Increased Size Inequality Research Articles

Plant growth-promoting rhizobacteria promote plant size inequality

The uniformity of crop yield is extremely important for consumers and of as much relevance to the grower as overall yield. However, size inequality within a plant population is rarely measured and has never before been considered in relation to the use of beneficial microbes for yield enhancement. For the first time, we show that addition of soil bacteria to calabrese plants significantly increased size inequality. These effects were usually more apparent in above-ground biomass. This was caused by some (but not all) plants growing very large when inoculated with bacteria, while control plants were mostly small. We suggest that the main reason is the incompatibility of the inoculated bacteria with those already present in the rhizosphere. In some cases the inoculum matched the indigenous community, providing a benefit to plant growth, while often it did not and plants remained relatively small. We conclude that analyses of size inequality should be an integral part of experiments using microbial soil amendments. These analyses can help to inform the production of more effective microbial products and to ensure that the integration of beneficial microbes into sustainable production systems does not impair uniformity in yield.

Relationship between size inequality and stand productivity is modified by self-thinning, age, site and planting density in Sassafras tzumu plantations in central China

Greater size inequality has been associated with lower forest productivity. Density-dependent mortality can reduce size inequality and potentially modify relationships between size inequality and productivity. We examined the effect of density-dependent mortality, age, planting density and site quality on this relationship using two Sassafras tzumu plantations in central China with three planting density levels (4.0 m × 5.0 m, 3.0 m × 4.0 m, and 2.0 m × 3.0 m) replicated at two sites. The stands were measured in winter every other year for 20 years, from 1996 to 2016. Generalized linear mixed effect models were used to examine the relationship between stand age, planting density, site quality, and size inequality (assessed using the Gini coefficient). Gini coefficients, and hence size inequality, increased with stand age, planting density and site quality. The onset of self-thinning reduced the rate at which size inequality increased. Productivity was negatively correlated with size inequality, and self-thinning modified this relationship such that productivity declined at a greater rate (with increasing size inequality) after self-thinning had begun to occur. The productivity-size inequality relationship was also modified by planting density, age and site quality, such that with all other things being equal, the negative effect of size inequality is greatest in young stands, dense stands and on low quality sites. The manipulation of stand size inequality (e.g. by thinning or mortality) influences growth more than simply by changing stand density. An understanding of the mechanisms driving the effect of density-dependent mortality on the relationship between productivity and size inequality may facilitate the further development of silvicultural regimes.

Effect of vegetation control, harvest intensity, and soil disturbance on 20-year jack pine stand development

Currently, there are uncertainties regarding the impacts and (or) efficacy of biomass harvesting and silvicultural practices on stand production on coarse-textured boreal soils. Replicated factorial field experiments examining effects of complete vegetation control (repeated glyphosate application) following operational stem-only harvest with disc trenching (SOT), operational whole-tree harvest with (WTT) and without (WT) disc trenching, and whole-tree harvest with complete forest floor removal by blading (WTB) and blading followed by compaction (WTBC) were installed on four sandy northern Ontario jack pine (Pinus banksiana Lamb.) sites. Over 20 years, WTB improved planted-tree survival but decreased longer term stand productivity in comparison with other harvest intensity – soil disturbance treatments. Vegetation control improved tree growth and stand productivity initially, but over time, benefits declined substantially. SOT and WTT had similar impacts on stand production. Disc trenching improved initial planted-tree growth (WTT vs. WT), particularly without vegetation control. Jack pine natural regeneration was greatest with SOT, accounting for 25% of stand biomass at year 20. Stand structure effects included increased size inequality of naturals with WTB and reduced size inequality and asymmetry of naturals with vegetation control. Overall, impacts of forest floor removal and natural regeneration on stand development have become increasingly important over time compared with those of vegetation control.

Modelling the effect of size-asymmetric competition on size inequality: Simple models with two plants

The concept of size asymmetry in resource competition among plants, in which larger individuals obtain a disproportionate share of contested resources, appears to be very straightforward, but the effects of size asymmetry on growth and size variation among individuals have proved to be controversial. It has often been assumed that competition among individual plants in a population has to be size-asymmetric to result in higher size inequality than in the absence of competition, but here we question this inference. Using very simple, individual-based models, we investigate how size symmetry of competition affects the development in size inequality between two competing plants and show that increased size inequality due to competition is not always strong evidence for size-asymmetric competition. Even absolute symmetric competition, in which all plants receive the same amount of resources irrespective of their sizes, can, under some assumptions, result in higher size inequality than when competition is absent. We demonstrate our approach by applying it to data from a greenhouse experiment investigating the size symmetry of belowground competition between pairs of Triticum aestivum (wheat) plants. The effects of size symmetry/asymmetry on size inequality are dependent on (1) the individual plant growth model, (2) the parameters of the growth model that are affected by competition and (3) the initial sizes and growth rates. Across a range of reasonable assumptions, very general patterns that have been considered evidence for or against size-asymmetric competition do not always hold. Our results emphasize the need for explicit growth models, even very simple ones, for making inferences about the effects of competition on plant growth and size inequality.

Common mycorrhizal networks amplify competition by preferential mineral nutrient allocation to large host plants.

Arbuscular mycorrhizal (AM) fungi interconnect plants in common mycorrhizal networks (CMNs) which can amplify competition among neighbors. Amplified competition might result from the fungi supplying mineral nutrients preferentially to hosts that abundantly provide fixed carbon, as suggested by research with organ-cultured roots. We examined whether CMNs supplied (15) N preferentially to large, nonshaded, whole plants. We conducted an intraspecific target-neighbor pot experiment with Andropogon gerardii and several AM fungi in intact, severed or prevented CMNs. Neighbors were supplied (15) N, and half of the target plants were shaded. Intact CMNs increased target dry weight (DW), intensified competition and increased size inequality. Shading decreased target weight, but shaded plants in intact CMNs had mycorrhizal colonization similar to that of sunlit plants. AM fungi in intact CMNs acquired (15) N from the substrate of neighbors and preferentially allocated it to sunlit, large, target plants. Sunlit, intact CMN, target plants acquired as much as 27% of their nitrogen from the vicinity of their neighbors, but shaded targets did not. These results suggest that AM fungi in CMNs preferentially provide mineral nutrients to those conspecific host individuals best able to provide them with fixed carbon or representing the strongest sinks, thereby potentially amplifying asymmetric competition below ground.

Do biodiversity indices behave as expected from traits of constituent species in simulated scenarios?

It is believed that diversity of plant communities has a positive effect on their productivity. The benefits of diversity are described by “biodiversity indices”, comparing yield of mixtures with yields of monocultures of constituent species. These indices are supposed to capture also the main mechanisms leading to increased yield. We have constructed a spatially explicit individual based model, simulating even-aged stand development, and compared the behaviour of selected biodiversity indices (overyielding, selectivity and complementarity) with expectations based on life history traits of constituent species. The results are based on comparisons of two species mixtures with corresponding monocultures. We designed three scenarios of changes in the two species life history differentiation, and compared the behaviour of the indices with expectation based on it. In the first scenario, selectivity was driven by increasing size inequality of the two species, mostly in accordance with expectations. The second scenario presents increasing shade tolerance of the smaller species that increased complementarity, again mostly as expected. In the last scenario, shortening of length of stress tolerance of the weaker species surprisingly increased values of the biodiversity indices. For each setting, we varied sowing density and spatial pattern of the constituent species. The behaviour of the indices was influenced by both factors, but the effect of density was more pronounced. In particular, at high sowing densities, the most important interactions happened in the very early stages of mixture development, and the behaviour of the indices was often counter-intuitive.

Kin recognition, not competitive interactions, predicts root allocation in young Cakile edentula seedling pairs

• Recent studies have demonstrated sibling vs stranger differences in group root allocation in plants, suggesting that plants have the potential for kin discrimination in competition. However, morphology differences could potentially be generated by competition-based mechanisms. Here, we tested these hypotheses for the sibling vs stranger differences in root allocation in Cakile edentula. • Seeds were planted in pairs of either kin (siblings) or strangers, from all combinations of eight families, to give eight kin (sibling) and 28 stranger pair identities. Because the species has a seed dimorphism, the 10 replicates of each pair identity included both seed types. Root allocation, size inequality between seedlings in a pair, and competitive ability were derived from measures of biomass and height. • Cakile edentula seedlings demonstrated the same kin recognition response previously observed in juvenile plants, with lower root allocation in kin pairs than stranger pairs. The seed dimorphism was not associated with root allocation. • The two competitive mechanisms, genetic differences in competitive ability and increased size inequality in stranger groups, did not explain the root allocation differences in these seedlings. Kin recognition offered the most probable explanation for the differences in root allocation between sibling and stranger pairs.

Positive interactions can increase size inequality in plant populations

Summary1. 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.

Plant responses to elevated CO2 concentration at different scales: leaf, whole plant, canopy, and population

AbstractElevated CO2 enhances photosynthesis and growth of plants, but the enhancement is strongly influenced by the availability of nitrogen. In this article, we summarise our studies on plant responses to elevated CO2. The photosynthetic capacity of leaves depends not only on leaf nitrogen content but also on nitrogen partitioning within a leaf. In Polygonum cuspidatum, nitrogen partitioning among the photosynthetic components was not influenced by elevated CO2 but changed between seasons. Since the alteration in nitrogen partitioning resulted in different CO2‐dependence of photosynthetic rates, enhancement of photosynthesis by elevated CO2 was greater in autumn than in summer. Leaf mass per unit area (LMA) increases in plants grown at elevated CO2. This increase was considered to have resulted from the accumulation of carbohydrates not used for plant growth. With a sensitive analysis of a growth model, however, we suggested that the increase in LMA is advantageous for growth at elevated CO2 by compensating for the reduction in leaf nitrogen concentration per unit mass. Enhancement of reproductive yield by elevated CO2 is often smaller than that expected from vegetative growth. In Xanthium canadense, elevated CO2 did not increase seed production, though the vegetative growth increased by 53%. As nitrogen concentration of seeds remained constant at different CO2 levels, we suggest that the availability of nitrogen limited seed production at elevated CO2 levels. We found that leaf area development of plant canopy was strongly constrained by the availability of nitrogen rather than by CO2. In a rice field cultivated at free‐air CO2 enrichment, the leaf area index (LAI) increased with an increase in nitrogen availability but did not change with CO2 elevation. We determined optimal LAI to maximise canopy photosynthesis and demonstrated that enhancement of canopy photosynthesis by elevated CO2 was larger at high than at low nitrogen availability. We also studied competitive asymmetry among individuals in an even‐aged, monospecific stand at elevated CO2. Light acquisition (acquired light per unit aboveground mass) and utilisation (photosynthesis per unit acquired light) were calculated for each individual in the stand. Elevated CO2 enhanced photosynthesis and growth of tall dominants, which reduced the light availability for shorter subordinates and consequently increased size inequality in the stand.

Asymmetric, two-sided competition in even-aged monocultures of red pine

A framework for analysis of even-aged plant monocultures was developed and used to examine hypotheses on the process of competition, using data obtained from three independently established red pine (Pinusresinosa Ait.) stand-density experiments. Growth rate and relative production rate (a measure of tree vigour) were computed for the individual trees in stands of differing density at points in time when these stands had the same mean stem volume. Competition is shown to be both asymmetric (small trees lose vigour more rapidly than large trees as density increases) and two-sided (not only do larger trees affect smaller trees, but to some degree small trees also cause growth reduction in larger neighbours). Increasing density results in increased size inequality as smaller trees lose vigour and become suppressed. The asymmetric, two-sided competition process in red pine stands is considered ecologically inherent for a species adapted to existence in even-aged monocultures.