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1.
Striped patterns are often observed on fish skin. Such patterns have been accounted for by reaction-diffusion (RD) Turing-type models, in which two substances can spontaneously form a spatially heterogeneous pattern in a homogeneous field. Among the striped patterns generated by Turing-type models, some are "straight-striped patterns," with many stripes running in parallel, while others are "labyrinthine patterns," in which the stripes often change direction, merge with each other, and frequently branch out. RD models differ in terms of their tendency to generate either labyrinthine or straight-striped patterns. Here, we studied the conditions under which either a labyrinthine or straight-striped pattern would emerge. First, we defined an index for stripe clearness, Sh. Straight-striped patterns (large Sh) are formed if only a narrow range of spatial periods corresponds to an unstable mode. Labyrinthine patterns (small Sh) are formed when a wide range of spatial periods is unstable. More specifically, labyrinthine patterns are formed when the maximum spatial period of unstable modes is more than twice that of the minimum spatial period of unstable modes; otherwise, straight-striped patterns are formed. We then examined RD models with nonlinear reaction terms, including both activator-inhibitor and substrate-depletion models, and we demonstrated that the same conclusions hold with respect to the conditions required for labyrinthine versus straight-striped patterns.  相似文献   

2.
A substantial literature treats the dynamics of populations in response to spatial patterning of underlying habitats, the most common formulations being source/sink populations and metapopulations living in a patchwork of habitats. A separate and growing literature on the self‐organization of spatial pattern focuses on how local interactions give rise to regional patterns that can be described with scale‐free distributions. Motivated by the potential ubiquity of the coupling of these processes (spatial self organization and the dynamics of organisms in spatially structured habitats), we link these two lines of thought into a theory of populations in fragmented habitats. Using a combined analytical and computational approach, we show that self organization can generate a background into which an independent population fits, and that the scale‐free nature of such habitat creates conditions that influence both the persistence versus extinction properties of the embedded population and where it lies on the continuum between source/sink populations and metapopulations. Both the analytical framework and the computer simulations demonstrate the potential for populations to persist on either end of the metapopulation–source/sink continuum while failing to persist under intermediate conditions. These results provide a new perspective on ecological theory related to habitat fragmentation and population persistence, suggesting that under some conditions intermediate states between these two extremes may maximize risk of extinction of the population. These implications could be of particular importance for managing self‐organized systems of conservation concern.  相似文献   

3.
Spatial self‐organisation of ecosystems is the process by which large‐scale ordered spatial patterns emerge from disordered initial conditions through local feedbacks between organisms and their environment. Such process is considered important for ecosystem functioning, providing increased productivity, resistance and resilience against environmental change. Although spatial self‐organisation has been found for an increasing number of ecosystems, it has never been shown so far for aquatic river vegetation. Here we explore the existence of spatial self‐organisation of freshwater macrophyte patches in a typical lowland river (Belgium), showing that the underlying mechanisms for pattern formation are scale‐dependent feedbacks between plant growth, water flow and local river bed erosion and sedimentation. The mapping of vegetation patches showed that the frequency distribution of patch sizes is governed by a power‐law function, suggesting that the patches are self‐organised. Scale‐dependent feedbacks, likely to lead to this self‐organised pattern, were demonstrated with a mimic experiment. Both positive and negative feedbacks on plants were confirmed by a transplantation experiment. Placing vegetation patch mimics in the river showed experimentally that on a short range (within and behind the mimics) flow reduction and increased sedimentation occurred, while on a larger range (next to patches) the flow was accelerated and decreased sedimentation took place. By transplanting macrophytes within, next to and further away from existing patches, it was proven that the conditions within the patches favoured the survival and growth of transplants (i.e. short‐range positive feedback), while the conditions just next to patches led to decreased survival and growth (i.e. long‐range negative feedback).  相似文献   

4.
The mysterious ‘fairy circles’ are vegetation‐free discs that cover vast areas along the pro‐Namib Desert. Despite 30 yr of research their origin remains unknown. Here we adopt a novel approach that focuses on analysis of the spatial patterns of fairy circles obtained from representative 25‐ha aerial images of north‐west Namibia. We use spatial point pattern analysis to quantify different features of their spatial structures and then critically inspect existing hypotheses with respect to their ability to generate the observed circle patterns. Our working hypothesis is that fairy circles are a self‐organized vegetation pattern. Finally, we test if an existing partial‐differential‐equation model, that was designed to describe vegetation pattern formation, is able to reproduce the characteristic features of the observed fairy circle patterns. The model is based on key‐processes in arid areas such as plant competition for water and local resource‐biomass feedbacks. The fairy circles showed at all three study areas the same regular spatial distribution patterns, characterized by Voronoi cells with mostly six corners, negative correlations in their size up to a distance of 13 m, and remarkable homogeneity over large spatial scales. These results cast doubts on abiotic gas‐leakage along geological lines or social insects as causal agents of their origin. However, our mathematical model was able to generate spatial patterns that agreed quantitatively in all of these features with the observed patterns. This supports the hypothesis that fairy circles are self‐organized vegetation patterns that emerge from positive biomass‐water feedbacks involving water transport by extended root systems and soil‐water diffusion. Future research should search for mechanisms that explain how the different hypotheses can generate the patterns observed here and test the ability of self‐organization to match the birth‐ and death dynamics of fairy circles and their regional patterns in the density and size with respect to environmental gradients.  相似文献   

5.
Aggregative groups entail costs that must be overcome for the evolution of complex social interactions. Understanding the mechanisms that allow aggregations to form and restrict costs of cheating can provide a resolution to the instability of social evolution. Aggregation in Tetrahymena thermophila is associated with costs of reduced growth and benefits of improved survival through “growth factor” exchange. We investigated what mechanisms contribute to stable cooperative aggregation in the face of potential exploitation by less‐cooperative lines using experimental microcosms. We found that kin recognition modulates aggregative behavior to exclude cheaters from social interactions. Long‐distance kin recognition across patches modulates social structure by allowing recruitment of kin in aggregative lines and repulsion in asocial lines. Although previous studies have shown a clear benefit to social aggregation at low population densities, we found that social aggregation has very different effects at higher densities. Lower growth rates are a cost of aggregation, but also present potential benefits when restricted to kin aggregations: slow growth and crowd tolerance allow aggregations to form and permit longer persistence on ephemeral resources. Thus in highly dynamic metapopulations, kin recognition plays an important role in the formation and stability of social groups that increase persistence through cooperative consumptive restraint.  相似文献   

6.
Pamela Graff  Martín R. Aguiar 《Oikos》2011,120(7):1023-1030
Since many arid ecosystems are overstocked with domestic herbivores, biotic stress could have a stronger influence in modulating the balance of species interactions than expected from the stress gradient hypothesis (SGH). Here we tested a priori predictions about the effect of grazing on species interactions and fine scale spatial structure of grasses in water‐limited ecosystems. We used detailed vegetation mapping and spatial analysis, and performed a field experiment where the direct and indirect components of positive interactions were disentangled to provide evidence of links between process and pattern. We found associational resistance (biotic refuge) to be the dominant process in grazing situations, while competition, instead of direct facilitation, seemed to govern grass spatial patterns when herbivore pressure was relaxed. These results suggest that facilitation between grasses in arid communities may be related to herbivory rather than nurse plant effects. Associational resistance tends to have the strongest effect on spatial aggregation of species at intermediate grazing pressure. Results suggest that contrary to SGH, this physical clustering of species decreased when grazing pressure reached their maximum levels. Positive associations remained significant only when palatability differences between neighbours is large, suggesting that managing stocking rate is a key factor determining the persistence of herbivory refuges. These refuges are potential foci to initiate population recovery of high quality forage species in arid degraded areas.  相似文献   

7.
Cooperation with neighbours may be crucial for the persistence of populations in stressful environments. Yet, cooperation is often not evolutionarily stable, since non-cooperative individuals can reap the benefits of cooperation without having to pay the costs associated with cooperation. Here we show that active aggregation leading to self-organized spatial pattern formation can promote the evolution of cooperativeness. To this end, we study the effect of movement strategies on the evolution of cooperation in mussel beds. Mussels cooperate by attaching themselves to neighbours via byssal threads, thereby providing mutual protection. Using an individual-based model for mussel bed formation, we first demonstrate that the spatial pattern and the corresponding number of neighbours strongly depends on the movement strategies of the mussels. With an evolutionary model, we then show that this has important implications for the evolution of cooperation, since the evolved level of cooperativeness (the number of byssus threads produced) strongly depends on the number of neighbours and on the harshness and variability of the environment. Our results suggest that spatial aggregation, abundantly found in self-organized ecosystems, may promote the evolution of cooperation.  相似文献   

8.
To provide histological foundation for studying the genetic mechanisms of color‐pattern polymorphisms, we examined light reflectance profiles and cellular architectures of pigment cells that produced striped, nonstriped, and melanistic color patterns in the snake Elaphe quadrivirgata. Both, striped and nonstriped morphs, possessed the same set of epidermal melanophores and three types of dermal pigment cells (yellow xanthophores, iridescent iridophores, and black melanophores), but spatial variations in the densities of epidermal and dermal melanophores produced individual variations in stripe vividness. The densities of epidermal and dermal melanophores were two or three times higher in the dark‐brown‐stripe region than in the yellow background in the striped morph. However, the densities of epidermal and dermal melanophores between the striped and background regions were similar in the nonstriped morph. The melanistic morph had only epidermal and dermal melanophores and neither xanthophores nor iridophores were detected. Ghost stripes in the shed skin of some melanistic morphs suggested that stripe pattern formation and melanism were controlled independently. We proposed complete‐ and incomplete‐dominance heredity models for the stripe‐melanistic variation and striped, pale‐striped, and nonstriped polymorphisms, respectively, according to the differences in pigment‐cell composition and its spatial architecture. J. Morphol. 274:1353–1364, 2013. © 2013 Wiley Periodicals, Inc.  相似文献   

9.
Niall P. Hanan 《Biotropica》2012,44(2):189-196
This paper examines the feasibility of applying self‐thinning concepts to savannas and how competition with herbaceous vegetation may modify self‐thinning patterns among woody plants in these ecosystems. Competition among woody plants has seldom been invoked as a major explanation for the persistence of herbaceous vegetation in mixed tree–grass ecosystems. On the contrary, the primary resource‐based explanations for tree–grass coexistence are based on tree–grass competition (niche‐separation) that assumes that trees are inferior competitors unless deeper rooting depths provide them exclusive access to water. Alternative nonresource‐based hypotheses postulate that trees are the better competitors, but that tree populations are suppressed by mortality related to fire, herbivores, and other disturbances. If self‐thinning of woody plants can be detected in savannas, stronger evidence for resource‐limitation and competitive interactions among woody plants would suggest that the primary models of savannas need to be adjusted. We present data from savanna sites in South Africa to suggest that self‐thinning among woody plants can be detected in low‐disturbance situations, while also showing signs that juvenile trees, more so than adults, are suppressed when growing with herbaceous vegetation in these ecosystems. This finding we suggest is evidence for size‐asymmetric competition in savannas.  相似文献   

10.
Spatial patterns are a subfield of spatial ecology, and these patterns modify the temporal dynamics and stability properties of population densities at a range of spatial scales. Localized ecological interactions can generate striking large-scale spatial patterns in ecosystems through spatial self-organization. Possible mechanisms include oscillating consumer–resource interactions, localized disturbance–recovery processes, and scale-dependent feedback. However, in this paper, our main aim is to study the effect of tide on the pattern formation of a spatial plant-wrack model. We discuss the changes of the wavelength, wave speed, and the conditions of the spatial pattern formation, according to the dispersion relation formula. Both the mathematical analysis and numerical simulations reveal that the tide has great influence on the spatial pattern. More specifically, typical traveling spatial patterns can be obtained. Our obtained results are consistent with the previous observation that wracks exhibit traveling patterns, which is useful to help us better understand the dynamics of the real ecosystems.  相似文献   

11.
Spatial pattern and process in forest stands within the Virginia piedmont   总被引:1,自引:0,他引:1  
Abstract. Question: Underlying ecological processes have often been inferred from the analysis of spatial patterns in ecosystems. Using an individual‐based model, we evaluate whether basic assumptions of species’life‐history, drought‐susceptibility, and shade tolerance generate dynamics that replicate patterns between and within forest stands. Location: Virginia piedmont, USA. Method: Model verification examines the transition in forest composition and stand structure between mesic, intermediate and xeric sites. At each site, tree location, diameter, and status were recorded in square plots ranging from 0.25 to 1.0 ha. Model validation examines the simulated spatial pattern of individual trees at scales of 1–25 m within each forest site using a univariate Ripley's K function. Results: 7512 live and dead trees were surveyed across all sites. All sites exhibit a consistent, significant shift in pattern for live trees by size, progressing from a clumped understorey (trees ± 0.1 m in diameter) to a uniform overstorey (trees > 0.25 m). Simulation results reflect not only the general shift in pattern of trees at appropriate scales within sites, but also the general transition in species composition and stand structure between sites. Conclusions: This shift has been observed in other forest ecosystems and interpreted as a result of competition; however, this hypothesis has seldom been evaluated using simulation models. These results support the hypothesis that forest pattern in the Virginia piedmont results from competition involving species’life‐history attributes driven by soil moisture availability between sites and light availability within sites.  相似文献   

12.
Mediterranean ecosystems are hotspots of species richness where fire is one of the key processes influencing their structure, composition and function. Post‐fire seedling emergence constitutes a crucial event in the life cycle of plants and species‐specific temporal and spatial patterns of seedling emergence have been hypothesized to contribute to the high diversity in these ecosystems. Here we study the temporal and spatial patterns of seedling emergence observed for the four dominant species (Cistus albidus, Ulex parviflorus, Helianthemum marifolium, Ononis fruticosa) after an experimental fire in a Mediterranean gorse shrubland. In a first analysis we compared the timing of emergence of each species using the Kaplan‐Meier method. The spatial component of seedling emergence and the spatiotemporal relationship between different cohorts of the same species were analyzed using recent techniques of spatial point pattern analyses. We found a bimodal temporal pattern of emergence. Emergence of Cistaceae species (H. marifolium and C. albidus) occurred predominantly early after the fire while Fabaceae (O. fruticosa and U. parviflorus) emerged mainly during the following autumn. Individually, all species showed an aggregated spatial pattern and, when testing for pair interactions, we found that the clusters of individual species were spatially segregated. Additionally, the clusters of individual species showed an internal spatial structure where seedlings of different cohorts were spatially segregated. Theoretical models predict that these patterns will promote species coexistence. We identified a number of mechanisms that all have the potential to contribute to the observed pattern formation. However, the potential interaction among these mechanisms are complex and not easy to predict. Our analyses take a significant step forward in studying seedling emergence in fire prone ecosystems since, to our knowledge, this is the first time that both spatial and temporal patterns of all dominant species have been studied together.  相似文献   

13.
The persistence of mutualisms in host‐microbial – or holobiont – systems is difficult to explain because microbial mutualists, who bear the costs of providing benefits to their host, are always prone to being competitively displaced by non‐mutualist ‘cheater’ species. This disruptive effect of competition is expected to be particularly strong when the benefits provided by the mutualists entail costs such as reduced competitive ability. Using a metacommunity model, we show that competition between multiple cheaters within the host's microbiome, when combined with the spatial structure of host–microbial interactions, can have a constructive rather than a disruptive effect by allowing the emergence and maintenance of mutualistic microorganisms within the host. These results indicate that many of the microorganisms inhabiting a host's microbiome, including those that would otherwise be considered opportunistic or even potential pathogens, play a cryptic yet critical role in promoting the health and persistence of the holobiont across spatial scales.  相似文献   

14.
The segmentation of many animals ranging from insects to mammals involves the sequential formation of stationary stripes of gene expression that are perpendicular to the growth axis of the developing embryo. This process has been accounted for by a variety of theoretical “clock-and-wavefront” type models that involve the arrest of an oscillation (the clock) at a moving boundary (the wavefront). Here, we demonstrate experimentally that progressive arrest of a homogeneous oscillation can control the symmetry as well as the wavelength of spatial structures in a chemical system. We show how a spontaneously formed, labyrinthine pattern can be converted into a pattern composed of ordered, parallel stripes and confirm a previously predicted proportionality between the wavelength and the period of the homogeneous oscillation. Our experiments provide the first experimental demonstration of a general mechanism for the control of pattern formation that has been hypothesized to operate in the context of biological morphogenesis.  相似文献   

15.
Understanding the dynamics and underlying mechanism of carbon exchange between terrestrial ecosystems and the atmosphere is one of the key issues in global change research. In this study, we quantified the carbon fluxes in different terrestrial ecosystems in China, and analyzed their spatial variation and environmental drivers based on the long‐term observation data of ChinaFLUX sites and the published data from other flux sites in China. The results indicate that gross ecosystem productivity (GEP), ecosystem respiration (ER), and net ecosystem productivity (NEP) of terrestrial ecosystems in China showed a significantly latitudinal pattern, declining linearly with the increase of latitude. However, GEP, ER, and NEP did not present a clear longitudinal pattern. The carbon sink functional areas of terrestrial ecosystems in China were mainly located in the subtropical and temperate forests, coastal wetlands in eastern China, the temperate meadow steppe in the northeast China, and the alpine meadow in eastern edge of Qinghai‐Tibetan Plateau. The forest ecosystems had stronger carbon sink than grassland ecosystems. The spatial patterns of GEP and ER in China were mainly determined by mean annual precipitation (MAP) and mean annual temperature (MAT), whereas the spatial variation in NEP was largely explained by MAT. The combined effects of MAT and MAP explained 79%, 62%, and 66% of the spatial variations in GEP, ER, and NEP, respectively. The GEP, ER, and NEP in different ecosystems in China exhibited ‘positive coupling correlation’ in their spatial patterns. Both ER and NEP were significantly correlated with GEP, with 68% of the per‐unit GEP contributed to ER and 29% to NEP. MAT and MAP affected the spatial patterns of ER and NEP mainly by their direct effects on the spatial pattern of GEP.  相似文献   

16.
Eva Banda  Guillermo Blanco 《Oikos》2009,118(7):991-1000
Nest‐site limitation may have different implications in the spatial distribution of breeding pairs depending on the availability of suitable habitat and the types of nest‐sites. Distribution of cavities suitable as nest sites may allow circumstantial aggregation or active choice of colonial nesting, which may have different implications on breeding performance through effects on breeding density, with variable costs and benefits depending on the consequences of intraspecific competition, social interactions and predation. We evaluated the effects of breeding density derived from nesting site limitation on breeding performance and predation at different spatial scales and considering multiple social, population and environmental limiting factors in the red‐billed chough Pyrrhocorax pyrrhocorax. The results indicate that variable breeding density may arise within the population depending on the availability and spatial distribution of nest‐sites. Nest‐site availability and distribution may also determine social breeding systems (isolated or aggregated) at variable densities, thus resembling differences found at different spatially distant populations under contrasting environmental conditions. Breeding performance was related to density‐dependent processes of population regulation, especially density‐dependent nest predation due to predator attraction to nest clusters. Results also indicate that predation pressure depend on density patterns at large scales. This suggest that predation may have important consequences on population dynamics of spatially structured populations depending on the strength of this kind of density dependence, which in turn may depend on habitat features affecting the prey but also the spatially variable guild of predators. Because habitat and nesting site availability may vary spatially depending on multiple human influences, understanding the strength and form in which breeding density and nest predation at different spatial scales may influence the size and persistence of populations can help to manage them more adequately.  相似文献   

17.
Marine reserves are widely used to protect species important for conservation and fisheries and to help maintain ecological processes that sustain their populations, including recruitment and dispersal. Achieving these goals requires well‐connected networks of marine reserves that maximize larval connectivity, thus allowing exchanges between populations and recolonization after local disturbances. However, global warming can disrupt connectivity by shortening potential dispersal pathways through changes in larval physiology. These changes can compromise the performance of marine reserve networks, thus requiring adjusting their design to account for ocean warming. To date, empirical approaches to marine prioritization have not considered larval connectivity as affected by global warming. Here, we develop a framework for designing marine reserve networks that integrates graph theory and changes in larval connectivity due to potential reductions in planktonic larval duration (PLD) associated with ocean warming, given current socioeconomic constraints. Using the Gulf of California as case study, we assess the benefits and costs of adjusting networks to account for connectivity, with and without ocean warming. We compare reserve networks designed to achieve representation of species and ecosystems with networks designed to also maximize connectivity under current and future ocean‐warming scenarios. Our results indicate that current larval connectivity could be reduced significantly under ocean warming because of shortened PLDs. Given the potential changes in connectivity, we show that our graph‐theoretical approach based on centrality (eigenvector and distance‐weighted fragmentation) of habitat patches can help design better‐connected marine reserve networks for the future with equivalent costs. We found that maintaining dispersal connectivity incidentally through representation‐only reserve design is unlikely, particularly in regions with strong asymmetric patterns of dispersal connectivity. Our results support previous studies suggesting that, given potential reductions in PLD due to ocean warming, future marine reserve networks would require more and/or larger reserves in closer proximity to maintain larval connectivity.  相似文献   

18.
Cell‐intrinsic metabolic reprogramming is a hallmark of cancer that provides anabolic support to cell proliferation. How reprogramming influences tumor heterogeneity or drug sensitivities is not well understood. Here, we report a self‐organizing spatial pattern of glycolysis in xenograft colon tumors where pyruvate dehydrogenase kinase (PDK1), a negative regulator of oxidative phosphorylation, is highly active in clusters of cells arranged in a spotted array. To understand this pattern, we developed a reaction–diffusion model that incorporates Wnt signaling, a pathway known to upregulate PDK1 and Warburg metabolism. Partial interference with Wnt alters the size and intensity of the spotted pattern in tumors and in the model. The model predicts that Wnt inhibition should trigger an increase in proteins that enhance the range of Wnt ligand diffusion. Not only was this prediction validated in xenograft tumors but similar patterns also emerge in radiochemotherapy‐treated colorectal cancer. The model also predicts that inhibitors that target glycolysis or Wnt signaling in combination should synergize and be more effective than each treatment individually. We validated this prediction in 3D colon tumor spheroids.  相似文献   

19.
The distribution of genetic variation in species is governed by factors that act differently across spatial scales. To tease apart the contribution of different processes, especially at intermediate spatial scales, it is useful to study simple ecosystems such as those on sub‐Antarctic oceanic islands. In this study, we characterize spatial genetic patterns of two keystone plant species, Azorella selago on sub‐Antarctic Marion Island and Azorella macquariensis on sub‐Antarctic Macquarie Island. Although both islands experience a similar climate and have a similar vegetation structure, they differ significantly in topography and geological history. We genotyped six microsatellites for 1,149 individuals from 123 sites across Marion Island and 372 individuals from 42 sites across Macquarie Island. We tested for spatial patterns in genetic diversity, including correlation with elevation and vegetation type, and clines in different directional bearings. We also examined genetic differentiation within islands, isolation‐by‐distance with and without accounting for direction, and signals of demographic change. Marion Island was found to have a distinct northwest–southeast divide, with lower genetic diversity and more sites with a signal of population expansion in the northwest. We attribute this to asymmetric seed dispersal by the dominant northwesterly winds, and to population persistence in a southwestern refugium during the Last Glacial Maximum. No apparent spatial pattern, but greater genetic diversity and differentiation between sites, was found on Macquarie Island, which may be due to the narrow length of the island in the direction of the dominant winds and longer population persistence permitted by the lack of extensive glaciation on the island. Together, our results clearly illustrate the implications of island shape and geography, and the importance of direction‐dependent drivers, in shaping spatial genetic structure.  相似文献   

20.
Robust estimates of dispersal are critical for understanding population dynamics and local adaptation, as well as for successful spatial management. Genetic isolation by distance patterns hold clues to dispersal, but understanding these patterns quantitatively has been complicated by uncertainty in effective density. In this study, we genotyped populations of a coral reef fish (Amphiprion clarkii) at 13 microsatellite loci to uncover fine‐scale isolation by distance patterns in two replicate transects. Temporal changes in allele frequencies between generations suggested that effective densities in these populations are 4–21 adults/km. A separate estimate from census densities suggested that effective densities may be as high as 82–178 adults/km. Applying these effective densities with isolation by distance theory suggested that larval dispersal kernels in A. clarkii had a spread near 11 km (4–27 km). These kernels predicted low fractions of self‐recruitment in continuous habitats, but the same kernels were consistent with previously reported, high self‐recruitment fractions (40–60%) when realistic levels of habitat patchiness were considered. Our results suggested that ecologically relevant larval dispersal can be estimated with widely available genetic methods when effective density is measured carefully through cohort sampling and ecological censuses, and that self‐recruitment studies should be interpreted in light of habitat patchiness.  相似文献   

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