Root system architecture and gravity perception of a mangrove plant,Sonneratia alba J. Smith

We describe the features of the root system and the gravitropism of roots produced bySonneratia alba. The root system consists of four root types with different growth directions: (a) Pneumatophores, which are negatively orthogravitropic and their statocytes are very large (922 μm²) and the statolith is located near the proximal wall, (b) Cable roots and (c) Feeding roots which are both diagravitropic and their statoliths are settled along the longitudinal wall, and (d) Anchor roots which are positively orthogravitropic. The statocyte cells are the smallest (420 μm²) and statoliths settled at the distal wall. We found that all roots with marked gravitropism have statoliths that settle along different walls of the statocyte. This implies that the statoliths sensing of gravity is done by gravity on mass, and that they are denser than surrounding cytoplasm and this position is related to root growth direction. This finding matches the statoliths sediment under the effect of gravity. Irrespective of statolith, position and direction of growth may be stable.     

Mechanotransduction in root gravity sensing cells

The analysis of the dose-response curve of the gravitropic reaction of lentil seedling roots has shown that these organs are more sensitive when they have been grown in microgravity than when they have been grown on a 1 g centrifuge in space before gravistimulation. This difference of gravisensitivity is not due to the volume or the density of starch grains of statoliths, which are about the same in both conditions (1 g or microgravity). However, the distribution of statoliths within the statocyte may be responsible for this differential sensitivity, since the dispersion of these organelles is greater in microgravity than in 1 g. When lentil roots grown in microgravity or in 1 g are stimulated at 0.93 g for 22 min, the amyloplasts sediment following two different trajectories. They move from the proximal half of the statocytes toward the lower longitudinal wall in the microgravity grown sample and from the distal half toward the longitudinal wall in the 1 g grown sample. At the end of the stimulation, they reach a similar position within the statocytes. If the roots of both samples are left in microgravity for 3 h, the amyloplasts move toward the cell centre in a direction that makes an average angle of 40 degrees with respect to the lower longitudinal wall. The actin filaments, which are responsible for this movement, may have an overall orientation of 40 degrees with respect to this wall. Thus, when roots grown in microgravity are stimulated on the minicentrifuge the amyloplasts slide on the actin filaments, whereas they move perpendicular to them in 1 g grown roots. Our results suggest that greater sensitivity of seedling roots grown in microgravity should be due to greater dispersion of statoliths, to better contacts between statoliths and the actin network and to greater number of activated mechanoreceptors. One can hypothesize that stretch activated ion channels (SACs) located in the plasma membrane are responsible for the transduction of gravistimulus. These SACs may be connected together by elements of the cytoskeleton lining the plasma membrane and to the actin filaments. They could be stimulated by the action of statoliths on the actin network and/or on these elements of the cytoskeleton which link the mechanoreceptors (SACs).     

Role of endodermal cell vacuoles in shoot gravitropism

In higher plants, shoots and roots show negative and positive gravitropism, respectively. Data from surgical ablation experiments and analysis of starch deficient mutants have led to the suggestion that columella cells in the root cap function as gravity perception cells. On the other hand, endodermal cells are believed to be the statocytes (that is, gravity perceiving cells) of shoots. Statocytes in shoots and roots commonly contain amyloplasts which sediment under gravity. Through genetic research with Arabidopsis shoot gravitropism mutants, sgr1/scr and sgr7/shr, it was determined that endodermal cells are essential for shoot gravitropism. Moreover, some starch biosynthesis genes and EAL1 are important for the formation and maturation of amyloplasts in shoot endodermis. Thus, amyloplasts in the shoot endodermis would function as statoliths, just as in roots. The study of the sgr2 and zig/sgr4 mutants provides new insights into the early steps of shoot gravitropism, which still remains unclear. SGR2 and ZIG/SGR4 genes encode a phospholipase-like and a v-SNARE protein, respectively. Moreover, these genes are involved in vacuolar formation or function. Thus, the vacuole must play an important role in amyloplast sedimentation because the sgr2 and zig/sgr4 mutants display abnormal amyloplast sedimentation.     

Tip-growing cells of the moss Ceratodon purpureus Are gravitropic in high-density media

Gravity sensing in plants and algae is hypothesized to rely upon either the mass of the entire cell or that of sedimenting organelles (statoliths). Protonemata of the moss Ceratodon purpureus show upward gravitropism and contain amyloplasts that sediment. If moss sensing were whole-cell based, then media denser than the cell should prevent gravitropism or reverse its direction. Cells that were inverted or reoriented to the horizontal displayed distinct negative gravitropism in solutions of iodixanol with densities of 1.052 to 1.320 as well as in bovine serum albumin solutions with densities of 1.037 to 1.184 g cm(-3). Studies using tagged molecules of different sizes and calculations of diffusion times suggest that both types of media penetrate through the apical cell wall. Estimates of the density of the apical cell range from 1.004 to 1.085. Because protonemata grow upward when the cells have a density that is lower than the surrounding medium, gravitropic sensing probably utilizes an intracellular mass in moss protonemata. These data provide additional support for the idea that sedimenting amyloplasts function as statoliths in gravitropism.     

Contributions of space experiments to the study of gravitropism

The study of gravitropism in space has permitted the discovery that statoliths are not completely free to sediment in the gravisensing cells of roots. These organelles are attached to actin filaments via motor proteins (myosin) which are responsible for their displacement from the distal pole of the cell toward the proximal pole when the seedlings are transferred from a 1g centrifuge in space to microgravity. On the ground, the existence of the link between the statoliths and the actin network could not be established because the gravity force is much greater than the force exerted by the motor proteins. This finding led to a new hypothesis on gravisensing. It has been proposed that statoliths can exert tensions in the actin network which become asymmetrical when the root is stimulated in the horizontal position on the ground. The space experiments have confirmed to some extent the results obtained on gravisensitivity with clinostats, although these devices do not simulate microgravity correctly. Reexamination of the means of estimating gravisensitivity has led to the conclusion that the perception and the transduction phases could be very short (that is, within a second). This data is consistent with the fact that the statoliths are attached to the actin filament and do not have to move a long distance to exert forces on the actin network. It has also been demonstrated that gravity regulates the gravitropic bending in order to avoid the overshooting of the vertical direction on the ground. The roots, which are stimulated and placed in microgravity, are not subjected to this regulation and curve more than roots stimulated continuously. However, the curvature of roots or of coleoptiles that takes place in microgravity can be greatly reduced by straightening the extremity of the organs.     

The RHG Gene Is Involved in Root and Hypocotyl Gravitropism in Arabidopsis thaliana

In higher plants, shoots show negative gravitropism and rootsshow positive gravitropism. To elucidate the molecular mechanismsof root and hypocotyl gravitropism, we segregated the secondmutation from the original phyB-1 mutant line which impairedboth root and hypocotyl gravitropism and characterized thisnovel mutation named rhg (for root and hyzypocotyl gravitropism).The rhg is a single recessive nuclear mutation and it is mappedon the lower part of the chromosome 1. Analyses on the gravitropicresponses of the rhg mutant indicate that root and hypocotylgravitropism are severely impaired but inflorescence stem gravitropismis not affected by the rhg mutation. In the rhg mutant seedlings,amyloplasts (statoliths for gravity-perception) were presentin the presumptive statocytes of roots and hypocotyls. Phototropismby roots and hypocotyls was not impaired in the rhg mutant.These results suggest that the RHG gene product probably actson the gravity-perception and/or the gravity-signal transductionin root and hypocotyl gravitropism. This is the first reportabout the genetic locus specifically involved in both root andhypocotyl gravitropism but not inflorescence stem gravitropism,supporting our hypothesis that the mechanisms of gravitropismare genetically different between hypocotyls and inflorescencestems. (Received March 11, 1997; Accepted April 17, 1997)     

The Mechanism of Geoperception in Lentil Roots

The sediment of the amyloplasts in the statocytes of lentilroots was analysed in two different samples of seedlings. Theroots of the first sample (HP) were grown in a horizatal position,whereas thoes of the second sample (VP+20) were grown in a horizontalposition and then exposure to give a response). It is demonstratedthat the statolish can almost enter into contact with the plasmamembrane lining the longitudinal wall of the statocytes of theHP sample. However, these organelles in the VP+20 sample areable to provoke a geotropic stimulation though they are sedimentedat a distance of 0.1350–0.2600 µm from the plasmalemma.From these data it is concluded that the cytoplasmic structureswhich may play a role in the geotropic stimulation are: (1)the endoplasmic reticulum located along the longitudianl wallof the statocytes and (2) the microtubules or the plasmalemmaif the action of the statoliths is transmitted by the parietalcytoplasm. The large aggregations of endoplasmic reticulum whichare situated in the distal part of the central root cap cellswould not have any role in the perception of gravity by roots.These results are discussed in the light of recent work showingthe action of a growth inibitor in the geotropic reaction ofroots.     

Early development and gravitropic response of lateral roots in Arabidopsis thaliana

Root system architecture plays an important role in determining nutrient and water acquisition and is modulated by endogenous and environmental factors, resulting in considerable developmental plasticity. The orientation of primary root growth in response to gravity (gravitropism) has been studied extensively, but little is known about the behaviour of lateral roots in response to this signal. Here, we analysed the response of lateral roots to gravity and, consistently with previous observations, we showed that gravitropism was acquired slowly after emergence. Using a lateral root induction system, we studied the kinetics for the appearance of statoliths, phloem connections and auxin transporter gene expression patterns. We found that statoliths could not be detected until 1 day after emergence, whereas the gravitropic curvature of the lateral root started earlier. Auxin transporters modulate auxin distribution in primary root gravitropism. We found differences regarding PIN3 and AUX1 expression patterns between the lateral root and the primary root apices. Especially PIN3, which is involved in primary root gravitropism, was not expressed in the lateral root columella. Our work revealed new developmental transitions occurring in lateral roots after emergence, and auxin transporter expression patterns that might explain the specific response of lateral roots to gravity.     

A Morphometric Analysis of the Redistribution of Organelles in Columella Cells of Horizontally-oriented Roots of Zea mays

In order to determine what structural changes in graviperceptivecells are associated with the onset of root gravicurvature,the redistribution of organelles in columella cells of horizontally-oriented,graviresponding roots of Zea mays has been quantified. Rootgravicurvature began by 15 min after reorientation, and didnot involve significant changes in the (i) volume of individualcolumella cells or amyloplasts, (ii) relative volume of anycellular organelle, (iii) number of amyloplasts per columellacell, or (iv) surface area or cellular location of endoplasmicreticulum. Sedimentation of amyloplasts began within 1 to 2min after reorientation, and was characterized by an intenselystaining area of cytoplasm adjacent to the sedimenting amyloplasts.By 5 min after reorientation, amyloplasts were located in thelower distal corner of columella cells, and, by 15 min afterreorientation, overlaid the entire length of the lower cellwall. No consistent contact between amyloplasts and any cellularstructure was detected at any stage of gravicurvature. Centrally-locatednuclei initially migrated upward in columella cells of horizontally-orientedroots, after which they moved to the proximal ends of the cellsby 15 min after reorientation. No significant pattern of redistributionof vacuoles, mitochondra, dictyosomes, or hyaloplasm was detectedthat correlated with the onset of gravicurvature. These resultsindicate that amyloplasts and nuclei are the only organelieswhose movements correlate positively with the onset of gravicurvatureby primary roots of this cultivar of Zea mays. Zea mays, root gravitropism, ultrastructure, morphometry, graviperception     

Lentil root statoliths reach a stable state in microgravity

 The kinetics of the movement of statoliths in gravity-perceiving root cap cells of Lens culinaris L. and the force responsible for it have been analysed under 1 g and under microgravity conditions (S/MM-03 mission of Spacehab 1996). At the beginning of the experiment in space, the amyloplasts were grouped at the distal pole of the statocytes by a root-tip-directed 1-g centrifugal acceleration. The seedlings were then placed in microgravity for increasing periods of time (13, 29, 46 or 122 min) and chemically fixed. During the first 29 min of microgravity there were local displacements (mean velocity: 0.154 μm min⁻¹) of some amyloplasts (first at the front of the group and then at the rear). Nevertheless, the group of amyloplasts tended to reconstitute. After 122 min in microgravity the bulk of amyloplasts had almost reached the proximal pole where further movement was blocked by the nucleus. After a longer period in microgravity (4 h; experiment carried out 1994 during the IML 2 mission) the statoliths reached a stable position due to the fact that they were stopped by the nucleus. The position was similar to that observed in roots grown continuously in microgravity. Treatment with cytochalasin D (CD) did not stop the movement of the amyloplasts but slowed down the velocity of their displacement (0.019 μm min⁻¹). Initial movement patterns were the same as in control roots in water. Comparisons of mean velocities of amyloplast movements in roots in space and in inverted roots on earth showed that the force responsible for the movement in microgravity (F_c) was about 86% less (F_c = 0.016 pN) than the gravity force (F_g = 0.11 pN). Treatment with CD reduced F_c by two-thirds. The apparent viscosity of the statocyte cytoplasm was found to be 1 Pa s or 3.3 Pa s for control roots or CD treated roots, respectively. Brownian motion or elastic forces due to endoplasmic reticulum membranes do not cause the movement of the amyloplasts in microgravity. It is concluded that the force transporting the statoliths is caused by the actomyosin system. Received: 22 March 1999 / Accepted: 18 December 1999     

水稻无侧根突变体的根向重力性异常

用化学诱变剂 (NaN3 )处理粳稻品种大力 (O ryzasativaL .cv .Oochikara) ,得到具有 2 ,4 D抗性、无侧根和根向重力性异常的突变体RM 10 9。对原品种为父本和突变体为母本的杂交后代F1、F2 根向重力性的遗传分离进行了研究。结果表明 :突变体的根向重力性异常 ,其性状是单显性基因控制且不受光照和黑暗培养的影响。通过对根冠组织切片观察发现 :突变体根冠中含淀粉体的细胞数量比大力少 ,根冠细胞中淀粉体的直径为原品种的 5 0 %且集中排列于细胞内的一角 ,其排列沉积方向与重力方向相同。推测 :突变体的根向重力性异常与淀粉体直径变小有关     

Amyloplasts are necessary for full gravitropic sensitivity in roots of Arabidopsis thaliana

The observation that a starchless mutant (TC7) of Arabidopsis thaliana (L.) Heynh. is gravitropic (T. Caspar and B.G. Pickard, 1989, Planta 177, 185–197) raises questions about the hypothesis that starch and amyloplasts play a role in gravity perception. We compared the kinetics of gravitropism in this starchless mutant and the wild-type (WT). Wild-type roots are more responsive to gravity than TC7 roots as judged by several parameters: (1) Vertically grown TC7 roots were not as oriented with respect to the gravity vector as WT roots. (2) In the time course of curvature after gravistimulation, curvature in TC7 roots was delayed and reduced compared to WT roots. (3) TC7 roots curved less than WT roots following a single, short (induction) period of gravistimulation, and WT, but not TC7, roots curved in response to a 1-min period of horizontal exposure. (4) Wild-type roots curved much more than TC7 roots in response to intermittent stimulation (repeated short periods of horizontal exposure); WT roots curved in response to 10 s of stimulation or less, but TC7 roots required 2 min of stimulation to produce a curvature. The growth rates were equal for both genotypes. We conclude that WT roots are more sensitive to gravity than TC7 roots. Starch is not required for gravity perception in TC7 roots, but is necessary for full sensitivity; thus it is likely that amyloplasts function as statoliths in WT Arabidopsis roots. Furthermore, since centrifugation studies using low gravitational forces indicated that starchless plastids are relatively dense and are the most movable component in TC7 columella cells, the starchless plastids may also function as statoliths.Abbreviations S2 story two - S3 story three - WT wild-type     

Graviperception of lentil seedling roots grown in space (Spacelab Dl Mission)

The growth and graviresponsiveness of roots were investigated in lentil seedlings (Lens culinaris L. cv. Verte du Puy) grown (1) in microgravity, (2) on a 1 g centrifuge in space, (3) in microgravity and then placed on the 1 g centrifuge for 3 h, (4) on the ground. Dry seeds were hydrated in space (except for the ground control) and incubated for 25 h at 22°C in darkness. At the end of the experiment, the seedlings were photographed and fixed in glutaraldehyde in a Biorack glove box. Root length was similar for seedlings grown in space and for the ground and the 1 g centrifuge controls. The direction of root growth in the microgravity sample deviated strongly from the initial orientation of the roots of the dry seeds. This deviation could be due to spontaneous curvatures similar to those observed on clinostats. When lentil seedlings were first grown in microgravity for 25 h and then placed on the 1 g centrifuge for 3 h, their roots bent strongly under the effect of the centrifugal acceleration. The amplitude of root curvature on the centrifuge was not significantly different from that observed on ground controls growing in the vertical position and placed in the horizontal position for 3 h. The gravisensitivity of statocytes differentiated in microgravity was similar to that of statocytes differentiated on earth. There were no qualitative differences in the ultrastructural features of the gravisensing cells in microgravity and in the 1 g centrifuge and ground controls. However, the distribution of statoliths in the gravisensing cells was different in microgravity: most of them were observed in the proximal part of these cells. Thus, these organelles were not distributed at random, which is in contradiction with results obtained with clinostats. The distal complex of endoplasmic reticulum in the statocytes was not in contact with the amyloplasts. Contact and pressure of amyloplasts on the tubules were not prerequisites for gravisensing. The results obtained are not in agreement with the hypothesis that the distal endoplasmic reticulum would be the transducer of the action of the statoliths.     

Hydrotropism interacts with gravitropism by degrading amyloplasts in seedling roots of Arabidopsis and radish

In response to a moisture gradient, roots exhibit hydrotropism to control the orientation of their growth. To exhibit hydrotropism, however, they must overcome the gravitropism that is dominant on Earth. We found that moisture gradient or water stress caused immediate degradation of the starch anchors, amyloplasts, in root columella cells of Arabidopsis and radish (Raphanus sativus). Namely, development of hydrotropic response was accompanied by a simultaneous reduction in starch content in columella cells. Rapid degradation of amyloplasts in columella cells also occurred in the water-stressed roots with sorbitol or mannitol. Both hydrotropically stimulated and water-stressed roots showed a reduced responsiveness to gravity. Roots of a starchless mutant, pgm1-1, showed an enhanced hydrotropism compared with that of the wild type. These results suggest that the reduced responsiveness to gravity is, at least in part, attributable to the degradation of amyloplasts in columella cells. Thus, the reduction in gravitropism allows the roots to exhibit hydrotropism.     

Roles of amyloplasts and water deficit in root tropisms

Directed growth of roots in relation to a moisture gradient is called hydrotropism. The no hydrotropic response (nhr1) mutant of Arabidopsis lacks a hydrotropic response, and shows a stronger gravitropic response than that of wild type (wt) in a medium with an osmotic gradient. Local application of abscisic acid (ABA) to seeds or root tips of nhr1 increased root downward growth, indicating the critical role of ABA in tropisms. Wt roots germinated and treated with ABA in this system were strongly gravitropic, even though they had almost no starch amyloplasts in the root-cap columella cells. Hydrotropically stimulated nhr1 roots, with or without ABA, maintained starch in the amyloplasts, as opposed to those of wt. Hence, the near-absence (wt) or abundant presence (nhr1) of starch granules does not influence the extent of downward gravitropism of the roots in an osmotic gradient medium. Starch degradation in the wt might help the root sustain osmotic stress and carry out hydrotropism, instead of reducing gravity responsiveness. nhr1 roots might be hydrotropically inactive because they maintain this starch reserve in the columella cells, sustaining both their turgor and growth, and in effect minimizing the need for hydrotropism and at least partially disabling its mechanism. We conclude that ABA and water stress are critical regulators of root tropic responses.     

Root gravitropism: a complex response to a simple stimulus?

Roots avoid depleting their immediate environment of essential nutrients by continuous growth. Root growth is directed by environmental cues, including gravity. Gravity sensing occurs mainly in the columella cells of the root cap. Upon reorientation within the gravity field, the root-cap amyloplasts sediment, generating a physiological signal that promotes the development of a curvature at the root elongation zones. Recent molecular genetic studies in Arabidopsis have allowed the identification of genes that play important roles in root gravitropism. Among them, the ARG1 gene encodes a DnaJ-like protein involved in gravity signal transduction, whereas the AUX1 and AGR1 genes encode proteins involved in polar auxin transport. These studies have important implications for understanding the intra- and inter-cellular signaling processes that underlie root gravitropism.     

Agravitropic behaviour of roots of rapeseed (Brassica napus L.) transformed by Agrobacterium rhizogenes

Transgenic hairy roots of Brassica napus (cv. Omega) have been developed, using Agrobacterium rhizogenes strain AR 25, for use as a model system in the investigation of physiological and morphological differences between transgenic and normal roots. The basic parameters of growth and normal or altered gravitropical behaviour of hairy roots are for the first time presented in this paper together with an ultrastructural and morphological analysis of the root statocytes. The results obtained also represented the basis for the TRANSF0RM-experiment on the IML-2 mission performed onboard the Space Shuttle Columbia. Typical hairy root traits such as hormone-autonomous growth high growth rate, lateral branching, and changed/absence of gravitropism were detected. The transformed nature of the roots was confirmed by Southern blot analyses. The gravitropical behaviour of apices from hairy root cultures of this clone has been compared with root tips from normal seedlings. While the wild type roots curved progressively with increasing stimulation angles, the transformed roots showed no curvature when stimulated at 45 degrees, 90 degrees or 135 degrees on the ground. The morphology and ultrastructure of the root tip regions were examined by light microscopy and transmission electron microscopy. At the ultrastructural level no major differences could be detected between the roots studied. There was, however, a slight reduction in the starch content of most of the amyloplasts of the transgenic root tips, and the root cap was more V-shaped in the transgenic roots than in the wild type. Preliminary results from the Shuttle experiment TRANSFORM show a random distribution of amyloplasts in the root cells of both transformed and wild type root caps after 14 h on a 1xg centrifuge followed by 37 h in microgravity.     

A Morphometric Analysis of the Redistribution of Organelles in Columella Cells in Primary Roots of Normal Seedlings and Agravitropic Mutants of Hordeum vulgare

Moore, R. 1985. A morphometric analysis of the redistributionof organellcs in columella cells in primary roots of normalseedlings and agravitropic mutants of Hordeum vulgare.—J.exp. Bot. 36:1275–1286. The redistribution of organeUes m columella cells of horizontally-orientedroots of Hordeum vulgare was quantified in order to determinewhat structural changes in graviperceptive (i.e, columella)cells are associated with the onset of root gravicurvature.The sedimentation of amyloplasts is the only major change incellular structure that correlates positively with the onsetof root gravicurvature, which begins within 15 min after re-orientation.There is no consistent contact between sedimented amyloplastsand any other organelles. Nuclei are restricted to the proximalends of columella cells in vertically-oriented roots, and remainthere throughout gravicurvature after roots are oriented horizontally.Root gravicurvature does not involve significant changes in(1) the volume of columella cells, (2) the relative or absolutevolumes of organelles in columella cells, or (3) the distributionof endoplasmic reticulum (ER). The size, number and sedimentationrates of amyloplasts in columella cells of non-graviresponsiveroots of mutant seedlings are not significantly different fromthose of graviresponsive roots of normal seedlings. Similarly,there is no significant difference in (1) cellular volume, (2)distribution or surface area of ER, (3) patterns or rates oforganelle redistribution in horizontally-oriented roots, or(4) relative or absolute volumes of organelles in columellacells of graviresponsive and non-graviresponsive roots. Theseresults suggest that the lack of gravi-responsiveness by rootsof mutant seedlings is probably not due to either (1) structuraldifferences in columella cells, or (2) differences in patternsor rates of organelle redistribution as compared to that characteristicof graviresponsive roots. Thus, the basis of non-graviresponsivenessin this mutant is probably different from other agravitropicmutants so far studied. Key words: Agravitropic mutant, barley, columella cell, gravitropism (root), Hordeum vulgare, ultrastructure     

Gravity-regulated formation of the peg in developing cucumber seedlings

It has been proposed that peg formation in the vascular transition region (TR zone) between the hypocotyl and the root in Cucurbitaceae seedlings is a gravimorphogenetic phenomenon. Initiation of the peg became visible 36 h after imbibition when cucumber (Cucumis sativus L. cv. Burpee Hybrid II) seeds were germinated in a horizontal position at 24°C in the dark. Simultaneously, sedimented amyloplasts (putative statoliths) were apparent in the sheath cells surrounding the vascular strands, and in the cortical cells immediately adjacent to them, in the TR zone. In contrast, the other cortical cells, some of which were destined to develop into the peg, contained amyloplasts which were not sedimented. These results suggest that the graviperception mechanism for peg formation may be like that of statoliths in shoot gravitropism. By 48 h following imbibition, the cells of the TR zone still had sedimented amyloplasts but had lost their sensitivity to gravity, possibly because of their maturation.     

Root Graviresponsiveness and Cellular Differentiation in Wild-type and a Starchless Mutant of Arabidopsis thaliana

Primary roots of a starchless mutant of Arabidopsis thalianaL. are strongly graviresponsive despite lacking amyloplastsin their columella cells. The ultrastructures of calyptrogenand peripheral cells in wild-type as compared to mutant seedlingsare not significantly different. The largest difference in cellulardifferentiation in caps of mutant and wild-type roots is therelative volume of plastids in columella cells. Plastids occupy12.3% of the volume of columella cells in wild-type seedlings,but only 3.69% of columella cells in mutant seedlings. Theseresults indicate that: (1) amyloplasts and starch are not necessaryfor root graviresponsiveness; (2) the increase in relative volumeof plastids that usually accompanies differentiation of columellacells is not necessary for root graviresponsiveness; and (3)the absence of starch and amyloplasts does not affect the structureof calyptrogen (i.e. meristematic) and secretory (i.e. peripheral)cells in root caps. These results are discussed relative toproposed models for root gravitropism. Arabidopsis thaliana, gravitropism (root), plastids, root cap, stereology, ultrastructure