Primary roots of the mutant 'Ageotropic' cultivar of Zea mays are nonresponsive to gravity. Their root caps secrete little or no mucilage and touch the root only at the extreme apex. A gap separates the cap and root at the periphery of the cap. Applying mucilage from normal roots or substances with a consistency similar to that of mucilage to tips of mutant roots causes these roots to become strongly graviresponsive. Gravicurvature stops when these substances are removed. Caps of some mutants secrete small amounts of mucilage and are graviresponsive. These results indicate that (a) the lack of graviresponsiveness in the mutant results from disrupting the transport pathway between the cap and root, (b) movement of the growth-modifying signal from the cap to the root occurs via an apoplastic pathway, and (c) gravity (18). This indicates that the lesion for gravitropism is downstream from gravity-detection, perhaps in transmitting the signal from the cap to the root. We studied this by examining the nature of the cap/root interface of mutant roots.
MATERIALS AND METHODSAlmost all models for root gravitropism involve a similar sequence of events. Detection of gravity occurs in the root cap and probably involves sedimentation of amyloplasts in columella cells (2,5,15). This change in cellular structure somehow generates a chemical and/or electrical signal in the cap (2,4,5). This signal moves from the root cap to the elongating zone of the root, which is 2 to 6 mm behind the cap. There, the signal preferentially inhibits cellular elongation along the lower side of the root, thus causing the root to curve down (4,5,15).Although we cannot identify the gravitropic signal, much evidence indicates that it moves from the cap to the root before gravicurvature. For example, decapped roots are nonresponsive to gravity (7,23,24), and removing half of the cap of vertically oriented roots induces curvature toward the remaining half-cap (6). Inserting barriers into one side of a root between the root cap and elongating zone causes curvature away from the barrier (26). Taken together, these results suggest that root gravitropism involves movement of a growth-modifying signal from the root cap to the root.The objective of this research was to use mutants to characterize the pathway by which the gravitropic signal moves from the root cap to the root in primary roots of Zea mays.'