Chiral and non-chiral nutations in Arabidopsis roots grown on the random positioning machine.

Arabidopsis thaliana roots grown on a vertically set plate do not elongate straight down the gravitational vector, but by making waves and coils, and by conspicuously slanting towards the right-hand. This behaviour, in a previous paper, was ascribed to the simultaneous effect of three processes: circumnutation, positive gravitropism and negative thigmotropism. However, when the plants are grown on the Random Positioning Machine (RPM), in conditions that are believed to simulate space microgravitational conditions closely, the roots do not show the usual pattern. In the wild type, the roots make large loops to the right-hand side, whereas in the gravitropic and auxinic mutants aux1, eir1, rha1, they just move randomly around the initial direction. Therefore, if the movements made on the RPM are those produced by the exclusion of gravitropism and negative thigmotropism, as is apparent, the conclusion is that Arabidopsis roots are animated by a form of chiral circumnutation, that is lacking in the auxinic and gravitropic mutants aux1, eir1 and rha1. In addition, the 1 g condition appears to reduce the scatter among the circumnutating tracks produced by the roots of the wild types, but not among those of the mutants. Because there is a scarcity of literature regarding circumnutation in roots, it is not known how widely root chiral circumnutation is spread, but it is known that, in some previously studied species, just random nutations are observed. Two kinds of nutating movements seem to exist in plant roots and, whereas the random process does not seem to be connected with auxin physiology and transport, the chiral process appears to be connected in the same way as gravitropism is.

A new Arabidopsis thaliana root gravitropism and chirality mutant.

A new Arabidopsis thaliana (L.) Heynh. root mutant, named clg1, was isolated from the Feldmann-Du Pont T-DNA insertional mutagenesis collection. It is characterized by primary roots that make early right-handed coils, show increased right-handed slanting, and reduced positive gravitropism. The mutant roots however did not exhibit increased resistance to IAA, but only a modest increment of resistance to the auxins 2,4 D, and NAA, and to the auxin transport inhibitors TIBA and NPA. By contrast, the mutant roots showed a notable resistance to plant hormone ethylene (given as ACC). clg1 appears to be new, since it complements the most known auxin and gravitropism mutants, maps to chromosome 5, and shows a phenotype largely different from that of the known ethylene mutants. The increased right-handed slanting (chirality) can possibly be a consequence of the reduced gravitropic response, since gravitropism and slanting are competitive growth-direction leading forces. The increased resistance to ethylene, seems to indicate that this phythormone plays a role in the gravitropic response of roots (as already proposed for shoots), and possibly in the regulation of the connected signal transduction pathway. The gene involved in the clg1 mutation, which is recessive, was mapped, as above reported, to chromosome 5, close to the visible marker tt3.

The right-handed slanting of Arabidopsis thaliana roots is due to the combined effects of positive gravitropism, circumnutation and thigmotropism.

When primary Arabidopsis roots grow down a tilted agar plate, they do not elongate following the gravitational vector along a straight line, but instead they slant noticeably to the right-hand. This process is seen mostly in the ecotypes Wassilewskjia and Landsberg, whereas it is attenuated in the ecotype Columbia, and in some mutants is even inverted. The origin of the slanting of Arabidopsis roots, that evidently constitutes a form of chirality, has so far not been sufficiently clarified. In the present paper we describe it as the general result of the cumulative effects of positive gravitropism, circumnutation and a thigmotropic obstacle-avoiding movement, and in particular, as the consequence of the alternating movement of circumnutation of the root to the right and to the left of the gravitational vector. This movement, which does not appear symmetrical in its nature, since the waves made to the right-hand are complete whereas those made to the left-hand are reduced or aborted, appears to be the reason for the observed slanting. In addition, evidence is furnished supporting the hypothesis that the strong left-handed cell-files torsion, seen in right-handed coiling roots, is not the consequence of a primary process, but of an artifact, and is due to the adjustment of the three dimensional root circumnutating helix to the flat two dimensional agar surface.