The root economics spectrum (RES) hypothesis predicts that fast‐growing tree species have short‐lived roots with high specific root length (SRL) to allow rapid resource uptake, and opposite trait expressions for slow‐growing species. Yet, the mixed support for this hypothesis suggests that trees can adopt alternative strategies to increase resource uptake, besides an increase in SRL. We combined a novel mechanistic whole‐tree model and empirical fine‐root data of 10 tree species to test the effects of one of these alternative strategies, notably increasing fine‐root mass, on the tree's net C gain (used here as a proxy for tree performance), and to assess how fine‐root life span influences the relative importance of SRL and fine‐root mass for the C balance of trees. Our results indicate that accounting for the short life span of high‐SRL roots has important implications for explaining tree performance and the role of roots herein. Without considering their faster turnover, high‐SRL roots and low fine‐root mass resulted in the highest performance as predicted from the RES. Yet, when their higher turnover rates were accounted for, a high fine‐root mass and low SRL lead to the highest performance. Both our model outcomes and field data further show a negative relationship between SRL and fine‐root mass through which species aim to realize a similar root length density. This trade‐off further indicates how high a SRL and low fine‐root mass as well as opposite trait values can both lead to a positive C balance in a similar environment. Our study may explain why high‐SRL roots do not necessarily lead to the fastest tree growth as often hypothesized and demonstrates the importance of fine‐root mass in combination with fine‐root life span for explaining interspecific differences in tree performance. More generally, our work demonstrates the value of identifying and investigating different below‐ground strategies across species from a whole‐plant modelling perspective, and identifies the relationship between SRL, fine‐root biomass and life span as an important functional dimension to variation in species’ performance. A free Plain Language Summary can be found within the Supporting Information of this article.

Background and AimsPlants usually compete with neighbouring plants for resources such as light as well as defend themselves against herbivorous insects. This requires investment of limiting resources, resulting in optimal resource distribution patterns and trade-offs between growth- and defence-related traits. A plant’s competitive success is determined by the spatial distribution of its resources in the canopy. The spatial distribution of herbivory in the canopy in turn differs between herbivore species as the level of herbivore specialization determines their response to the distribution of resources and defences in the canopy. Here, we investigated to what extent competition for light affects plant susceptibility to herbivores with different feeding preferences.MethodsTo quantify interactions between herbivory and competition, we developed and evaluated a 3-D spatially explicit functional–structural plant model for Brassica nigra that mechanistically simulates competition in a dynamic light environment, and also explicitly models leaf area removal by herbivores with different feeding preferences. With this novel approach, we can quantitatively explore the extent to which herbivore feeding location and light competition interact in their effect on plant performance.Key ResultsOur results indicate that there is indeed a strong interaction between levels of plant–plant competition and herbivore feeding preference. When plants did not compete, herbivory had relatively small effects irrespective of feeding preference. Conversely, when plants competed, herbivores with a preference for young leaves had a strong negative effect on the competitiveness and subsequent performance of the plant, whereas herbivores with a preference for old leaves did not.ConclusionsOur study predicts how plant susceptibility to herbivory depends on the composition of the herbivore community and the level of plant competition, and highlights the importance of considering the full range of dynamics in plant–plant–herbivore interactions.

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Plants defend themselves against diverse communities of herbivorous insects. This requires an investment of limited resources, for which plants also compete with neighbours. The consequences of an investment in defence are determined by the metabolic costs of defence as well as indirect or ecological costs through interactions with other organisms. These ecological costs have a potentially strong impact on the evolution of defensive traits, but have proven to be difficult to quantify. We aimed to quantify the relative impact of the direct and indirect or ecological costs and benefits of an investment in plant defence in relation to herbivory and intergenotypic competition for light. Additionally, we evaluated how the benefits of plant defence balance its costs in the context of herbivory and intergenotypic competition. To this end, we utilised a functional‐structural plant (FSP) model of Brassica nigra that simulates plant growth and development, morphogenesis, herbivory and plant defence. In the model, a simulated investment in defences affected plant growth by competing with other plant organs for resources and affected the level and distribution of herbivore damage. Our results show that the ecological costs of intergenotypic competition for light are highly detrimental to the fitness of defended plants, as it amplifies the size difference between defended and undefended plants. This leads to herbivore damage counteracting the effects of intergenotypic competition under the assumption that herbivore damage scales with plant size. Additionally, we show that plant defence relies on reducing herbivore damage rather than the dispersion of herbivore damage, which is only beneficial under high levels of herbivore damage. We conclude that the adaptive value of plant defence is highly dependent on ecological interactions and is predominantly determined by the outcome of competition for light. plain language summary is available for this article.

Recent studies show that the variation in root functional traits can be explained by a twodimensional trait framework, containing a 'collaboration' axis in addition to the classical fastslow 'conservation' axis. This collaboration axis spans from thin and highly branched roots that employ a 'do-it-yourself' strategy to thick and sparsely branched roots that 'outsource' nutrient uptake to symbiotic arbuscular mycorrhizal fungi (AMF).Here, we explore the functionality of this collaboration axis by quantifying how interactions with AMF change the impact of root traits on plant performance. To this end, we developed a novel functional-structural plant (FSP) modelling approach that simulates plants competing for light and nutrients in the presence or absence of AMF.Our simulation results support the notion that in the absence of AMF, plants rely on thin, highly branched roots for their nutrient uptake. The presence of AMF, however, promotes thick, unbranched roots as an alternative strategy for uptake of immobile phosphorus, but not for mobile nitrogen.This provides further support for a root trait framework that accommodates for the interactive effect of roots and AMF. Our modelling study offers unique opportunities to incorporate soil microbial interactions into root functionality as it integrates consequences of belowground trait expression.