Control of gravitropic orientation. I. Non-vertical orientation by primary roots of maize results from decay of competence for orthogravitropic induction

Plant organs may respond to gravity by vertical (orthogravitropic), oblique (plagiogravitropic) or horizontal (diagravitropic) growth. Primary roots of maize (Zea mays L.) provide a good system for studying such behaviours because they are reportedly capable of displaying all three responses. In current work using maize seedlings of the Silver Queen cultivar, stabilisation of growth at an oblique orientation was commonplace. Hypothetically, plagiogravitropism may be accomplished either by a process we call graded orthogravitropism or by hunting about a sensed non-vertical setpoint. In graded orthotropism primary bending is unidirectional and depends on facilitative stimuli that determine its extent. The hallmark of the setpoint mechanism is restorative curvature of either sign following a displacement; both diagravitropism and orthogravitropism are based on setpoints. Roots settled in a plagiogravitropic orientation were tested with various illumination and displacement protocols designed to distinguish between these two hypotheses. The tests refuted the setpoint hypothesis and supported that of graded orthotropism. No evidence of diagravitropism could be found, thus, earlier claims were likely based on inadequately controlled observations of graded orthotropism. We propose that orthotropism is graded by the sequential action of dual gravity receptors: induction of a vectorial gravitropic response requires gravitational induction of a separate facilitative response, whose decay in the absence of fresh stimuli can brake gravitropism at plagiotropic angles.