(J.L.M., R.P.H.)The interaction of tropisms is important in determining the final growth form of the plant body. In roots, gravitropism is the predominant tropistic response, but phototropism also plays a role in the oriented growth of roots in flowering plants. In blue or white light, roots exhibit negative phototropism that is mediated by the phototropin family of photoreceptors. In contrast, red light induces a positive phototropism in Arabidopsis roots. Because this red-light-induced response is weak relative to both gravitropism and negative phototropism, we used a novel device to study phototropism without the complications of a counteracting gravitational stimulus. This device is based on a computer-controlled system using real-time image analysis of root growth and a feedback-regulated rotatable stage. Our data show that this system is useful to study root phototropism in response to red light, because in wild-type roots, the maximal curvature detected with this apparatus is 30°to 40°, compared with 5°to 10°without the feedback system. In positive root phototropism, sensing of red light occurs in the root itself and is not dependent on shoot-derived signals resulting from light perception. Phytochrome (Phy)A and phyB were severely impaired in red-light-induced phototropism, whereas the phyD and phyE mutants were normal in this response. Thus, PHYA and PHYB play a key role in mediating red-light-dependent positive phototropism in roots. Although phytochrome has been shown to mediate phototropism in some lower plant groups, this is one of the few reports indicating a phytochrome-dependent phototropism in flowering plants.Plants have evolved selective and sensitive mechanisms to deal with the constant sensory input they receive from the environment. In roots, gravity is the most critical signal for growth and development, and, thus, gravitropism has been well-characterized in this organ (Sack, 1991;Kiss, 2000). However, it has become increasingly clear that gravitropism interacts with a number of other tropistic responses including phototropism, thigmotropism, and hydrotropism in determining the final growth form of the primary root and the entire root system (Hangarter, 1997;Correll and Kiss, 2002).Phototropism in roots was extensively reviewed in a classical paper by Hubert and Funke (1937) but has received increased attention since the report by Okada and Shimura (1992), who isolated mutants in root phototropism that were later shown to be deficient in the blue-light receptor PHOT1 (Briggs and Christie, 2002). Roots are typically negatively phototropic in response to white and blue light (Okada and Shimura, 1992;Vitha et al., 2000) and use the same photoreceptors that are involved in phototropism in stems and stem-like organs (Sakai et al., 2000). Furthermore, similar to root gravisensing (Blancaflor et al., 1998), sensing of blue light for phototropism occurs in the root cap .We have recently identified a red-light-induced positive phototropism in primary roots of Arabidopsis ). This tropistic response ...