Complex dynamics and phase synchronization in spatially extended ecological systems.

Population cycles that persist in time and are synchronized over space pervade ecological systems, but their underlying causes remain a long-standing enigma. Here we examine the synchronization of complex population oscillations in networks of model communities and in natural systems, where phenomena such as unusual '4- and 10-year cycle' of wildlife are often found. In the proposed spatial model, each local patch sustains a three-level trophic system composed of interacting predators, consumers and vegetation. Populations oscillate regularly and periodically in phase, but with irregular and chaotic peaks together in abundance-twin realistic features that are not found in standard ecological models. In a spatial lattice of patches, only small amounts of local migration are required to induce broad-scale 'phase synchronization, with all populations in the lattice phase-locking to the same collective rhythm. Peak population abundances, however, remain chaotic and largely uncorrelated. Although synchronization is often perceived as being detrimental to spatially structured populations, phase synchronization leads to the emergence of complex chaotic travelling-wave structures which may be crucial for species persistence.     

Spatial synchronization of vole population dynamics by predatory birds

Northern vole populations exhibit large-scale, spatially synchronous population dynamics. Such cases of population synchrony provide excellent opportunities for distinguishing between local intrinsic and regional extrinsic mechanisms of population regulation. Analyses of large-scale survey data and theoretical modelling have indicated several plausible synchronizing mechanisms. It is difficult, however, to determine the most important one without detailed data on local demographic processes. Here we combine results from two field studies in southeastern Norway--one identifies local demographic mechanisms and landscape-level annual synchrony among 28 enclosed experimental populations and the other examines region-level multi-annual synchrony in open natural populations. Despite fences eliminating predatory mammals and vole dispersal, the growth rates of the experimental populations were synchronized and moreover, perfectly linked with vole abundance in the region. The fates of 481 radio-marked voles showed that bird predation was the synchronizing mechanism. A higher frequency of risky dispersal movements in slowly growing populations appeared to accelerate predation rate. Thus, dispersal may induce a feedback-loop between predation and population growth that enhances synchrony.     

Spatial clumping of sexually receptive females induces space sharing among male voles

The spatial organization of individuals in populations (their spacing system) can be highly variable even among populations of the same species. As spacing systems have important consequences for ecological processes such as population regulation, competition and mating systems, there have been many attempts to explore factors that may cause this variation. For mammals, it has been argued that the spatial distribution of sexually receptive females is the most important factor determining the spacing system of males, whereas habitat characteristics are most important to females. This has been difficult to test experimentally as it requires manipulations of the spatial distribution of the opposite sex without changing other properties of the environment. Here, I present a novel experimental procedure that can achieve this and demonstrate that the spatial distribution of the opposite sex in a population of voles is indeed an important determinant of the spacing system of males, but not of females. However, the effects on males are different from those predicted by many theoretical studies.     

生境片断化对植物的遗传影响

生境片断化指大片连续的生境被分割成空间隔离的小生境的现象。通常认为，生境片断化使植物残遗种群由于经历随机遗传漂变和高水平的近交以及基因流的下降，而发生遗传侵蚀，遗传多样性下降，遗传结构改变，变得更加分化。但近期的研究证明，并不是所有的片断化事件都导致植物种群遗传变异丧失，且种群问的基因流甚至有可能增加了。片断化对植物种群的遗传作用还受到诸如世代长短的种间差异、片断化前的丰富度、有性和无性繁殖系统的多样性、种子库以及和传粉媒介及种子传播者的相互作用等的影响。     

Neutral metacommunity models predict fish diversity patterns in Mississippi-Missouri basin

River networks, seen as ecological corridors featuring connected and hierarchical dendritic landscapes for animals and plants, present unique challenges and opportunities for testing biogeographical theories and macroecological laws. Although local and basin-scale differences in riverine fish diversity have been analysed as functions of energy availability and habitat heterogeneity, scale-dependent environmental conditions and river discharge, a model that predicts a comprehensive set of system-wide diversity patterns has been hard to find. Here we show that fish diversity patterns throughout the Mississippi-Missouri River System are well described by a neutral metacommunity model coupled with an appropriate habitat capacity distribution and dispersal kernel. River network structure acts as an effective template for characterizing spatial attributes of fish biodiversity. We show that estimates of average dispersal behaviour and habitat capacities, objectively calculated from average runoff production, yield reliable predictions of large-scale spatial biodiversity patterns in riverine systems. The success of the neutral theory in two-dimensional forest ecosystems and here in dendritic riverine ecosystems suggests the possible application of neutral metacommunity models in a diverse suite of ecosystems. This framework offers direct linkage from large-scale forcing, such as global climate change, to biodiversity patterns.     

Local dispersal promotes biodiversity in a real-life game of rock-paper-scissors

One of the central aims of ecology is to identify mechanisms that maintain biodiversity. Numerous theoretical models have shown that competing species can coexist if ecological processes such as dispersal, movement, and interaction occur over small spatial scales. In particular, this may be the case for non-transitive communities, that is, those without strict competitive hierarchies. The classic non-transitive system involves a community of three competing species satisfying a relationship similar to the children's game rock-paper-scissors, where rock crushes scissors, scissors cuts paper, and paper covers rock. Such relationships have been demonstrated in several natural systems. Some models predict that local interaction and dispersal are sufficient to ensure coexistence of all three species in such a community, whereas diversity is lost when ecological processes occur over larger scales. Here, we test these predictions empirically using a non-transitive model community containing three populations of Escherichia coli. We find that diversity is rapidly lost in our experimental community when dispersal and interaction occur over relatively large spatial scales, whereas all populations coexist when ecological processes are localized.     

Predators increase the risk of catastrophic extinction of prey populations.

There has been considerable research on both top-down effects and on disturbances in ecological communities; however, the interaction between the two, when the disturbance is catastrophic, has rarely been examined. Predators may increase the probability of prey extinction resulting from a catastrophic disturbance both by reducing prey population size and by changing ecological traits of prey individuals such as habitat characteristics in a way that increases the vulnerability of prey species to extinction. We show that a major hurricane in the Bahamas led to the extinction of lizard populations on most islands onto which a predator had been experimentally introduced, whereas no populations became extinct on control islands. Before the hurricane, the predator had reduced prey populations to about half of those on control islands. Two months after the hurricane, we found only recently hatched individuals--apparently lizards survived the inundating storm surge only as eggs. On predator-introduction islands, those hatchling populations were a smaller fraction of pre-hurricane populations than on control islands. Egg survival allowed rapid recovery of prey populations to pre-hurricane levels on all control islands but on only a third of predator-introduction islands--the other two-thirds lost their prey populations. Thus climatic disturbance compounded by predation brought prey populations to extinction.     

生境破坏的空间结构对集合种群动态的影响

构建任意生境破坏结构下集合种群动态的元胞自动机模型,研究生境破坏的空间结构对集合种群全局密度及局部密度的影响.该模型不仅包括了生境破坏的空间结构,而且能够描述集合种群的空间分布模式.模型结果表明:生境破坏的空间结构复杂地影响着集合种群的全局密度和局部密度;生境破坏的混合分布增大了集合种群灭绝的风险;生境破坏的聚集分布降低了集合种群灭绝的风险.在生境破坏比例较小或较大时,集合种群全局密度和局部密度随生境破坏比例的增加而加大,随生境破坏聚集度的增加而减小,且生境破坏比例对集合种群动态的影响要大于生境破坏聚集度的影响.但在特定的情况下,即生境破坏比例在55%左右时,生境聚集度的增加可能会使集合种群的全局密度和局部密度都减小.此外,发现集合种群的局部密度存在一个阈值,当集合种群局部密度低于此值时,集合种群将无法续存.这意味着在生物保护中不能仅仅考虑生境的恢复、斑块的质量,还需要考虑生境破坏的空间结构和被保护物种的局部密度.     

Evolutionary biology: adaptive developmental plasticity in snakes

The morphology of organisms is generally well matched to their environment, presumably because expression of their genes is tailored either at the population or the individual level to suit local conditions: for example, snake populations that persistently encounter large prey may accumulate gene mutations that specify a large head size, or head growth may be increased in individual snakes to meet local demands (adaptive developmental plasticity). Here we test the relative contributions of genetics and environment to the jaw sizes of two tiger snake populations: one that consumes small prey on the mainland, and an island population that relies on larger prey and has a larger jaw size. Although the idea of adaptive plasticity in response to environmental pressures is controversial, we find that both factors influence the difference in jaw size between the two populations, and the influence of developmental plasticity is greater in the island population.     

Adaptation varies through space and time in a coevolving host-parasitoid interaction

One of the central challenges of evolutionary biology is to understand how coevolution organizes biodiversity over complex geographic landscapes. Most species are collections of genetically differentiated populations, and these populations have the potential to become adapted to their local environments in different ways. The geographic mosaic theory of coevolution incorporates this idea by proposing that spatial variation in natural selection and gene flow across a landscape can shape local coevolutionary dynamics. These effects may be particularly strong when populations differ across productivity gradients, where gene flow will often be asymmetric among populations. Conclusive empirical tests of this theory have been particularly difficult to perform because they require knowledge of patterns of gene flow, historical population relationships and local selection pressures. We have tested these predictions empirically using a model community of bacteria and bacteriophage (viral parasitoids of bacteria). We show that gene flow across a spatially structured landscape alters coevolution of parasitoids and their hosts and that the resulting patterns of adaptation can fluctuate in both space and time.     

The effect of migration on local adaptation in a coevolving host-parasite system

Antagonistic coevolution between hosts and parasites in spatially structured populations can result in local adaptation of parasites; that is, the greater infectivity of local parasites than foreign parasites on local hosts. Such parasite specialization on local hosts has implications for human health and agriculture. By contrast with classic single-species population-genetic models, theory indicates that parasite migration between subpopulations might increase parasite local adaptation, as long as migration does not completely homogenize populations. To test this hypothesis we developed a system-specific mathematical model and then coevolved replicate populations of the bacterium Pseudomonas fluorescens and a parasitic bacteriophage with parasite only, with host only or with no migration. Here we show that patterns of local adaptation have considerable temporal and spatial variation and that, in the absence of migration, parasites tend to be locally maladapted. However, in accord with our model, parasite migration results in parasite local adaptation, but host migration alone has no significant effect.     

一类具有强食饵和弱食饵的捕食系统分析

提出并研究了一类具有两个相互竞争的食饵种群与一个以这两个物种为食的捕食者种群的捕食系统.首先通过比较原理研究了该系统解的正有界性及系统平衡点的存在性,并运用特征值方法对可能存在的平衡点的局部渐近稳定性进行了讨论.其次利用微分方程的基本理论,得到了系统持久性的充分条件.然后通过构造Lyapunov函数讨论了系统平衡点的全局稳定性.最后,利用数值模拟验证本文的主要结论.     

3物种食物链的集合群落中捕食者追赶和猎物逃逸对空间同步性的影响

空间同步性能够导致区域物种灭绝从而降低集合种群的续存.在集合群落的框架内建立了两斑块3种群食物链联立模型来研究捕食者追赶和猎物逃逸对空间同步性的影响.由于种群存在较大的群落中,从集合种群向集合群落扩展是很自然的.表明了在集合种群中猎物逃逸降低空间同步性的结论在集合群落中有所改变.得到一个有悖直觉的结果:3物种食物链基层物种的逃逸并没有保护自己,反而增加了所有这3物种集合种群的空间同步性,因此也增加了灭绝的风险.这个结果说明在制定关于种群同步性也即续存的保护措施时,物种间的相互作用也必须考虑.     

多辐溲疏居群过氧化物酶同工酶分化的研究

运用聚丙烯酰胺凝胶电泳，测试了采自四川南川县金佛山不同生境的多辐溲疏５个居群５０株个体的过氧化物酶同工酶，对酶谱带位置进行了二态编码，然后采用Ｒｏｇｅｒｓ－Ｔａｎｉｍｏｔｏ，Ｓｏｋａｌ－Ｍｉｃｈ－ｅｎｅｒ，Ｓｏｋａｌ－Ｓｎｅａｔｈ，Ｊａｃｃａｒｄ等结合系数用组平均法（ＵＰＧＭＡ）对各谱带位置进行了聚类分析．结果表明，相同生境同一居群内的个体间表现出较大的遗传同质性，而来源于不同生境的不同居群个体间却呈现出较大的遗传趋异性；同时，５个居群不但具有许多共同谱带，而且整体相聚的结合系数大多超过０．５，这表明５个居群在很早以前可能是属于同一大居群，在长期进化过程中，由于微生境的地理与气候因子选择，使得多辐溲疏现仅在金佛山海拔６００～１２００ｍ的约ｋｍ２范围内呈岛状分布；又由于其特殊的繁殖方式——居群内个体虽能杂交，但因该种仅能正常开花传粉，大部分种子败育，使得居群内个体遗传变异性不能通过有性生殖方式固定下来，仅能通过营养繁殖方式繁殖后代，而正是营养繁殖导致了居群内较大遗传同质性以及居群间的遗传趋异性．在整个进化过程中生境只不过是对那些遗传变异起到了选择固定作用     

Speciation along environmental gradients

Traditional discussions of speciation are based on geographical patterns of species ranges. In allopatric speciation, long-term geographical isolation generates reproductively isolated and spatially segregated descendant species. In the absence of geographical barriers, diversification is hindered by gene flow. Yet a growing body of phylogenetic and experimental data suggests that closely related species often occur in sympatry or have adjacent ranges in regions over which environmental changes are gradual and do not prevent gene flow. Theory has identified a variety of evolutionary processes that can result in speciation under sympatric conditions, with some recent advances concentrating on the phenomenon of evolutionary branching. Here we establish a link between geographical patterns and ecological processes of speciation by studying evolutionary branching in spatially structured populations. We show that along an environmental gradient, evolutionary branching can occur much more easily than in non-spatial models. This facilitation is most pronounced for gradients of intermediate slope. Moreover, spatial evolutionary branching readily generates patterns of spatial segregation and abutment between the emerging species. Our results highlight the importance of local processes of adaptive divergence for geographical patterns of speciation, and caution against pitfalls of inferring past speciation processes from present biogeographical patterns.     

Strong effect of dispersal network structure on ecological dynamics

A central question in ecology with great importance for management, conservation and biological control is how changing connectivity affects the persistence and dynamics of interacting species. Researchers in many disciplines have used large systems of coupled oscillators to model the behaviour of a diverse array of fluctuating systems in nature. In the well-studied regime of weak coupling, synchronization is favoured by increases in coupling strength and large-scale network structures (for example 'small worlds') that produce short cuts and clustering. Here we show that, by contrast, randomizing the structure of dispersal networks in a model of predators and prey tends to favour asynchrony and prolonged transient dynamics, with resulting effects on the amplitudes of population fluctuations. Our results focus on synchronization and dynamics of clusters in models, and on timescales, more appropriate for ecology, namely smaller systems with strong interactions outside the weak-coupling regime, rather than the better-studied cases of large, weakly coupled systems. In these smaller systems, the dynamics of transients and the effects of changes in connectivity can be well understood using a set of methods including numerical reconstructions of phase dynamics, examinations of cluster formation and the consideration of important aspects of cyclic dynamics, such as amplitude.     

一类捕食与被捕食系统的持续生存性与周期解

讨论了一类一个捕食者种群和两个食饵种群构成的非自治生态系统．系统中食饵种群有各自的避难所，它们在避难所内外可以迁移．应用常微分方程定性和稳定性的方法，得到了在一定条件下系统是持续生存的；进一步，若系统是一个周期系统，则在合适的条件下，系统存在唯一的、全局渐近稳定的周期解     

集合种群受生境破坏影响的新模型及其生态效应研究

从两物种竞争模型出发,提出了集合种群上3物种的食物网机制,构造了一个捕食者和两个猎物(似然竞争)受生境破坏影响的新模型;基于微分动力系统的数值模拟和网格模型的元胞自动机模拟,研究了生境破坏对集合种群动态的影响.结果发现:集合种群似然竞争模型系统对生境破坏非常敏感,生境破坏导致了更不稳定的动态,而且发现具有较小捕食者侵占率的猎物受生境破坏影响较大,进而表明了猎物对捕食者的敏感度与受生境破坏的影响之间存在负藕联关系;适当的生境破坏有利于劣势物种的生存;捕食者通过猎物受到生境破坏的间接影响,而且在生境破坏生态过程中捕食者是脆弱的.该研究结果为物种续存和生物多样性保护提供了理论依据.     

车辆编队系统的多目标分布式控制

针对车辆编队系统,通过引入空间变量和空间移动算子,建立多维的空间关联系统模型.采用多目标分布式控制方式,同时考虑系统的动态过程和能量消耗,设计了与被控对象有着相同关联结构的分布式反馈H2/H∞控制器,在系统具有较好动态性能的同时使能量消耗尽可能地少.以弹簧质点模型来模拟车辆的编队模型,进行仿真实验,仿真结果说明方法的有效性和可行性.     

Evolution driven by differential dispersal within a wild bird population

Evolutionary theory predicts that local population divergence will depend on the balance between the diversifying effect of selection and the homogenizing effect of gene flow. However, spatial variation in the expression of genetic variation will also generate differential evolutionary responses. Furthermore, if dispersal is non-random it may actually reinforce, rather than counteract, evolutionary differentiation. Here we document the evolution of differences in body mass within a population of great tits, Parus major, inhabiting a single continuous woodland, over a 36-year period. We show that genetic variance for nestling body mass is spatially variable, that this generates different potential responses to selection, and that this diversifying effect is reinforced by non-random dispersal. Matching the patterns of variation, selection and evolution with population ecological data, we argue that the small-scale differentiation is driven by density-related differences in habitat quality affecting settlement decisions. Our data show that when gene flow is not homogeneous, evolutionary differentiation can be rapid and can occur over surprisingly small spatial scales. Our findings have important implications for questions of the scale of adaptation and speciation, and challenge the usual treatment of dispersal as a force opposing evolutionary differentiation.