The distribution of leaf photosynthetic activity in a mixed grass-legume pasture canopy

The distribution of net photosynthetic activity of leaves was measured in a mixed grass (Setaria sphacelata var. sericea)-legume (Desmodium intortum) pasture stand using a method based on concurrent measurement of the rate of CO₂ exchange, and ¹⁴CO₂ dosing followed by rapid harvesting according to height strata. Comparisons were also made between plots which differed in the period of regrowth following defoliation. The usual superiority of leaf net photosynthetic rates of a C₄ grass, compared with C₃ legume leaves, was found in the upper, well illuminated strata. These rates were, however, much lower than those usually described for horizontally exposed leaves, primarily because leaves in the pasture stand were inclined to the horizontal. At greater depth in the canopies, the superiority of rates in the grass was less evident, and consequently the relative contributions of grass and legume to canopy photosynthesis became more dependent on their leaf area indices. Attention is drawn to the relative simplicity of the method for examining the contribution of leaves, which may differ according to species or position in the canopy, to productivity of the whole stand.     

The distribution of leaf photosynthetic activity in a mixed grass-legume pasture canopy

The distribution of net photosynthetic activity of leaves was measured in a mixed grass (Setaria sphacelata var. sericea)-legume (Desmodium intortum) pasture stand using a method based on concurrent measurement of the rate of CO₂ exchange, and ¹⁴CO₂ dosing followed by rapid harvesting according to height strata. Comparisons were also made between plots which differed in the period of regrowth following defoliation. The usual superiority of leaf net photosynthetic rates of a C₄ grass, compared with C₃ legume leaves, was found in the upper, well illuminated strata. These rates were, however, much lower than those usually described for horizontally exposed leaves, primarily because leaves in the pasture stand were inclined to the horizontal. At greater depth in the canopies, the superiority of rates in the grass was less evident, and consequently the relative contributions of grass and legume to canopy photosynthesis became more dependent on their leaf area indices. Attention is drawn to the relative simplicity of the method for examining the contribution of leaves, which may differ according to species or position in the canopy, to productivity of the whole stand.     

Complex dynamics in a model microbial system

The forced double-Monod model (for a chemostat with a predator, a prey and periodically forced inflowing substrate) displays quasiperiodicity, phase locking, period doubling and chaotic dynamics. Stroboscopic sections reveal circle maps for the quasiperiodic regimes and noninvertible maps of the interval for the chaotic regimes. Criticality in the circle maps sets the stage for chaos in the model. This criticality may arise with an increase in the period or amplitude of forcing.     

Complex dynamics occur in a single-locus, multiallelic model of general frequency-dependent selection

We examine the characteristics of non-equilibrium dynamics produced by a simple well-known model of frequency-dependent selection at a single diploid locus. An examination of the parameter space of this “pairwise-interaction model” (PIM) revealed non-equilibrium dynamics for polymorphisms of 3, 4 and 5 alleles; both allele-frequency cycling and aperiodic trajectories were detected. We measured the number, cycle length and domains of attraction of the various attractors produced by the model. The domains of attraction tended to be smaller, and the cycles longer, for systems with larger number of alleles. Fitnesses that parametrized negative frequency-dependent selection were more likely to allow cycling, and these cycles also had larger domains of attraction. Aperiodic trajectories were detected only in cases with 4 or 5 alleles. The genetic cycles produced by the model do not have periods as short as those predicted in ecological models with cycling (such as predator–prey population cycles, etc.). Consequently, in a real-world system, PIM allele-frequency cycling is likely to be indistinguishable from stable equilibria when observed over short time scales.     

Chaos and order in plankton dynamics. Complex behavior of a simple model

The role of the diffusive interaction between fish-populated and fish-free habitats in a patchy environment in plankton pattern formation is studied by means of a minimal reaction-diffusion model of the nutrient-plankton-fish food chain. It is shown that such interaction can give rise to spatio-temporal plankton patterns. The fractal dimension of the patterns is shown dependent on the fish predation rate. The spatially averaged plankton dynamics depending on both fish predation rate and distance between fish-populated habitats can exhibit chaotic and regular behavior. The chaotic plankton dynamics is characteristic of a wide parameter range.     

Complex dynamics occur in a single-locus,multiallelic model of general frequency-dependent selection

We examine the characteristics of non-equilibrium dynamics produced by a simple well-known model of frequency-dependent selection at a single diploid locus. An examination of the parameter space of this “pairwise-interaction model” (PIM) revealed non-equilibrium dynamics for polymorphisms of 3, 4 and 5 alleles; both allele-frequency cycling and aperiodic trajectories were detected. We measured the number, cycle length and domains of attraction of the various attractors produced by the model. The domains of attraction tended to be smaller, and the cycles longer, for systems with larger number of alleles. Fitnesses that parametrized negative frequency-dependent selection were more likely to allow cycling, and these cycles also had larger domains of attraction. Aperiodic trajectories were detected only in cases with 4 or 5 alleles. The genetic cycles produced by the model do not have periods as short as those predicted in ecological models with cycling (such as predator–prey population cycles, etc.). Consequently, in a real-world system, PIM allele-frequency cycling is likely to be indistinguishable from stable equilibria when observed over short time scales.     

Effects of large herbivores on wood pasture dynamics in a European wetland system

Whether self-regulating large herbivores play a key role in the development of wood-pasture landscapes remains a crucial unanswered question for both ecological theory and nature conservation. We describe and analyse how a ‘partly self-regulating’ population of cattle, horses and red deer affected the development of the woody vegetation in the Oostvaardersplassen nature reserve (Netherlands). Using aerial photographs from 1980 to 2011, we analysed the development of shrubs and trees. Before the large herbivores were introduced in the Oostvaardersplassen in 1983, the woody vegetation increased and vegetation type significantly affected the number of establishments. Cover of woody species increased further from 1983 to 1996, not only by canopy expansion but also by new establishments. After 1996, cover of the woody vegetation decreased from 30% to <1% in 2011 and no new establishments were seen on the photographs. Survival of Sambucus nigra and Salix spp. increased with increasing distance to grassland, which is the preferred foraging habitat of the herbivores. These results support the hypothesis of Associational Palatability. In addition, our results show that the relative decline in cover of S. nigra and Salix spp. over a certain period was negatively correlated with the cover of S. nigra in the beginning of this period, presenting some evidence for the Associational Resistance and Aggregational Resistance hypothesis. Our research shows aspects necessary for the woodland–grassland cycle, such as a strong decline of woody vegetation at high numbers of large herbivores and regeneration of shrubs and trees at low densities. Thorny shrubs, which are important for the cycle, have not yet established in the grasslands. It seems that a temporary decline in herbivore numbers is necessary to create a window of opportunity for the establishment of these woody species.     

Mediterranean pasture dynamics: the role of germination

Abstract. Meteorological patterns have a decisive influence on the inter-annual dynamics of therophyte pastures under Mediterranean climatic conditions. The germination behaviour of annual pasture species was studied by subjecting two collections of seeds taken from plants and soil-seed banks to two phytotron-simulated weather patterns: early and late autumn rains. Species from these pastures were arranged along a gradient of sensitivity to temperature on the arrival of the first persistent rain. This sensitivity was manifested in both the total germination success of the species and the germination time profile. The different germination patterns of the species can provide competitive advantages depending on the autumn weather conditions.     

Complex dynamics of a neuron model with discontinuous magnetic induction and exposed to external radiation

The last two decades have seen many literatures on the mathematical and computational analysis of neuronal activities resulting in many mathematical models to describe neuron. Many of those models have described the membrane potential of a neuron in terms of the leakage current and the synaptic inputs. Only recently researchers have proposed a new neuron model based on the electromagnetic induction theorem, which considers inner magnetic fluctuation and external electromagnetic radiation as a significant missing part that can participate in neural activity. While the flux coupling of the membrane is considered equivalent to a memductance function of a memristor, standard memductance model of \(\alpha + 3\beta \phi^{2}\) has been used in the literatures, but in this paper we propose a new memductance function based on discontinuous flux coupling. Various dynamical properties of the neuron model with discontinuous flux coupling are studied and interestingly the proposed model shows hyperchaotic behavior which was not identified in the literatures. Furthermore, we consider a ring network of the proposed model and investigate whether the chimera state can emerge. The chimera state relates to the state with simultaneously coherence and incoherence in oscillatory networks and has received much attention in recent years.     

Seasonal dynamics of Mediterranean pasture seed banks along environmental gradients

The size and dynamics of seed banks were studied in grazed and ungrazed Mediterranean pastures at different altitudes and topography positions. The soil samples were collected in autumn and spring and the seed banks composition was determined by greenhouse germination over a 9-month period. The percentage of bare ground and the presence of new seedlings were recorded monthly from October to July in the field. A fall in seed density and species richness in the banks and a tendency for seeds to remain in the banks were linked to a rise in altitude. Germination in lower pastures mainly occurred in October in the numerous gaps left by the summer drought. At higher altitudes, the scarcity of gaps and the harsh climate led to an autumn–spring segregation of germination. On a local scale, the low slope positions and the ungrazed plots had a larger number of persistent seed bank species and a lower percentage of bare ground where seeds could germinate than their respective plots in the upper positions and grazed plots. A higher seed density in ungrazed than grazed plots was only detected in the three highest plots. No seed bank species richness trend was detected. In populations of the same species in different types of environments, the seasonal variation of seed numbers was environment-dependent for the majority of the species. In general, perennial grassland and its related low gaps availability appear to favour persistent seed banks.     

Complex dynamics in a three-level trophic system with intraspecies interaction

In this paper, we present a three-level trophic food chain, including intraspecies interaction. In contrast with other analyses, we consider the effect on the third trophic level by the first-level parameters. The model shows complex, as well as, chaotic oscillations. Bifurcation diagrams show period doubling route to chaos and crises. Also from the forward and backwards sections of the bifurcation diagrams, we find hysteresis. This result implies the coexistence of attractors for the same parameter values. In particular, we consider the coexistence of a chaotic and a P1 attractors. Our results show that the regulation in the food chain is not exclusive to either a food-prey or prey-predator interaction, but to a more subtle food-prey-predator interaction, where, for some parameter values, a food-prey or a prey-predator regulation may dominate the system's dynamics. Finally, we consider the impact of the intraspecies interaction in the overall dynamics of the food chain.     

Complex dynamics of a stochastic uni-directional consumer-resource mutualism system

Like predation and competition, mutualism is recognized as a consumer-resource interaction, which includes bi-directional and uni-directional mutualisms. In this paper, we firstly propose a stochastic uni-directional consumer-resource system of two species in which the consumer has both positive and negative effects on the resource, while the resource has only a positive effect on the consumer. We then mathematically analyze the system, to demonstrate the existence, uniqueness, asymptotic pathwise behavior and stochastically ultimately boundedness of the global positive solution, and to establish sufficient conditions for the global attractivity and the existence of ergodic stationary distribution of the system. We also establish sufficient conditions for the extinction and persistence in mean of the resource, the consumer or the entire system. Numerical simulations are carried out to demonstrate the analytical results.     

Plasmid dynamics in a model soil column

Continuous-flow column reactors were used to study the dynamics of plasmid exchange in a structured, thermodynamically open system containing either Enterobacter cloacae or Pseudomonas cepacia , both carrying the transmissible recombinant plasmid R388::Tn1721. Plasmid transfer rates were higher in vermiculite and sterile soil columns supplied with nutrient solution than those in sterile and non-sterile soil columns without input of nutrient solution. For both species, donor and recipient strains took about 5 days to reach their maximum densities in effluents from the columns supplied with nutrient solution. After about 8 day s the donor and transconjugant populations of P. cepacia in the effluent solution decreased exponentially, whereas E. cloacae donor, recipient and transconjugant strains maintained steady-state concentrations. The difference between plasmid stability in the two species may have significant consequences in terms of releasing plasmid-bearing genetically modified microorganisms into the natural environment. The plasmid is persistent in E. cloacae in non-sterile soil even though its transfer to the marked recipient in non-sterile soil was minimal.     

Overshooting dynamics in a model adaptive radiation

The history of life is punctuated by repeated periods of unusually rapid evolutionary diversification called adaptive radiation. The dynamics of diversity during a radiation reflect an overshooting pattern with an initial phase of exponential-like increase followed by a slower decline. Much attention has been paid to the factors that drive the increase phase, but far less is known about the causes of the decline phase. Decreases in diversity are rarely associated with climatic changes or catastrophic events, suggesting that they may be an intrinsic consequence of diversification. We experimentally identify the factors responsible for losses in diversity during the later stages of the model adaptive radiation of the bacterium Pseudomonas fluorescens. Proximately, diversity declines because of the loss of biofilm-forming niche specialist morphotypes. We show that this loss occurs despite the presence of strong divergent selection late in the radiation and is associated with continued adaptation of resident niche specialists to both the biotic and abiotic environments. These results suggest that losses of diversity in the latter stages of an adaptive radiation may be a general consequence of diversification through competition and lends support to the idea that the conditions favouring the emergence of diversity are different from those that ensure its long-term maintenance.     

禾-豆混播草地种间竞争与共存

以羊草分别与沙打旺、兴安胡枝子、花苜蓿、紫花苜蓿、山野豌豆5种豆科牧草在混播数量比为1:0、2:1、1:2、0:1的条件下建立两物种混播草地,以相对产量、相对密度和相对产量总值为指标,比较各个混播草地中种间竞争的相对激烈程度;各个物种组合的种间竞争优势以及是否发生氮素资源分离;并探索不同禾-豆混播群落达到共存状态的可能途径.研究结果表明,各个禾-豆组合的相对产量总值分别在不同收获时期大于1,禾草与豆科牧草的生态位发生了不同程度的分离.沙打旺和紫花苜蓿对羊草具有显著的竞争优势,即使其种内竞争大于种间竞争时,混生的羊草亦受到强烈的种间竞争压力.与此相反,羊草对兴安胡枝子、花苜蓿和山野豌豆具有种间竞争优势.刈割对竞争双方的优劣地位产生很大影响,减少强竞争力物种的混播比例,可促进混播物种双方均受益,形成共存格局.实验采用的相对密度指标在预测未来混播种群组成上比相对产量更为可行,并且具有维持低个体大小、高构件密度能力是竞争关系中忍耐型物种能够长期存在的可能原因之一.     

Emergent dynamics in a model of visual cortex

This paper proposes that the network dynamics of the mammalian visual cortex are highly structured and strongly shaped by temporally localized barrages of excitatory and inhibitory firing we call ‘multiple-firing events’ (MFEs). Our proposal is based on careful study of a network of spiking neurons built to reflect the coarse physiology of a small patch of layer 2/3 of V1. When appropriately benchmarked this network is capable of reproducing the qualitative features of a range of phenomena observed in the real visual cortex, including spontaneous background patterns, orientation-specific responses, surround suppression and gamma-band oscillations. Detailed investigation into the relevant regimes reveals causal relationships among dynamical events driven by a strong competition between the excitatory and inhibitory populations. It suggests that along with firing rates, MFE characteristics can be a powerful signature of a regime. Testable predictions based on model observations and dynamical analysis are proposed.     

Evidence for a periodic excretion of nitrogen by roots of grass-legume associations

Diurnal variation in ion content of the solution bathing roots of two plants growing together in sand culture was analysed for three pairs of grass-legume species (Lolium multiflorum andTrifolium pratense; Zea mays andGlycine hispida; Avena sativa andVicia sativa) and their monospecific controls. Biomass and nitrogen content of plants were determined. Ion concentration (NO ₃ ⁻ , NO ₂ ⁻ , NH ₄ ⁺ , and K⁺) and pH of root solutions were measured for Lolium-Trifolium plant pairs and controls at 6 hours intervals over 36 h, starting at 8 am within a circadian cycle. Root solutions were regularly depleted in NO ₃ ⁻ by the grasses (Lolium-Lolium control) throughout the cycle. For associations involving the legume (Lolium-Trifolium and Trifolium-Trifolium), NO ₃ ⁻ depletion was followed by NO ₃ ⁻ enrichment at night, from late afternoon to early morning; the enrichment was more marked for the Lolium-Trifolium association. Solutions which did not contain NO ₂ ⁻ ions, were enriched by trace amounts of NH ₄ ⁺ ions, largely depleted in K⁺ and alkalanized for all associations throughout the cycle. Repeating the experiment with the three pairs of species at the vegetative phase of development confirmed the previous results: NO ₃ ⁻ enrichment during the night for associations with legumes. When the experiment was repeated with older plants which had almost completed their flowering stage, depletion only was observed and no NO ₃ ⁻ enrichment. These data suggest that NO ₃ ⁻ enrichment results from N excretion from active nodulated roots of the legume, accounting for the increase in both biomass and nitrogen content of the companion grass in grass-legume association. The quantitative importance and periodicity of nitrogen excretion as well as the origin of nitrate enrichment are discussed.     

Complex biotic interactions drive long-term vegetation dynamics in a subarctic ecosystem

Predicting impacts of global warming requires understanding of the extent to which plant biomass and production are controlled by bottom-up and top-down drivers. By annually monitoring community composition in grazed control plots and herbivore-free exclosures at an Arctic location for 15 years, we detected multiple biotic interactions. Regular rodent cycles acted as pulses driving synchronous fluctuations in the biomass of field-layer vegetation; reindeer influenced the biomass of taller shrubs, and the abundance of plant pathogenic fungi increased when densities of their host plants increased in exclosures. Two outbreaks of geometrid moths occurred during the study period, with contrasting effects on the field layer: one in 2004 had marginal effects, while one in 2012 severely reduced biomass in the control plots and eliminated biomass that had accumulated over 15 years in the exclosures. The latter was followed by a dramatic decline of the dominant understory dwarf-shrub Empetrum hermaphroditum, driven by an interaction between moth herbivory on top buds and leaves, and increased disease severity of a pathogenic fungus. We show that the climate has important direct and indirect effects on all these biotic interactions. We conclude that long time series are essential to identify key biotic interactions in ecosystems, since their importance will be influenced by climatic conditions, and that manipulative treatments are needed in order to obtain the mechanistic understanding needed for robust predictions of future ecosystem changes and their feedback effects.