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PubMed1992

A conceptual framework for investigating plant growth movements, with special reference to root gravitropism, utilizing a microgravity environment.

Microgravity (micro-g) has become established as an important environment in which to conduct experiments that aid in the understanding of gravity-responsive processes in plant growth and development. The gravitropic movement of roots is one example of a gravity-driven differential growth process that can be so investigated, autonomous nutational movements may be another. Although comparison of root movements of agravitropic mutants and their isogenic graviresponsive wild-types in l-g can answer many questions about their gravity-driven processes, they often relate only to gravity perception. Micro-g, however, can help define other constitutive processes which are also influenced by gravity. Examples of the latter are the so-called "tonic" effects, where organ or protoplast weight influences root growth and cellular polarity. A flow-chart is presented which orders the processes of differential growth in roots for analysis using both agravitropic mutants insensitive to gravity and mutant and wild-type roots grown in the micro-g environment. This approach addresses the question whether gravitropism and nutation of root tips are independent or co-existent processes, and also whether gravity determines any features of the nutation, e.g. its amplitude and period. While agravitropic mutants at 1-g can partially answer these questions, micro-g can validate the conclusions and indicate requirements for postulating additional influences such as tonic effects on signal transmission. Gravitropism of roots normally depends on the presence of the root cap, but embryonic tissue and its connection with the seed or grain can also influence gravitropic growth. A hypothetical scheme, based on experimental observations, is proposed for this grain-root-cap interaction. Testing the scheme could be achieved by physically dissociating the compartments presumed to interact in bringing about the tropism. The use of micro-g and agravitropic mutants to compare results with those obtained in 1-g would be of great value in further defining this system.

Gravitation↗

PubMed1996

Structure of cress root statocytes in microgravity (Bion-10 mission).

Experiments on primary roots of Lepidium sativum L. have been performed on board the Bion-10 satellite. The experimental set-up was extremely miniaturized and completely automatic. The results demonstrate the effectiveness of the instrumentation. The spatial orientation, growth, root cap differentiation and statocyte structure of roots grown under microgravity (MG) have been compared with control roots grown on the ground (GC) and in a 1G-reference centrifuge in space (RC). Root length and cap shape did not differ between MG and control samples. Under MG, the mean distance of the statoliths from the distal cell wall of the statocytes increased significantly, the mean distance of the mitochondria decreased and the nucleus did not change its position in comparison to both controls. The number and the shape of the amyloplasts (statoliths) were not influenced by the space flight factors, but their size as well as their relative area in the cell decreased. The number of starch grains per statolith as well as their size and shape changed under MG. In MG and RC samples the number of lipid bodies in the statocytes was higher and the relative area larger than in GC samples. The relative area occupied by vacuoles was greater in RC statocytes than in GC and MG statocytes. These results partly confirm and, in addition, extend the data from earlier experiments in space.

Microscopy, Electron↗

PubMed1984

Ultrastructure and movements of cell organelles in the root cap of agravitropic mutants and normal seedlings of Arabidopsis thaliana.

The root anatomy and ultrastructure of the agravitropic Arobidopsis thaliana L. mutants Dwf and aux-1 were compared with the gravitropic mutant aux-2 and the wild type (WT) in an attempt to find an explanation for the lack of response to gravity. No differences were found in the organization of the root cap. The central part of the cap (columella) contains 5 storeys of developing, functioning and degenerating statocytes. Their ultrastructure is very similar in all four types of plant. Particular attention was paid to the distribution of rough endoplasmic reticulum (ER). Both in the WT and the mutants the ER is concentrated in the distal part at the "floor" of the cell. Light micrographs were used to compare the sedimentation rates of movable cell structures in normal and agravitropic root statocytes. A longitudinal movement of amyloplasts and nuclei was observed when the roots were inverted. In WT and aux-2 the rates were on average 6.3 micrometers h-1 (amyloplasts) and 2.1 micrometers h-1 (nucleus). In aux-1 the sedimentation rates were significantly lower: 2.4 and 0.6 micrometers h-1, respectively. Based on magnified electron micrographs of normal and inverted statocytes a morphometrical analysis of the distribution and redistribution of amyloplasts, nuclei, mitochondria, vacuoles and ER was made. The only significant difference was found in the redistribution of amyloplasts between aux-1 and the gravitropical normal types.

Arabidopsis↗

PubMed1999

Statoliths motions in gravity-perceiving plant cells: does actomyosin counteract gravity?

Statocytes from plant root caps are characterized by a polar arrangement of cell organelles and sedimented statoliths. Cortical microtubules and actin microfilaments contribute to development and maintenance of this polarity, whereas the lack of endoplasmic microtubules and prominent bundles of actin microfilaments probably facilitates sedimentation of statoliths. High-resolution video microscopy shows permanent motion of statoliths even when sedimented. After immunofluorescence microscopy using antibodies against actin and myosin II the most prominent labeling was observed at and around sedimented statoliths. Experiments under microgravity have demonstrated that the positioning of statoliths depends on the external gravitational force and on internal forces, probably exerted by the actomyosin complex, and that transformation of the gravistimulus evidently occurs in close vicinity to the statoliths. These results suggest that graviperception occurs dynamically within the cytoplasm via small-distance sedimentation rather than statically at the lowermost site of sedimentation. It is hypothesized that root cap cells are comparing randomized motions with oriented motions of statoliths and thereby perceiving gravity.

Actomyosin↗

PubMed2005

Auxin and ethylene interactions control mitotic activity of the quiescent centre, root cap size, and pattern of cap cell differentiation in maize.

Root caps (RCs) are the terminal tissues of higher plant roots. In the present study the factors controlling RC size, shape and structure were examined. It was found that this control involves interactions between the RC and an adjacent population of slowly dividing cells, the quiescent centre, QC. Using the polar auxin transport inhibitor 1-N-naphthylphthalamic acid (NPA), the effects of QC activation on RC gene expression and border cell release was characterized. Ethylene was found to regulate RC size and cell differentiation, since its addition, or the inhibition of its synthesis, affected RC development. The stimulation of cell division in the QC following NPA treatment was reversed by ethylene, and quiescence was re-established. Moreover, inhibition of both ethylene synthesis and auxin polar transport triggered a new pattern of cell division in the root epidermis and led to the appearance of supernumerary epidermal cell files with cap-like characteristics. The data suggest that the QC ensures an ordered internal distribution of auxin, and thereby regulates not only the planes of growth and division in both the root apex proper and the RC meristem, but also regulates cell fate in the RC. Ethylene appears to regulate the auxin redistribution system that resides in the RC. Experiments with Arabidopsis roots also reveal that ethylene plays an important role in regulating the auxin sink, and consequently cell fate in the RC.

Amino Acids, Cyclic↗

PubMed1987

The role of calcium ions in gravity signal perception and transduction.

Ca2+ is implicated as a messenger in coupling various environmental stimuli, such as gravity and light, to response. In recent years, it has become evident that Ca2+ plays a central role in all three phases of gravitropism--perception, transduction and response. The root cap, which is known to contain high amounts of Ca2+ and calmodulin, is the primary site of gravity perception. The possible role of phosphoinositide turnover and Ca(2+) - and Ca(2+) -calmodulin-dependent enzymes such as Ca(2+) -ATPase and protein kinases in gravitropism is discussed. A model is proposed to describe the role of Ca2+ in both normal and light-dependent gravity response in roots.

Calcium↗

PubMed1976

[Geoperception in the lentil root cap].

Previous analysis showed that, in its initial phase, the geotropic response of Lens culinaris L. roots cannot be explained by a simple action by sliding, pressure or contact of amyloplasts on a sensitive surface located along the longitudinal wall. In this study another mode of action is tested by considering the following parameters as functions of the roots inclination: (1) the distance (d) which the amyloplasts move; (2) their number of contacts (mean c) with parietal cytoplasm; (3) the variable (sin alpha) of the transversal component of the statolith weight (mean M x g sin alpha). It is shown that the initial rate of curvature (mean V), at the various angles, is related to the sedimentation of the amyloplasts by the equation mean V = a log b mean d mean c sin alpha (where a and b are constants). The results obtained prove that the geotropic stimulation is dependent upon the sine of the angle (alpha) of the root inclination and explain the sine rule deviation. The role of statoliths is discussed in the light of recent literature on growth inhibitors which are involved in the geotropic reaction.

Fabaceae↗

PubMed2001

Two tomato expansin genes show divergent expression and localization in embryos during seed development and germination.

Expansins are plant proteins that can induce extension of isolated cell walls and are proposed to mediate cell expansion. Three expansin genes were expressed in germinating tomato (Lycopersicon esculentum Mill.) seeds, one of which (LeEXP4) was expressed specifically in the endosperm cap tissue enclosing the radicle tip. The other two genes (LeEXP8 and LeEXP10) were expressed in the embryo and are further characterized here. LeEXP8 mRNA was not detected in developing or mature seeds but accumulated specifically in the radicle cortex during and after germination. In contrast, LeEXP10 mRNA was abundant at an early stage of seed development corresponding to the period of rapid embryo expansion; it then decreased during seed maturation and increased again during germination. When gibberellin-deficient (gib-1) mutant seeds were imbibed in water, LeEXP8 mRNA was not detected, but a low level of LeEXP10 mRNA was present. Expression of both genes increased when gib-1 seeds were imbibed in gibberellin. Abscisic acid did not prevent the initial expression of LeEXP8 and LeEXP10, but mRNA abundance of both genes subsequently decreased during extended incubation. The initial increase in LeEXP8, but not LeEXP10, mRNA accumulation was blocked by low water potential, but LeEXP10 mRNA amounts fell after longer incubation. When seeds were transferred from abscisic acid or low water potential solutions to water, abundance of both LeEXP8 and LeEXP10 mRNAs increased in association with germination. The tissue localization and expression patterns of both LeEXP8 and LeEXP10 suggest developmentally specific roles during embryo and seedling growth.

Cell Wall↗

PubMed1997

Gravitropic reaction in the growth of tea roots.

In Japan, tea (Camellia sinenis (L.) Kuntze) seedlings are propagated by cutting. A root system of clonal plants by cutting consists of adventitious roots and lateral roots. Most of the roots grow horizontally, which results in a shallow distribution of the root system. Such a shallow root system could be one of the factors contributing to the deterioration of nutrient uptake and resistance to water stress. Gravitropism of the roots is considered to be a decisive factor that controls the depth of a root system. The authors have investigated changes in the growth direction of roots to gravitative stimulus, using several kinds of roots (seminal roots, lateral roots and adventitious roots). Furthermore, amyloplasts in the root-cap cells, which are considered to be an equipment sensing gravistimulus, were observed. Seminal roots prominently showed orthogravitropism and contained many amyloplast particles in their root cap cells. Most lateral and adventitious roots showed plagiogravitropism, growing in an angle to gravistimulus, and lacked observable amyloplast particles in their root cap cells. The results suggest that the shallowing of root systems of elonal tea plants could be attributed to a gravitropic reaction of the adventitious and lateral roots composing the root system. There could also be a close relationship between the growth direction of roots and the presence of amyloplasts in root-cap cells.

Gravitropism↗

PubMed1999

Genetic ablation of root cap cells in Arabidopsis.

The root cap is increasingly appreciated as a complex and dynamic plant organ. Root caps sense and transmit environmental signals, synthesize and secrete small molecules and macromolecules, and in some species shed metabolically active cells. However, it is not known whether root caps are essential for normal shoot and root development. We report the identification of a root cap-specific promoter and describe its use to genetically ablate root caps by directing root cap-specific expression of a diphtheria toxin A-chain gene. Transgenic toxin-expressing plants are viable and have normal aerial parts but agravitropic roots, implying loss of root cap function. Several cell layers are missing from the transgenic root caps, and the remaining cells are abnormal. Although the radial organization of the roots is normal in toxin-expressing plants, the root tips have fewer cytoplasmically dense cells than do wild-type root tips, suggesting that root meristematic activity is lower in transgenic than in wild-type plants. The roots of transgenic plants have more lateral roots and these are, in turn, more highly branched than those of wild-type plants. Thus, root cap ablation alters root architecture both by inhibiting root meristematic activity and by stimulating lateral root initiation. These observations imply that the root caps contain essential components of the signaling system that determines root architecture.

Arabidopsis↗

PubMed1990

Comparison of the ultrastructure of conventionally fixed and high pressure frozen/freeze substituted root tips of Nicotiana and Arabidopsis.

To circumvent the limitations of chemical fixation (CF) and to gain more reliable structural information about higher plant tissues, we have cryofixed root tips of Nicotiana and Arabidopsis by high pressure freezing (HPF). Whereas other freezing techniques preserve tissue to a relatively shallow depth, HPF in conjunction with freeze substitution (FS) resulted in excellent preservation of entire root tips. Compared to CF, in tissue prepared by HPF/FS: (1) the plasmalemma and all internal membranes were much smoother and often coated on the cytoplasmic side by a thin layer of stained material, (2) the plasmalemma was appressed to the cell wall, (3) organelle profiles were rounder, (4) the cytoplasmic, mitochondrial, and amyloplast matrices were denser, (5) vacuoles contained electron dense material, (6) microtubules appeared to be more numerous and straighter, with crossbridges observed between them, (7) cisternae of endoplasmic reticulum (ER) were wider and filled with material, (8) Golgi intercisternal elements were more clearly resolved and were observed between both Golgi vesicles and cisternae, and (9) larger vesicles were associated with Golgi stacks. This study demonstrates that HPF/FS can be used to successfully preserve the ultrastructure of relatively large plant tissues without the use of intracellular cryoprotectants.

Arabidopsis↗

PubMed1994

Vicia faba chromosomal aberration assay.

A collaborative study involving laboratories in six countries was initiated under the sponsorship of the International Programme on Chemical Safety (IPCS) to determine the sensitivity, efficiency and reliability of the Vicia faba root tip meristem chromosomal aberration assay using a standardized protocol. The six laboratories that participated in this study were located in the Slovak Republic, India, Japan, Poland, Sweden and the USA. All laboratories adhered to a standardized protocol for the Vicia faba chromosomal aberration assay. Four coded chemicals, azidoglycerol (AG), N-methyl-N-nitrosourea (MNU), sodium azide (NaN3) and maleic hydrazide (MH) were tested with the Vicia faba chromosomal aberration assay. Of the four chemicals, three (MH, AG and MNU) were found to be clastogenic and gave a concentration related response. However, the results of NaN3 were equivocal which might be explained by the stability of NaN3. The conclusions from this study suggest that the Vicia faba chromosomal aberration bioassay is an efficient and reliable short-term bioassay for the rapid screening of chemicals for clastogenicity.

Azides↗

PubMed1994

Effects of altered gravity on plant cell processes: results of recent space and clinostatic experiments.

Space and clinostatic experiments revealed that plant cell structure and metabolism rearrangements depend on taxonomical position and physiological state of objects, growth phase and real or simulated microgravity influence duration. It was shown that clinostat conditions reproduce only a part of microgravity biological effects. It is established that various responses occur in microgravity: 1) rearrangements of cytoplasmic organelles ultrastructure and calcium balance; 2) physical-chemical properties of the plasmalemma are changed; 3) enzymes activity is often enhanced. These events provoke the acceleration of growth and differentiation of cells and their aging as a result; at the same time some responses can be considered as cell adaptation to microgravity.

Adaptation, Physiological↗

PubMed2005

Application of GFP technique for cytoskeleton visualization onboard the International Space Station.

Cytoskeleton recently attracted wide attention of cell and molecular biologists due to its crucial role in gravity sensing and trunsduction. Most of cytoskeletal research is conducted by the means of immunohistochemical reactions, different modifications of which are beneficial for the ground-based experiments. But for the performance onboard the space vehicles, they represent quite complicated technique which requires time and special skills for astronauts. In addition, immunocytochemistry provides only static images of the cytoskeleton arrangement in fixed cells while its localization in living cells is needed for the better understanding of cytoskeletal function. In this connection, we propose a new approach for cytoskeletal visualization onboard the ISS, namely, application of green fluorescent protein (GFP) from Aequorea victoria, which has the unique properties as a marker for protein localization in vivo. The creation of chimerical protein-GFP gene constructs, obtaining the transformed plant cells possessed protein-GFP in their cytoskeletal composition will allow receiving a simple and efficient model for screening of the cytoskeleton functional status in microgravity.

Actin Cytoskeleton↗

PubMed2001

Stomatal and leaf growth responses to partial drying of root tips in willow.

Root tips of intact willow (Salix dasyclados Wimm., Clone 81-090) plants were partially dried by exposure to ambient greenhouse air and then kept in water-vapor-saturated air for up to 3 days. The drying treatment increased abscisic acid (ABA) concentrations in both the root tips subjected to drying and in the xylem sap, while it reduced leaf stomatal conductance and leaf extension rate. Despite the decrease in stomatal conductance, leaf water potentials were unaffected by the root drying treatment, indicating that the treatment reduced hydraulic conductivity between roots and foliage. After roots subjected to drying were returned to a nutrient solution or excised, ABA concentrations in the remaining roots and in the xylem sap, stomatal conductance of mature leaves and extension rate of unfolding leaves all returned to values observed in control plants. The 4-fold increase in xylem sap ABA concentration following the root drying treatment was not solely the result of reduced sap flow, and thus may be considered a potential cause, not merely a consequence, of the observed reduction in stomatal conductance.

Abscisic Acid↗

PubMed1992

How well does the clinostat mimic the effect of microgravity on plant cells and organs?

The effect of clinostatting and microgravity on plant cells and organs is considered on the basis of distinguishing two types of gravistimulation: static and dynamic. The former is switched off both by clinostatting and microgravity, the latter is switched off by microgravity but occurs inevitably during clinostatting and may be perceived by cells if the rotation is not fast enough. Effects of clinostatting and microgravity on different examples of static gravistimulation (tonic effects, formation of compression wood, growth of "grass nodes," compensation of epinasty, stabilization of cellular polarity) are considered. The mechanism of the dynamic stimulation is presented; it is related to the displacement of the gravity sensing masses in the cell containing them, and involves disturbance of cytoskeletal tension. The low threshold for gravity perception and short minimal time of dynamic stimulation are emphasized. Only a relatively fast rotating clinostat, on which the radial distance of the cells from the rotational axis is small enough to keep the centrifugal force low, can effectively compensate gravity. However, one must take into account the extreme sensitivity of plants to mechanical stresses that may appear during clinostatting at different levels of plant organization.

Cytoskeleton↗

PubMed1996

Microsomal membrane proteins and vanadate-sensitive ATPase from Vicia faba root tips after clinostat treatment.

Microsomal and soluble protein fractions from Vicia faba root tips were used for SDS-PAGE and Western-immunoblot analysis with anti-ubiquitin antibodies after 9 h clinostat treatment of the plants. In contrast to soluble proteins omnilateral gravistimulation (9 h) resulted in an enhanced proteolytic capacity for microsomal proteins. The increase of vanadate-sensitive ATPase activity was 83% after 9 h clinostat treatment, when the enzyme activity was measured directly after membrane preparation. Enhanced ATPase activity was correlated with the appearance of a polypeptide of about 100 kDa and its fragments (93 and 80 kDa). ATPases are not the only membrane bound proteins, which are changed during clinostat treatment, as several ubiquitinated polypeptides were also affected. A 1 h storage of microsomal fractions led to a shift of band intensities on ubiquitin-specific Western-blots. The demonstrated effect could not be observed, when fractions were isolated in the presence of protease inhibitors. In accordance with the polypeptide analysis omnilateral gravistimulation resulted in an enhanced capacity to degrade specific microsomal ubiquitin-conjugates, whereas the soluble ubiquitin-pool was not visibly affected.

Adenosine Triphosphatases↗

PubMed1995

Effects of clinorotation and microgravity on sweet clover columella cells treated with cytochalasin D.

The cytoskeleton of columella cells is believed to be involved in maintaining the developmental polarity of cells observed as a reproducible positioning of cellular organelles. It is also implicated in the transduction of gravitropic signals. Roots of sweet clover (Melilotus alba L.) seedlings were treated with a microfilament disrupter, cytochalasin D, on a slowly rotating horizontal clinostat (2 rpm). Electron micrographs of treated columella cells revealed several ultrastructural effects including repositioning of the nucleus and the amyloplasts and the formation of endoplasmic reticulum (ER) whorls. However, experiments performed during fast clinorotation (55 rpm) showed an accumulation (but no whorling) of a disorganized ER network at the proximal and distal pole and a random distribution of the amyloplasts. Therefore, formation of whorls depends upon the speed of clinorotation, and the overall impact of cytochalasin D suggests the necessity of microfilaments in organelle positioning. Interestingly, a similar drug treatment performed in microgravity aboard the US Space Shuttle Endeavour (STS-54, January 1993) caused a displacement of ER membranes and amyloplasts away from the distal plasma membrane. In the present study, we discuss the role of microfilaments in maintaining columella cell polarity and the utility of clinostats to simulate microgravity.

Actin Cytoskeleton↗