Plant movements: navigating the light environment.

Plants use light both as a resource for photosynthesis and as a signal about their environment. In response to light cues, plants can move their organs via directional growth driven by cell expansion. In dense vegetation where light is available in spatially heterogeneous patterns, plants need to navigate this space to improve the position of their photosynthetic tissues. In canopies blue light irradiance and red to far-red light ratio decrease due to absorption by chloroplasts, and these changes regulate distinct processes within the plant. Changes in light environment are detected by cryptochrome and phytochrome photoreceptors, both regulating phytochrome interacting factors (PIFs) and thereby enhancing elongation in hypocotyls, stems, and leaves and inducing upward leaf movement (hyponasty). An additional class of photoreceptors, phototropins, decodes horizontal light gradients to produce directional growth toward the light source (phototropism). Here we review the current state of knowledge on these differential growth responses to light cues, with specific emphasis on the regulatory pathways that translate light signaling into differential cell expansion. Downstream of the photoreceptors, the phytohormone auxin induces cell growth in shoot tissues, but also other phytohormones contribute to balancing light responses. Cell expansion is regulated primarily at the level of cell walls, and a comparison of different transcriptome datasets reveals that only a small group of cell wall-modifying genes are tightly regulated by shade cues. It remains poorly understood which cell layers are causal to the initiation of cellular expansion. Here we combine insights from different differential growth behaviors in different species and organs to generate different hypotheses for the cellular underpinnings of light-driven leaf movements.