity, disturbance, stochasticity, variation in species attributes, etc.) are regarded as regulatory mechanisms of species coexistence. However, none of the models so far proposed consider both size structure and variability simultaneously. A size-structured variation model for plant-community dynamics is proposed, which is based on the diffusion model for growth dynamics of plant populations. This model has four functions: (1) mean growth rate of individuals of size x at time t, G(t, x) (species-specific mean traits, e.g. competitive ability); (2) variance in growth rate of individuals of size x at time t, D(t, x) (stochastic factors due to genetic variation, environmental heterogeneity, spatial variation of individuals, etc.); (3) mortality rate of individuals of size x at time t, M(t, x); and (4) recruitment rate at time t, R(t), as a boundary condition. The interference function for individuals of size x at time t, C(t, x), is introduced, which expresses the degree of interactions between individuals and hence averaged effects of local neighbourhood competition; the G(t, x), D(t, x), M(t, x) and R(t) functions are given in terms of C(t, x). These four functions describe the growth dynamics of individuals of each species in the plant community. Effects of the G(t, x), D(t, x), M(t, x) and R(t) functions on species coexistence in plant communities were evaluated by simulation and the relative importance of the D(t, x) function as well as size structure was shown for species coexistence especially in plant communities where competition among species is non-transitive or niche limitation does not work.     

Consequences of behavioral dynamics for the population dynamics of predator-prey systems with switching

This article explores how different mechanisms governing the rate of change of the predators preference alter the dynamics of predator-prey systems in which the predator exhibits positive frequency-dependent predation. The models assume that individuals of the predator species adaptively adjust a trait that determines their relative capture rates of each of two prey species. The resulting switching behavior does not instantaneously attain the optimum for current prey densities, but instead lags behind it. Several mechanisms producing such lags are discussed and modeled. In all cases examined, our question is whether a realistic behavioral lag can significantly change the dynamics of the system relative to an analogous case in which the predators switching is effectively instantaneous. We also explore whether increasing the rate parameters of dynamic models of behavior results in convergence to the population dynamics of analogous models with instantaneous switching, and whether different behavioral models produce similar population dynamics. The analysis concentrates on systems that undergo endogenously generated predator-prey cycles in the absence of switching behavior. The average densities and the nature of indirect interactions are often sensitive to the rate of behavioral change, and are often qualitatively different for different classes of behavioral models. Dynamics and average densities can be very sensitive to small changes in parameters of either the prey growth or predator switching functions. These differences suggest that an understanding of switching in natural systems will require research into the behavioral mechanisms that govern lags in the response of predator preference to changes in prey density.     

Evolution of specialization and ecological character displacement of herbivores along a gradient of plant quality

We study the combined evolutionary dynamics of herbivore specialization and ecological character displacement, taking into account foraging behavior of the herbivores, and a quality gradient of plant types. Herbivores can adapt by changing two adaptive traits: their level of specialization in feeding efficiency and their point of maximum feeding efficiency along the plant gradient. The number of herbivore phenotypes, their levels of specialization, and the amount of character displacement among them are the result of the evolutionary dynamics, which is driven by the underlying population dynamics, which in turn is driven by the underlying foraging behavior. Our analysis demonstrates broad conditions for the diversification of a herbivore population into many specialized phenotypes, for basically any foraging behavior focusing use on highest gains while also including errors. Our model predicts two characteristic phases in the adaptation of herbivore phenotypes: a fast character-displacement phase and a slow coevolutionary niche-shift phase. This two-phase pattern is expected to be of wide relevance in various consumer-resource systems. Bringing together ecological character displacement and the evolution of specialization in a single model, our study suggests that the foraging behavior of herbivorous arthropods is a key factor promoting specialist radiation.     

Stability of the replicator equation for a single species with a multi-dimensional continuous trait space

The replicator equation model for the evolution of individual behaviors in a single species with a multi-dimensional continuous trait space is developed as a dynamics on the set of probability measures. Stability of monomorphisms in this model using the weak topology is compared to more traditional methods of adaptive dynamics. For quadratic fitness functions and initial normal trait distributions, it is shown that the multi-dimensional continuously stable strategy (CSS) of adaptive dynamics is often relevant for predicting stability of the measure-theoretic model but may be too strong in general. For general fitness functions and trait distributions, the CSS is related to dominance solvability which can be used to characterize local stability for a large class of trait distributions that have no gaps in their supports whereas the stronger neighborhood invader strategy (NIS) concept is needed if the supports are arbitrary.     

Effects of food supplies and water temperature on growth rates of two species of freshwater tropical shrimps

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Copepods of the genus Asterocheres (Copepoda: Siphonostomatoida) feeding on sponges: behavioral and ecological traits

Abstract. Four copepod species of the genus Asterocheres (A. echinicola, A. latus, A. suberitis and A. complexus ) are recorded for the first time as associated with Mediterranean sponges of the classes Calcarea and Demospongiae (orders Dictyoceratida, Dendroceratida, and Poecilosclerida). Individuals of A. suberitis are described for the first time as inhabiting sponges other than Suberites domuncula . The copepods seemed to use their trunk to suck the material (cuticles or excreted cells) produced at the sponge periphery (ectosome), which indicates that their feeding habits are similar to those observed in species of the same genus that feed on exfoliated tissues of echinoderms. In the laboratory, copepods fed on young sponges (rhagons) of the species Crambe crambe, Cacospongia mollior , and Dysidea avara without reducing the rhagon biomass. In contrast, rhagons of the species Haliclona sp., Hymedesmia spp., and Phorbas tenacior were consumed and their biomass reduced. This may be because the latter group does not produce external proteinaceous material. Copepod offspring were continuously released (∼6 nauplii every 48 h) and the larval stages were capable of limited swimming. We found significantly more copepods over the surface of the sponges than inside them. Copepod abundance was positively correlated with the sponge surface area while not with the sponge biomass. These results indicate that the copepods live preferentially over the sponge surface and that their abundance is surface-area dependent. The behavioral traits described here indicate that the interactions between asterocherid species and sponges are worthy of future study.     

The effects of adaptation to urea on feeding rates and growth in Drosophila larvae

A collection of forty populations were used to study the phenotypic adaptation of Drosophila melanogaster larvae to urea‐laced food. A long‐term goal of this research is to map genes responsible for these phenotypes. This mapping requires large numbers of populations. Thus, we studied fifteen populations subjected to direct selection for urea tolerance and five controls. In addition, we studied another twenty populations which had not been exposed to urea but were subjected to stress or demographic selection. In this study, we describe the differentiation in these population for six phenotypes: (1) larval feeding rates, (2) larval viability in urea‐laced food, (3) larval development time in urea‐laced food, (4) adult starvation times, (5) adult desiccation times, and (6) larval growth rates. No significant differences were observed for desiccation resistance. The demographically/stress‐selected populations had longer times to starvation than urea‐selected populations. The urea‐adapted populations showed elevated survival and reduced development time in urea‐laced food relative to the control and nonadapted populations. The urea‐adapted populations also showed reduced larval feeding rates relative to controls. We show that there is a strong linear relationship between feeding rates and growth rates at the same larval ages feeding rates were measured. This suggests that feeding rates are correlated with food intake and growth. This relationship between larval feeding rates, food consumption, and efficiency has been postulated to involve important trade‐offs that govern larval evolution in stressful environments. Our results support the idea that energy allocation is a central organizing theme in adaptive evolution.     

联峰山10种草本植物营养器官形态结构与生态适应性

利用常规石蜡制片技术对生长在联峰山上的10种植物营养器官进行解剖学观察。主要结果如下:(1)叶表皮细胞均为1层,绝大多数无表皮毛,气孔少;异面叶或等面叶;中脉维管束束数1至多个。(2)根:主要为次生构造,也有初生构造的。但大多数植物无周皮;皮层细胞的层数在不同植物中差异较大;有些植物次生韧皮部普遍不发达,次生木质部较韧皮部明显。(3)茎:表皮细胞均为1层,具角质膜;皮层细胞层数普遍较少,外皮层有机械组织存在;维管束数目多个并呈一圈排列,外韧或双韧;髓部发达,髓射线明显。该研究结果表明10种植物具有中生植物或阴性植物的结构特点,是植物长期适应环境条件的结果;该研究为更好地开发和利用植物提供了解剖学依据。     

Consequences of insect herbivory on grape fine root systems with different growth rates

Herbivory tolerance has been linked to plant growth rate where plants with fast growth rates are hypothesized to be more tolerant of herbivory than slower-growing plants. Evidence supporting this theory has been taken primarily from observations of aboveground organs but rarely from roots. Grapevines differing in overall rates of new root production, were studied in Napa Valley, California over two growing seasons in an established vineyard infested with the sucking insect, grape phylloxera (Daktulosphaira vitifoliae Fitch). The experimental vineyard allowed for the comparison of two root systems that differed in rates of new root tip production (a 'fast grower', Vitis berlandieri x Vitis rupestris cv. 1103P, and a slower-growing stock, Vitis riparia x Vitis rupestris cv. 101-14 Mgt). Each root system was grafted with a genetically identical shoot system (Vitis vinifera cv. Merlot). Using minirhizotrons, we did not observe any evidence of spatial or temporal avoidance of insect populations by root growth. Insect infestations were abundant throughout the soil profile, and seasonal peaks in phylloxera populations generally closely followed peaks in new root production. Our data supported the hypothesis that insect infestation was proportional to the number of growing tips, as indicated by similar per cent infestation in spite of a threefold difference in root tip production. In addition, infested roots of the fast-growing rootstock exhibited somewhat shorter median lifespans (60 d) than the slower-growing rootstock (85 d). Lifespans of uninfested roots were similar for the two rootstocks (200 d). As a consequence of greater root mortality of younger roots, infested root populations in the fast-growing rootstock had an older age structure. While there does not seem to be a trade-off between potential growth rate and relative rate of root infestation in these cultivars, our study indicates that a fast-growing root system may more readily shed infested roots that are presumably less effective in water and nutrient uptake. Thus, differences in root tip production may be linked to differences in the way plants cope with roots that are infested by sucking insects.     

月光花素对彩叶芋愈伤组织细胞生长和分化的影响

在含有NAA0.2mg/L,2.4-D 0.2mg/L的MS基础培养基中添加月光花素培养彩叶芋愈伤组织,结果显示,低浓度的月光花素能促进愈伤组织生长,高浓度则有明显的抑制作用,最适浓度为1.0~2.0mg/L;培养两个月后,添加适宜浓度的月光花素(0.2~2.0mg/L)的培养基能促进体细胞分化和植株再生,而高浓度(10.0mg/L)处理和对照均未能分化出绿苗。表明适宜浓度的月光花素在合适的生长素剂量的协同作用下,对植物愈伤组织细胞的生长和分化有较强的调节作用。