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Patterns of dispersal and dynamics among habitat patches varying in quality.

Both source-sink theory and extensions of optimal foraging theory ("balanced dispersal" theory) address dispersal and population dynamics in landscapes where habitat patches vary in quality. However, studying dispersal mechanisms empirically has proven difficult, and dispersal is rarely tied back to long-term spatial dynamics. We used a manipulable laboratory system consisting of bacteria and protozoa to investigate the ability of source-sink and optimal foraging theories to explain both dispersal and emergent spatial dynamics. Consistent with source-sink models and contrary to balanced dispersal models, there was a consistent net flux of protist individuals from high to low resource patches. However, unlike the simplest source-sink models, intermediate rates of dispersal led to highest abundances in low resource patches. Side experiments found strong density dependence in local population dynamics and differences in average protist body size in high and low resource patches. Parameterization and analysis of a two-patch model showed that high migration from high to low resource patches could have depressed population density in low resource patches, creating pseudosinks. The movement of individuals and biomass from sources to sinks (a form of ecosystem subsidy) resulted in the convergence of body size and population densities in sources and sinks. Our results indicate a need to carefully consider movement patterns and interaction with local dynamics in potential source-sink systems.

Animals↗

Aedes polynesiensis in the Society Islands: environmental correlates of isoenzyme differentiation.

Isoenzyme genetic differentiation of Aedes polynesiensis mosquitoes in Raiatea island, French Polynesia, was evaluated by two models of population structure based on seven gene-enzyme systems: Ak, Est, Got, Gpi, Hk, Mdh and Pgm. The ecological model tested whether genetic differentiation is congruent with habitat differences. The isolation model evaluated whether genetic differentiation is proportional to geographical distribution. The ecological model found no significant differentiation between populations of Ae.polynesiensis from beach and forest ecotopes, whereas the isolation model was consistent with the data. However, incipient speciation is opposed by the source-sink system of population dynamics in such small neighbouring islands, where Ae.polynensiensis extinction is readily followed by reinvasion causing considerable gene flow between island populations.

Aedes↗

Population Genetic Models of Source-Sink Metapopulations

The present study investigates the effect that different patterns of migration have on the genetic structure of source-sink metapopulations specifically modelling the dynamics of local populations. The model assumes a metapopulation consisting of a single source and s different sink populations and considers the expected number of nucleotide differences between two genes drawn at random from the source-sink metapopulation. The results show that a collection of interconnected sinks can maintain a substantial fraction of the genetic variability observed in the source population, particularly where migration from the source is continuous over time. The degree of genetic differentiation among the sinks might be large, especially if migration from the source is stochastic. Genetic differentiation is small when extinctions are frequent (because most of the sink populations are composed of recent migrants from the source) or extremely rare (because of the increased effect of migration among sinks), being maximized when the frequency of extinctions is between these two extremes.

Journal Article↗

Genetic variability in sensitivity to population density affects the dynamics of simple ecological models.

Many 1-dimensional discrete time ecological models contain a sensitivity parameter that does not affect the dynamic complexity of these models. We show that genetic variability in this parameter can have a strong effect on population dynamics. We incorporate ecological dynamics in two different population genetic models with one locus and two alleles. The first is the classical model of a randomly mating population in Hardy-Weinberg equilibrium, and the second is a model of differential selection in males and females. In populations in Hardy-Weinberg equilibrium, variability in the sensitivity parameter can be maintained by overdominance. In this case, the dynamics of the polymorphic population tend to be much simpler than those of monomorphic populations. In the model with different selection in males and females, polymorphisms can be maintained in various ways, e.g., by opposing directional selection in males and females. Polymorphism in the sensitivity parameter tends to simplify population dynamics in the model with different selection in males and females as well. A number of interesting dynamic effects can be observed, e.g., multiple attractors with complicated basins of attraction. Then the final state of the system after a successful invasion by mutant alleles may depend on the mutation rate and on the distribution of mutational steps. In addition, there are situations in which genetic variability destabilizes a stable population dynamic equilibrium in the monomorphic model. There is an analogy between genetic variability and variability imposed by the environment. If differences in sensitivity are caused by the environment, dynamic effects similar to those in the genetic models can be observed. In addition, source-sink structures that are known to occur in spatially structured models can be seen in the genetic model if one of the genotypes is inviable. The results suggest that combining ecological and population genetic models can lead to a number of new insights. More work is needed, e.g., with fertility models, in which fitnesses are not assigned to individuals, but to mating pairs.

Alleles↗

The dynamics of grazed woodlands in southwest Queensland, Australia and their effect on greenhouse gas emissions.

This study outlines the development of an approach to evaluate the sources, sinks, and magnitudes of greenhouse gas emissions from a grazed semiarid rangeland dominated by mulga (Acacia aneura) and how these emissions may be altered by changes in management. This paper describes the modification of an existing pasture production model (GRASP) to include a gas emission component and a dynamic tree growth and population model. An exploratory study was completed to investigate the likely impact of changes in burning practices and stock management on emissions. This study indicates that there is a fundamental conflict between maintaining agricultural productivity and reducing greenhouse gas emissions on a given unit of land. Greater agricultural productivity is allied with the system being an emissions source while production declines and the system becomes a net emissions sink as mulga density increases. Effective management for sheep production results in the system acting as a net source (approximately 60-200 kg CO2 equivalents/ha/year). The magnitude of the source depends on the management strategies used to maintain the productivity of the system and is largely determined by starting density and average density of the mulga over the simulation period. Prior to European settlement, it is believed that the mulga lands were burnt almost annually. Simulations indicate that such a management approach results in the system acting as a small net sink with an average net absorption of greenhouse gases of 14 kg CO2 equivalents/ha/year through minimal growth of mulga stands. In contrast, the suppression of fire and the introduction of grazing results in thickening of mulga stands and the system can act as a significant net sink absorbing an average of 1000 kg CO2 equivalents/ha/year. Although dense mulga will render the land largely useless for grazing, land in this region is relatively inexpensive and could possibly be developed as a cost-effective carbon offset for greenhouse gas emissions elsewhere. These results also provide support for the hypothesis that changes in land management, and particularly, suppression of fire is chiefly responsible for the observed increases in mulga density over the past century.

Acacia↗

Using L-systems for modeling source-sink interactions, architecture and physiology of growing trees: the L-PEACH model.

Functional-structural plant models simulate the development of plant structure, taking into account plant physiology and environmental factors. The L-PEACH model is based on the development of peach trees. It demonstrates the usefulness of L-systems in constructing functional-structural models. L-PEACH uses L-systems both to simulate the development of tree structure and to solve differential equations for carbohydrate flow and allocation. New L-system-based algorithms are devised for simulating the behavior of dynamically changing structures made of hundreds of interacting, time-varying, nonlinear components. L-PEACH incorporates a carbon-allocation model driven by source-sink interactions between tree components. Storage and mobilization of carbohydrates during the annual life cycle of a tree are taken into account. Carbohydrate production in the leaves is simulated based on the availability of water and light. Apices, internodes, leaves and fruit grow according to the resulting local carbohydrate supply. L-PEACH outputs an animated three-dimensional visual representation of the growing tree and user-specified statistics that characterize selected stages of plant development. The model is applied to simulate a tree's response to fruit thinning and changes in water stress. L-PEACH may be used to assist in horticultural decision-making processes after being calibrated to specific trees.

Biological Transport, Active↗

Effective size in management and conservation of subdivided populations.

A numerical method for computing the eigenvalue variance effective size of a subdivided population connected by any fixed pattern of migration is described. Using specific examples it is shown that total effective size of a subdivided population can become less than the sum of the subpopulation sizes as a result of directionalities in the pattern of migration. For an extension of the model with threshold harvesting and local deterministic logistic population dynamic we consider the problem of maximizing the total harvesting yield with constraints on the total effective size. For some simple source-sink systems and more complicated population structures where subpopulations differ in their degree of isolation, it is shown to be optimal, for a given total effective size, to raise the harvesting thresholds relatively more in small and in isolated populations. Finally, we show how the method applies to populations which are supplemented, either intentionally or unintentionally. It is shown that the total effective size can be reduced by several orders of magnitude if the captive component of a population is much smaller than the wild component, even with symmetric backward migration.

Animals↗

Fine root branch orders respond differentially to carbon source-sink manipulations in a longleaf pine forest.

Fine roots are a key component of carbon (C) flow and nitrogen (N) cycling in forest ecosystems. However, the complexity and heterogeneity of the fine root branching system have hampered the assessment and prediction of C and N dynamics at ecosystem scales. We examined how root morphology, biomass, and chemistry differed with root branch orders (1-5 with root tips classified as first order roots) and how different root orders responded to increased C sink strength (via N fertilization) and reduced carbon source strength (via canopy scorching) in a longleaf pine (Pinus palustris L.) ecosystem. With increasing root order, the diameter and length of individual roots increased, whereas the specific root length decreased. Total root biomass on an areal basis was similar among the first four orders but increased for the fifth order roots. Consequently, total root length and total root surface area decreased systematically with increasing root order. Fine root N and lignin concentrations decreased, while total non-structural carbohydrate (TNC) and cellulose concentrations increased with increasing root order. N addition and canopy disturbance did not alter root morphology, but they did influence root chemistry. N fertilization increased fine root N concentration and content per unit area in all five orders, while canopy scorching decreased root N concentration. Moreover, TNC concentration and content in fifth order roots were also reduced by canopy scorching. Our results indicate that the small, fragile, and more easily overlooked first and second order roots may be disproportionately important in ecosystem scale C and N fluxes due to their large proportions of fine root biomass, high N concentrations, relatively short lifespans, and potentially high decomposition rates.

Biomass↗

Habitat-specific demography: evidence for source-sink population structure in a mammal, the pika.

Theory suggests that populations may persist in sink habitats that cannot support replacement-level birth rates. Although it is commonly believed that organisms that can actively select habitat should rarely occur in sinks, the frequency of use of sinks in free-ranging species is not well-documented. We found that a population of American pikas ( Ochotona princeps, Lagomorpha) inhabiting distinct alpine habitats (meadow and snowbed) in Wyoming, USA, had habitat-specific demographic rates that produced a source-sink population structure. Population size increased in both habitats in summer and declined in both habitats in winter, with populations in snowbeds increasing more during summer and decreasing more over winter. Birth rates were consistently higher in meadows and populations in meadows had a consistently higher finite rate of increase (lambda, from life tables) than did those in snowbeds, for which lambda was far below that needed for replacement. Patterns of immigration, population structure, and temporal variation in population size were as expected if meadows were functional sources and snowbeds functional sinks. Patterns of snowmelt differed between habitats, predicted the critical difference in birth rates between habitats, and are a likely primary cause of the differences in habitat-specific birth rates that we observed. This study provides a clear example of source-sink population structure for a mammal.

Animals↗

Water quality functioning of lowland permeable catchments: inferences from an intensive study of the RIVER KENNEt and upper River Thames.

This paper brings together information on the water quality functioning of the River Kennet and other parts of the upper River Thames in the south east of England. The Kennet represents a groundwater fed riverine environment impacted by agricultural and sewage sources of nutrient pollution. Descriptions of the general water quality of the area, nutrient sources, sinks and within river processes are provided together with biological responses to driving issues of agriculture, sewage treatment and climatic change. Models are developed and applied to assess the key processes involved for a highly dynamic system and to provide initial estimates of the likely responses to environmental change. Furthermore, the economic aspects of pollution control are reviewed, together with legislation issues, which are presented within the context of a landmark case known as the 'Axford Inquiry', the implications of which extend to regional and national dimensions. The paper concludes with a discussion on the present state of knowledge, key issues and future research on the science and management of groundwater fed nutrient impacted riverine systems.

Agriculture↗

Spatial and temporal modeling of microbial contaminants on grazing farmlands.

This paper introduces an integrated spatial and temporal modeling system developed mathematically for assessing microbial contaminants on animal-grazed farmlands. The model uses fecal coliform, specifically Escherichia coli, as an indicator of fecal contamination and describes the sources, sinks, transport processes, and fate of E. coli contaminants in catchments and associated streams. Spatial features include grazing location, land topography, distance to a nearby stream, and distance through the stream network to the outlet. Temporal features are population dynamics on the land surface, in flow, and on streambeds. The model applies the principles of conservation of mass balance on two different types of pools: grid cells on land surfaces and networked stream segments. The model aims to improve the prediction of the effects of different land management strategies on the fecal contamination of waterways. This is achieved by characterizing the movement of fecal contaminants from land to streams and in-stream mobilization. Processes of attenuation, diffusion, and transport govern the movement. Our study site is a hill land catchment with an area of 140 ha and is used exclusively for animal grazing. The model was calibrated with previous research results, and then tested using the data collected at the outlet of the catchment. The sensitivity of the model predictions was analyzed for different scenarios: effect of stock rate, attenuation rate, and flow volumes. The similar pattern between monitored and predicted E. coli concentration proved that the model captures the key features that control the population dynamics of fecal contaminants. Further experiments are required to expand the model's functionality for covering more mitigation options.

Animal Husbandry↗

Protein dynamics, activity and cellular localization of soybean lipoxygenases indicate distinct functional roles for individual isoforms.

Vegetative lipoxygenases (VLXs) in soybean are hypothesized to function in nitrogen storage and partitioning. Isoform-specific antibodies for four of the five known VLX isoenzymes were used to investigate the influence of source-sink status on protein levels, as well as to analyze the tissue and subcellular localization of the different isoforms. VLXD responded most strongly to sink limitation, although the levels of VLXA, B and C increased as well. After sink limitation, VLXD and the vegetative storage protein, VSPalpha, accumulated in the vacuoles of bundle sheath and paraveinal mesophyll cells, while VLXA, B and C localized to the cytosol of these cells. All five known VLX isoenzymes were active with both linoleic and linolenic acid substrates after expression in Escherichia coli. The strong upregulation of VLXD levels after sink limitation as well as the localization of this isoform to the vacuoles of paraveinal mesophyll and bundle sheath cells (where VSPs are found) strongly suggest that VLXD should be considered as a major storage protein in soybean leaves. Furthermore, since VLXA, B and C also accumulate in sink-limited soybean leaves, are located in the cytosol of paraveinal mesophyll cells and are active at pH values typically found in this compartment, their activities may well contribute to lipid metabolism in this tissue. This multi-gene family is thus ideally poised to play a pivotal role in the balance of N deposition relative to lipid-based storage, defense or signaling, by modulating contributions to these processes in the transient storage cells of the paraveinal mesophyll.

Amino Acid Sequence↗

Spatial variation and density-dependent dispersal in competitive coexistence.

It is well known that dispersal from localities favourable to a species' growth and reproduction (sources) can prevent competitive exclusion in unfavourable localities (sinks). What is perhaps less well known is that too much emigration can undermine the viability of sources and cause regional competitive exclusion. Here, I investigate two biological mechanisms that reduce the cost of dispersal to source communities. The first involves increasing the spatial variation in the strength of competition such that sources can withstand high rates of emigration; the second involves reducing emigration from sources via density-dependent dispersal. I compare how different forms of spatial variation and modes of dispersal influence source viability, and hence source-sink coexistence, under dominance and pre-emptive competition. A key finding is that, while spatial variation substantially reduces dispersal costs under both types of competition, density-dependent dispersal does so only under dominance competition. For instance, when spatial variation in the strength of competition is high, coexistence is possible (regardless of the type of competition) even when sources experience high emigration rates; when spatial variation is low, coexistence is restricted even under low emigration rates. Under dominance competition, density-dependent dispersal has a strong effect on coexistence. For instance, when the emigration rate increases with density at an accelerating rate (Type III density-dependent dispersal), coexistence is possible even when spatial variation is quite low; when the emigration rate increases with density at a decelerating rate (Type II density-dependent dispersal), coexistence is restricted even when spatial variation is quite high. Under pre-emptive competition, density-dependent dispersal has only a marginal effect on coexistence. Thus, the diversity-reducing effects of high dispersal rates persist under pre-emptive competition even when dispersal is density dependent, but can be significantly mitigated under dominance competition if density-dependent dispersal is Type III rather than Type II. These results lead to testable predictions about source-sink coexistence under different regimes of competition, spatial variation and dispersal. They identify situations in which density-independent dispersal provides a reasonable approximation to species' dispersal patterns, and those under which consideration of density-dependent dispersal is crucial to predicting long-term coexistence.

Animals↗

The plant stress hypothesis and variable responses by blue grama grass (Bouteloua gracilis) to water, mineral nitrogen, and insect herbivory.

Acting simultaneously or sequentially, plants encounter multiple stresses from combined abiotic and biotic factors that result in decreased growth and internal reallocation of resources. The plant stress hypothesis predicts that environmental stresses on plants decrease plant resistance to insect herbivory by altering biochemical source-sink relationships and foliar chemistry, leading to more palatable food. Such changes in the nutritional landscape for insects may facilitate insect population outbreaks during periods of moderate stress on host plants. We examined the plant stress hypothesis with field experiments in continental grassland (USA) using the C4 grass Bouteloua gracilis. Water, nitrogen fertilizer, and herbivory from the grass-feeding grasshopper Ageneotettix deorum were manipulated. Combined stresses from water and mineral-N in the soil decreased plant growth and altered foliar percent total N (TN) and percent total nonstructural carbohydrate (TNC) concentrations in an additive fashion. Grasshopper herbivory affected final biomass only in dry years; plants compensated for tissue loss when rainfall was abundant. Foliar TN and TNC concentrations were dynamic with respect to variable climatic conditions and treatment combinations, showing significant interactions. Grasshopper herbivory had its greatest impact on TN or TNC in dry years, interacting with other forms of stress. Herbivory as a single factor had strong effects on TNC in years with normal precipitation, but not in a dry year. Performance (developmental rate and survival) by the grasshoppers Phoetaliotes nebrascensis and A. deorum were not greatly affected by plant stress in a manner consistent with the plant stress hypothesis.

Animals↗

Effects of metapopulation processes on measures of genetic diversity.

Many species persist as a metapopulation under a balance between the local extinction of subpopulations or demes and their recolonization through dispersal from occupied patches. Here we review the growing body of literature dealing with the genetic consequences of such population turnover. We focus our attention principally on theoretical studies of a classical metapopulation with a 'finite-island' model of population structure, rather than on 'continent-island' models or 'source-sink' models. In particular, we concern ourselves with the subset of geographically subdivided population models in which it is assumed that all demes are liable to extinction from time to time and that all demes receive immigrants. Early studies of the genetic effects of population turnover focused on population differentiation, such as measured by F(ST). A key advantage of F(ST) over absolute measures of diversity is its relative independence of the mutation process, so that different genes in the same species may be compared. Another advantage is that F(ST) will usually equilibrate more quickly following perturbations than will absolute levels of diversity. However, because F(ST) is a ratio of between-population differentiation to total diversity, the genetic effects of metapopulation processes may be difficult to interpret in terms of F(ST) on its own, so that the analysis of absolute measures of diversity in addition is likely to be informative. While population turnover may either increase or decrease F(ST), depending on the mode of colonization, recurrent extinction and recolonization is expected always to reduce levels of both within-population and species-wide diversity (piS and piT, respectively). One corollary of this is that piS cannot be used as an unbiased estimate of the scaled mutation rate, theta, as it can, with some assumptions about the migration process, in species whose demes do not fluctuate in size. The reduction of piT in response to population turnover reflects shortened mean coalescent times, although the distribution of coalescence times under extinction colonization equilibrium is not yet known. Finally, we review current understanding of the effect of metapopulation dynamics on the effective population size.

Genetic Techniques↗

Community patterns in source-sink metacommunities.

We present a model of a source-sink competitive metacommunity, defined as a regional set of communities in which local diversity is maintained by dispersal. Although the conditions of local and regional coexistence have been well defined in such systems, no study has attempted to provide clear predictions of classical community-wide patterns. Here we provide predictions for species richness, species relative abundances, and community-level functional properties (productivity and space occupation) at the local and regional scales as functions of the proportion of dispersal between communities. Local (alpha) diversity is maximal at an intermediate level of dispersal, whereas between-community (beta) and regional (gamma) diversity decline as dispersal increases because of increased homogenization of the metacommunity. The relationships between local and regional species richness and the species rank abundance distributions are strongly affected by the level of dispersal. Local productivity and space occupation tend to decline as dispersal increases, resulting in either a hump-shaped or a positive relationship between species richness and productivity, depending on the scale considered (local or regional). These effects of dispersal are buffered by decreasing species dispersal success. Our results provide a niche-based alternative to the recent neutral-metacommunity model and have important implications for conservation biology and landscape management.

Animals↗

The effects of habitat fragmentation on persistence of source-sink metapopulations in systems with predators and prey or apparent competitors.

We consider systems with one predator and one prey, or a common predator and two prey species (apparent competitors) in source and sink habitats. In both models, the predator species is vulnerable to extinction, if productivity in the source is insufficient to rescue demographically deficient sink populations. Conversely, in the model with two prey species, if the source is too rich, one of the prey species may be driven extinct by apparent competition, since the predator can maintain a large population because of the alternative prey. Increasing the rate of predator movement from the source population has opposite effects on prey and predator persistence. High emigration rate exposes the predator population to danger of extinction, reducing the number of individuals that breed and produce offspring in the source habitat. This may promote coexistence of prey by relaxing predation pressure and apparent competition between the two prey species. The number of sinks and spatial arrangement of patches, or connectivity between patches, also influence persistence of the species. More sinks favor the prey and fewer sinks are advantageous to the predator. A linear pattern with the source at one end is profitable for the predator, and a centrifugal pattern in which the source is surrounded by sinks is advantageous to the prey. When the dispersal rate is low, effects of the spatial structure may exceed those of the number of sinks. In brief, productivity in patches and patterns of connectivity between patches differentially influence persistence of populations in different trophic levels.

Animals↗

Age at flowering differentially affects vegetative and reproductive responses of a determinate annual plant to elevated carbon dioxide.

Plant population and community dynamics may be altered by increasing atmospheric CO(2) concentrations [[CO(2)]] through intraspecific variation in the responses of vegetative and reproductive growth. Although these responses may be regulated by age at flowering, little is known about the direct effects of age at flowering on growth responses to elevated [CO(2)]. In this study, we examined the interactive effects of elevated [CO(2)] and age at flowering on absolute and relative allocation to vegetative and reproductive growth in the determinate, short-day species Xanthium strumarium L. (common cocklebur). Six cohorts were planted at 5-day intervals in chambers maintained at either 365 or 730 micro mol mol(-1) CO(2), with an 18-h photoperiod and a non-limiting nutrient supply. All plants were simultaneously induced to flower by switching the photoperiod to 12 h for 2 days, then switching back to an 18-h photoperiod for the remainder of the experiment. All plants were harvested 15 days after the onset of flowering. Total plant biomass increased 11-41% with increasing [CO(2)] and 45% from the youngest to the oldest cohort. Vegetative growth responses to elevated [CO(2)] significantly increased with increasing age at flowering, associated with increasing sink relative to source capacity. In contrast, total fruit mass decreased 32% from the youngest to the oldest cohort and was not significantly affected by CO(2) supply. Relative biomass allocation to fruit decreased 47% from the youngest to the oldest cohort, reflecting decreased numbers of fruit, and 6-28% with increasing [CO(2)], reflecting decreased mean mass per mature fruit. Our findings suggest that elevated [CO(2)] may increase vegetative growth in Xanthium without increasing reproductive biomass, and that age at flowering may influence these responses through effects on source:sink balance. Further, changes in the allometric relationship between vegetative and reproductive growth associated with growth in elevated [CO(2)] suggest that long-term population and community-level responses to elevated [CO(2)] may differ substantially from predictions based on vegetative responses.

Carbon Dioxide↗