A role of microtubules in the polarity of statocytes from roots of Lepidium sativum L.

When roots of Lepidium sativum L. are immersed in a colchicine solution (10^-4 mol l^-1), the cortical microtubules of statocytes are affected such that the dense network ofmicrotubules at the distal cell edges, between the endoplasmic reticulum and the plasma membrane, disappears almost completely, whereas the microtubules, lining the anticlinal cell walls are reduced only to a limited extent. Upon inversion of colchicine-pretreated roots, the distal complex of endoplasmic reticulum sinks into the interior of the statocyte. Germination of seeds in the cold (3–4°C) leads to a retardation of statocyte development; the elaborated system of endoplasmic reticulum is lacking, and only a few microtubules are observable, lining the plasma membrane along the anticlinal cell walls. During an additional 4 h at 24°C, groups of microtubules develop near the plasma membrane in the distal one-third of the statocytes, coaligning with newly synthesized cisternae of the endoplasmic reticulum. It is proposed that, particularly at the distal statocyte pole, microtubules in coordination with cross-bridging structures, act in stabilizing the polar arrangement of the distal endoplasmic reticulum and, in turn, facilitate an integrated function of amyloplasts, endoplasmic reticulum and plasma membrane in graviperception.Abbreviations ER endoplasmic reticulum - MT microtubule     

Restitution of polarity in statocytes from centrifuged roots*

Abstract The structural polarity of statocytes from cress roots is changed by centrifugation. Upon low- dose centrifugation (3000 g min), the extent of stratification depends on statocyte position, i.e., central statocytes are affected more than lateral ones. Upon higher doses of centrifugation (60,000 and 360,000 g min), a uniform density gradient is established in all statocytes. If, after centrifugation, the roots are exposed to gravity again, the endoplasmic reticulum (ER) cisternae are relocated parallel to the periclinal cell walls within a few minutes; this relocation is independent of the direction of gravity in relation to the root axis, and independent of the previously applied centrifugation dose. This supports the notion that polarity is determined genetically. Cytochalasin B treatment, before and during centrifugation, totally inhibits the relocation of ER. After removing the drug by rinsing the roots, the statocytes restore cell polarity and relocate ER. These results indicate that relocation of ER cisternae may be mediated by microfilaments. When centrifuged roots are exposed to 1 g in the horizontal position, the latent period of gravitropism increases by 8–10 min relative to controls, regardless of the previously applied centrifugation doses. The kinetics of curvature are virtually identical. Since the increase in the latent period coincides with the time needed for most statocytes to restore the distal cell pole, it is evident that perception of gravity is correlated to the integrity of the distal cell pole.     

Effects of prolonged omnilateral gravistimulation on the ultrastructure of statocytes and on the graviresponse of roots

Statocytes of vertically growing roots of Lepidium sativum L. exhibit a strict polarity: The nucleus is positioned near the proximal periclinal cell wall, amyloplasts are sedimented on a complex of rough endoplasmic reticulum (ER) consisting of parallel cisternae near the distal periclinal cell wall.When 24 h old, vertically grown roots are rotated for an additional 20 h on a horizontal clinostat, this polarity is destroyed. Furthermore, the prolonged omnilateral stimulation leads to a damage of the statocytes, which in some cases ends in the self-destruction of the sensitive cells. The different components of the ultrastructural respones of the statocytes are: Displacement of the nucleus; changes in amount and distribution of the ER; loss of amyloplast starch; confluence of lipid droplets to large aggregates: a considerable increase of the lytic compartment. In addition, even anticlinal cell walls may be lysed up to small stumps. As all these effects are clearly restricted to the statocytes, only these cells are able to respond to the continuously changing direction of the gravity vector, thus perceiving gravity as such.After being exposed horizontally, the graviresponse of rotated roots is delayed as compared to the controls. About 20% of the rotated roots do not respond (curve) at all, but grow perpendicular in relation to the gravity vector. Perception of gravity is inevitably correlated with the polarity and the integrity of the statocytes.Abbreviation ER endoplasmic reticulum A preliminary report was presented at the Fall Meeting of the German Society for Cell Biology in Salzburg, Austria, September 1979 (Hensel and Sievers 1979)This paper represents part of a dissertation (D 5) of W. H.     

Cytochalasin B affects the structural polarity of statocytes from cress roots (Lepidium sativum L.)

Summary The effect of cytochalasin B (CB; 25 ·ml^–1 in 1% dimethylsulfoxide, DMSO) upon the structural polarity of statocytes in cress roots is demonstrated. If normal, vertically grown roots are incubated in CB, the structural polarity of the statocytes is altered according to the developmental stage of the root. Statocytes from young roots (13 or 17 hours, additionally 7 hours CB) are characterized by proximal ER cisternae and a sparsely developed distal ER-complex. Statocytes from older roots (24 hours, additionally 7 hours CB) still accumulate distal ER, as in control roots, but at the proximal cell pole in the vicinity of the nucleus additional ER is found. These effects are reversed by washing out the drug in DMSO. Growth of the roots under a continuous supply of CB yields statocytes with sedimented nuclei, proximal ER and almost no distal ER. Together with quantitative data from morphometric studies, a dynamic model of the expression of inherent cell polarity in structural polarity is proposed.Abbreviations CB cytochalasin B - DMSO dimethylsulfoxide - ER endoplasmic reticulum Preliminary results were presented at the joint Annual Meeting of the Belgian and German Society for Cell Biology, Bonn, 18–22 March 1985; Eur. J. Cell Biol. 36 (Suppl. 7), 1985, 25.Dedicated to Professor Dr. A.Betz on the occasion of his 65th birthday.     

Cytodifferentiation of polar plant cells Use of anti-microtubular agents during the differentiation of statocytes from cress roots (Lepidium sativum L.)

The development of the structural polarity of statocytes from cress roots (Lepidium sativum L.) was studied in a time- and stage-dependent manner. Outgrowing radicles had statocytes with abundant lipid droplets, sparsely developed endoplasmic reticulum (ER) and nuclei located at the proximal cell poles. During differentiation, coincidentally the lipid droplets disappeared, while rough ER increased in length. The ER was translocated into the distal cell pole to establish a complex of stacked ER. Microtubules occurred first at the distal cell edges. As a second step, ER was produced in the vicinity of the nucleus and was also translocated distally. By application of the antimicrotubular agents heavy water (90%), colchicine (10^-4 mol·l^-1) and triethyl lead chloride (20 mol·l^-1), the involvement of microtubules in these events was studied. Triethyl lead chloride led to a complete cessation of differentiation; root-cap cells remained at a stage without polar arrangement of the ER. Colchicine affected the development of structural polarity slightly, as shown by a higher density of cortical ER cisternae. Heavy water inhibited the translocation of ER almost completely and yielded ER located also in the cell center. All anti-microtubular agents inhibited cell division and the differentiation of the distal cell layer of the dermatocalyptrogen into statocytes. It is hypothesized that microtubules serve as anchoring sites for microfilaments, which actually mediate the translocation of the ER. Hence, an intact system of microtubules and microfilaments is necessary for the expression of structural polarity.Abbreviations DC dermatocalyptrogen - ER endoplasmic reticulum - M meristem cell layer - MT microtubule - pI prospective story I - TrEl triethyl lead chloride     

Polarity of statocytes in lentil seedling roots grown in space (Spacelab D1 Mission)

A morphometric analysis of root statocytes was performed on seedlings of lentil ( Lens culinaris L., cv. Verte du Puy) in order to determine the effects of microgravity on the polarity of these cells. Seedlings were grown: (1) on the ground, (2) in microgravity, (3) on a 1 g centrifuge in space, (4) first in microgravity and then placed on a 1 g centrifuge for 3 h. Dry seeds were hydrated in space (except for the ground control) for 25 h in darkness at 22°C in the Biorack facility developed by the European Space Agency. At the end of the experiment, the seedlings were photographed and fixed in glutaraldehyde in the Biorack glove box. The average shape of the statocytes and the location of endoplasmic reticulum, amyloplasts and nucleus in the cells were analysed in the four samples. By considering the cell shape, it appears that the morphology of the statocytes on the ground was different from that observed in the space samples. Cell polarity was similar in microgravity and in the centrifuged samples except for the distribution of the amyloplasts. These organelles were not distributed at random in near zero gravity, and they were more numerous in the proximal than in the distal half. Moreover, the statoliths were more voluminous in microgravity than in the centrifuged samples. The nucleus was closer to the cell center in the statocytes of roots grown in microgravity than in statocytes of roots grown in microgravity and then placed on the 1 g centrifuge for 3 h. It is hypothesized that the nucleus is attached to the cell periphery and that its location is dependent upon gravity.     

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     

The effect of uni-axial clinostat rotation on germination and root anatomy of Phaseolus vulgaris L.

ABSTRACT

Phaseolus vulgaris L. seed germination and seedling root anatomy were investigated on a slowly rotating clinostat in 1g. Clinostat rotating seeds were oriented as follows: the first group with the longer axis parallel to the rotation pole (horizontal), the other with the longer axis normal to the rotation pole with due attention to the position of the root apex primordium in the dry seeds (vertical). Germination time, percent germination and curvature of developing roots were monitored. Furthermore, the anatomy of the root apex was quantitatively analysed. Seeds placed on the clinostat germinated earlier than controls, and columella cells of roots developed while rotating lost the strict polarity with the nucleus positioned near the proximal periclinal cell wall and amyloplasts sedimented on the distal periclinal wall. Irrespective of seed orientation on the rotation axis, loss of cell polarity occurred as well as a decrease in starch content, modification in cell size, and damage to statocytes whose walls appeared partially digested. Cell size in the elongation zone was also larger in roots rotating on the clinostat than in controls, both in vertically and horizontally placed specimens. Our results demonstrate that prolonged rotation has an effect on the statocyte that continuously perceives gravity from ever-changing directions, although this effect is irrespective of seed position on the rotating axis in P. vulgaris.     

Induction of gravity-dependent plasmatic responses in root statocytes by short time contact between amyloplasts and the distal endoplasmic reticulum complex

Statocytes from roots of Lepidium sativum L., which developed after a 2-min soaking on a horizontal clinostat (2 rotations per min) for 44 h, exhibit the same polarity as in vertically grown roots, as indicated by a complex of endoplasmic reticulum (ER) cisternae at the distal cell pole. Amyloplasts are distributed randomly. The kinetics of graviresponse (=curvature) of such roots are identical to those of normally grown roots. Ten-minute exposure of the root, after 24 h development on the clinostat with gravity acting towards the root's basis (inversion), induces no changes in statocyte ultrastructure. However, corresponding exposure in normal orientation leads to subsequent disintegration of the distal ER complex, loss of amyloplast starch, confluence of lipid droplets, and an increase of the lytic compartment. These ultrastructural events thus appear to be induced by a physical contact — however short — between amyloplasts and the distal ER complex.Abbreviation ER endoplasmic reticulum Dedicated to Professor Dr. W. Haupt on the occasion of his 60th birthday     

The polarity of statocytes and the gravisensitivity of roots are dependent on the concentration of calcium in statocytes

In order to examine a possible role of calcium in graviperception, the calcium ionophore A23187 was used to elevate the concentration of free cytoplasmic calcium in statocytes of the roots of Lepidium sativum L. After a brief incubation (30 min) in a medium that contained 10 micromoles A23187 and 5.5 micromoles CaCl2, 50% of the roots bent gravitropically during a subsequent 2 h of horizontal exposure, with an angle of curvature that varied from 5 degrees to 70 degrees. The corresponding statocytes exhibited a polar arrangement of cell organelles as did the controls. However, in statocytes from 50% of the roots which were not curved after gravistimulation a portion of the distal endoplasmic reticulum (ER) complex was displaced in the direction of gravity within 30 min of horizontal exposure. After washing of the briefly treated roots for 24 h with 1% dimethylsulfoxide the percentage of gravitropically bending roots increased to approximately 80%, but the angle of curvature amounted to only 5 degrees-10 degrees. Longer treatment (2 h) with A23187 caused a complete loss of graviresponsiveness which was accompanied by disintegration of statocyte polarity. We concluded from these results that i) calcium is involved in graviperception and ii) gravisensitivity depends on the integrity of statocytes.     

The effect of centrifugal accelerations on the polarity of statocytes and on the graviperception of cress roots

The structural polarity of statocytes of Lepidium sativum L. is converted to a physical stratification by a root-tip-directed centrifugal acceleration. Sedimentation of amyloplasts and nucleus to the centrifugal (distal) cell pole and the lateral displacement of the distal endoplasmic reticulum (ER) complex occur after centrifugation for 20 min at an acceleration of 50 g. With higher doses (20 min, 100-2,000 g), smaller organelles become increasingly displaced. From the centrifugal to the centripetal cell pole, the following stratification is observed: 1) amyloplasts with mitochondria; 2) nucleus with mitochondria and a few dictyosomes, as well as laterally located ER; 3) dictyosomes with a few mitochondria; 4) vacuoles; and 5) lipid droplets. Within the first 7.5 min, after the roots have been returned to 1 g, the original arrangement of the amyloplasts sedimented on the underlying ER complex is reestablished in 66% of the statocytes. When roots previously centrifuged in an apical direction are exposed in a horizontal position to 1 g, the latent period of the graviresponse is increased by 7.5 min relative to the non-centrifuged controls. The kinetics of the response are identical to the controls. Roots centrifuged first in an apical direction and then for 2 h in a lateral direction (1,000 g) have statocytes with a physical stratification perpendicular to the root axis. A gravitropic curvature does not take place during the lateral centrifugation. These results support the hypothesis that the distal ER complex is necessary and sufficient for graviperception.     

A role for gibberellic acid in orienting microtubules and regulating cell growth polarity in the maize root cortex

The role of gibberellins and cortical microtubules in determining the polarity of cell growth in the root cortex of maize (Zea mays L.) was examined. Inhibition of gibberellin biosynthesis, either naturally through mutation (d5 mutant) or by means of chemicals such as 2S,3S paclobutrazol, caused thickening of root apices and increased their starch content. Immunofluorescence microscopy of cortical microtubules, coupled with a comparison of cell widhts, lengths and shapes, indicated that the meristem and immediate post-mitotic zone were the targets of gibberellin deficiency. Cortical cells in these regions were impaired in their ability to develop highly ordered transversal arrays of cortical microtubules. Consequently, the cells became wider and shorter. Application of gibberellic acid re-established the arrangements of cortical microtubules and the polarity of cell growth characteristic for roots having normal levels of gibberellins, it also decreased the starch content. These results indicate that gibberellins are morphogenetically active substances, not only in shoots but also in roots of maize.Abbreviations CMT cortical microtubule - GA gibberellin - GA₃ gibberellic acid - MT microtubule - PIG postmitotic isodiametric growth The authors acknowledge the support to F.B. from the Royal Society (London UK). We also thank Dr. J. Lenton (University of Bristol, Long Ashton Research Station) who kindly supplied us with 2S,3S paclobutrazol and grains of the GA-deficient d5 mutant of maize.     

Tissue slices from living root caps as a model system in which to study cytodifferentiation of polar cells

Tissue slices of living root caps of cress (Lepidium sativum L.), two to three cell layers in thickness, were prepared by a microsurgical procedure. The viability, cellular structures and cytoplasmic movement of the cells were examined in the light microscope. Nuclei, amyloplasts, vacuoles and endoplasmic reticulum were identified and their positions confirmed after fixation and observation of the same cells in the electron microscope. The distribution of microtubules was shown by immunocytochemistry. During germination, microtubules appear first at the distal edges of the statocytes, while in mature statocytes a distal domain of criss-crossed microtubules could be distinguished from a proximal domain with transversally oriented microtubules. Microfilaments in young statocytes form a nuclear enclosure; in mature statocytes bundles of microfilaments fan out into the cell cortex. The transition from statocytes to secretion cells is accompanied by a more pronounced cortical network of microfilaments, while the nucleus-associated microfilaments remain visible. It is suggested that these microfilaments play a role in the positioning of the nucleus and the translocation of endoplasmic reticulum.Abbreviations ER endoplasmic reticulum - MF microfilament - MT microtubule     

Changes in calcium signalling, gravitropism, and statocyte ultrastructure in pea roots induced by calcium channel blockers

The effect of Ca2+ channel blockers (D600 and nicardipine) were investigated in our experiments on 5-day seedlings of pea. Nicardipine had more inhibiting effect on root elongation than D600. The Ca2+ channel blockers (CCB) depressed gravitropic response in roots. In root statocytes, the destruction of the polar arrangement of cell organelles and other changes were induced by 10(-5) M D600 or nicardipine treatment for 12 h. At ultrastructural level, there were observed a lack of polarity, pronounced vacuolization, and changes in dictyosome structure in treated statocytes. Cytochemical study indicated that Ca2+ ions were concentrated in the intracellular organelles and cell walls in statocytes treated with CCB similar to untreated control. The data suggest that the effects of the CCB that demonstrated the correlation between the loss of polarity in statocytes and altered root gravitropism may be functionally related to systems that regulate Ca2+ homeostasis, particularly Ca2+ channels.     

Microtubules in statocytes from roots of cress (Lepidium sativum L.)

Summary Statocytes in root caps ofLepidium sativum L. were examined by means of ultrathin serial sections to evaluate the amount and distribution of cortical microtubules. The microtubules encircle the cell, oriented normal to the root length axis. In the distal cell edges, microtubules form a network, separating the distal complex of endoplasmic reticulum from the plasmalemma. Preprophase bands in meristem cells are observable rarely, structures which can be regarded as nucleating sites for microtubules are lacking. During ageing of the root cap cells, the number of microtubules increases in combination with a decrease of microtubule length. Development of the roots on a horizontal clinostat preserves a younger developmental stage of the microtubule system regarding amount and length of the individual microtubules. Evidence for an involvement of microtubules in graviperception is low, whereas their role in orienting cellulose microfibrils cannot be ruled out. Compression of the distal network of microtubules after centrifugation of the roots indicates that microtubules in statocytes ofLepidium sativum L. roots might function in stabilizing the distal complex of endoplasmic reticulum.     

杜仲花粉不对称分裂的超微结构观察

运用透射电镜对杜仲花粉发育进程进行了观察研究。结果显示,杜仲小孢子的第一次分裂为不等分裂,形成小的生殖细胞和大的营养细胞。分裂开始前小孢子的营养极形成许多小液泡,建立细胞极性;然后随着核膜的解体核周围的细胞器逐渐向纺锤体区靠近,围绕在纺锤体周围。花粉第一次有丝分裂完成后,生殖细胞所获得的细胞器开始分布在细胞的两侧,后来移向生殖细胞的营养极,而紧贴花粉壁的生殖极无细胞器分布。这种生殖细胞早期的细胞极性,可能为进一步分裂形成精细胞奠定基础。     

Expression of Polarity during early Development of Microspore-derived and Zygotic Embryos of Brassica napus L. cv. Topas

Microspore derived (MS-)embryogenesis and zygotic embryogenesis of Brassica napus L. cv. Topas were investigated by light and scanning electron microscopy to reveal the expression of polarity during the transition phase from globular to heart and torpedo shape. During the first 5 days of MS-embryo formation, the cell wall of the former microspores remained intact and a globular mass of cells developed within. Pollen walls ruptured after 5 days of culture; embryos proceeded through heart-shape and torpedo-shape stages within 15 days in a way comparable to, but faster than observed during zygotic embryogenesis. Expression of polarity in globular and elongating MS-embryos was analyzed by detection of the distribution of activated calmodulin as well as of free cytosolic calcium by using confocal scanning laser microscopy, and by the detection of starch. Calmodulin was evenly distributed in globular embryos and only exhibited clear polar distribution in elongated embryos. Free cytosolic Ca²⁺ accumulated in the protoderm of globular embryos and in the central cylinder of torpedo shaped embryos, but never showed polar distribution. Accumulation of starch granules at the root poles of both sexual as well as MS-embryos, however, indicated polar distribution before the transition from globular to heart shape stage. Since the local rupture of the pollen wall of 6-day-old MS-embryos was never preceded by the decrease of starch at that site, it is likely that the rupture of the pollen wall plays an important role in the local activation of cell metabolism and thus in the determination of the polarity axis in MS-embryos.     

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     

In cress roots (Lepidium sativum L.), abscisic acid prevents the development of the central cap cells into statocytes

Summary If seeds of Lepidium sativum are soaked in a 10^-4M solution of abscisic acid (ABA) for 30 min, the structural development of the central cap cells (C.C.) deviates strongly from the norm. ABA inhibits the elongation of the C.C., the sedimentation of the amyloplasts which may be caused by a higher viscosity of the cytoplasm, the formation of the stacked ER-cisternae at the distal cell pole, and the polar arrangement of the cell organelles along the gravity vector. The statistical distribution of the organelles and the arrangement of lipid droplets near the plasmalemma indicate that ABA prevents the normal development of the C.C. into statocytes.Abbreviations ABA abscisic acid - C.C. central cap cells     

Actin filaments responsible for the location of the nucleus in the lentil statocyte are sensitive to gravity

The location of the nucleus in statocytes or lentil roots grown: 1), at 1 g on the ground, 2), on a 1 g centrifuge in space, 3), in simulated microgravity on a slowly rotating clinostat (0.9 rmp) 4), in microgravity in space was investigated and statistically evaluated. In cells differentiated at 1 g on the ground, the nuclear membrane was almost in contact with the plasmalemma lining the proximal cell wall, whereas in statocytes of roots crown on the clinostat there was a distance of 0.47 micrometers (horizontal clinorotation) and or 0.76 micrometers (vertical clinorotation) between these membranes. However, in microgravity the nucleus was the most displaced, 0.87 micrometers from the proximal cell wall. Centrifugation of vertically grown roots in the root-tip direction showed that the threshold of centrifugal force to detach all nuclei from the proximal cell wall was about 40 g. In statocytes developed in the presence of cytochalasin B at 1 g the nuclei were sedimented on the amyloplasts at the distal cell pole, demonstrating that the location of the nucleus depends on actin filaments. The results obtained are in agreement with the hypothesis that gravity causes a tension of actin filaments and that this part of the cytoskeleton undergoes a relaxation in microgravity.