Dancing in the dark: darkness as a signal in plants

Daily cycles of light and dark provide an organizing principle and temporal constraints under which life on Earth evolved. While light is often the focus of plant studies, it is only half the story. Plants continuously adjust to their surroundings, taking both dawn and dusk as cues to organize their growth, development and metabolism to appropriate times of day. In this review, we examine the effects of darkness on plant physiology and growth. We describe the similarities and differences between seedlings grown in the dark versus those grown in light–dark cycles, and the evolution of etiolated growth. We discuss the integration of the circadian clock into other processes, looking carefully at the points of contact between clock genes and growth‐promoting gene‐regulatory networks in temporal gating of growth. We also examine daily starch accumulation and degradation, and the possible contribution of dark‐specific metabolic controls in regulating energy and growth. Examining these studies together reveals a complex and continuous balancing act, with many signals, dark included, contributing information and guiding the plant through its life cycle. The extraordinary interconnection between light and dark is manifest during cycles of day and night and during seedling emergence above versus below the soil surface. Daily cycles of light and darkness provide structure that plants use to temporally organize aspects of their physiology and growth. Plants use distinct modes of growth in continuous dark versus light–dark cycles but these programmes share many components. In this review, we examine progress toward understanding how plants coordinate light‐sensitive and time‐sensitive processes to optimize growth and resource allocation across the day–night cycle, and how darkness changes the function of gene regulatory networks involved in seedling growth, timing and metabolism.