Intra-plant versus Inter-plant Root Competition in Beans: avoidance,resource matching or tragedy of the commons

Root competition inhibits root proliferation. All else equal, a plant should invest roots in a nutrient patch devoid of roots rather than one already occupied by roots. Less clear is how a plant should respond to intra-plant versus inter-plant root competition. We consider three responses for how a plant may select habitats based on intra-versus inter-plant root competition: inter-plant avoidance, resource matching, or intra-plant avoidance. The first assumes that plants prefer to have their own space and preferentially proliferate roots away from neighboring plants. The second response, based on the ideal free distribution, assumes that plants invest so as to equalize average returns from roots, regardless of the identity of the neighboring roots. The third, based on game theory, assumes that the plant proliferates roots so as to maximize whole-plant fitness, in which case it is better to proliferate plants among a neighbor's roots than to continue proliferating amongst one's own roots. To test among these models we grew beans (Phaseolus varigaris, var. Kenya) in a greenhouse under two planting scenarios. Both scenario were tested under 0.5 and 0.1 strength of nutrient solution. Under scenario A (fence-sitters), two split-root plants each shared two patches by virtue of having roots in each. Under scenario B (owners) two plants each had their own patch. The results supported the game theory model of intra-plant avoidance (whole plant habitat selection). Fence-sitters produced 150% more root mass per individual than owners. Owners produced 90% more yield (dry mass of pods) than fence-sitters. Furthermore, owners had significantly higher shoot-root ratios than fence-sitters. These effects did not vary with high or low nutrient levels. The over-proliferation of roots under inter-plant competition (fence-sitters) was manifest by the tenth day after planting. In short, the fence-sitters engaged in a tragedy of the commons in which they competed with each other through root proliferation. At the ESS, the fitness maximizing strategy of the individual is to sacrifice collective yield in a quest to `steal' nutrients from its neighbor. The research has three implications. First, plants may be able to assess and respond to local opportunities in a manner that maximizes the good of the whole plant. Second, nutrient foraging as a game may provide a fresh perceptive for viewing root competition either intra-specifically or inter-specifically. Third, it may be possible to increase the yield of certain crop species by breeding more `docile' cultivars that do not overproduce roots in response to inter-plant competition.     

The ecological significance of rapid wound-induced changes in plants: insect grazing and plant competition

Summary This paper investigates the hypothesis that a rapidly induced phytochemical response to grazing damage, such as that seen in tomato, serves to deflect insect herbivores away from leaves soon after damaging them (the grazing dispersal hypothesis). As a result, grazing damage is more dispersed than it otherwise would be, and young leaves, which may be of particular importance to a plant in competition for light, are not damaged excessively. In the first experiment, artificial removal of c. 15% of leaf area led to a significant reduction in plant performance compared with undamaged controls, but only when the plants were grown together in competition for light. The second experiment demonstrated that the distribution of grazing damage within the plant was an important factor in the outcome of competition; in those plants in which grazing was applied to the lower leaves there was no effect of damage upon performance compared with undamaged controls, whereas grazing to the upper leaves significantly reduced plant performance. A third experiment provided some insight into how this interaction between damage and competition comes about. It was shown that damage to leaves led to a rapid drop in the rate of extension growth of the main shoot, especially when the upper leaves were damaged, and normal rates of growth were not resumed for at least 3 days. It is argued that in a rapidly growing canopy, such an effect may mean that a damaged plant loses its position in the height hierarchy. The final experiment showed that previous damage to plants can affect the distribution of subsequent grazing by larvae of Spodoptera littoralis, apparently through a wound-induced reduction in leaf palatability. Plants which had been artificially damaged 48 h previously were grazed significantly less than controls, and the avoidance effect was greatest in the young leaves. These results are consistent with the grazing dispersal hypothesis, and suggest that rapid wound-induced responses may be of greatest significance in species characteristic of fertile environments where competition for light is particularly intense.     

A mixed strategy model for the emergence and intensification of social learning in a periodically changing natural environment

Based on a population genetic model of mixed strategies determined by alleles of small effect, we derive conditions for the evolution of social learning in an infinite-state environment that changes periodically over time. Each mixed strategy is defined by the probabilities that an organism will commit itself to individual learning, social learning, or innate behavior. We identify the convergent stable strategies (CSS) by a numerical adaptive dynamics method and then check the evolutionary stability (ESS) of these strategies. A strategy that is simultaneously a CSS and an ESS is called an attractive ESS (AESS). For certain parameter sets, a bifurcation diagram shows that the pure individual learning strategy is the unique AESS for short periods of environmental change, a mixed learning strategy is the unique AESS for intermediate periods, and a mixed learning strategy (with a relatively large social learning component) and the pure innate strategy are both AESS's for long periods. This result entails that, once social learning emerges during a transient era of intermediate environmental periodicity, a subsequent elongation of the period may result in the intensification of social learning, rather than a return to innate behavior.     

Mate choice in the face of costly competition

Studies of mate choice commonly ignore variation in preferencesand assume that all individuals should favor the highest-qualitymate available. However, individuals may differ in their matepreferences according to their own age, experience, size, orgenotype. In the present study, we highlight another simplereason why preferences may differ: if there is costly competitionfor mates, the poorest competitors might be better off avoidingthe highest-quality partners and instead targeting low-qualitypartners, so that they minimize the costs they incur. We presenta game-theoretical model of mate choice in which males of differingquality compete for access to females and try to retain themtill the time of mating. Our model predicts that high-qualitymales, who are better competitors, have a preference for thebest females that is typically several times stronger than thatof low-quality males. Early in the competitive period, the lattermay even prefer low-quality females over high-quality females.Thus, variation in competitive ability generates variation inboth the strength and direction of preferences. Differencesin competitive ability result in assortative mating with respectto quality, which is reinforced by variation in preferences.As the time of mating draws near and there is an increased riskof ending up unpaired, all males become indifferent to the qualityof potential mates. Our findings are equally applicable to femalechoice for males, and offer a new explanation for adaptive variationin mating preferences based on differing abilities to cope withthe costs of mate choice.     

Effect of plant age and turfgrass competition on the efficacy of the Sclerotinia minor granular bioherbicide on broadleaf plantain and prostrate knotweed

Broadleaf plantain and prostrate knotweed are important weeds of turfgrass systems. The fungus Sclerotinia minor Jagger (IMI 344141) has been registered as a biological herbicide (Sarritor?) for dandelion (Taraxacum officinale) in Canadian turfgrass habitats. The objective of this study was to evaluate the effect of plant age and turfgrass environment on the efficacy of S. minor against two additional weeds; broadleaf plantain and prostrate knotweed. The turfgrass environment alone exerted significant reduction of above and below ground biomass of broadleaf plantain, to the same magnitude as the S. minor treatment in a grass-free environment. Prostrate knotweed biomass, however, was not reduced to this extent by competition with turfgrass. In the presence of grass, S. minor caused a significant biocontrol effect on all studied variables with more than 90% above ground damage on both weed species. Severe damage occurred on 3–6-week-old plantains with 100% above and below ground reduction, although smaller dry weight reductions were observed on older plantains. Treatment with S. minor reduced the dry matter of 3–5-week-old prostrate knotweed by 65–85%, but less damage occurred on older prostrate knotweed. The bioherbicide fungus is destructive for both species, but variation in area of contact due to different growth forms, growth rates and resource allocation patterns due to different life forms resulted in different biocontrol efficacy on the two species. Control of broadleaf plantain was effective – similar to that previously reported for dandelion – but control of prostrate knotweed was only partial.     

The ideal free distribution: a review and synthesis of the game-theoretic perspective

The Ideal Free Distribution (IFD), introduced by Fretwell and Lucas in [Fretwell, D.S., Lucas, H.L., 1970. On territorial behavior and other factors influencing habitat distribution in birds. Acta Biotheoretica 19, 16-32] to predict how a single species will distribute itself among several patches, is often cited as an example of an evolutionarily stable strategy (ESS). By defining the strategies and payoffs for habitat selection, this article puts the IFD concept in a more general game-theoretic setting of the “habitat selection game”. Within this game-theoretic framework, the article focuses on recent progress in the following directions: (1) studying evolutionarily stable dispersal rates and corresponding dispersal dynamics; (2) extending the concept when population numbers are not fixed but undergo population dynamics; (3) generalizing the IFD to multiple species.For a single species, the article briefly reviews existing results. It also develops a new perspective for Parker’s matching principle, showing that this can be viewed as the IFD of the habitat selection game that models consumer behavior in several resource patches and analyzing complications involved when the model includes resource dynamics as well. For two species, the article first demonstrates that the connection between IFD and ESS is now more delicate by pointing out pitfalls that arise when applying several existing game-theoretic approaches to these habitat selection games. However, by providing a new detailed analysis of dispersal dynamics for predator-prey or competitive interactions in two habitats, it also pinpoints one approach that shows much promise in this general setting, the so-called “two-species ESS”. The consequences of this concept are shown to be related to recent studies of population dynamics combined with individual dispersal and are explored for more species or more patches.     

Stability of a Ricker-type competition model and the competitive exclusion principle

Our main objective is to study a Ricker-type competition model of two species. We give a complete analysis of stability and bifurcation and determine the centre manifolds, as well as stable and unstable manifolds. It is shown that the autonomous Ricker competition model exhibits subcritical bifurcation, bubbles, period-doubling bifurcation, but no Neimark–Sacker bifurcations. We exhibit the region in the parameter space where the competition exclusion principle applies.     

The evolution of conformist transmission in social learning when the environment changes periodically

Conformity is often observed in human social learning. Social learners preferentially imitate the majority or most common behavior in many situations, though the strength of conformity varies with the situation. Why has such a psychological tendency evolved? I investigate this problem by extending a standard model of social learning evolution with infinite environmental states (Feldman, M.W., Aoki, K., Kumm, J., 1996. Individual versus social learning: evolutionary analysis in a fluctuating environment. Anthropol. Sci. 104, 209-231) to include conformity bias. I mainly focus on the relationship between the strength of conformity bias that evolves and environmental stability, which is one of the most important factors in the evolution of social learning. Using the evolutionarily stable strategy (ESS) approach, I show that conformity always evolves when environmental stability and the cost of adopting a wrong behavior are small, though environmental stability and the cost of individual learning both negatively affect the strength of conformity.     

Size asymmetry in intraspecific competition and the density-dependence of inbreeding depression in a natural plant population: a case study in cassava (Manihot esculenta Crantz, Euphorbiaceae)

The effects of competition on the genetic composition of natural populations are not well understood. We combined demography and molecular genetics to study how intraspecific competition affects microevolution in cohorts of volunteer plants of cassava (Manihot esculenta) originating from seeds in slash-and-burn fields of Palikur Amerindians in French Guiana. In this clonally propagated crop, genotypic diversity is enhanced by the incorporation of volunteer plants into farmers' stocks of clonal propagules. Mortality of volunteer plants was density-dependent. Furthermore, the size asymmetry of intraspecific competition increased with local clustering of plants. Size of plants was correlated with their multilocus heterozygosity, and stronger size-dependence of survival in clusters of plants, compared with solitary plants, increased the magnitude of inbreeding depression when competition was severe. The density-dependence of inbreeding depression of volunteer plants helps explain the high heterozygosity of volunteers that survive to harvest time and thus become candidates for clonal propagation. This effect could help favour the maintenance of sex in this 'vegetatively' propagated crop plant.     

国际植物基因组组织与计划概况及我国的应对策略

本文简要介绍了植物基因组领域主要的组织和计划,简单分析了国际植物基因组计划的特点和我国存在的问题,并初步提出我国的应对策略。     

Carbon gain in a multispecies canopy: the role of specific leaf area and photosynthetic nitrogen-use efficiency in the tragedy of the commons

Models have been formulated for monospecific stands in which canopy photosynthesis is determined by the vertical distribution of leaf area, nitrogen and light. In such stands, resident plants can maximize canopy photosynthesis by distributing their nitrogen parallel to the light gradient, with high contents per unit leaf area at the top of the vegetation and low contents at the bottom. Using principles from game theory, we expanded these models by introducing a second species into the vegetation, with the same vertical distribution of biomass and nitrogen as the resident plants but with the ability to adjust its specific leaf area (SLA, leaf area:leaf mass). The rule of the game is that invaders replace the resident plants if they have a higher plant carbon gain than those of the resident plants. We showed that such invaders induce major changes in the vegetation. By increasing their SLA, invading plants could increase their light interception as well as their photosynthetic nitrogen-use efficiency (PNUE, the rate of photosynthesis per unit organic nitrogen). By comparison with stands in which canopy photosynthesis is maximized, those invaded by species of high SLA have the following characteristics: (1) the leaf area index is higher; (2) the vertical distribution of nitrogen is skewed less; (3) as a result of the supra-optimal leaf area index and the more uniform distribution of nitrogen, total canopy photosynthesis is lower. Thus, in dense canopies we face a classical tragedy of the commons: plants that have a strategy to maximize canopy carbon gain cannot compete with those that maximize their own carbon gain. However, because of this strategy, individual as well as total canopy carbon gain are eventually lower. We showed that it is an evolutionarily stable strategy to increase SLA up to the point where the PNUE of each leaf is maximized.     

Modeling the growth of individuals in plant populations: local density variation in a strand population of Xanthium strumarium (Asteraceae)

We studied the growth of individual Xanthium strumarium plants growing at four naturally occurring local densities on a beach in Maine: (1) isolated plants, (2) pairs of plants ≤1 cm apart, (3) four plants within 4 cm of each other, and (4) discrete dense clumps of 10-39 plants. A combination of nondestructive measurements every 2 wk and parallel calibration harvests provided very good estimates of the growth in aboveground biomass of over 400 individual plants over 8 wk and afforded the opportunity to fit explicit growth models to 293 of them. There was large individual variation in growth and resultant size within the population and within all densities. Local crowding played a role in determining plant size within the population: there were significant differences in final size between all densities except pairs and quadruples, which were almost identical. Overall, plants growing at higher densities were more variable in growth and final size than plants growing at lower densities, but this was due to increased variation among groups (greater variation in local density and/or greater environmental heterogeneity), not to increased variation within groups. Thus, there was no evidence of size asymmetric competition in this population. The growth of most plants was close to exponential over the study period, but half the plants were slightly better fit by a sigmoidal (logistic) model. The proportion of plants better fit by the logistic model increased with density and with initial plant size. The use of explicit growth models over several growth intervals to describe stand development can provide more biological content and more statistical power than "growth-size" methods that analyze growth intervals separately.     

Fisher’s fundamental theorem of inclusive fitness and the change in fitness due to natural selection when conspecifics interact

Competition and cooperation is fundamental to evolution by natural selection, both in animals and plants. Here, I investigate the consequences of such interactions for response in fitness due to natural selection. I provide quantitative genetic expressions for heritable variance and response in fitness due to natural selection when conspecifics interact. Results show that interactions among conspecifics generate extra heritable variance in fitness, and that interacting with kin is the key to evolutionary success because it translates the extra heritable variance into response in fitness. This work also unifies Fisher’s fundamental theorem of natural selection (FTNS) and Hamilton’s inclusive fitness (IF). The FTNS implies that natural selection maximizes fitness, whereas Hamilton proposed maximization of IF. This work shows that the FTNS describes the increase in IF, rather than direct fitness, at a rate equal to the additive genetic variance in fitness. Thus, Hamilton’s IF and Fisher’s FTNS both describe the maximization of IF.