We have screened a large population of M2 seeds ofArabidopsis thaliana for plants which are resistant to exogenously applied indole-acetic acid (IAA). One of the resistant lines identified in this screen carries a dominant mutation which we have namedaxr2. Linkage analysis indicates that theaxr2 gene lies on chromosome 3. Plants carrying theaxr2 mutation are severe dwarfs and display defects in growth orientation of both the shoot and root suggesting that the mutation affects some aspect of gravitropic growth. In addition, the roots ofaxr2 plants lack root hairs. Growth inhibition experiments indicate that the roots ofaxr2 plants are resistant to ethylene and abscisic acid as well as auxin.

A dominant mutation in Arabidopsis confers resistance to auxin, ethylene and abscisic acid.

Aspects of Plant Intelligence

For a better understanding of Al inhibition of root elongation, knowledge of the morphological and functional organization of the root apex is a prerequisite. We developed a polyvinyl chloride-block technique to supply Al (90 μm monomeric Al) in a medium containing agarose to individual 1-mm root zones of intact seedlings of maize (Zea mays L. cv Lixis). Root elongation was measured during a period of 5 h. After Al treatment, callose (5 h) and Al (1 h) contents of individual 1-mm apical root segments were determined. For comparison, callose and Al levels were also measured in root segments after uniform Al supply in agarose blocks to the 10-mm root apex. Only applying Al to the three apical 1-mm root zones inhibited root elongation after 1 h. The order of sensitivity was 1 to 2 > 0 to 1 > 2 to 3 mm. In the 1- to 2-mm root zone high levels of Al-induced callose formation and accumulation of Al was found, independently of whether Al was applied to individual apical root zones or uniformly to the whole-root apex. We conclude from these results that the distal part of the transition zone of the root apex, where the cells are undergoing a preparatory phase for rapid elongation (F. Baluska, D. Volkmann, P.W. Barlow [1996] Plant Physiol 112: 3–4), is the primary target of Al in this Al-sensitive maize cultivar.

The Distal Part of the Transition Zone Is the Most Aluminum-Sensitive Apical Root Zone of Maize

Membrane flow and interconversions among endomembranes

It is well accepted that lateral redistribution of the phytohormone auxin underlies the bending of plant organs towards light. In monocots, photoreception occurs at the shoot tip above the region of differential growth. Despite more than a century of research, it is still unresolved how light regulates auxin distribution and where this occurs in dicots. Here, we establish a system in Arabidopsis thaliana to study hypocotyl phototropism in the absence of developmental events associated with seedling photomorphogenesis. We show that auxin redistribution to the epidermal sites of action occurs at and above the hypocotyl apex, not at the elongation zone. Within this region, we identify the auxin efflux transporter ATP-BINDING CASSETTE B19 (ABCB19) as a substrate target for the photoreceptor kinase PHOTOTROPIN 1 (phot1). Heterologous expression and physiological analyses indicate that phosphorylation of ABCB19 by phot1 inhibits its efflux activity, thereby increasing auxin levels in and above the hypocotyl apex to halt vertical growth and prime lateral fluxes that are subsequently channeled to the elongation zone by PIN-FORMED 3 (PIN3). Together, these results provide new insights into the roles of ABCB19 and PIN3 in establishing phototropic curvatures and demonstrate that the proximity of light perception and differential phototropic growth is conserved in angiosperms.

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phot1 Inhibition of ABCB19 Primes Lateral Auxin Fluxes in the Shoot Apex Required For Phototropism

The magnitude of acceleration required to induce growth responses in Avena seedlings grown in the absence of tropic response to earth gravity has been investigated. For this purpose, a clinostat was developed that imposes accelerations from about 10−9 g to 3 g upon the seedling; simultaneously, it nullifies, or compensates for, response to the directional component of the gravitational-force vector by rotating the seedling on a horizontal axis. When accelerations less than 10−3 g are applied in either the acropetal or the basipetal direction, the growth in length and weight of the various organs is not materially different from that of compensated seedlings to which no longitudinal force is applied. At accelerations between 10−3 and 10−2 g, differences in growth become highly significant. When the centrifugal forces are transverse to the seedling during compensation, the threshold acceleration range for geoperception, as manifest by shoot reorientation, is again between 10−3 and 10−2 g. Geotropic reorientation of the root becomes apparent after exposures between 10−4 and 10−3 g.

Thresholds for georesponse to acceleration in gravity-compensated Avena seedlings.

The effect of phototropic stimulation of Zea coleoptile tips on the distribution of both endogenous indoleacetic acid (IAA) and applied C(14)-labeled IAA was determined. The tips rested on bisected agar blocks. More IAA was found in the blocks under the shaded side of the coleoptile tips than those under the irradiated side. However, no significant difference was observed between the total amounts of IAA, endogenous or labeled, in the irradiated and shaded sides of the experimental system. In addition, less endogenous auxin was found in the shaded tissues than in their irradiated counterparts. It is suggested that phototropism following unilateral irradiation with first positive radiant densities might be a consequence of lateral inequalities in the ability of the irradiated and shaded tissues to transport auxin basipetally.

Hormonal Relations in the Phototropic Response: III. The Movement of C14-labeled and Endogenous Indoleacetic Acid in Phototropically Stimulated Zea Coleoptiles

Avena seedlings were germinated and grown while continuously rotated on the horizontal axis of a clinostat. The coleoptiles of these gravity-compensated plants were phototropically more responsive than those of plants rotated on a vertical axis. When the plants were compensated after unilateral irradiation, phototropic curvature of the shoot progressed for the next 6 hours, with the rate of curving decreasing about 3 hours after irradiation. The decrease in rate was less in the plants gravity-compensated before irradiation than in those vertically rotated. In the period 70 to 76 hours after planting, the growth rate of the compensated coleoptiles was significantly less than that of the vertically rotated seedlings. The greater phototropic curvature, the decreased growth rate, and the slower rate of straightening of the curved, compensated shoot can be correlated with several consequences of compensation: an increase in sensitivity to auxin, a lowering of auxin content in the coleoptile tip, and possibly, from an interaction between compensation and phototropic stimulation, an enhanced difference in auxin transport between the illuminated and shaded halves of the unilaterally irradiated shoot. The phototropic response of the vertically rotated seedling was significantly different from that of the vertical stationary, indicating the importance of vertically rotated controls in clinostat experiments.

Gravitational Compensation and the Phototropic Response of Oat Coleoptiles

We find a differential distribution of dictyosomes in the avascular tip cells of the oat (Avena sativa) coleoptile upon geostimulation. A differential activation (increased vesicle production) of the dictyosomes with respect to gravity also occurs, but only in the tip cells of the lower tissues. Similar differences in distribution and activation of dictyosomes occur also in cells subjacent to the avascular tip (1st and 5th millimeter from the apex) of both the upper and lower half-tissues. When only the outer epidermal cells below the apex are considered, the differential distribution and activation of dictyosomes occur only in the lower outer epidermis. The changes in distribution of dictyosomes begin at 6 minutes, or sooner, from the start of geostimulation, before the onset of geotropism. The kinetics of amyloplast sedimentation and Golgi movement do not appear to differ in the cells of the avascular tip. We suggest that the Golgi participates in, and possibly initiates, the differential elongation of cells of geotropically stimulated coleoptiles.

http://www.plantphysiol.org/content/plantphysiol/49/4/634.full.pdf

Distribution and Activation of the Golgi Apparatus in Geotropism

(14)C-Indoleacetic acid was applied to coleoptiles of corn (Zea mays) and oat (Avena sativa). The coleoptiles were detached from the endosperms at 6-minute intervals after indoleacetic acid application, and the radioactivity was determined in successive 2-millimeter regions. The rate (per cent per minute) of basipetal transport of indoleacetic acid is periodic in various regions of the coleoptile, with a period of about 20 minutes. The possible relation of this cyclic phenomenon to other rhythmic processes of similar periodicities is discussed. A distinct acropetal transport (against the concentration gradient) from the subapical region to the apical 2-millimeter region of the coleoptile was detected.The velocity of indoleacetic acid transport differs in different regions of the coleoptile. Within an entire coleoptile the velocity can be divided into three classes for corn, 41 millimeters per hour (apical), 13 millimeters per hour (mid), and 34 millimeters per hour (base), and 2 classes for oats, 28 millimeters per hour (apical) and 14 millimeters per hour (remainder). An inverse relationship between the velocity of indoleacetic acid transport, and the growth rate of the coleoptile is discussed. Corn coleoptiles exceed oat coleoptiles both in rate and in velocity of IAA transport.

J. Shen-Miller

Papers

Thresholds for georesponse to acceleration in gravity-compensated Avena seedlings.

Hormonal Relations in the Phototropic Response: III. The Movement of C14-labeled and Endogenous Indoleacetic Acid in Phototropically Stimulated Zea Coleoptiles

Gravitational Compensation and the Phototropic Response of Oat Coleoptiles

Distribution and Activation of the Golgi Apparatus in Geotropism

Rhythmicity in the Basipetal Transport of Indoleacetic Acid through Coleoptiles