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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.more » « less
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Abstract Background and Aims Understanding shifts in the demographic and functional composition of forests after major natural disturbances has become increasingly relevant given the accelerating rates of climate change and elevated frequency of natural disturbances. Although plant demographic strategies are often described across a slow–fast continuum, severe and frequent disturbance events influencing demographic processes may alter the demographic trade-offs and the functional composition of forests. We examined demographic trade-offs and the shifts in functional traits in a hurricane-disturbed forest using long-term data from the Luquillo Forest Dynamics Plot (LFPD) in Puerto Rico.
Methods We analysed information on growth, survival, seed rain and seedling recruitment for 30 woody species in the LFDP. In addition, we compiled data on leaf, seed and wood functional traits that capture the main ecological strategies for plants. We used this information to identify the main axes of demographic variation for this forest community and evaluate shifts in community-weighted means for traits from 2000 to 2016.
Key Results The previously identified growth–survival trade-off was not observed. Instead, we identified a fecundity–growth trade-off and an axis representing seedling-to-adult survival. Both axes formed dimensions independent of resprouting ability. Also, changes in tree species composition during the post-hurricane period reflected a directional shift from seedling and tree communities dominated by acquisitive towards conservative leaf economics traits and large seed mass. Wood specific gravity, however, did not show significant directional changes over time.
Conclusions Our study demonstrates that tree demographic strategies coping with frequent storms and hurricane disturbances deviate from strategies typically observed in undisturbed forests, yet the shifts in functional composition still conform to the expected changes from acquisitive to conservative resource-uptake strategies expected over succession. In the face of increased rates of natural and anthropogenic disturbance in tropical regions, our results anticipate shifts in species demographic trade-offs and different functional dimensions.
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Predicting drought responses of individual trees in tropical forests remains challenging, in part because trees experience drought differently depending on their position in spatially heterogeneous environments. Specifically, topography and the competitive environment can influence the severity of water stress experienced by individual trees, leading to individual-level variation in drought impacts. A drought in 2015 in Puerto Rico provided the opportunity to assess how drought response varies with topography and neighborhood crowding in a tropical forest. In this study, we integrated 3 years of annual census data from the El Yunque Chronosequence plots with measurements of functional traits and LiDAR-derived metrics of microsite topography. We fit hierarchical Bayesian models to examine how drought, microtopography, and neighborhood crowding influence individual tree growth and survival, and the role functional traits play in mediating species’ responses to these drivers. We found that while growth was lower during the drought year, drought had no effect on survival, suggesting that these forests are fairly resilient to a single-year drought. However, growth response to drought, as well as average growth and survival, varied with topography: tree growth in valley-like microsites was more negatively affected by drought, and survival was lower on steeper slopes while growth was higher in valleys. Neighborhood crowding reduced growth and increased survival, but these effects did not vary between drought/non-drought years. Functional traits provided some insight into mechanisms by which drought and topography affected growth and survival. For example, trees with high specific leaf area grew more slowly on steeper slopes, and high wood density trees were less sensitive to drought. However, the relationships between functional traits and response to drought and topography were weak overall. Species sorting across microtopography may drive observed relationships between average performance, drought response, and topography. Our results suggest that understanding species’ responses to drought requires consideration of the microenvironments in which they grow. Complex interactions between regional climate, topography, and traits underlie individual and species variation in drought response.more » « less
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Abstract Individual‐level demographic outcomes should be predictable upon the basis of traits. However, linking traits to tree performance has proven challenging likely due to a failure to consider physiological traits (i.e. hard‐traits) and the failure to integrate organ‐level and whole plant‐level trait information.
Here, we modelled the survival rate and relative growth rate of trees while considering crown allocation, hard‐traits and local‐scale biotic interactions, and compared these models to more traditional trait‐based models of tree performance.
We found that an integrative trait, total tree‐level photosynthetic mass (estimated by multiplying specific leaf area and crown area) results in superior models of tree survival and growth. These models had a lower AIC than those including the effect of initial tree size or any other combination of the traits considered. Survival rates were positively related to higher values of crown area and photosynthetic mass, while relative growth rates were negatively related to the photosynthetic mass. Relative growth rates were negatively related to a neighbourhood crowding index. Furthermore, none of the hard‐traits used in this study provided an improvement in tree performance models.
Synthesis . Overall, our results highlight that models of tree performance can be greatly improved by including crown area information to generate a better understanding of plant responses to their environment. Additionally, the role of the hard‐traits in improving models of tree performance is likely dependent upon the level of stress (e.g. drought stress), micro‐environmental conditions or short‐term climatic variations that a particular forest experiences. -
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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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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.
