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Theory questions the persistence of nonreciprocal interactions in which one plant has a positive net effect on a neighbor that, in return, has a negative net impact on its benefactor - a phenomenon known as antagonistic facilitation. We develop a spatially explicit consumer-resource model for belowground plant competition between ecosystem engineers, plants able to mine resources and make them available for any other plant in the community, and exploiters. We use the model to determine in what environmental conditions antagonistic facilitation via soil-resource engineering emerges as an optimal strategy. Antagonistic facilitation emerges in stressful environments where ecosystem engineers' self-benefits from mining resources outweigh the competition with opportunistic neighbors. Among all potential causes of stress considered in the model, the key environmental parameter driving changes in the interaction between plants is the proportion of the resource that becomes readily available for plant consumption in the absence of any mining activity. Our results align with theories of primary succession and the stress gradient hypothesis. However, we find that the total root biomass and its spatial allocation through the root system, often used to measure the sign of the interaction between plants, do not predict facilitation reliably.

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The allocation pattern of plant biomass presents the strategy of the plant community to adopt environmental changes, while the driver of biomass allocation is still unclear in degraded alpine grassland ecosystems. To explore the issue, this study investigated the shoot-to-root (R/S) ratio, plant aboveground traits, and root competition of three functional groups (i.e., grasses, sedges, and forbs) at three degradation levels (i.e., no obvious degradation, ND; moderate degradation, MD; and severe degradation, SD) in an alpine meadow in the eastern Qinghai-Tibetan Plateau. The relationships among plant aboveground traits, root competition, and R/S ratio were tested using the structural equation model (SEM). The results showed that the shoot and root biomass tended to decrease, but the R/S ratio of the plant community did not change along the degradation gradient. Plant height, lateral spread, and leaf length of most plant functional groups reduced, while leaf width and leaf area of most plant functional groups did not change along the degradation gradients. The root competition ability (presented as the fraction of root biomass in total biomass) of sedges in MD was the lowest, while that of grasses was the highest. The effects of aboveground competition on the R/S ratio were non-linear because of the different roles of plant height, lateral spread, and leaf area in regulating the R/S ratio along the degradation gradient. In contrast, the effects of belowground competition on the R/S ratio were linear because belowground competition promoted the R/S ratio, and the strength of this effect reduced along the degradation gradient. These results indicate that plant competition might be a critical factor to maintain the high R/S ratio in degraded alpine meadows.

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Root competition is a key factor determining plant performance, community structure and ecosystem productivity. To adequately estimate the extent of root proliferation of plants in response to neighbours independently of nutrient availability, one should use a set-up that can simultaneously control for both nutrient concentration and soil volume at plant individual level. With a mesh-divider design, which was suggested as a promising solution for this problem, we conducted two intraspecific root competition experiments: one with soybean (Glycine max) and the other with sunflower (Helianthus annuus). We found no response of root growth or biomass allocation to intraspecific neighbours, i.e. an 'ideal free distribution' (IFD) norm, in soybean; and even a reduced growth as a negative response in sunflower. These responses are all inconsistent with the hypothesis that plants should produce more roots even at the expense of reduced fitness in response to neighbours, i.e. root over-proliferation. Our results suggest that neighbour-induced root over-proliferation is not a ubiquitous feature in plants. By integrating the findings with results from other soybean studies, we conclude that for some species this response could be a genotype-dependent response as a result of natural or artificial selection, or a context-dependent response so that plants can switch from root over-proliferation to IFD depending on the environment of competition. We also critically discuss whether the mesh-divider design is an ideal solution for root competition experiments.

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Farmers expect yield reduction of cash crops like cocoa when growing in agroforestry systems compared to monocultures, due to competition for resources, e.g. nutrients and water. However, complementarities between species in the use of resources may improve resource use efficiency and result in higher system performance. Cocoa trees have a shallow rooting system while the rooting characteristics of the associated trees are mainly unknown. This work investigates fine root distribution and production in five cocoa production systems: two monocultures and two agroforestry systems under conventional and organic farming, and a successional agroforestry system. In the organic systems a perennial leguminous cover crop was planted and compost was added, while herbicides and chemical fertilizers were applied in the conventional ones. We measured cocoa fine root parameters in the top 10cm of soil and annual total fine root production at 0-25 and 25-50cm depth. We related the root data with both the aboveground performance (tree and herbaceous biomass), and the cocoa and system yields. Cocoa fine roots were homogenously distributed over the plot area. Around 80% of the total fine roots were located in the upper 25cm of soil. The total fine root production was 4-times higher in the agroforestry systems and the organic monoculture than in the conventional monoculture. The roots of the associated tree species were located in the same soil space as the cocoa roots and, in principle, competed for the same soil resources. The cocoa yield was lower in the agroforestry systems, but the additional crops generated a higher system yield and aboveground biomass than the conventional cocoa monocultures, implying effective resource exploitation. The leguminous cover crop in the organic monoculture competed with the cocoa trees for nutrients, which may explain the lower cocoa yield in this system in contrast with the conventional monoculture.

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There is little direct evidence for effects of soil heterogeneity and root plasticity on the competitive interactions among plants. In this study, we experimentally examined the impacts of temporal nutrient heterogeneity on root growth and interactions between two plant species with very different rooting strategies: Liquidambar styraciflua (sweet gum), which shows high root plasticity in response to soil nutrient heterogeneity, and Pinus taeda (loblolly pine), a species with less plastic roots. Seedlings of the two species were grown in sandboxes in inter- and intraspecific combinations. Nutrients were applied in a patch either in a stable (slow-release) or in a variable (pulse) manner. Plant aboveground biomass, fine root mass, root allocation between nutrient patch and outside the patch, and root vertical distribution were measured. L. styraciflua grew more aboveground (40% and 27% in stable and variable nutrient treatment, respectively) and fine roots (41% and 8% in stable and variable nutrient treatment, respectively) when competing with P. taeda than when competing with a conspecific individual, but the growth of P. taeda was not changed by competition from L. styraciflua. Temporal variation in patch nutrient level had little effect on the species' competitive interactions. The more flexible L. styraciflua changed its vertical distribution of fine roots in response to competition from P. taeda, growing more roots in deeper soil layers compared to its roots in conspecific competition, leading to niche differentiation between the species, while the fine root distribution of P. taeda remained unchanged across all treatments. Synthesis. L. styraciflua showed greater flexibility in root growth by changing its root vertical distribution and occupying space of not occupied by P. taeda. This flexibility gave L. styraciflua an advantage in interspecific competition.

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Understanding how forested ecosystems respond to climatic changes is a challenging problem as forest self-organization occurs simultaneously across multiple scales. Here, we explore the hypothesis that soil water availability shapes above-ground competition and gap dynamics, and ultimately alters the dominance of shade tolerant and intolerant species along the moisture gradient. We adapt a spatially explicit individual-based model with simultaneous crown and root competitions. Simulations show that the transition from xeric to mesic soils is accompanied by an increase in shade-tolerant species similar to the patterns documented in the North American forests. This transition is accompanied by a change from water to sunlight competitions, and happens at three successive stages: (i) mostly water-limited parkland, (ii) simultaneously water- and sunlight-limited closed canopy forests featuring a very sparse understory, and (iii) mostly sunlight-limited forests with a populated understory. This pattern is caused by contrasting successional dynamics that favour either shade-tolerant or shade-intolerant species, depending on soil moisture and understory density. This work demonstrates that forest patterns along environmental gradients can emerge from spatial competition without physiological trade-offs between shade and growth tolerance. Mechanistic understanding of population processes involved in the forest-parkland-desert transition will improve our ability to explain species distributions and predict forest responses to climatic changes.

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Plant-soil feedback is receiving increasing interest as a factor influencing plant competition and species coexistence in grasslands. However, we do not know how spatial distribution of plant-soil feedback affects plant below-ground interactions. We investigated the way in which spatial heterogeneity of soil biota affects competitive interactions in grassland plant species. We performed a pairwise competition experiment combined with heterogeneous distribution of soil biota using four grassland plant species and their soil biota. Patches were applied as quadrants of 'own' and 'foreign' soils from all plant species in all pairwise combinations. To evaluate interspecific root responses, species-specific root biomass was quantified using real-time PCR. All plant species suffered negative soil feedback, but strength was species-specific, reflected by a decrease in root growth in own compared with foreign soil. Reduction in root growth in own patches by the superior plant competitor provided opportunities for inferior competitors to increase root biomass in these patches. These patterns did not cascade into above-ground effects during our experiment. We show that root distributions can be determined by spatial heterogeneity of soil biota, affecting plant below-ground competitive interactions. Thus, spatial heterogeneity of soil biota may contribute to plant species coexistence in species-rich grasslands.

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Evidence suggests that belowground self-recognition in clonal plants can be disrupted between sister ramets by the loss of connections or long distances within a genet. However, these results may be confounded by severing connections between ramets in the setups. Using Potentilla reptans, we examined severance effects in a setup that grew ramet pairs with connections either intact or severed. We showed that severance generally reduced new stolon mass but had no effect on root allocation of ramets. However, it did reduce root mass of younger ramets of the pairs. We also explored evidence for physiological self-recognition with another setup that avoided severing connections by manipulating root interactions between closely connected ramets, between remotely connected ramets and between disconnected ramets within one genet. We found that ramets grown with disconnected neighbors had less new stolon mass, similar root mass but higher root allocation as compared to ramets grown with connected neighbors. There was no difference in ramet growth between closely connected- and remotely connected-neighbor treatments. We suggest that severing connections affects ramet interactions by disrupting their physiological integration. Using the second setup, we provide unbiased evidence for physiological self-recognition, while also suggesting that it can persist over long distances.

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Worldwide, exotic invasive grasses have caused numerous ecosystem perturbations. Rangelands of the western USA have experienced increases in the size and frequency of wildfires largely due to invasion by the annual grass Bromus tectorum. Rehabilitation of invaded rangelands is difficult; but long-term success is predicated on establishing healthy and dense perennial grass communities, which suppress B. tectorum. This paper reports on two experiments to increase our understanding of soil factors involved in suppression. Water was not limiting in this study. Growth of B. tectorum in soil conditioned by and competing with the exotic perennial Agropyron cristatum was far less relative to its growth without competition. When competing with A. cristatum, replacing a portion of conditioned soil with fresh soil before sowing of B. tectorum did not significantly increase its growth. The ability of conditioned soil to suppress B. tectorum was lost when it was separated from growing A. cristatum. Soil that suppressed B. tectorum growth was characterized by low mineral nitrogen (N) availability and a high molar ratio of [Formula: see text] in the solution-phase pool of [Formula: see text] Moreover, resin availability of [Formula: see text] explained 66 % of the variability in B. tectorum above-ground mass, attesting to the importance of A. cristatum growth in reducing N availability to B. tectorum. Trials in which B. tectorum was suppressed the most were characterized by very high shoot/root mass ratios and roots that have less root hair growth relative to non-suppressed counterparts, suggesting co-opting of biological soil space by the perennial grass as another suppressive mechanism. Greater understanding of the role of biological soil space could be used to breed and select plant materials with traits that are more suppressive to invasive annual grasses.

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Root-root interaction research gained more and more attention over the past few years. Roots are pivotal for plant survival because they ensure uptake of water and nutrients. Therefore, detection of adjacent roots might lead to competitive advantages. Several lines of experimental evidence suggest that roots have ways to discriminate non-related roots, kin, and-importantly-that they can sense self/non-self roots to avoid intra-plant competition. In this mini-review, the existence of self/non-self recognition in plant roots will be discussed and the current knowledge on the mechanisms that could be involved will be summarized. Although the process of identity recognition is still not completely understood, interesting data are available and emerging new technologies will certainly aid to better understand this research field that can have an important biological, ecological, and agricultural impact.

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BACKGROUND AND AIMS: Considerable variation in seed size commonly exists within plants, and is believed to be favoured under natural selection. This study aims to examine the extent to which seed size distribution depends on the presence of competing neighbour plants. METHODS: Phaseolus vulgaris plants rooting with or without a conspecific neighbour were grown in soil with high or low nutrient availability. Seeds were harvested at the end of the growth cycle, the total nitrogen and phosphorus invested in seed production were measured and within-plant seed size distribution was quantified using a set of statistical descriptors. KEY RESULTS: Exposure to neighbours' roots induced significant changes in seed size distribution. Plants produced proportionally more large seeds and fewer small ones, as reflected by significant increases in minimal seed size, mean seed size, skewness and Lorenz asymmetry coefficient. These effects were different from, and in several cases opposite to, the responses when the soil nutrient level was reduced, and were significant after correction for the amount of resources invested in seed production. CONCLUSIONS: Below-ground neighbour presence affects within-plant seed size distribution in P. vulgaris. This effect appears to be non-resource-mediated, i.e. to be independent of neighbour-induced effects on resource availability. It implies that, based on current environmental cues, plants can make an anticipatory adjustment of their investment strategy in offspring as an adaptation to the local environment in the future.

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The effects of long-term carbon dioxide enrichment on competition for nutrients and light in a ryegrass/clover association were determined for simulated swards of perennial ryegrass (Lolium perenne L. cv. Parcour) and white clover (Trifolium repens L. cv. Karina), which were grown as monocultures and in three mixtures (25/75, 50/50, 75/25), according to the replacement design, at two levels of nitrogen (N) supply (no additional N and 200 kg N ha-1 ) and at season-long ambient (380 ppm) and elevated (670 ppm) CO2 concentrations, in open-top chambers. Stands were cut four times, at about monthly intervals, to a height of 5 cm. Plant material was separated into different species, fresh and dry weights were determined and the content of macroelements (N, P, K, S, Mg) in both species was measured. In addition, plant height of both species at harvest dates and during several regrowth periods was monitored. Results indicate that both species made demand on different resources and profited from growth in a mixed sward. Co2 related yield increase amounted to 16-4-2 % for white clover whereas the effect of high CO2 on ryegrass yield ranged between -33% and +9% depending on N supply, mixture and year. As a result the contribution of white clover to total yield in mixed swards was significantly enhanced by CO2 enrichment at many harvests in both N supply treatments. Without additional N supply, shoot competition for light was intensified by CO2 enrichment to the disadvantage of ryegrass, since clover petioles grew longer and ryegrass was shorter at elevated CO2 With N fertilization, no marked effect of CO2 enrichment on interspecific competition could be observed. Since clover and total yield were increased by CO2 enrichment, nutrient requirements were also increased and potassium deficiency and increased intraspecific competition of clover for K was observed in the mixtures under elevated CO2 which had the highest nutrient withdrawal. Although white clover profited much more from CO2 enrichment in both N fertilization treatments, the suppression of ryegrass in mixed swards could only be observed under low N conditions. Generally, the effect of N fertilization on competitive interference between both species was much greater than the effect of CO2 enrichment and it is suggested that the effect of elevated CO2 on the balance of species and the outcome of competition in a grass/clover sward is mainly dependent on the N status.

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Plant demographic and root exclusion approaches were used to examine the influence of roots of adult Artemisia tridentata, Agropyron desertorum, and Agropyron spicatum individuals on seedling survival of four C3 semiarid species, three perennials, Ar. tridentata, Ag. desertorum, Ag. spicatum, and an annual, Bromus tectorum. Furthermore, height of Ar. tridentata seedlings and seed production of B. tectorum were assessed. The probability of a seedling being alive significantly depended on the seedling species, the neighboring adult species, and on the depth to which root competition was excluded. As seedlings, survival of Agropyron species did not differ, whereas survival of Ar. tridentata seedlings was higher than Ag. desertorum and was similar to Ag. spicatum. Bromus tectorum maintained significantly higher survival rates than perennial seedlings. Established individuals of Ar. tridentata reduced seedling survival more than established individuals of either Agropyron species. Seedling survival significantly increased with greater depth of root exclusion for the perennials but did not significantly affect seedling survival of B. tectorum. Height of Ar. tridentata seedlings and seed production of B. tectorum significantly increased with depth of root exclusion. Seed production of B. tectorum was highest when competing with Ag. desertorum and was lowest with Ar. tridentata. Root competition decreased the seed population of B. tectorum in the next generation even though it had no impact on survival. Competition in the upper soil horizon occurs between seedlings and established adults early in the growing season and potentially restricts root growth of seedlings. In arid and semiarid ecosystems, soil moisture is depleted from the upper horizons first, resulting in the death of seedlings that do not have access to moisture.