Plasticity and the role of mass-scaling in allocation, morphology, and anatomical trait responses to above- and belowground resource limitation in cultivated sunflower (Helianthus annuus L.).

In the face of resource limitations, plants show plasticity in multiple trait categories, including biomass allocation, morphology, and anatomy, yet inevitably also grow less. The extent to which passive mass-scaling plays a role in trait responses that contribute to increased potential for resource acquisition is poorly understood. Here, we assessed the role of mass-scaling on the direction, magnitude, and coordination of trait plasticity to light and/or nutrient limitation in cultivated sunflower (Helianthus annuus). We grew seedlings of 10 sunflower genotypes for 3 weeks in a factorial of light (50% shade) and nutrient (10% supply) limitation in the greenhouse and measured a suite of allocational, morphological, and anatomical traits for leaves, stems, fine roots, and tap roots. Under resource limitation, plants were smaller and more biomass was allocated to the organ capturing the most limiting resource, as expected. Traits varied in the magnitude of plasticity and the extent to which the observed response was passive (scaled with plant mass) and/or had an additional active component. None of the allocational responses were primarily passive. Plastic changes to specific leaf area and specific root length were primarily active, and adjusted toward more acquisitive trait values under light and nutrient limitation, respectively. For many traits, the observed response was a mixture of active and passive components, and for some traits, the active adjustment was antagonistic to the direction of passive adjustment, for example, stem height, and tap root and stem theoretical hydraulic conductance. Passive scaling with size played a major role in the coordinated response to light, but correcting for mass clarified that the active responses to both limitations were more similar in magnitude, although still resource and organ specific. Our results demonstrate that both passive plasticity and active plasticity can contribute to increased uptake capacity for limiting resources in a manner that is resource, organ, and trait specific. Indeed, passive adjustments (scaling with mass) of traits due to resource stress extend well beyond just mass allocation traits. For a full understanding of plants' response to environmental stress, both passive and active plasticity need to be taken into account.