Extensive studies in both lower and higher plants indicate that plant phytochrome photoreceptors signal not only by regulating transcription in the nucleus but also by acting within the cytoplasm, the latter signaling routes acting within minutes or even seconds and also providing directional information. Directional signals seem to arise from phytochromes attached anisotropically to the plasma membrane. Neochromes-phytochrome-phototropin hybrid photoreceptors probably attached to the plasma membrane-provide this signal in various ferns and perhaps certain algae but are absent from other groups. In mosses and probably higher plants too, a subpopulation of canonical phytochromes interact with phototropins at the plasma membrane and thereby steer directional responses. Phytochromes also seem able to regulate translation in the cytoplasm. This review discusses putative phytochrome functions in these contexts.