Plant water relations and control of cell elongation at low water potentials

Recent developments in water status measurement techniques using the psychrometer, the pressure probe, the osmometer and pressure chamber are reviewed, and the process of cell elongation from the viewpoint of plant-water relations is discussed for plants subjected to various environmental stress conditions. Under water-deficient conditions, cell elongation of higher plants can be inhibited by interruption of water flow from the xylem to the surrounding elongating cells. The process of growth inhibition at low water potentials could be reversed by increasing the xylem water potential by means of pressure application in the root region, allowing water to flow from the xylem to the surrounding cells. This finding confirmed that a water potential field associated with growth process,i.e., the growth-induced water potential, is an important regulating factor for cell elongation other than metabolic factors. The concept of the growth-induced water potential was found to be applicable for growth retardation caused by cold stress, heat stress, nutrient deficiency and salinity stress conditions. In the present review, the fact that the cell elongation rate is primarily associated with how much water can be absorbed by elongating cells under water-deficiency, nutrient deficiency, salt stress, cold stress and heat stress conditions is suggested.     

Osmotic adjustment in Spirulina platensis

Unilateral irradiation of maize (Zea mays L.) seedlings results in a fluence-rate gradient, and hence below saturation, a gradient of the far-red-absorbing form of phytochrome (Pfr). The Pfr-gradients established by blue, red and far-red light were spectrophotometrically measured in the mesocotyl. Based on these Pfr-gradients and the fluence-response curves of phytochrome photoconversion the fluence-rate gradients were calculated. The fluence-rate gradient in the blue (460 nm) was steeper than that in the red (665 nm), which in turn was steeper than that in the far-red light (725 nm). The fluence-rate ratios front to rear were 1:0.06 (460 nm), 1:0.2 (665 nm), and 1:0.33 (725 nm). The assumption that phytochrome-mediated phototropism of maize mesocotyls is caused by local phytochrome-mediated growth inhibition was tested in the following manner. Firstly, the Pfr response curve for growth inhibition was calculated; these calculations were based on measurements of Pfr-gradients and data from red-light-induced phototropism. Secondly, the Pfr response curve for growth inhibition was used as a basis for calculating fluence-response curves for blue-and far-red-light-induced phototropism. Finally, these calculated results were compared with experimental data. It was concluded that the threshold for phytochrome-mediated phototropism of maize mesocotyls reflects the apparent photoconversion cross section of phytochrome whereas the maximal inducable curvature depends on the steepness of the light (Pfr) gradient across the mesocotyl.Abbreviations Pfr far-red-absorbing form of phytochrome - Ptot total phytochrome - Fr far-red light     

Control of the rate of cell enlargement: Excision,wall relaxation,and growth-induced water potentials

A new guillotine thermocouple psychrometer was used to make continuous measurements of water potential before and after the excision of elongating and mature regions of darkgrown soybean (Glycine max L. Merr.) stems. Transpiration could not occur, but growth took place during the measurement if the tissue was intact. Tests showed that the instrument measured the average water potential of the sampled tissue and responded rapidly to changes in water potential. By measuring tissue osmotic potential ( _s ), turgor pressure ( _p ) could be calculated. In the intact plant, _s and _p were essentially constant for the entire 22 h measurement, but _s was lower and _p higher in the elongating region than in the mature region. This caused the water potential in the elongating region to be lower than in the mature region. The mature tissue equilibrated with the water potential of the xylem. Therefore, the difference in water potential between mature and elongating tissue represented a difference between the xylem and the elongating region, reflecting a water potential gradient from the xylem to the epidermis that was involved in supplying water for elongation. When mature tissue was excised with the guillotine, _s and _p did not change. However, when elongating tissue was excised, water was absorbed from the xylem, whose water potential decreased. This collapsed the gradient and prevented further water uptake. Tissue _p then decreased rapidly (5 min) by about 0.1 MPa in the elongating tissue. The _p decreased because the cell walls relaxed as extension, caused by _p , continued briefly without water uptake. The _p decreased until the minimum for wall extension (Y) was reached, whereupon elongation ceased. This was followed by a slow further decrease in Y but no additional elongation. In elongating tissue excised with mature tissue attached, there was almost no effect on water potential or _p for several hours. Nevertheless, growth was reduced immediately and continued at a decreasing rate. In this case, the mature tissue supplied water to the elongating tissue and the cell walls did not relax. Based on these measurements, a theory is presented for simultaneously evaluating the effects of water supply and water demand associated with growth. Because wall relaxation measured with the psychrometer provided a new method for determining Y and wall extensibility, all the factors required by the theory could be evaluated for the first time in a single sample. The analysis showed that water uptake and wall extension co-limited elongation in soybean stems under our conditions. This co-limitation explains why elongation responded immediately to a decrease in the water potential of the xylem and why excision with attached mature tissue caused an immediate decrease in growth rate without an immediate change in _p Abbreviations and symbols L tissue conductance for water - m wall extensibility - Y average yield threshold (MPa) - _o water potential of the xylem - _p turgor pressure - _s osmotic potential - _w water potential of the elon gating tissue     

Acclimation of photosynthesis in Zea mays to low water potentials involves alterations in protoplast volume reduction

Effects of water-stress treatment of Zea mays L. plants on protoplast volume and photosynthesis in leaf slices exposed to solutions of different osmotic potential ( _s) were studied. Decreased photosynthetic capacity in the leaf slices at low tissue _w was associated with dehydration-induced protoplast-volume reduction. Leaf slices from plants exposed to in-situ water deficits exhibited greater photosynthetic capacity and relative protoplast volume at low water potential ( _w) invitro than tissue from control plants.In-situ water stress induced osmotic adjustment of the leaf tissue as determined by pressure/volume analysis. It is concluded that plant acclimation to low leaf _w may involve a reduced degree of cell shrinkage at a given _w. This acclimation would allow for the maintenance of relatively higher photosynthetic capacity at low water protentials.Symbols _s Osmotic potential - _w water potential New Jersey Agricultural Experiment Station Publication No. 12149-6-87     

Effect of water stress on growth, osmotic adjustment, cell wall elasticity and water-use efficiency in Spartina alterniflora

In the northern spring–summer season of 2004–2005, vegetative propagated plants of Spartina alterniflora were grown under control and water stress conditions on the Mediterranean sea shore of the south-east of Tunis. Control plants were irrigated every week and water stress plants were irrigated until the soil achieved 50% (mild stress) and 25% (severe stress) field capacity (FC). Dry and fresh weight at the whole plant level (g plant⁻¹), shoot to root ratio on dry and fresh weight, photosynthesis (A), transpiration rate (E), instantaneous water-use efficiency (WUE_i), leaf water potential (Ψw), leaf water content (WC), osmotic potential at full turgor (Ψs¹⁰⁰), osmotic potential at turgor loss point (Ψs⁰), osmotic adjustment (OA), proline, sugars, inorganic compounds and cell wall elasticity (CWE) were evaluated during a period of 6 days period between 82 and 90 days after the beginning of treatment (DAT). Plants grown under severe and mild-water stress showed lower Ψw than in control plants with values that averaged −3.1, −1.6 and −0.9 MPa, respectively. S. alterniflora plants submitted to mild-water stress exhibited OA and a decrease in CWE. However, under severe water stress the OA was not observed and CWE also decreased, but it was higher than in the mild-water stress. OA was mainly explained by the accumulation of nitrates, sugars and at a lesser degree, proline. S. alterniflora had a strong decline of the dry and fresh weight of the whole plant associated to a marked decrease of photosynthesis (A) and transpiration (E) in response to water stress, although WUE_i was increased. These results suggest that OA and WUE_i can be important components of the water stress adaptation mechanism in this species, but they are not sufficient enough to contribute to resistance to water stress.     

Importance of the cap in maize root growth

A large population of primary roots of Zea mays (cv. LG 11) was selected for uniform length at zero time. Their individual growth rates were measured over an 8-h period in the vertical position (in humid air, darkness). Three groups of these roots with significantly different growth rates were then chosen and their cap length was measured. It was found that slowly growing roots had long caps whereas rapidly growing roots had short caps. The production by the cap cells of basipetally transported growth inhibitors was tested (biologically by the curvature of half-decapped roots) and found to be significantly higher for longer root caps than that for shorter ones.     

Acclimation of leaf growth to low water potentials in sunflower

Abstract Leaf growth is one of the most sensitive of plant processes to water deficits and is frequently inhibited in field crops. Plants were acclimated for 2 weeks under a moderate soil water deficit to determine whether the sensitivity of leaf growth could be altered by sustained exposure to low water potentials. Leaf growth under these conditions was less than in the controls because expansion occurred more slowly and for less of the day than in control leaves. However, acclimated leaves were able to grow at leaf water potentials (Ψ₁) low enough to inhibit growth completely in control plants. This ability was associated with osmotic adjustment and maintenance of turgor in the acclimated leaves. Upon rewatering, the growth of acclimated leaves increased but was less than the growth of controls, despite higher concentrations of cell solute and greater turgor in the acclimated leaves than in controls. Therefore, factors other than turgor and osmotic adjustment limited the growth of acclimated leaves at high ψ₁ Four potentially controlling factors were investigated and the results showed that acclimated leaves were less extensible and required more turgor to initiate growth than control leaves. The slow growth of acclimated leaves was not due to a decrease in the water potential gradient for water uptake, although changes in the apparent hydraulic conductivity for water transport could have occurred. It was concluded that leaf growth acclimated to low ψ₁, by adjusting osmotically, and the concomitant maintenance of turgor permitted growth where none otherwise would occur. However, changes in the extensibility of the tissue and the turgor necessary to initiate growth caused generally slow growth in the acclimated leaves.     

Osmotic responses of maize roots

Water and solute relations of excised seminal roots of young maize (Zea mays L) plants, have been measured using the root pressure probe. Upon addition of osmotic solutes to the root medium, biphasic root pressure relaxations were obtained as theoretically expected. The relaxations yielded the hydraulic conductivity Lp _r) the permeability coefficient (P _sr), and the reflection coefficient (σ _sr) of the root. Values of Lp _r in these experiments were by nearly an order of magnitude smaller than Lp _r values obtained from experiments where hydrostatic pressure gradients were used to induce water flows. The value of P _sr was determined for nine different osmotica (electrolytes and nonelectrolytes) which resulted in rather variable values (0.1·10^-8–1.7·10^-8m·s^-1). The reflection coefficient σ _sr of the same solutes ranged between 0.3 and 0.6, i.e. σ _sr was low even for solutes for which cell membranes exhibit a σ _s≈1. Deviations from the theoretically expected biphasic responses occured which may have reflected changes of either P _sr or of active pumping induced by the osmotic change. The absolute values of Lp _r, P _sr, and σ _sr have been critically examined for an underestimation by unstirred layer effecs. The data indicate a considerable apoplasmic component for radial movement of water in the presence of hydrostatic gradients and also some solute flow byppassing root protoplasts. In the presence of osmotic gradients, however, there was a substantial cell-to-cell transport of water. Cutting experiments demonstrated that the hydraulic resistance for the longitudinal movement of water was much smaller than for radial transport except for the apical ends of the segments (length=5 to 20 mm). The differences in Lp _r as well as the low σ _sr values suggest that the simple osmometer model of the root with a single osmotic barrier exhibiting nearly semipermeable properties should be extended for a composite membrane model with hydraulic and osmotic barriers arranged in series and in parallel.     

Cell water potential,osmotic potential,and turgor in the epidermis and mesophyll of transpiring leaves

Water potential, osmotic potential and turgor measurements obtained by using a cell pressure probe together with a nanoliter osmometer were compared with measurements obtained with an isopiestic psychrometer. Both types of measurements were conducted in the mature region of Tradescantia virginiana L. leaves under non-transpiring conditions in the dark, and gave similar values of all potentials. This finding indicates that the pressure probe and the osmometer provide accurate measurements of turgor, osmotic potentials and water potentials. Because the pressure probe does not require long equilibration times and can measure turgor of single cells in intact plants, the pressure probe together with the osmometer was used to determine in-situ cell water potentials, osmotic potentials and turgor of epidermal and mesophyll cells of transpiring leaves as functions of stomatal aperture and xylem water potential. When the xylem water potential was-0.1 MPa, the stomatal aperture was at its maximum, but turgor of both epidermal and mesophyll cells was relatively low. As the xylem water potential decreased, the stomatal aperture became gradually smaller, whereas turgor of both epidermal and mesophyll cells first increased and afterward decreased. Water potentials of the mesophyll cells were always lower than those of the epidermal cells. These findings indicate that evaporation of water is mainly occurring from mesophyll cells and that peristomatal transpiration could be less important than it has been proposed previously, although peristomatal transpiration may be directly related to regulation of turgor in the guard cells.     

Transmembrane electrical potentials in growing maize roots

The anti-auxin 4-chlorophenoxyisobutyric acid (PCIB) applied at a concentration of 10^-2 mol m^-3 to maize root segments was found to induce a transmembrane electrical potential of up to-130 mV (pd of 30 mV). The kinetics of this response were comparable to the time scale for PCIB-stimulated H⁺-extrusion. Both effects are eliminated by the addition of p-fluoromethoxycarbonyl cyanide phenylhydrazone (FCCP). Treatment with fusicoccin (FC) and PCIB together does not result in a hyperpolarization greater than with FC alone. Benzoic acid (10^-2 mol m^-3) had no effect on the transmembrane electrical potentials. These results are discussed in relation to a possible electrogenic proton pump which may be regulated by perturbations in the cellular auxin content or activity.Abbreviations ATPase adenosine triphosphatase - FC fusicoccin - FCCP p-fluoromethoxy carbonylcyanide phenylhydrazone - IAA indole-3yl-acetic acid - NAA naphthyl-lylacetic acid - PCIB 4-chlorophenoxyisobutyric acid - PD potential difference     

Auxin increases the hydraulic conductivity of auxin-sensitive hypocotyl tissue

The ability of water to enter the cells of growing hypocotyl tissue was determined in etiolated soybean (Glycine max (L.) Merr.) seedlings. Water uptake was restricted to that for cell enlargement, and the seedlings were kept intact insofar as possible. Tissue water potentials ( _w) were measured at thermodynamic equilibrium with an isopiestic thermocouple psychrometer. _wwas below the water potential of the environment by as much as 3.1 bars when the tissue was enlarging rapidly. However, _w was similar to the water potential of the environment when cell enlargement was not occurring. The low _w in enlarging tissue indicates that there was a low conductivity for water entering the cells.The ability of water to enter the enlarging cells was defined as the apparent hydraulic conductivity of the tissue (Lp). Despite the low Lp of growing cells, Lp decreased further as cell enlargement decreased when intact hypocotyl tissue was deprived of endogenous auxin (indole-3-acetic acid) by removal of the hypocotyl hook. Cell enlargement resumed and Lp increased when auxin was resupplied exogenously. The auxin-induced increase in Lp was correlated with the magnitude of the growth enhancement caused by auxin, and it was observed during the earliest phase of the growth response to auxin. The increase in Lp appeared to be caused by an increase in the hydraulic conductivity of the cell protoplasm, since other factors contributing to Lp remained constant. The rapidity of the response is consistent with a cellular site of action at the plasmalemma, although other sites are not precluded.Because the experiments involved only short times, auxin-induced changes in cell enlargement could not be attributed to changes in cell osmotic potentials. Neither could they be attributed to changes in turgor, which increased when the rate of enlargement decreased. Rather, auxin appeared to act by altering the extensibility of the cell walls and by simultaneously altering the ability of water to enter the growing cells under a given water potential gradient. The hydraulic conductivity and extensibility of the cell walls appeared to contribute about equally to the control of the growth rate of the hypocotyls.     

Effect of water stress on growth, Na+ and K+ accumulation and water use efficiency in relation to osmotic adjustment in two populations of Atriplex halimus L.

The effect of water stress on growth, Na⁺ and K⁺ accumulation and water utilization was investigated in plants of two populations of Atriplex halimus L. originating from Kairouan (Tunisia) and Tensift (Morocco). Water deficit was applied by withholding water for 22 days. All plants remained alive until the end of the treatment although growth was strongly reduced in both populations. Water stress decreased CO₂ assimilation in saturating conditions, mainly in the population obtained from Kairouan, suggesting an impact of drought on the dark phase of photosynthesis, beside a decrease in stomatal conductance which was recorded mainly in the population obtained from Tensift. The two studied populations did not differ in their water consumption, as indicated by similar soil gravimetric water content and plant transpiration. However, water use efficiency increased under stress conditions in the population from Tensift but not in the population from Kairouan. Thelatter population displayed a larger capacity for osmotic adjustment. A drought-induced specific increase in Na⁺ concentration was also reported in both populations. It is concluded that in A. halimus, water stress resistance estimated in terms of biomass production, could be associated with higher WUE rather than with with a greater osmotic adjustment and that sodium may assume a specific physiological function in this xerohalophytic C₄ species.     

The kinetics of bidirectional growth of stem sections from etiolated pea seedlings in response to acid,auxin and fusicoccin

Growth in length and diameter of abraded stem sections from etiolated pea (Pisum sativum L.) seedlings was monitored continuously using a double laser optical level auxanometer system. Acidic solutions (pH 4.0–4.5) induced rapid elongation accompanied by lateral shrinkage (up to 8% of the initial diameter). The shrinkage phase lasted for 30–45 min. Pretreatment with permeant solutes (KCl, NaCl, sucrose or glucose) prevented lateral shrinkage, while pretreatment with the impermeant solute, polyethylene glycol, did not block lateral contraction in response to acid. A slight turgor step-up given during the shrinkage phase inhibited lateral shrinkage and increased the elongation rate. Visual observation confirmed that shrinkage occurred and that the same region of the stem that contracted in diameter also elongated. It is proposed that lateral shrinkage results from a decrease in turgor pressure during acid-stimulated elongation. Elongation induced by auxin and fusicoccin (FC) was also accompanied by a decrease in the diameter; this decrease could be prevented by pretreatment with KCl or glucose. Thus, the early phase of auxin and FC action is acid-like. However, the shrinkage is of shorter duration (14–20 min) and it is less drastic (ca. 2%). In addition, FC caused lateral expansion after a 20-min lag period in stems pretreated with KCl. The results are consistent with an acid-growth mechanism during the early phase (first 20–40 min) of the responses to both auxin and FC. It is suggested that enhanced osmoregulation subsequently inhibits further lateral shrinkage and helps to maintain steady-state growth. FC, unlike auxin, may alter the anisotropic character of the wall.Abbreviations FC fusicoccin - IAA indole-3-acetic acid - LOLA laser optical levar auxanometer - PEG polyethyleneglycol 600     

Leaf water relations characteristics of Lupinus angustifolius and L. cosentinii

Summary Lupins (Lupinus angustifolius and L. cosentinii) growing in 321 containers in a glasshouse were exposed to drought by withholding water. Leaf water potential (₁), and leaf osmotic potential (_s) were measured daily as soil water became depleted. Leaf water relations were further assessed by a pressure-volume technique and by measuring _s and relative water content of leaves after rehydration. Analysis by pressure-volume or cryoscopic techniques showed that leaf osmotic potential at saturation (_s100) decreased from -0.6 MPa in well watered to -0.9 MPa in severely droughted leaves, and leaf water potential at zero turgor (_zt) decreased from about -0.7 to -1.1 MPa in well watered and droughted plants, respectively. Relative water content at zero turgor (RWC_zt) was high (88%) and tended to be decreased by drought. The ratio of turgid leaf weight to dry weight was not influenced by drought and was high at about 8.0. The bulk elastic modulus () was approximately halved by drought when related to leaf turgor potential (_p) and probably mediated turgor maintenance during drought. The latter was found to be negatively influenced by rate of drought. Supplying the plants with high levels of K salts did not promote adjustment or turgor maintenance.     

Rapid,localized light effect on root growth in maize

A method using optical microfibers permitted localized exposure of the cap or the elongating part of growing maize (Zea mays L.) roots to white light. When the cap was illuminated, a strong and very rapid inhibition of the elongation rate of the roots was found. When the light microbeam was directed at the elongating region, the roots continued to grow at the same rate as before the illumination.     

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.     

Change in XET activities,cell wall extensibility and hypocotyl elongation of soybean seedlings at low water potential

In dark-grown soybean (Glycine max [L.] Merr.) seedlings, exposing the roots to water-deficient vermiculite (_w=–0.36 MPa) inhibited hypocotyl (stem) elongation. The inhibition was associated with decreased extensibility of the cell walls in the elongation zone. A detailed spatial analysis showed xyloglucan endotransglucosylase (XET; EC 2.4.1.207) activity on the basis of unit cell wall dry weight was decreased in the elongation region after seedlings were transplanted to low _w. The decrease in XET activity was at least partially due to an accumulation of cell wall mass. Since cell number was only slightly altered, wall mass had increased per cell and probably led to increased wall thickness and decreased cell wall extensibility. Alternatively, an increase in cell wall mass may represent a mechanism for regulating enzyme activity in cell walls, XET in this case, and therefore cell wall extensibility. Hypocotyl elongation was partially recovered after seedlings were grown in low-_w vermiculate for about 80 h. The partial recovery of hypocotyl elongation was associated with a partial recovery of cell wall extensibility and an enhancement of XET activity in the hypocotyl elongation zone. Our results indicate XTH proteins may play an important role in regulating cell wall extensibility and thus cell elongation in soybean hypocotyls. Our results also showed an imperfect correlation of spatial elongation and XET activity along the hypocotyls. Other potential functions of XTH and their regulation in soybean hypocotyl growth are discussed.     

Biomass production,photosynthesis, and leaf water relations of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> under moderate water stress

The perennial smooth cordgrass, Spartina alterniflora, has been successfully introduced in salty ecosystems for revegetation or agricultural use. However, it remains unclear whether it can be introduced in arid ecosystems. The aim of this study was to investigate the physiological response of this species to water deficiency in a climate-controlled greenhouse. The experiment consisted of two levels of irrigation modes, 100 and 50% field capacities (FC). Although growth, photosynthesis, and stomatal conductance of plants with 50% FC were reduced at 90 days from the start of the experiment, all of the plants survived. The water-stressed plants exhibited osmotic adjustment and an increase in the maximum elastic modulus that is assumed to be effective to enhance the driving force for water extraction from the soil with small leaf water loss. An increase in the water use efficiency was also found in the water-stressed plants, which could contribute to the maintenance of leaf water status under drought conditions. It can be concluded that S. alterniflora has the capacity to maintain leaf water status and thus survive in arid environment.     

Gravitropism of the primary root of maize: a complex pattern of differential cellular growth in the cortex independent of the microtubular cytoskeleton

The spatio-temporal sequence of cellular growth within the post-mitotic inner and outer cortical tissue of the apex of the primary root of maize (Zea mays L.) was investigated during its orthogravitropic response. In the early phase (0–30 min) of the graviresponse there was a strong inhibition of cell lengthening in the outer cortex at the lower side of the root, whereas lengthening was only slightly impaired in the outer cortex at the upper side. Initially, inhibition of differential cell lengthening was less pronounced in the inner cortex indicating that tissue tensions which, in these circumstances, inevitably develop at the outer-inner cortex interface, might help to drive the onset of the root bending. At later stages of the graviresponse (60 min), when a root curvature had already developed, cells of the inner cortex then exhibited a prominent cell length differential between upper and lower sides, whereas the outer cortex cells had re-established similar lengths. Again, tissue tensions associated with the different patterns of cellular behaviour in the inner and outer cortex tissues, could be of relevance in terminating the root bending. The perception of gravity and the complex tissue-specific growth responses both proceeded normally in roots which were rendered devoid of microtubules by colchicine and oryzalin treatments. The lack of involvement of microtubules in the graviresponse was supported by several other lines of evidence. For instance, although taxol stabilized the cortical microtubules and prevented their re-orientation in post-mitotic cortical cells located at the lower side of gravistimulated roots, root bending developed normally. In contrast, when gravistimulated roots were physically prevented from bending, re-oriented arrays of cortical microtubules were seen in all post-mitotic cortical cells, irrespective of their position within the root.Abbreviations CMTs cortical microtubules - CW Cholodny-Went - FF form factor - MT microtubule The research was supported by a fellowship from the Alexander von Humboldt Stiftung (Bonn, Germany) to F.B. Financial support to AGRAVIS by Deutsche Agentur für Raumfahrtangelegenheiten (DARA, Bonn) and Ministerium für Wissenschaft und Forschung (Düsseldorf) is gratefully acknowledged. IACR receives grant-aided support from the Biotechnology and Biological Sciences Research Council of the United Kingdom.     

Effect of applied and endogenous indol-3-yl-acetic acid on maize root growth

The level of endogenous Indol-3-yl-acetic acid (IAA) measured by gas chromatography-mass spectrometry in the elongating zone of intact primary roots of Zea mays showed a good linear correlation with the growth rate of these roots. When they were treated with IAA, their relative elongation decreased; this indicates a supraoptimal content of endogenous IAA. However, the growth of some of the relatively rapidly extending roots was enhanced by such treatment. Interactions between endogenous and applied IAA in the control of root growth are discussed.Abbreviations GC-MS gas chromatography-mass spectrometry - IAA Indol-3-yl-acetic acid