Intracellular magnetophoresis of amyloplasts and induction of root curvature

High-gradient magnetic fields (HGMFs) were used to induce intracellular magnetophoresis of amyloplasts. The HGMFs were generated by placing a small ferromagnetic wedge into a uniform magnetic field or at the gap edge between two permanent magnets. In the vicinity of the tip of the wedge the dynamic factor of the magnetic field, (H²/2), was about 10⁹ Oe² · cm^–1, which subjected the amyloplasts to a force comparable to that of gravity. When roots of 2-d-old seedlings of flax (Linum usitatissimum L.) were positioned vertically and exposed to an HGMF, curvature away from the wedge was transient and lasted approximately 1 h. Average curvature obtained after placing magnets, wedge and seedlings on a 1-rpm clinostat for 2 h was 33 ± 5 degrees. Roots of horizontally placed control seedlings without rotation curved about 47 ± 4 degrees. The time course of curvature and changes in growth rate were similar for gravicurvature and for root curvature induced by HGMFs. Microscopy showed displacement of amyloplasts in vitro and in vivo. Studies with Arabidopsis thaliana (L.) Heynh. showed that the wild type responded to HGMFs but the starchless mutant TC7 did not. The data indicate that a magnetic force can be used to study the gravisensing and response system of roots.Abbreviations HGMF high-gradient magnetic field - emu electromagnetic units - Oe Oersted We thank Dr. John Kiss, Miami University, Ohio for providing the Arabidopsis seeds. This work was supported by NASA grant NAGW-3656     

Effects of submergence on development and gravitropism in the coleoptile of Oryza sativa L.

Caryopses of rice (Oryza sativa L. cv. Sasanishiki) were germinated in air or under water. In submerged seedlings a twofold increase in coleoptile growth rate and an inhibition of root growth was observed. The amount of starch in the amyloplasts of submerged coleoptiles was substantially reduced compared to the air-grown control plants and plastids had a proplastidic character. During the rapid elongation of coleoptiles under water, the osmotic concentration of the press sap remained constant, whereas in air-grown coleoptiles a decrease was measured. Determination of curvature of gravistimulated air-grown and submerged shoots was carried out by placing the coleoptiles horizontally in air of 98% relative humidity. Air-grown coleoptiles reached a vertical orientation within 5 h after onset of gravistimulation. In coleoptiles germinated under water the first signs of consistent negative gravitropic bending occurred after 4–5 h and curvature was complete after 24 h. During the first 5 h of gravistimulation the water-grown coleoptiles grew at an average rate of 0.39 mm·h^–1, whereas in air-grown coleoptiles a rate of 0.27 mm·h^–1 was measured. Concomitant with the delayed onset of gravitropic bending of the water-grown coleoptiles, a change in plastid ultrastructure and an increase in starch content was observed. We conclude that the gravitropic responsiveness of the rice coleoptile depends on the presence of starch-filled amyloplasts.We wish to thank H.-J. Ensikat for technical assistance with the scanning electron microscopy. Supported by the Bundesminister für Forschung und Technologie and the Deutsche Forschungsgemeinschaft.     

Gravitropism and starch statoliths in an Arabidopsis mutant

The mutant TC 7 of Arabidopsis thaliana (L.) Heynh. has been reported to be starch-free and still exhibit root gravitropism (T. Caspar and B. G. Pickard 1989, Planta 177, 185–197). This is not consistent with the hypothesis that plastid starch has a statolith function in gravity perception. In the present study, initial light microscopy using the same mutant showed apparently starch-free statocytes. However, ultrastructural examination detected residues of amyloplast starch grains in addition to the starch-depleted amyloplasts. Applying a point-counting morphometric method, the starch grains in the individual amyloplasts in the mutant were generally found to occupy more than 20% and in a few cases up to 60% of the amyloplast area. In the wild type (WT) the starch occupied on average 98 % of the amyloplast area and appeared as densely packed grains. The amyloplasts occupied 13.9% of the area of the statocyte in the mutant and 23.3% of the statocyte area in the WT. Sedimentation of starch-depleted amyloplasts in the mutant was not detected after 40 min of inversion while in the WT the amyloplasts sedimented at a speed of 6 m · h^-1. The gravitropic reactivity and the curvature pattern were also examined in the WT and the mutant. The time-courses of root curvature in the WT and the mutant showed that when cultivated under standard conditions for 60 h in darkness, the curvatures were 83° and 44°, respectively, after 25 h of continuous stimulation in the horizontal position. The WT roots curved significantly more rapidly and with a more normal gravitropic pattern than those of the mutant. These results are discussed in relation to the results previously obtained with the mutant and with respect to the starch-statolith hypothesis.Abbreviation WT wild type This work was supported by grants from Norwegian Research Council for Science and the Humanities (NAVF) which we gratefully acknowledge. We would also like to thank Dr. Timothy Caspar, Michigan State University, East Lansing, USA, for providing us with the seeds of TC 75.     

Gravitropism in roots of intermediate-starch mutants of Arabidopsis

Gravitropism was studied in roots of wild type (WT) Arabidopsis thaliana (L.) Heynh. (strain Wassilewskija) and three starch-deficient mutants that were generated, by T-DNA insertional mutagenesis. One of these mutants was starchless while the other two were intermediate mutants, which had 51% and 60%, respectively, of the WT amount of starch as. determined by light and electron microscopy. The four parameters used to assay gravitropism were: orientation during vertical growth, time course of curvature, induction, and intermittent stimulation experiments. WT roots were much more responsive to gravity than were roots of the slarchless mutant, and the intermediate starch mutants exhibited an intermediate graviresponse. Our data suggest that lowered starch content in the mutants primarily affects gravitropism rather than differential growth because both phototropic curvature and growth rates were approximately equal among all four genotypes. Since responses of intermediate-starch mutants were closer to the WT response than to that of the starchless mutant, it appears that 51–60% of the WT level of starch is near the threshold amount needed for full gravitropic sensitivity. While other interpretations are possible, the data are consistent with the starch statolith hypothesis for gravity perception in that the degree of graviresponsiveness is proportional to the total mass of plastids per cell.     

Kinetics of amyloplast sedimentation in gravistimulated maize coleoptiles

Inner mesophyll cells from coleoptiles of Zea mays L. cv. Merit were fixed after varying periods of gravistimulation. A statistically significant amount (17–21%) of amyloplast sedimentation occurred in these cells after 30 s of gravistimulation. The presentation time is approx. 40 s or less. The accumulation of amyloplasts near the new lower wall shows a linear relationship to the logarithm of the gravistimulation time (r=0.92 or higher). The intercept of this line with the baseline value of amyloplasts in vertical coleoptiles indicates that the number of amyloplasts on the new lower wall begins increasing 11–15 s after the onset of gravistimulation. Direct observations of living cells confirm that many amyloplasts sediment within less than 15–30 s. These rapid kinetics are consistent with the classical statolith hypothesis of graviperception involving the sedimentation of amyloplasts to the vicinity of the new lower wall.     

Rootcap structure in wild type and in a starchless mutant of Arabidopsis.

Rootcaps of the wild type (WT) and of a starchless, gravitropic mutant (TC7) of Arabidopsis thaliana L. were examined by electron microscopy to identify cellular polarities with respect to gravity. In columella cells, nuclei are located proximally, and the nuclear envelope is continuous with endoplasmic reticulum (ER) that is in turn connected to nearby plasmodesmata. Impregnation of ER with osmium ferricyanide revealed numerous contacts between columella plastids and ER in both genotypes. ER is present mostly in the outer regions of the columella protoplast except in older columella cells that are developing into peripheral cells. In vertical roots, only columella cells that are intermediate in development (story 2 cells) have a higher surface density (S) of ER in the distal compared to proximal regions of the cell. The distal but not the proximal S of the ER is constant throughout columella development. Plastids are less sedimented in TC7 columella cells compared to those of the WT. It is hypothesized that plastid contact with the ER plays a role in gravity perception in both genotypes.     

Amyloplast sedimentation kinetics in gravistimulated maize roots

Amyloplast sedimentation in gravistimulated maize (Zea mays L.) roots was measured using the change in angle from the center of the cell to each amyloplast as an index of sedimentation. Using tissue fixed after gravistimulation, the relationship between mean amyloplast angle and the duration of gravistimulation was found to be linear when plotted on a logarithmic time scale. Extrapolated values for the onset of angular change are 5.9 s after the start of gravistimulation for the entire population of amyloplasts and 11.8 s for lead amyloplasts. By multiplying the instantaneous angular velocity (in radians) by the cell center to amyloplast radius, it is possible to calculate the initial sedimentation velocity to be 19.1 m min^-1 at 5.9 s. During sedimentation, the mean amyloplast angles surpass the calculated cell corner angle of 123° at 2.2 min for all amyloplasts and at 19 s for lead amyloplasts near the new lower wall. Thus, substantial sedimentation occurs within the presentation time, calculated to be 4.1 min. These kinetics are consistent with several hypotheses of graviperception.Symbol t_p presentation time     

Gravitropism in a starchless mutant of Arabidopsis

The starch-statolith theory of gravity reception has been tested with a mutant of Arabidopsis thaliana (L.) Heynh. which, lacking plastid phosphoglucomutase (EC 2.7.5.1) activity, does not synthesize starch. The hypocotyls and seedling roots of the mutant were examined by light and electron microscopy to confirm that they did not contain starch. In upright wild-type (WT) seedlings, starch-filled plastids in the starch sheath of the hypocotyl and in three of the five columellar layers of the root cap were piled on the cell floors, and sedimented to the ceilings when the plants were inverted. However, starchless plastids of the mutant were not significantly sedimented in these cells in either upright or inverted seedlings. Gravitropism of light-grown seedling roots was vigorous: e.g., 10^o curvature developed in mutants rotated on a clinostat following a 5 min induction at 1 · g, compared with 14^o in the WT. Curvatures induced during intervals from 2.5 to 30 min were 70% as great in the mutant as the WT. Thus under these conditions the presence of starch and the sedimentation of plastids are unnecessary for reception of gravity by Arabidopsis roots. Gravitropism by hypocotyls of light-grown seedlings was less vigorous than that by roots, but the mutant hypocotyls exhibited an average of 70–80% as much curvature as the WT. Roots and hypocotyls of etiolated seedlings and flower stalks of mature plants were also gravitropic, although in these cases the mutant was generally less closely comparable to the WT. Thus, starch is also unnecessary for gravity reception in these tissues.Abbreviations PAR photosynthetically active radiation - PAS periodic acid-Schiff's reagent - PGM phosphoglucomutase - WT wild-type     

The Role of Starch in the Gravitropic Response of the Lentil Root

It is well accepted that the amyloplasts of the cap are responsible for gravisensing in primary roots. However, roots with starch-depleted plastids are able to respond to gravistimulus, but their curvature is slower than that of roots containing amyloplasts. The goal of our experiment was to analyse the effects of natural variations of statolith starch in the gravitropic response of lentil roots to a stimulation in the horizontal position. In lentil seedlings grown in the vertical position for 26 h, the volume of the amyloplasts in the statocytes differed between individual roots. The amount of starch in the cap was determined parallel to the rate of gravitropic curvature. There was no statistical correlation between the intensity of the gravitropic response and the starch content in the statocytes. Lentil roots were treated with gibberellic acid (GA₃) at 32°C in order to reduce the volume of starch in the statoliths. There was 53% less starch in the cap of GA₃treated roots as compared to the cap of control roots. But there was no relationship between starch content in the cap and the responsiveness of the root to a gravistimulus, except when the amount of starch was small.     

Geotropism and the lateral transport of auxin in the corn mutant amylomaize

Summary In coleoptiles of the amylomaize corn mutant (AM), the amyloplasts are much reduced in size in comparison with the wild type corn (WT), permitting a comparison of geotropic responsiveness as related to lateral displacement of amyloplasts and lateral transport of auxin. The amyloplasts of AM showed 30–40% lesser lateral redistribution in response to horizontal exposure in comparison with WT. With geotropic stimulation, the lateral transport of auxin in the direction of growth was 40–80% less, and the geotropic curvature by the coleoptiles was also significantly less in the mutant as compared with WT. These correlations support the hypothesis that the starch plastids serve as gravity sensors in the geotropic responses of coleoptiles.     

Amyloplasts as possible statoliths in gravitropic protonemata of the moss Ceratodon purpureus

The kinetics of gravitropism and of amyloplast sedimentation were studied in dark-grown protonemata of the moss Ceratodon purpureus (Hedw.) Brid. The protonemata grew straight up at a rate of 20–25 m·h^– in nutrient-supplemented agar. After they were oriented to the horizontal, upward curvature was first detected after 1–1.5 h and reached 84° by 24 h. The tip cells exhibited an amyloplast zonation, with a tip cluster of nonsedimenting amyloplasts, an amyloplast-free zone, and a zone with pronounced amyloplast sedimentation. This latter zone appears specialized more for lateral than for axial sedimentation since amyloplasts sediment to the lower wall in horizontal protonemata but do not fall to the basal wall in vertical protonemata. Amyloplast sedimentation started within 15 min of gravistimulation; this is within the 12–17-min presentation time. The data support the hypothesis that some amyloplasts function as statoliths in these cells.This work was supported by the National Aeronautics and Space Administration grant NAGW-780. We thank Professor E. Hartmann and J. Schwuchow for providing Ceratodon cultures, Dr. John Z. Kiss and Jeff Young for valuable discussions, and Professor Rainer Hertel (University of Freiburg, FRG) for bringing this material to our attention.     

Inhibition of polar calcium movement and gravitropism in roots treated with auxin-transport inhibitors

Primary roots of maize (Zea mays L.) and pea (Pisum sativum L.) exhibit strong positive gravitropism. In both species, gravistimulation induces polar movement of calcium across the root tip from the upper side to the lower side. Roots of onion (Allium cepa L.) are not responsive to gravity and gravistimulation induces little or no polar movement of calcium across the root tip. Treatment of maize or pea roots with inhibitors of auxin transport (morphactin, naphthylphthalamic acid, 2,3,5-triiodobenzoic acid) prevents both gravitropism and gravity-induced polar movement of calcium across the root tip. The results indicate that calcium movement and auxin movement are closely linked in roots and that gravity-induced redistribution of calcium across the root cap may play an important role in the development of gravitropic curvature.Abbreviations 9-HFCA 9-hydroxyfluorenecarboxylic acid - NPA naphthylphthalamic acid - TIBA 2,3,5-triiodobenzoic acid - IAA indole-3-acetic acid     

Correlations between Gravitropic Curvature and Auxin Movement across Gravistimulated Roots of Zea mays

We compared the kinetics of auxin redistribution across the caps of primary roots of 2-day-old maize (Zea mays, cv Merit) seedlings with the time course of gravitropic curvature. [³H] indoleacetic acid was applied to one side of the cap in an agar donor and radioactivity moving across the cap was collected in an agar receiver applied to the opposite side. Upon gravistimulation the roots first curved upward slightly, then returned to the horizontal and began curving downward, reaching a final angle of about 67°. Movement of label across the caps of gravistimulated roots was asymmetric with preferential downward movement (ratio downward/upward = ca. 1.6, radioactivity collected during the 90 min following beginning of gravistimulation). There was a close correlation between the development of asymmetric auxin movement across the root cap and the rate of curvature, with both values increasing to a maximum and then declining as the roots approached the final angle of curvature. In roots preadapted to gravity (alternate brief stimulation on opposite flanks over a period of 1 hour) the initial phase of upward curvature was eliminated and downward bending began earlier than for controls. The correlation between asymmetric auxin movement and the kinetics of curvature also held in comparisons between control and preadapted roots. Both downward auxin transport asymmetry and downward curvature occurred earlier in preadapted roots than in controls. These findings are consistent with suggestions that the root cap is not only the site of perception but also the location of the initial redistribution of effectors that ultimately leads to curvature.     

A Role for the TOC Complex in Arabidopsis Root Gravitropism

Arabidopsis (Arabidopsis thaliana) roots perceive gravity and reorient their growth accordingly. Starch-dense amyloplasts within the columella cells of the root cap are important for gravitropism, and starchless mutants such as pgm1 display an attenuated response to gravistimulation. The altered response to gravity1 (arg1) mutant is known to be involved with the early phases of gravity signal transduction. arg1 responds slowly to gravistimulation and is in a genetically distinct pathway from pgm1, as pgm1 mutants enhance the gravitropic defect of arg1. arg1 seeds were mutagenized with ethylmethane sulfonate to identify new mutants that enhance the gravitropic defect of arg1. Two modifier of arg1 mutants (mar1 and mar2) grow in random directions only when arg1 is present, do not affect phototropism, and respond like the wild type to application of phytohormones. Both have mutations affecting different components of the Translocon of Outer Membrane of Chloroplasts (TOC) complex. mar1 possesses a mutation in the TOC75-III gene; mar2 possesses a mutation in the TOC132 gene. Overexpression of TOC132 rescues the random growth phenotype of mar2 arg1 roots. Root cap amyloplasts in mar2 arg1 appear ultrastructurally normal. They saltate like the wild type and sediment at wild-type rates upon gravistimulation. These data point to a role for the plastidic TOC complex in gravity signal transduction within the statocytes.     

Membrane-potential responses following gravistimulation in roots of Lepidium sativum L.

Membrane potentials were measured in lateral statocytes of vertically and nonvertically growing roots of Lepidium sativum L. using conventional glass-microelectrode techniques. Statocytes in vertically growing roots showed a stable resting potential of-118±5.9 mV without spontaneous fluctuations. Upon tilting the root 45° from the vertical, an electrical asymmetry was observed. Statocytes on the physically lower side of the root depolarized by approx. 25 mV. This depolarization occurred following a latent period of 8 s reaching a minimum (approx.-93 mV) after 170 s. This depolarization is the earliest event in graviperception ever recorded. After this depolarization, the cell repolarized within 60 s to a potential approx. 10 mV more positive than the original resting potential. Statocytes on the upper flank showed a slow hyperpolarization (t _1/2h=half time for hyperpolarization=168 s) reaching a final, stable potential at a level 10 mV more negative. These effects of gravistimulation were statenchyma-specific, since cells in the cortex and rhizodermis showed no similar effects. The gravi-electrical responses were observed in 25% of all roots tested. Roots which showed no gravi-electrical response had a reduced elongation growth, lacked gravity-induced bending and lacked the typical structural polarity in punctured statocytes. This observed transition from a symmetrical pattern of resting potential in the statenchyma to an asymmetrical pattern following gravistimulation supports the results observed with external current measurements (Behrens et al., Plant Physiol. 70, 1079–1083, 1982) and extends these results to the cellular level and to considerably improved temporal resolution. The asymmetry in the gravi-electrical response extends the graviperception model of Sievers and Volkmann (Planta 102, 160–172, 1972) which comprises an asymmetrical sedimentation of the amyloplasts on the distal endoplasmic reticulum of statocytes. This generates an intraorgan signal which then must be transmitted to the growth zone.Abbreviation ER endoplasmic reticulum Preliminary reports were presented at the 11^th International Conference on Plant Growth Substances, Aberystwyth, July 1982, and International Symposium, Membranes and Compartimentation in Regulation of Plant Functions, Toulouse, September 1983     

Reduced Gravitropism in Hypocotyls of Starch-Deficient Mutants of Arabidopsis

Gravitropism was examined in dark- and light-grown hypocotylsof wild-type (WT), two reduced starch mutants (ACG 20 and ACG27), and a starchless mutant (ACG 21) of Arabidopsis. In addition,the starch content of these four strains was studied with lightand electron microscopy. Based on time course of curvature andorientation studies, the graviresponse in hypocotyls is proportionalto the amount of starch in a genotype. Furthermore, starch mutationsseem to primarily affect gravitropism rather than differentialgrowth since both phototropic curvature and growth rates amongthe four genotypes are approximately equal. Our results suggestthat gravity perception may require a greater plastid mass inhypocotyls compared to roots. The kinetics of gravitropic curvaturealso was compared following reorientation at 45°, 90°,and 135°. As has been reported for other plant species,the optimal angle of reorientation is 135° for WT Arabidopsisand the two reduced starch mutants, but the magnitude of curvatureof the starchless mutant appears to be independent of the initialangle of displacement. Taken together, the results of the presentstudy and our previous experiments with roots of the same fourgenotypes [Kiss et al. (1996) Physiol. Plant. 97: 237] supporta plastid-based hypothesis for gravity perception in plants. (Received December 16, 1996; Accepted February 7, 1997)     

Gravitropic bending of cress roots without contact between amyloplasts and complexes of endoplasmic reticulum

The polar arrangement of cell organelles in Lepidium root statocytes is persistently converted to a physical stratification during lateral centrifugation (the centrifugal force acts perpendicular to the root long axis) or by apically directed centrifugation combined with cytochalasin-treatment. Lateral centrifugation (10 min, 60 min at 10\g or 50\g) causes displacement of amylplasts to the centrifugal anticlinal cell wall and shifting of the endoplasmic reticulum (ER) complex to the centripetal distal cell edge. After 60 min of lateral centrifugation at 10\g or 50\g all roots show a clear gravitropic curvature. The average angle of curvature is about 40° and corresponds to that of roots stimulated gravitropically in the horizontal position at 1\g in spite of the fact that the gravistimulus is 10-or 50-fold higher. Apically directed centrifugation combined with cytochalasin B (25 g\ml^-1) or cytochalasin D (2.5 g\ml^-1) incubation yields statocytes with the amyloplasts sedimented close to the centrifugal periclinal cell wall and ER cisternae accumulated at the proximal cell pole. Gravitropic stimulation for 30 min in the horizontal position at 1\g and additional 3 h rotation on a clinostat result in gravicurvature of cytochalasin B-treated centrifuged (1 h at 50\g) roots, but because of retarded root growth the angle of curvature is lower than in control roots. Cytochalasin D-treatment during centrifugation (20 min at 50\g) does not affect either root growth or gravicurvature during 3 h horizontal exposure to 1\g relative to untreated roots. As lateral centrifugation enables only short-term contact between the amyloplasts and the distal ER complex at the onset of centrifugation and apically directed centrifugation combined with cytochalasin-treatment even exclude any contact the integrity of the distal cell pole need not necessarily be a prerequisite for graviperception in Lepidium root statocytes.Abbreviations CB cytochalasin B - CD cytochalasin D - ER endoplasmic reticulum - g gravitational acceleration     

Restoration of gravitropic sensitivity in starch-deficient mutants of Arabidopsis by hypergravity

Despite the extensive study of plant gravitropism, there have been few experiments which have utilized hypergravity as a tool to investigate gravisensitivity in flowering plants. Previous studies have shown that starch-deficient mutants of Arabidopsis are less sensitive to gravity compared to the wild-type (WT). In this report, the question addressed was whether hypergravity could restore the sensitivity of starch-deficient mutants of Arabidopsis. The strains examined include a WT, a starchless mutant and a reduced-starch mutant. Vertical orientation studies with dark-grown seedlings indicate that increased centrifugal acceleration improves orientation relative to the acceleration vector for all strains, even the WT. For starchless roots, growth of seedlings under constant 5 g acceleration was required to restore orientation to the level of the WT at 1 g. In contrast, approximately 10 g was required to restore the orientation of the starchless mutant hypocotyls to a WT level at 1 g. Examination of plastid position in root cap columella cells of the starchless mutant revealed that the restoration of gravitropic sensitivity was correlated with the sedimentation of plastids toward the distal cell wall. Even in WT plants, hypergravity caused greater sedimentation of plastids and improved gravitropic capability. Collectively, these experiments support the hypothesis of a statolith-based system of gravity perception in plants. As far as is known, this is the first report to use hypergravity to study the mechanisms of gravitropism in Arabidopsis.     

The action of gravity in agravitropic Zea primary roots: Effect of gravistimulation on the extracellular free-Ca2+ content in the 1-mm apical root tip in the dark

The action of gravity stimulation in darkness was examined in agravitropic primary roots of Zea mays L. (cv. Golden Cross Bantam 70). Contents of diffusible and nitric-acid-extractable Ca²⁺ in 1-mm apical tips of roots gravistimulated in the dark were measured by flowinjection analysis as free Ca²⁺ and bound Ca²⁺, respectively. The free-Ca²⁺ content increased transiently, reaching a maximum 0.5 h after gravistimulation. This transient increase was also observed when gravistimulation was applied by changing the orientation of the roots back from horizontal to vertical again. On the other hand, the bound-Ca²⁺ content decreased transiently following gravistimulation. Furthermore, when the root caps were treated with 10 mM 2-(N-morpholino) ethanesulfonic acid buffer, the elevation of free Ca²⁺ following gravistimulation was prevented. These results indicate that gravity perception and the initial transduction steps proceed in the dark, and moreover that the elevation of free Ca²⁺ brought about by the interaction of Ca²⁺/H ⁺ in the apoplast of root tips may be involved in transmission of the gravity signal.Abbreviations FIA flow-injection analysis - Mes 2-(N-morpholino) ethanesulfonic acid - Pipes 1,4-piperazinediethanesulfonic acid - Quin 2 2-[(2-bis-[carboxymethyl]amino-5-methylphenoxy)methyl]-6-methoxy-8-bis(carboxymethyl) aminoquinoline     

Plastid Distribution in Columella Cells of a Starchless Arabidopsis Mutant Grown in Microgravity

Wild-type and starchless Arabidopsis thaliana mutant seedlings(TC7) were grown and fixed in the microgravity environment ofa U.S. Space Shuttle spaceflight. Computer image analysis oflongitudinal sections from columella cells suggest a differentplastid positioning mechanism for mutant and wild-type in theabsence of gravity. (Received September 24, 1996; Accepted January 21, 1997)