Green light added to blue light has been proposed to shift cryptochromes from their semireduced active form to the reduced, inactive state. Whether the increased proportion of green light observed under leaf canopies compared to open places reduces cryptochrome-mediated effects remained to be elucidated. Here we report that the length of the hypocotyl of Arabidopsis (Arabidopsis thaliana) seedlings grown under controlled conditions decreased linearly with increasing blue/green ratios of the light within the range of ratios found in natural environments. This effect was stronger under higher irradiances. We developed a model, parameterized on the basis of field experiments including photoreceptor mutants, where hypocotyl growth of seedlings exposed to different natural radiation environments was related to the action and interaction of phytochromes and cryptochromes. Adding the blue/green ratio of the light in the term involving cryptochrome activity improved the goodness of fit of the model, thus supporting a role of the blue/green ratio under natural radiation. The blue/ green ratio decreased sharply with increasing shade by green grass leaves to one-half of the values observed in open places. The impact of blue/green ratio on cryptochrome-mediated inhibition of hypocotyl growth was at least as large as that of irradiance. We conclude that cryptochrome is a sensor of blue irradiance and blue/green ratio.

Phototropism is an asymmetric growth response enabling plants to optimally position their organs. In flowering plants, the phototropin (phot) blue light receptors are essential to detect light gradients. In etiolated seedlings, the phototropic response is enhanced by the red/far-red (R/FR)-sensing phytochromes (phy) with a predominant function of phyA. In this study, we analyzed the influence of the phytochromes on phototropism in green (de-etiolated) Arabidopsis seedlings. Our experiments in the laboratory and outdoors revealed that, in open environments (high R/FR ratio), phyB inhibits phototropism. In contrast, under foliar shade, where access to direct sunlight becomes important, the phototropic response was strong. phyB modulates phototropism, depending on the R/FR ratio, by controlling the activity of three basic-helix-loop-helix (bHLH) transcription factors of the PHYTOCHROME INTERACTING FACTORs (PIFs) family. Promotion of phototropism depends on PIF-mediated induction of several members of the YUCCA gene family, leading to auxin production in the cotyledons. Our study identifies PIFs and YUCCAs as novel molecular players promoting phototropism in photoautotrophic, but not etiolated, seedlings. Moreover, our findings reveal fundamental differences in the phytochrome-phototropism crosstalk in etiolated versus green seedlings. We propose that in natural conditions where the light environment is not homogeneous, the uncovered phytochrome-phototropin co-action is important for plants to adapt their growth strategy to optimize photosynthetic light capture.