Effects of positive interactions, size symmetry of competition and abiotic stress on self-thinning in simulated plant populations

Background and Aims

Competition drives self-thinning (density-dependent mortality) in crowded plant populations. Facilitative interactions have been shown to affect many processes in plant populations and communities, but their effects on self-thinning trajectories have not been investigated.

Methods

Using an individual-based ‘zone-of-influence’ model, we studied the potential effects of the size symmetry of competition, abiotic stress and facilitation on self-thinning trajectories in plant monocultures. In the model, abiotic stress reduced the growth of all individuals and facilitation ameliorated the effects of stress on interacting individuals.

Key Results

Abiotic stress made the log biomass – log density relationship during self-thinning steeper, but this effect was reduced by positive interactions among individuals. Size-asymmetric competition also influenced the self-thinning slope.

Conclusions

Although competition drives self-thinning, its course can be affected by abiotic stress, facilitation and competitive symmetry.     

Facilitation among plants can accelerate density‐dependent mortality and steepen self‐thinning lines in stressful environments

The speed and slope of plant self‐thinning are all affected by plant–plant interactions across environmental gradients. Possible mechanisms driving the self‐thinning dynamics include the relative strength of root versus shoot competition, and the interplay between competition and facilitation. Although these mechanisms often act in concert, their relative importance has not yet been fully explored. We used both a one‐layer and a two‐layer zone‐of‐influence (ZOI) model to examine how competition and facilitation drive self‐thinning across stress gradients. As a development of the traditional ZOI model, the two‐layer version explicitly models shoot and root growth and neighbor interactions, and thus the overall size‐symmetry of competition is regulated by the relative strength of root versus shoot competition. One‐layer model simulations revealed that increasingly asymmetric competition accelerated thinning, and steepened (slope ranged from about –1 to –4/3) and lowered self‐thinning lines. Stress slowed down density‐dependent mortality considerably when competition was not completely symmetric. Stress significantly decreased the self‐thinning intercept, while facilitation simply counteracted stress effects. Both stress and facilitation showed little effect on the slope. In the two‐layer model, both stress and facilitation affected mortality in the same way as in the one‐layer version when competition was not completely symmetric. Different from the one‐layer model, the two‐layer version showed that the effects of stress and facilitation on the self‐thinning slope were mediated by the asymmetry of competition. As stress increased, the overall asymmetry of competition shifted from asymmetric to symmetric due to increased relative strength of root competition. High stress thus dramatically flattened self‐thinning lines, whereas the inclusion of facilitation counteracted stress and led to steeper self‐thinning lines. Our two‐layer model is based on the current knowledge of plant–plant interactions, and better represents ecological realities. It can help elaborate experiments for testing the role of competition and facilitation in driving plant population dynamics.     

Asymmetric facilitation can reduce size inequality in plant populations resulting in delayed density‐dependent mortality

Size inequality in plant populations is a ubiquitous feature that has received much attention due to ecological and evolutionary implications. The mechanisms driving size inequality were mainly attributed to different modes of competition (symmetric versus asymmetric), while the potential effects of different modes of facilitation (symmetric versus asymmetric) to this pattern have not yet been fully explored. We employed an individual‐based model to explore the relative roles of both competition and facilitation simultaneously along an environmental stress gradient. Special emphasis was given to the assessment of symmetric facilitation (plants receive benefit from each other equally or proportionally to benefactors’ sizes) and asymmetric facilitation (beneficiary plants receive benefits from benefactor plants that are higher than proportional to the benefactors’ size) in altering plant size inequality. We found that independent of the particular mode of competition, symmetric facilitation generally increased size inequality, whereas asymmetric facilitation decreased it. This pattern was consistent along the stress gradient. Because of their different effects on size inequality, symmetric facilitation accelerated self‐thinning, whereas asymmetric facilitation delayed the onset of density‐dependent mortality, promoting survival under intermediate stress conditions. We compared our model predictions with both 1) a previous modelling study focusing on the effect of (symmetric) facilitation on the size inequality, and 2) re‐analysed data from a published experiment generating asymmetric facilitation of plants against enhanced ultraviolet‐B (UV‐B). Whereas our model predictions and the results of the empirical experiment were consistent, we found that previous theoretical results that solely relied on symmetric facilitation need to be re‐adjusted. Our study showed that combinations of different modes of competition and facilitation can alter size inequality in different ways and with important consequences for the onset of density‐dependent mortality during population development. Explicitly considering different modes and mechanisms of interactions (both facilitation and competition) will improve mechanistic understanding in plant ecology.     

Effects of diversity on diversity: consequences of competition and facilitation

Diversity in one group of species or genotypes is often correlated with diversity in a second group – prominent examples including native vs exotic species, and genetic diversity in a focal species vs species diversity in the rest of the community. I used simulation models to investigate the roles of competition and facilitation among species or genotypes in creating diversity–diversity relationships, with a focus on facilitation, which has received little theoretical attention. When competitive interactions dominate, increasing diversity in one group reduces diversity in the second group via filling of available niche space. Facilitation can create positive diversity–diversity relationships via a sampling effect, whereby a strong facilitator of the second group is more likely to be present as diversity increases in the first group, and also via one group acting as a source of biotic heterogeneity (i.e. diversifying selection) on the second group. However, the biotic heterogeneity effect is expected only under restricted conditions – with asymmetric facilitation, only during a transient period, or only over a small range of species diversity levels – and therefore seems unlikely to operate within trophic levels in natural communities. More generally, the simultaneous operation of competition and facilitation results in several different diversity–diversity relationships and underlying mechanisms. The results clarify the potential roles of positive and negative interactions in creating diversity–diversity relationships, and in determining the outcome of community dynamics in general. This study also highlights some important difficulties in incorporating facilitation into ecological theory for communities with many species.     

The role of natural selection in evolution

To evaluate properly a role of natural selection, its effect should be considered in relation to different phases of the evolutionary cycle postulated earlier by the author. At the first stage of the cycle natural selection is directed towards organism's persistence to detrimental external factors and leads to an increased fitness (that is viability and fecundity) in every generation. At the next stage of the cycle natural selection occurs under conditions of intraspecific competition and is directed towards a more efficient utilization of food resources. At this stage natural selection leads to formation and divergence of intraspecific races and is carried out by "single" selection actions occurring now and then and consisting of the survival of rare mutants with an altered ecological potential. Such a strict selection for certain mutants occurs again during the periods of acute competition for food, the selected mutants being characterized by a decrease of fitness, the latter to have been restored by means of the "ordinary" selection within the intervals between crises. According to the model suggested, homozygotes for "detrimental" recessive alleles could be selected in diploids, as the mutants mentioned with altered ecological potential. At the end of the cycle, there is a kind of selection for hybrids in which ecological potential of specialized intraspecific races is combined. The genetic drift is considered as an inevitable consequence of the postulated mechanism of natural selection.     

植物邻体间的正相互作用

植物间的正负相互作用是构建植被群落的重要因素,也是群落生态学研究的中心内容之一。近20a来,植物间正相互作用的研究得到快速发展。综述了正相互作用的定义,不同植物群落中的直接、间接正相互作用及其发生机制,正相互作用研究的实验和模型方法,正负相互作用随胁迫梯度的变化及正相互作用对群落构建的影响。探讨了正相互作用研究前景:(1)进一步理解正负相互作用的平衡及其对群落构建的影响;(2)加深对全球变暖背景下的正相互作用的认识;(3)需把正相互作用研究同进化联系起来;(4)充分发挥正相互作用在生态系统中的推动力作用,把正相互作用应用到生态恢复中,为恢复退化生态系统服务。     

Geomorphological disturbance affects ecological driving forces and plant turnover along an altitudinal stress gradient on alpine slopes

The altitudinal gradient is considered as a stress gradient for plant species because the development and fitness of plant communities tend to decrease as a result of the extreme environmental conditions present at high elevations. Abiotic factors are predicted to be the primary filter for species assemblage in high alpine areas, influencing biotic interactions through both competition for resources and positive interactions among species. We hypothesised that the relative importance of the ecological driving forces that affect the biotic interactions within plant communities changes along an elevation gradient on alpine debris slopes. We used multiple gradient analyses of 180 vegetation plots along an altitudinal range from ~1,600 to 2,600 m and single 100 m-bands in the Adamello-Presanella Group (Central Alps) to investigate our hypothesis; we measured multiple environmental variables related to different ecological driving forces. Our results illustrate that resource limitations at higher elevations affect not only the shift from competition to facilitation among species. A geomorphological disturbance regime along alpine slopes favours the resilience of the high-altitude species within topographic/geomorphological traps. An understanding of the ecological driving forces and positive interactions as a function of altitude may clarify the mechanisms underlying plant responses to present and future environmental changes.     

Intraspecific competition drives increased resource use diversity within a natural population

Resource competition is thought to play a major role in driving evolutionary diversification. For instance, in ecological character displacement, coexisting species evolve to use different resources, reducing the effects of interspecific competition. It is thought that a similar diversifying effect might occur in response to competition among members of a single species. Individuals may mitigate the effects of intraspecific competition by switching to use alternative resources not used by conspecific competitors. This diversification is the driving force in some models of sympatric speciation, but has not been demonstrated in natural populations. Here, we present experimental evidence confirming that competition drives ecological diversification within natural populations. We manipulated population density of three-spine sticklebacks (Gasterosteus aculeatus) in enclosures in a natural lake. Increased population density led to reduced prey availability, causing individuals to add alternative prey types to their diet. Since phenotypically different individuals added different alternative prey, diet variation among individuals increased relative to low-density control enclosures. Competition also increased the diet-morphology correlations, so that the frequency-dependent interactions were stronger in high competition. These results not only confirm that resource competition promotes niche variation within populations, but also show that this increased diversity can arise via behavioural plasticity alone, without the evolutionary changes commonly assumed by theory.     

Sex and aging: A comparison between two phenoptotic phenomena

Phenoptosis is a phenomenon that is genetically programmed and favored by natural selection, and that determines death or increased risk of death (fitness reduction) for the individual that manifests it. Aging, here defined as agerelated progressive mortality increase in the wild, if programmed and favored by natural selection, falls within the definition of phenoptosis. Sexual reproduction (sex), as for the involved individuals determines fitness reduction and, in some species, even certain death, also falls within the definition of phenoptosis. In this review, sex and aging are analyzed as phenoptotic phenomena, and the similarities between them are investigated. In particular, from a theoretical standpoint, the genes that cause and regulate these phenomena: (i) require analyses that consider both individual and supra-individual selection because they are harmful in terms of individual selection, but advantageous (that is, favored by natural selection) in particular conditions of supra-individual selection; (ii) determine a higher velocity of and greater opportunities for evolution and, therefore, greater evolutionary potential (evolvability); (iii) are advantageous under ecological conditions of K-selection and with finite populations; (iv) are disadvantageous (that is, not favored by natural selection) under ecological conditions of r-selection and with unlimited populations; (v) are not advantageous in all ecological conditions and, so, species that reproduce asexually or species that do not age are predicted and exist.     

From selection to complementarity: the shift along the abiotic stress gradient in a controlled biodiversity experiment

The effects of principal mechanisms (selection and complementarity) of biodiversity on ecosystem functionality have been well studied. However, it remains unknown how environmental conditions affect the relative strength of these two mechanisms. To answer this question, a controlled pot experiment was conducted in which species diversity was manipulated in low (natural soil) and high stress (mine tailing) plots, respectively. Our results demonstrate that the principal mechanism underlying the increasing biomass shifts from the selection to complementarity with increasing abiotic stress. The shift occurs because species interactions varied with increasing abiotic stress. Competition prevails in low stress plots, while facilitation dominates in high stress plots. In low stress plots, the monoculture biomass of a specific species is a good indicator of the competitive ability of that species in the mixture, and the dominant species significantly affects the plot biomass. In high stress plots, the tolerance indexes of all individual species increase with the manipulated species richness, providing clear evidence for the increasing role of facilitation.     

Positive interactions can increase size inequality in plant populations

1.  Large variation in the size of individuals is a ubiquitous feature of natural plant populations. While the role of competition in generating this variation has been studied extensively, the potential effects of positive interactions among plants, which are common in high-stress environments, have not been investigated.
2.  Using an individual-based 'zone-of-influence' model, we investigate the effects of competition, abiotic stress and facilitation on size inequality in plant monocultures. In the model, stress reduces the growth rate of plants, and facilitation ameliorates the effects of stress. Both facilitation and competition occur in overlapping zones of influence. We tested some of the model's predictions with a field experiment using the clonal grass Elymus nutans in an alpine meadow.
3.  Facilitation increased the size inequality of model populations when there was no density-dependent mortality. This effect decreased with density as competition overwhelmed facilitation. The lowest size inequality was found at intermediate densities both with the model and in the field.
4.  When density-dependent mortality was included in the model, stress delayed its onset and reduced its rate by reducing growth rates, so the number of survivors at any point in time was higher under harsh than under more benign conditions. Facilitation increased size inequality during self-thinning.
5.   Synthesis . Our results demonstrate that facilitation interacts with abiotic stress and competition to influence the degree of size inequality in plant populations. Facilitation increased size inequality at low to intermediate densities and during self-thinning.     

Natural selection drives the fine-scale divergence of a coevolutionary arms race involving a long-mouthed weevil and its obligate host plant

Background

One of the major recent advances in evolutionary biology is the recognition that evolutionary interactions between species are substantially differentiated among geographic populations. To date, several authors have revealed natural selection pressures mediating the geographically-divergent processes of coevolution. How local, then, is the geographic structuring of natural selection in coevolutionary systems?

Results

I examined the spatial scale of a "geographic selection mosaic," focusing on a system involving a seed-predatory insect, the camellia weevil (Curculio camelliae), and its host plant, the Japanese camellia (Camellia japonica). In this system, female weevils excavate camellia fruits with their extremely-long mouthparts to lay eggs into seeds, while camellia seeds are protected by thick pericarps. Quantitative evaluation of natural selection demonstrated that thicker camellia pericarps are significantly favored in some, but not all, populations within a small island (Yakushima Island, Japan; diameter ca. 30 km). At the extreme, camellia populations separated by only several kilometers were subject to different selection pressures. Interestingly, in a population with the thickest pericarps, camellia individuals with intermediate pericarp thickness had relatively high fitness when the potential costs of producing thick pericarps were considered. Also importantly, some parameters of the weevil - camellia interaction such as the severity of seed infestation showed clines along temperature, suggesting the effects of climate on the fine-scale geographic differentiation of the coevolutionary processes.

Conclusion

These results show that natural selection can drive the geographic differentiation of interspecific interactions at surprisingly small spatial scales. Future studies should reveal the evolutionary/ecological outcomes of the "fine scale geographic mosaics" in biological communities.     

Facilitation can maintain clustered spatial pattern of plant populations during density‐dependent mortality: insights from a zone‐of‐influence model

Recent years have seen a growing body of evidence showing that plant competition and facilitation usually operate simultaneously to drive population dynamics, community structure and ecosystem functions. However, the potential role of facilitation in spatial patterning of plant populations has rarely been explicitly examined. We used a ‘zone‐of‐influence’ model to explore how facilitation interacts with competition and abiotic stress to determine the spatial patterning of populations during density‐dependent mortality. Model simulations revealed that started with the same clustered pattern, the final pattern of simulated populations depended strongly on the interaction among facilitation, stress level and size‐symmetry of competition. Asymmetric competition consistently led to immediate and non‐random mortality towards regularity, thus rapidly decayed the initially clustered pattern to final patterns of small‐scale regularity and large‐scale randomness. The role of symmetric competition in decaying the clustered pattern increased with abiotic stress because stress‐induced reductions in plants’ growth rates can make individuals in high‐density clusters more likely to die even from symmetric competition. Facilitation played a clear role in counteracting the effect of stress, thus tended to maintain the degree of clustering of the pattern during density‐dependent mortality. This is because the amelioration of harsh conditions by neighboring plants relieved the reductions in plant growth due to competition, thus slowed down and reduced the mortality inside clusters (relative to that outside clusters). Moreover, the effect of facilitation appeared to increase with abiotic stress. Our results indicate that facilitation among neighboring plants should partially be responsible for clustered population spatial patterns observed in stressful environments, even though its contribution relative to other factors (e.g. local dispersal and environmental heterogeneity) remains to be evaluated. In addition, the potential influence of facilitation on self‐thinning trajectory should be explicitly examined in future modeling and experimental studies considering its effects on density‐dependent mortality.     

Whole-community facilitation regulates biodiversity on Patagonian rocky shores

Background

Understanding the factors that generate and maintain biodiversity is a central goal in ecology. While positive species interactions (i.e., facilitation) have historically been underemphasized in ecological research, they are increasingly recognized as playing important roles in the evolution and maintenance of biodiversity. Dominant habitat-forming species (foundation species) buffer environmental conditions and can therefore facilitate myriad associated species. Theory predicts that facilitation will be the dominant community-structuring force under harsh environmental conditions, where organisms depend on shelter for survival and predation is diminished. Wind-swept, arid Patagonian rocky shores are one of the most desiccating intertidal rocky shores ever studied, providing an opportunity to test this theory and elucidate the context-dependency of facilitation.

Methodology/Principal Findings

Surveys across 2100 km of southern Argentinean coastline and experimental manipulations both supported theoretical predictions, with 43 out of 46 species in the animal assemblage obligated to living within the matrices of mussels for protection from potentially lethal desiccation stress and predators having no detectable impact on diversity.

Conclusions/Significance

These results provide the first experimental support of long-standing theoretical predictions and reveal that in extreme climates, maintenance of whole-community diversity can be maintained by positive interactions that ameliorate physical stress. These findings have important conservation implications and emphasize that preserving foundation species should be a priority in remediating the biodiversity consequences of global climate change.     

Plant–plant facilitation increases with reduced phylogenetic relatedness along an elevation gradient

Environmental conditions can modify the intensity and sign of ecological interactions. The stress gradient hypothesis (SGH) predicts that facilitation becomes more important than competition under stressful conditions. To properly test this hypothesis, it is necessary to account for all (not a subset of) interactions occurring in the communities and consider that species do not interact at random but following a specific pattern. We aim to assess elevational changes in facilitation, in terms of species richness, frequency and intensity of the interaction as a function of the evolutionary relatedness between nurses and their associated species. We sampled nurse and their facilitated plant species in two 1000–2000 m. elevation gradients in Mediterranean Chile where low temperature imposes a mortality filter on seedlings. We first estimated the relative importance of facilitation as a mechanism adding new species to communities distributed along these gradients. We then tested whether the frequency and intensity of facilitation increases with elevation, taking into account the evolutionary relatedness of the nurse species and the facilitated species. We found that nurses increase the species richness of the community by up to 35%. Facilitative interactions are more frequent than competitive interactions (56% versus 44%) and facilitation intensity increased with elevation for interactions involving distantly related lineages. Our results highlight the importance of including an evolutionary dimension in the study of facilitation to have a clearer picture of the mechanisms enabling species to coexist and survive under stressful conditions. This knowledge is especially relevant to conserve vulnerable and threatened communities facing new climate scenarios, such as those located in Mediterranean-type ecosystems.     

Effects of Competition and Facilitation on Species Assemblage in Two Types of Tropical Cloud Forest

Competition and facilitation between tree individuals are two kinds of non-random processes influencing the structure and functioning of forest communities, but how these two plant-plant interactions change along gradient of resources or environments remains very much a matter of debate. We developed a null model to test the size-distance regression, and assessed the effects of competition and facilitation (including interspecific interactions, intraspecific interactions and overall species interactions) on each adult tree species assemblage [diameter at breast height (dbh) ≥5 cm] across two types of tropical cloud forest with different environmental and resource regimes. The null model test revealed that 17% to 27% tree species had positive dbh-distance correlations while 11% to 19% tree species showed negative dbh-distance correlations within these two forest types, indicating that both competition and facilitation processes existed during the community assembly. The importance of competition for heterospecific species, and the intensity of competition for both heterospecific and overall species increased from high to low resources for all the shared species spanning the two forests. The importance of facilitation for conspecific and overall species, as well as that the intensity of facilitation for both heterospecific and conspecific species increased with increasing low air temperature stress for all the shared species spanning the two forests. Our results show that both competition and facilitation processes simultaneously affect parts of species assemblage in the tropical cloud forests. Moreover, the fact that nearly 50% species assemblage is not detected with our approaches suggest that tree species in these tropical forest systems are assembled with multiple ecological processes, and that there is a need to explore the processes other than the two biotic interactions in further researches.     

Multilevel and sex‐specific selection on competitive traits in North American red squirrels

Individuals often interact more closely with some members of the population (e.g., offspring, siblings, or group members) than they do with other individuals. This structuring of interactions can lead to multilevel natural selection, where traits expressed at the group‐level influence fitness alongside individual‐level traits. Such multilevel selection can alter evolutionary trajectories, yet is rarely quantified in the wild, especially for species that do not interact in clearly demarcated groups. We quantified multilevel natural selection on two traits, postnatal growth rate and birth date, in a population of North American red squirrels (Tamiasciurus hudsonicus). The strongest level of selection was typically within‐acoustic social neighborhoods (within 130 m of the nest), where growing faster and being born earlier than nearby litters was key, while selection on growth rate was also apparent both within‐litters and within‐study areas. Higher population densities increased the strength of selection for earlier breeding, but did not influence selection on growth rates. Females experienced especially strong selection on growth rate at the within‐litter level, possibly linked to the biased bequeathal of the maternal territory to daughters. Our results demonstrate the importance of considering multilevel and sex‐specific selection in wild species, including those that are territorial and sexually monomorphic.     

Shifts from competition to facilitation between a foundation tree and a pioneer shrub across spatial and temporal scales in a semiarid woodland

* Theoretical and empirical research has supported the hypothesis that plant-plant interactions change from competition to facilitation with increasing abiotic stress. However, the consistency of such changes has been questioned in arid and semiarid ecosystems. * During a drought in the semiarid south-western USA, we used observations and a field experiment to examine the interactions between juveniles of a foundation tree (Pinyon pine, Pinus edulis) and a common shrub (Apache plume, Fallugia paradoxa) in replicated areas of high and low stress. * The presence of F. paradoxa reduced P. edulis performance at low-stress sites, but had the opposite effect at high-stress sites. However, the intensity of the interactions depended on temporal variation in climate and age of P. edulis. Both above- and below-ground factors contributed to competition, while only above-ground factors contributed to facilitation. * These results support the hypothesis that interactions can change from competition to facilitation as abiotic stress increases in semiarid environments. A shift from competition to facilitation may be important for the recovery of P. edulis and other foundation species that have experienced large-scale mortality during recent droughts.     

Strain and vegetation effects on local limiting resources explain the outcomes of biotic interactions

Positive interactions are hypothesized to increase with stress (stress-gradient hypothesis, “SGH”), which is defined in terms of standing biomass at the community level. However, recent evidence suggests that facilitation may decrease or remain constant as stress increases. Several reasons for this discrepancy are possible: (i) the outcomes of biotic interactions depend on the component of the fitness considered; (ii) they are influenced by how vegetation affects local limiting resources; (iii) within a particular community, only species that are deviated from their physiological optima are likely to be facilitated. In a removal experiment, we quantified the deviations of species in subalpine grassland from their physiological optima, defined here as species-level “strain”, and examined whether strain and vegetation effects on local resources can explain the outcome of biotic interactions. The experiment was performed along a gradient of standing biomass driven by contrasting land use and resource availability, and used five grass species with contrasting traits and ecological optima. Strain for each species was estimated by comparing growth without vegetation (target species only submitted to local abiotic factors) to growth in optimal conditions (under controlled conditions in an experimental garden). The outcomes of biotic interactions, recorded in terms of survival and growth, could be predicted from the data about strain and vegetation effects on local limiting resources (light and water). Only highly strained species were affected by facilitation, which occurred when the surrounding vegetation alleviated the constraining factors. On the other hand, standing biomass was poorly related with the outcomes of biotic interactions. The “SGH” was only partially validated with growth data when strain and vegetation effects co-varied with standing biomass. As a consequence, strain (at species level) represents a mechanistic basis which could improve the prediction of the outcomes of biotic interactions along ecological gradients.