Caught between two states: The compromise in acclimation of photosynthesis, transpiration and mesophyll conductance to different amplitudes of fluctuating irradiance.

While dynamic regulation of photosynthesis in fluctuating light is increasingly recognized as an important driver of carbon uptake, acclimation to realistic irradiance fluctuations is still largely unexplored. We subjected Arabidopsis thaliana (L.) wild-type and jac1 mutants to irradiance fluctuations with distinct amplitudes and average irradiance. We examined how irradiance fluctuations affected leaf structure, pigments and physiology. A wider amplitude of fluctuations produced a stronger acclimation response. Large reductions of leaf mass per area under fluctuating irradiance framed our interpretation of changes in photosynthetic capacity and mesophyll conductance as measured by three separate methods, in that photosynthetic investment increased markedly on a mass basis, but only a little on an area basis. Moreover, thermal imagery showed that leaf transpiration was four times higher under fluctuating irradiance. Leaves growing under fluctuating irradiance, although thinner, maintained their photosynthetic capacity, as measured through light- and CO-response curves; suggesting their photosynthesis may be more cost-efficient than those under steady light, but overall may incur increased maintenance costs. This is especially relevant for plant performance globally because naturally fluctuating irradiance creates conflicting acclimation cues for photosynthesis and transpiration that may hinder progress towards ensuring food security under climate-related extremes of water stress.