Plant-Herbivore Interaction and Its Consequences for Succession in Wetland Ecosystems: A Modeling Approach

Herbivore grazing is increasingly used as a management tool to prevent the dominance of vegetation by tall grasses or trees. In this report, a model is described that is used to analyze plant-herbivore interactions and their scaling up to landscape scale. The model can be used to predict effects of herbivory on vegetation development. The model is an ecosystem model including modules for carbon and nitrogen cycling through plants, soil organic matter, and atmosphere. Plants compete for light and nitrogen. An herbivory module is included that implements selective foraging by a herbivore in a spatially heterogeneous area. Simulations were done to analyze the effects of herbivore density on vegetation dynamics, to analyze the impact of soil fertility on maximum herbivore density, and to analyze effects of herbivore density on landscapes. Two important points come forward from the model. Maximum herbivore abundance shows a hump-shaped curve along a soil fertility gradient. At higher soil fertility, light competition becomes more important. Herbivory interferes with plant competition, giving the tall, less palatable species a competitive advantage and thereby reducing the food quality and availability and hence the carrying capacity of the area. At a landscape scale, herbivory leads to increased heterogeneity. This increased heterogeneity may increase carrying capacity. The implications of these points for nature management are discussed. Received 13 May 1998; accepted 23 November 1998.     

The impact of sea‐level rise on Snowy Plovers in Florida: integrating geomorphological,habitat, and metapopulation models

Sea‐level rise (SLR) is a projected consequence of global climate change that will result in complex changes in coastal ecosystems. These changes will cause transitions among coastal habitat types, which will be compounded by human‐made barriers to the gradual inland migration of these habitat types. The effect of these changes on the future viability of coastal species will depend on the habitat requirements and population dynamics of these species. Thus, realistic assessments of the impact of SLR require linking geomorphological models with habitat and population models. In this study, we implemented a framework that allows this linkage, and demonstrated its feasibility to assess the effect of SLR on the viability of the Snowy Plover population in Florida. The results indicate that SLR will cause a decline in suitable habitat and carrying capacity for this species, and an increase in the risk of its extinction and decline. The model projected that the population size will decline faster than the area of habitat or carrying capacity, demonstrating the necessity of incorporating population dynamics in assessing the impacts of SLR on coastal species. The results were most sensitive to uncertainties in survival rate and fecundity, and suggested that future studies on this species should focus on the average and variability of these demographic rates and their dependence on population density. The effect of SLR on this species’ viability was qualitatively similar with most alternative models that used the extreme values of each uncertain parameter, indicating that the results are robust to uncertainties in the model.     

Identification of embryo paternity using polymorphic DNA markers to assess fertilizing capacity of spermatozoa after heterospermic insemination in boars

Differences in sperm fertilizing capacity of males often remain undetected by routine semen parameters. Heterospermic insemination with equal numbers of spermatozoa from 2 males is an accurate method for assessing differences in fertility. Use of heterospermic insemination depends on a reliable, efficient assay to identify paternity of conceptuses or offspring. In this study, polymorphic DNA markers amplified by PCR were tested to determine paternity of Day 5 to 6 embryos. The fertilizing capacity of 2 boars (A and B) with similar semen parameters was compared after homospermic (n=14 gilts) and heterospermic (n=11 gilts) insemination. Single AI's were performed under suboptimal conditions using 1 x 10(9) spermatozoa at 12 to 24 h before ovulation to prompt differences in fertilization and to stimulate sperm competition. The fertilization rate and the number of accessory spermatozoa were determined in Day 5 to 6 embryos. Using 5 different polymorphic DNA markers, paternity could be determined in 95.8% of the embryos. Boar B sired significantly (P<0.05) more offspring than Boar A after insemination with pooled semen, and this was reflected by a significantly (P<0.05) higher number of accessory spermatozoa following homospermic insemination with semen from Boar B, although fertilization rates did not differ between the 2 boars after homospermic insemination. The results suggest that the viability of spermatozoa in the female reproductive tract contributes to differences in fertility rates of males with similar in vitro sperm quality parameters. The number of accessory spermatozoa is a more sensitive measure of boar fertility than the fertilization rate. Polymorphic DNA markers are suitable for verification of parentage even at a very early stage of embryonic development.     

Density dependent selection incorporating intraspecific competition 1. A haploid model

A haploid model is introduced and analyzed in which intraspecific competition is incorporated within a density dependent framework. It is assumed that each genotype has a unique carrying capacity corresponding to the equilibrium population size when fixed for that type. Each genotypic fitness at a single multi-allelic locus is a function of a distinctive effective population size formed by adding the numbers of each genotype present, weighted by an intraspecific competition coefficient. As a result, the fitnesses depend upon the relative frequencies of the various genotypes as well as the total population size. Intergenotypic interactions can have a profound effect upon the outcome of the population. In particular, when the density effect of one individual upon another depends upon their respective genotypes, a unique stable interior equilibrium is possible in which all alleles are present. This stands in contrast to the purely density dependent haploid system in which the only possible stable state corresponds to fixation for the type with the highest carrying capacity. In the present model selective advantage is determined by a balance between carrying capacity and sensitivity to density pressures from other genotypes. Fixation for the genotype with the highest carrying capacity, for instance, will not be stable if it exerts a sufficiently weak competitive effect upon the other genotypes. In the diallelic case, maintenance of both alleles at a stable equilibrium requires that the net intragenotypic competition between individuals of like genotype be stronger than that between unlike types. As for purely density regulated systems, there may be no stable equilibria and/or regular and chaotic cycling may occur. The results may also be interpreted in terms of a discrete time model of interspecific competition with each haplotype representing a different species.     

Limiting similarity revisited

We reinvestigate the validity of the limiting similarity principle via numerical simulations of the Lotka–Volterra model. A Gaussian competition kernel is employed to describe decreasing competition with increasing difference in a one-dimensional phenotype variable. The simulations are initiated by a large number of species, evenly distributed along the phenotype axis. Exceptionally, the Gaussian carrying capacity supports coexistence of all species, initially present. In case of any other, distinctly different, carrying capacity functions, competition resulted in extinction of all, but a few species. A comprehensive study of classes of fractal-like carrying capacity functions with different fractal exponents was carried out. The average phenotype differences between surviving species were found to be roughly equal to the competition width. We conclude that, despite the existence of exceptional cases, the classical picture of limiting similarity and niche segregation is a good rule of thumb for practical purposes.     

Effect of habitat degradation on competition,carrying capacity,and species assemblage stability

Changes in species’ trophic niches due to habitat degradation can affect intra‐ and interspecific competition, with implications for biodiversity persistence. Difficulties of measuring species’ interactions in the field limit our comprehension of competition outcomes along disturbance gradients. Thus, information on how habitat degradation can destabilize food webs is scarce, hindering predictions regarding responses of multispecies systems to environmental changes. Seagrass ecosystems are undergoing degradation. We address effects of Posidonia oceanica coverage reduction on the trophic organization of a macroinvertebrate community in the Tyrrhenian Sea (Italy), hypothesizing increased trophic generalism, niche overlap among species and thus competition and decreased community stability due to degraded conditions. Census data, isotopic analysis, and Bayesian mixing models were used to quantify the trophic niches of three abundant invertebrate species, and intra‐ and interspecific isotopic and resource‐use similarity across locations differing in seagrass coverage. This allowed the computation of (1) competition strength, with respect to each other and remaining less abundant species and (2) habitat carrying capacity. To explore effects of the spatial scale on the interactions, we considered both individual locations and the entire study area (“‘meadow scale”). We observed that community stability and habitat carrying capacity decreased as P. oceanica coverage declined, whereas niche width, similarity of resource use and interspecific competition strength between species increased. Competition was stronger, and stability lower, at the meadow scale than at the location scale. Indirect effects of competition and the spatial compartmentalization of species interactions increased stability. Results emphasized the importance of trophic niche modifications for understanding effects of habitat loss on biodiversity persistence. Calculation of competition coefficients based on isotopic distances is a promising tool for describing competitive interactions in real communities, potentially extendible to any subset of ecological niche axes for which specimens’ positions and pairwise distances can be obtained.     

Effects of ecological differentiation on Lotka-Volterra systems for species with behavioral adaptation and variable growth rates

We study the properties of a n2-dimensional Lotka-Volterra system describing competing species that include behaviorally adaptive abilities. We indicate as behavioral adaptation a mechanism, based on a kind of learning, which is not viewed in the evolutionary sense but is intended to occur over shorter time scales. We consider a competitive adaptive n species Lotka-Volterra system, n > or = 3, in which one species is made ecologically differentiated with respect to the others by carrying capacity and intrinsic growth rate. The symmetry properties of the system and the existence of a certain class of invariant subspaces allow the introduction of a 7-dimensional reduced model, where n appears as a parameter, which gives full account of existence and stability of equilibria in the complete system. The reduced model is effective also in describing the time-dependent regimes for a large range of parameter values. The case in which one species has a strong ecological advantage (i.e. with a carrying capacity higher than the others), but with a varying growth rate, has been analyzed in detail, and time-dependent behaviors have been investigated in the case of adaptive competition among four species. Relevant questions, as species survival/exclusion, are addressed focusing on the role of adaptation. Interesting forms of species coexistence are found (i.e. competitive stable equilibria, periodic oscillations, strange attractors).     

Relationship between thirty post-thaw spermatozoal characteristics and the field fertility of 11 high-use Australian dairy AI sires

This study determined the relationship between two measures of field fertility of 11 high-use Australian artificial insemination (AI) dairy bulls and thirty standard laboratory assessments of spermatozoal post-thaw viability.The two measures of field fertility used, conception rates (cCR) and non-return rates (cNRR), were both corrected for all major non-bull variables. Sperm viability assessments were conducted on semen collected within the same season as that used to derive the field fertility estimates. These assessments measured sperm concentration, motility, morphology and membrane integrity at thawing, after 2h incubation and after the swim-up sperm selection procedure. Derivations of these measures and in vitro embryo fertilizing and developmental capacity were also determined. The Genstat Statistical Package [Genstat 5 Release 4.2 Reference Manual, VSN International, Oxford, 2000] was used to conduct an analysis of variance on the viability parameters across semen straws and bulls, and to calculate the strength of correlation between each semen parameter, cNRR and cCR in a correlation matrix. Step forward multiple regression identified the combination of semen parameters that were most highly correlated with cCR and with cNRR.The sperm parameters identified as being most predictive of cCR were the percentage of morphologically normal sperm immediately post-thaw (zeroNorm), the number of morphologically normal sperm after the swim-up procedure (nSuNorm), and the rate of zygote cleavage in vitro (Clv); the predictive equation formed by these parameters accounted for 70% of variance. The predictive equation produced for cNRR contained the variables zeroNorm, the proportion of membrane intact sperm after 2h incubation at 37 degrees C (twoMem) and Clv and accounted for 76.5% of the variation. ZeroNorm was found to be consistent across straws and semen batches within-bull and the sperm parameter with the strongest individual predictive capacity for both cCR (P=0.1) and cNRR (P=0.001). Post-thaw sperm parameters can be used to predict field fertility of Australian dairy sires; the calculated predictive equations are particularly useful for identifying and monitoring bulls of very high and very low potential fertility within a group.     

虫害诱导的植株对小菜蛾取食和生长发育的影响

本文从菜田生态系统的角度出发,就黄曲条跳甲Phyllotreta striolata(Fabricius)取食诱导对小菜蛾Plutella xylostella(L.)造成的影响进行了研究。黄曲条跳甲取食诱导对小菜蛾取食的影响比较明显,一般来说,黄曲条跳甲少量或短时间的取食会刺激小菜蛾幼虫在相应叶片上的取食,反之则会抑制;而且不同的寄主,不同完整度的叶片(即是否被黄曲条跳甲取食过)上的承载能力不同,芥菜、白菜、菜心和萝卜的承载力弱于甘蓝和芥蓝,黄曲条跳甲取食过的叶片弱于未被黄曲条跳甲取食过的叶片;不同数量的黄曲条跳甲取食对小菜蛾的产卵量影响显著,对其余生物学参数影响不显著,少量的黄曲条跳甲取食会刺激小菜蛾的产卵,多数则会抑制。     

A lotka-volterra model of coexistence between a sexual population and multiple asexual clones

At carrying capacity, small advantages in competitive ability can compensate a sexual population for its two-fold disadvantage in growth capacity when facing invasion by asexual mutants. In this paper, we develop a generic analytical model to consider the ecology of a sexual population comprising equal numbers of males and females, competing for shared prey resources with multiple female-only clones. We assume that the clones arise from the sexual population and are distinguished from it only by having narrower resource niches and twice the growth capacity. For sexual populations, at density-dependent carrying capacity, intra-specific competition between clonal individuals prevents them from realizing their two-fold advantage in intrinsic growth. This prediction leads to three novel outcomes: (i) a sexual population can coexist with any number of clones, provided their combined competitive impact remains less than the impact of the clones on each other; (ii) a sexual species can immediately exclude asexual invaders if it is a fast growing and strong competitor of shared resources and also has refuge in an abundant alternative resource; (iii) the rate of accumulation of clones in a sexual population will be slowed by intra and inter-specific competition amongst the clones themselves, in addition to the competitive impact from the original sexual population.     

Neutron- and X-ray-induced mutations at the yellow, white, forked and vermilion loci of Drosophila melanogaster; a preliminary analysis

Neutrons and X-rays were used to induce mutations at the yellow, white, vermilion and forked loci of Drosophila melanogaster by irradiation of spermatozoa in males. The mutations were characterized for the presence and location of simultaneously induced rearrangements and recessive lethal mutations. F1 females carrying induced visible mutations were identified, described and tested for fertility. The data are given in this paper. The proportions of mutants at the 4 loci, the ratios of whole-body: mosaic mutations, and the fertility of the mutant-carrying F1 females were similar for both types of radiation. Differences were observed between the frequencies of induced visible mutations and the rates of coincident visible and lethal induction. Although the analysis of the mutant chromosomes has not yet been completed, our data can be interpreted as providing confirmation that there are qualitative differences between the genetic effects of neutrons and X-rays.     

Recent changes in colony sizes of two seabird species in Scotland are not strongly influenced by Birds Directive status of the colony

Special Protection Areas (SPAs) designated under the EU Birds Directive have improved the conservation status of many terrestrial bird species in Europe, but protecting breeding sites may be less effective for highly mobile birds such as seabirds. Colony census data for Great Skuas Stercorarius skua and Great Black-backed Gulls Larus marinus in Scotland show that breeding numbers have fared no better in sites where these species are SPA breeding features and, counter-intuitively, the evidence indicates better performance in non-SPA colonies, most likely because non-SPA colonies are generally smaller so are less subject to density-dependent competition. The main drivers of population change are widespread rather than colony-based in these two species with recent reductions in carrying capacity. Many other seabird species are vulnerable to similar widespread pressures so seabird conservation strategy needs to focus on mitigating these pressures, as designation of seabird breeding sites as SPAs is not enough to ensure effective seabird conservation.     

Identifying useable semen

The "predictors of useable semen" used in most commercial AI centers provide a very conservative estimate of the relative fertility of individual boars. Furthermore, the relatively high sperm numbers used in commercial AI practice (usually >3 x10(9) total sperm per dose of extended semen) usually compensate for reduced fertility, as can be demonstrated in some boars when lower numbers of sperm are used for AI. Differences in relative boar fertility are also masked by the widespread use of pooled semen for commercial AI in many countries. However, the need to continually improve the efficiency of pork production, suggests that commercial AI practice should involve increased use of boars with the highest genetic merit for important production traits. Necessarily, this must be linked to the use of fewer sperm per AI dose, fewer inseminations per sow bred, and hence more sows bred by these superior sires. In turn, this requires improved techniques for evaluating semen characteristics directly related to the fertilization process, such as IVM-IVF assays, analysis of seminal plasma protein markers, more discriminatory tests of sperm motility and morphology, with the goal of identifying high-index boars whose fertility is sustained when low numbers of sperm are used for AI. This paper reviews the current status of laboratory-based boar semen evaluation techniques that meet these criteria.     

Impacts of Biotic Resource Enrichment on a Predator–Prey Population

The environmental carrying capacity is usually assumed to be fixed quantity in the classical predator–prey population growth models. However, this assumption is not realistic as the environment generally varies with time. In a bid for greater realism, functional forms of carrying capacities have been widely applied to describe varying environments. Modelling carrying capacity as a state variable serves as another approach to capture the dynamical behavior between population and its environment. The proposed modified predator–prey model is based on the ratio-dependent models that have been utilized in the study of food chains. Using a simple non-linear system, the proposed model can be linked to an intra-guild predation model in which predator and prey share the same resource. Distinct from other models, we formulate the carrying capacity proportional to a biotic resource and both predator and prey species can directly alter the amount of resource available by interacting with it. Bifurcation and numerical analyses are presented to illustrate the system’s dynamical behavior. Taking the enrichment parameter of the resource as the bifurcation parameter, a Hopf bifurcation is found for some parameter ranges, which generate solutions that posses limit cycle behavior.     

The importance of porcine sperm parameters on fertility in vivo

It would be desirable to use semen parameters to predict the in vivo fertilizing capacity of a particular ejaculate. In animal production, an ejaculate is divided into multiple doses for artificial insemination (AI); therefore, it would be economically beneficial to know the functional quality (i.e., fertility) of the semen before it is inseminated. To identify a predictive assay of the fertilizing capacity of a porcine ejaculate, we performed 4 rapid assays of sperm quality (motility, viability, physiological status as assessed by chlortetracycline fluorescence, and ATP content) on samples from 9 ejaculates, before and after a thermal stress test (42.5 degrees C, 45 min). These parameters were subsequently correlated with in vivo fertility resulting from AI with 2 sperm doses, 3 x 10(9) or 0.3 x 10(9) motile cells in 70 mL (optimal or suboptimal sperm number per insemination, respectively) from these same ejaculates. No parameter was correlated to the fertility rates obtained after inseminating with the optimal semen doses, either before or after the thermal stress test (P > 0.05). However, with respect to the animals inseminated with the suboptimal semen dose, sperm motility (the percentage of motile spermatozoa as assessed visually by microscopy) prior to thermal stress was well-correlated to fertility rates (r = 0.783, P = 0.01). The percentage of spermatozoa displaying the chlortetracycline Pattern AR (acrosome reaction) was also statistically related to fertility (r = 0.05, P = 0.04), but the biological importance of this relationship is questionable given the small variation among ejaculates (range: 0 to 2%). No other sperm parameter was significantly related to fertility rates in this group (P > 0.05). These data, therefore, indicate that sperm motility is a useful indicator of sperm fertilizing capacity in vivo. Moreover, to identify a predictor of semen fertility it is critical that the number of spermatozoa used during insemination is sufficiently low to detect differences in sperm fertilizing efficiency.     

Seasonal drought, soil fertility and the species density of tropical forest plant communities

Recent comparisons of plant species densities in tropical forest make it possible to evaluate factors that govern species richness. Contrary to earlier predictions, plant species densities are not greater on soils of relatively low fertility. In fact, the opposite trend is often observed, although the relationship between species densities and soil fertility is highly variable. However, tropical forest plant species densities consistently increase with rainfall. Species coexistence in wetter tropical forests may be facilitated by the absence of competition for moisture combined with year-round pest pressure and low understory light levels, which reduce growth rates and the potential for competition for other resources.     

Competition between Mitochondrial Haplotypes in Distinct Nuclear Genetic Environments: Drosophila Pseudoobscura Vs. D. Persimilis

A test for coadaptation of nuclear and mitochondrial genomes was performed using the sibling species, Drosophila pseudoobscura and D. persimilis. Two lines of flies with ``disrupted' cytonuclear genotypes were constructed by repeated backcrossing of males from one species to females carrying mitochondrial DNA (mtDNA) from the other species. Each ``disrupted' strain was competed in population cages with the original stock of each species from which the recurrent males were obtained during the backcrossing. As such, the two species' mitochondrial types were competed reciprocally in the nuclear genetic environments of each species. The trajectories of mtDNA haplotypes were followed in discrete-generation population cages using a PCR-four-cutter approach. A significant increase in the frequency of D. pseudoobscura mtDNA was observed in each of four replicate cages with a D. pseudoobscura nuclear background. In the D. persimilis nuclear background, one cage actually showed an increase in frequency of D. pseudoobscura mtDNA, although together the four replicate cages show little change in frequency. These results were repeated after frequency perturbations and reinitiation of each cage. An analysis of fitness components revealed that fertility selection greatly outweighed viability selection in these cytonuclear competition experiments. The asymmetry of the fitnesses of the mtDNA haplotypes on the two genetic backgrounds is consistent in direction with the previously reported asymmetry of female fertility in backcrosses between these two species. While our experiments do not allow us to identify mtDNA as the sole source of fitness variation, at a minimum the data indicate a fitness association between nuclear fertility factors and the D. pseudoobscura mtDNA on its own genetic background.     

A cost of restoration of male fertility in a gynodioecious species,Lobelia siphilitica

Abstract.— Models allowing the coexistence of females and hermaphrodites in gynodioecious populations assume a simple genetic system of sex determination, a seed fitness advantage of females (compensation), and a negative pleiotropic effect of nuclear sex-determining genes on fitness (cost of restoration). In Lobelia siphilitica , sex is determined by both mitochondrial genes causing cytoplasmic male sterility (CMS) and nuclear genes that restore fertility when present with specific CMS haplotypes (nuclear restorers). I tested for a cost of restoration in L. siphilitica by measuring restored hermaphrodites for five fitness components and estimating the number of nuclear restorers by crosses with females carrying CMS1 and CMS2. A cost of restoration appears as a significant negative coefficient (B) in the regression model explaining fitness. I found that hermaphrodites carrying more nuclear restorer genes for CMS2 (or restorer genes of greater effect) have lower pollen viability (B =– 1.08, P = 0.001). This pollen viability cost of restoration in L. siphilitica supports the theoretical prediction that negative pleiotropic effects of restorers will exist in populations of gynodioecious species containing females. The existence of such a cost supports the view that gynodioecy can be a stable breeding system in nature.     

Biological equilibrium in ecosystems 1. A theory of biological equilibrium

In this paper the process of establishing equilibrium in ecosystems is explored, and compared to the process of transformation. This process changes the state of organisms from absent to present, and the reverse, and if functioning randomly on very large numbers of individuals, results in biological equilibrium which can be expressed by the ratio (e?x): 1, withx=1 at Natural Biological Equilibrium (NBE). A model of ecosystems shows that four basic factors are operative in ecosystems: the constraint, or habitat-type; the regulator, or carrying capacity of the substrate; the input, or organisms; and the operator, death. The Theory of Biological Equilibrium (TBE) proposed, postulates that Natural Biological Equilibrium is reached when the individuals in an ecosystem are in dynamic balance with the carrying capacity of the substrate of the ecosystem. The TBE provides the theoretical basis for the Canonical Hypothesis, which postulates that the parameters in eco- systems are fixed, and can be computed from the number of species in the system. The TBE is in agreement with many well-known ecological phenomena, provides the basis for several hypotheses, and forces the rejection of some traditional hypotheses, viz., the hypothesis of competition as a factor in ecosystems.     

The sterile release method for population control with interspecific competition

A differential equations model of competing species with the release of sterile individuals of one of the species is examined. The system is found to have two positive steady states for certain parameter values; one of these is stable and the other is unstable. The system is quite resilient around the stable steady state. The release of steriles causes the nontarget species to increase in numbers. There exists a value of the release rate above which the pest species collapses to extinction. The existence of the competitor species assists the sterile release program since the pest equilibrium at any release rate is lower with the competitor species present than without it; in addition the release rate required to cause collapse of the pest species is lower with the competitor species present than without it. The effects of the parameters on the ease of eradication were examined. It was found that the ideal competitor species should have a high rate of increase, a large carrying capacity and exert strong competitive depression on the pests. The ideal pest would have a low rate of increase, a low carrying capacity and be a poor competitor.