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  1. Abstract

    Trait‐based approaches have been extensively used in community ecology to provide a mechanistic understanding of the drivers of community assembly. However, a foundational assumption of the trait framework, traits relate to performance, has been mainly examined through univariate relationships that simplify the complex phenotypic integration of organisms. We evaluate a conceptual framework in which traits are organized hierarchically combining trait information at the individual‐ and species‐level from biomass allocation and organ‐level traits. We focus on photosynthetic traits and predict that the positive effects of increasing plant leaf mass on growth depend on species‐level leaf traits. We modeled growth data on more than 1,500 seedlings from 97 seedling species from a tropical forest in China. We found that seedling growth increases with allocation to leaves (high leaf area ratio and leaf mass fraction) and this effect is accentuated for species with high specific leaf area and leaf area. Also, we found that light has a significant effect on growth, and this effect is additive with leaf allocation traits. Our work offers an approach to gain further understanding of the effects of traits on the whole plant‐level growth via a hierarchical framework including organ‐level and biomass allocation traits at species and individual levels.

     
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  2. Abstract

    Plants allocate biomass to different organs in response to resource variation for maximizing performance, yet we lack a framework that adequately integrates plant responses to the simultaneous variation in above‐ and below‐ground resources. Although traditionally, the optimal partition theory (OPT) has explained patterns of biomass allocation in response to a single limiting resource, it is well‐known that in natural communities multiple resources limit growth. We study trade‐offs involved in plant biomass allocation patterns and their effects on plant growth under variable below‐ and above‐ground resources—light, soil N and P—for seedling communities.

    We collected information on leaf, stem and root mass fractions for more than 1,900 seedlings of 97 species paired with growth data and local‐scale variation in abiotic resources from a tropical forest in China.

    We identified two trade‐off axes that define the mass allocation strategies for seedlings—allocation to photosynthetic versus non‐photosynthetic tissues and allocation to roots over stems—that responded to the variation in soil P and N and light. Yet, the allocation patterns did not always follow predictions of OPT in which plants should allocate biomass to the organ that acquires the most limiting resource. Limited soil N resulted in high allocation to leaves at the expense of non‐photosynthetic tissues, while the opposite trend was found in response to limited soil P. Also, co‐limitation in above‐ and below‐ground resources (light and soil P) led to mass allocation to stems at the expense of roots. Finally, we found that growth increased under high‐light availability and soil P for seedlings that invested more in photosynthetic over non‐photosynthetic tissues or/and that allocated mass to roots at the expense of stem.

    Synthesis. Biomass allocation patterns to above‐ and below‐ground tissues are described by two independent trade‐offs that allow plants to have divergent allocation strategies (e.g. high root allocation at the expense of stem or high leaf allocation at the expense of allocation to non‐photosynthetic tissues) and enhance growth under different limiting resources. Identifying the trade‐offs driving biomass allocation is important to disentangle plant responses to the simultaneous variation in resources in diverse forest communities.

     
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  3. Tree performance depends on the coordinated functioning of interdependent leaves, stems and (mycorrhizal) roots. Integrating plant organs and their traits, therefore, provides a more complete understanding of tree performance than studying organs in isolation. Until recently, our limited understanding of root traits impeded such a whole‐tree perspective on performance, but recent developments in root ecology provide new impetuses for integrating the below‐ and aboveground. Here, we identify two key avenues to further develop a whole‐tree perspective on performance and highlight the conceptual and practical challenges and opportunities involved in including the belowground. First, traits of individual roots need to be scaled up to the root system as a whole to determine belowground functioning, e.g. total soil water and nutrient uptake, and hence performance. Second, above‐ and belowground plant organs need to be mechanistically connected to account for how they functionally interact and to investigate their combined impacts on tree performance. We further identify mycorrhizal symbiosis as the next frontier and emphasize several courses of actions to incorporate these symbionts in whole‐tree frameworks. By scaling up and mechanistically integrating (mycorrhizal) roots as argued here, the belowground can be better represented in whole‐tree conceptual and mechanistic models; ultimately, this will improve our estimates of not only the functioning and performance of individual trees, but also the processes and responses to environmental change of the communities and ecosystems they are part of. 
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