Nonrecirculating hydroponic system suitable for uptake studies at very low nutrient concentrations

Seedlings of maize (Zea mays L. cv WF9 × Mo 17) growing at low water potentials in vermiculite contained greatly increased proline concentrations in the primary root growth zone. Proline levels were particularly high toward the apex, where elongation rates have been shown to be completely maintained over a wide range of water potentials. Proline concentration increased even in quite mild treatments and reached 120 millimolal in the apical millimeter of roots growing at a water potential of −1.6 megapascal. This accounted for almost half of the osmotic adjustment in this region. Increases in concentration of other amino acids and glycinebetaine were comparatively small. We have assessed the relative contributions of increased rates of proline deposition and decreased tissue volume expansion to the increases in proline concentration. Proline content profiles were combined with published growth velocity distributions to calculate net proline deposition rate profiles using the continuity equation. At low water potential, proline deposition per unit length increased by up to 10-fold in the apical region of the growth zone compared to roots at high water potential. This response accounted for most of the increase in proline concentration in this region. The results suggest that osmotic adjustment due to increased proline deposition plays an important role in the maintenance of root elongation at low water potentials.     

Growth of the maize primary root at low water potentials : I. Spatial distribution of expansive growth

Seedlings of maize (Zea mays L. cv WF9 × Mo 17) were grown in vermiculite at various water potentials. The primary root continued slow rates of elongation at water potentials which completely inhibited shoot growth. To gain an increased understanding of the root growth response, we examined the spatial distribution of growth at various water potentials. Time lapse photography of the growth of marked roots revealed that inhibition of root elongation at low water potentials was not explained by a proportional decrease in growth along the length of the growing zone. Instead, longitudinal growth was insensitive to water potentials as low as − 1.6 megapascal close to the root apex, but was inhibited increasingly in more basal locations such that the length of the growing zone decreased progressively as the water potential decreased. Cessation of longitudinal growth occurred in tissue of approximately the same age regardless of spatial location or water status, however. Roots growing at low water potentials were also thinner, and analysis revealed that radial growth rates were decreased throughout the elongation zone, resulting in greatly decreased rates of volume expansion.     

Osmoregulation,solute distribution,and growth in soybean seedlings having low water potentials

Soybean (Glycine max (L.) Merr.) seedlings osmoregulate when the supply of water is limited around the roots. The osmoregulation involves solute accumulation (osmotic adjustment) by the elongating region of the hypocotyls. We investigated the relationship between growth, solute accumulation, and the partitioning of solutes during osmoregulation. Darkgrown seedlings were transplanted to vermiculite containing 1/8 (0.13 x) the water of the controls. Within 12–15 h, the osmotic potential of the elongating region had decreased to-12 bar, but it was-7 bar in the controls. This osmoregulation involved a true solute accumulation by the hypocotyls, since cell volume and turgor were virtually the same regardless of the water regime. The hypocotyls having low water potentials elongated slowly but, when deprived of their cotyledons, did not elongate or accumulate solute. This result indicated a cotyledonary origin for the solutes and a dependence of slow growth on osmotic adjustment. The translocation of nonrespired dry matter from the cotyledons to the seedling axis was unaffected by the availability of water, but partitioning was altered. In the first 12 h, dry matter accumulated in the elongating region of the 0.13 x hypocotyls, and osmotic adjustment occurred. The solutes involved were mostly free amino acids, glucose, fructose, and sucrose, and these accounted for most of the increased dry weight. After osmotic adjustment was complete, dry matter ceased to accumulate in the hypocotyls and bypassed them to accumulate in the roots, which grew faster than the control roots. The proliferation of the roots resulted in an increased root/shoot ratio, a common response of plants to dry conditions.Osmotic adjustment occurred in the elongating region of the hypocotyls because solute utilization for growth decreased while solute uptake continued. Adjustment was completed when solute uptake subsequently decreased, and uptake then balanced utilization. The control of osmotic adjustment was therefore the rate of solute utilization and, secondarily, the rate of solute uptake. Elongation was inhibited by unknown factors(s) despite the turgor and substrates associated with osmotic adjustment. The remaining slow elongation depended on osmotic adjustment and represented some optimum between the necessary inhibition for solute accumulation and the necessary growth for seedling establishment.     

Spatial distributions of potassium, solutes, and their deposition rates in the growth zone of the primary corn root

Densities of osmoticum and potassium were measured as a function of distance from the tip of the primary root of Zea mays L. (cv WF9 × mo17). Millimeter segments were excised and analyzed for osmotic potential by a miniaturized freezing point depression technique, and for potassium by flame spectrophotometry. Local deposition rates were estimated from the continuity equation with values for density and growth velocity. Osmotic potential was uniform, −0.73 ± 0.05 megapascals, throughout the growth zone of well-watered roots. Osmoticum deposition rate was 260 μosmoles per gram fresh weight per hour. Potassium density fell from 117 micromoles per gram in the first mm region to 48 micromoles per gram at the base of the growth zone. Potassium deposition rates had a maximum of 29 micromoles per gram per hour at 3.5 millimeters from the tip and were positive (i.e. potassium was being added to the tissue) until 8 millimeters from the tip. The results are discussed in terms of ion relations of the growing zone and growth physics.     

盐胁迫下盐芥渗透调节物质的积累及其渗透调节作用

用含有NaCl0、50、100、200、300、400mmol/L的Hoagland培养液处理盐芥幼苗一定时间后,分别测定其根和叶含水量、渗透势、几种无机和有机渗透调节物质含量,并计算了渗透调节物质在不同条件下的计算渗透势值(COP).结果表明:随盐处理浓度的增加,盐芥根和叶的含水量和渗透势逐渐降低;Na 和Cl-是根和叶积累的无机渗透调节物质;SS、OA和FAA是根积累的有机渗透调节物质,Pro是叶和根积累的有机渗透调节物质.Na X-ray微区分析表明液泡是积累Na 的主要部位.     

Symplast Volume, Turgor, Stomatal Conductance and Growth in Relation to Osmotic and Elastic Adjustment in Droughted Sugarcane1

Leaf water relations, stomatal conductance (g) and shoot growthrate (SGR) were monitored during a soil drying cycle in threesugarcane cultivars growing in pots in a greenhouse. The pressure-volumetechnique was used to evaluate diurnal and droughtinduced variationin leaf water relations characteristics. Leaf solute contentand bulk elasticity varied diurnally in both irrigated and droughtedplants and were highest at midday. Solute accumulation and increasedelasticity were also observed as leaf water deficits developedmore slowly during soil drying. This osmotic and elastic adjustmentmaintained symplast volume essentially constant both diurnallyand during soil drying, whereas turgor was only partially maintained.The extent of osmotic adjustment associated with drought wasnot reflected in the leaf osmotic potential at full turgor becausethe concurrent increase in tissue elasticity resulted in a largersymplast volume at full turgor. Cultivar responses over therange of leaf water deficits imposed did not provide conclusiveevidence for genotypic variation in osmotic and elastic adjustment.It appeared that behavioural differences in rates of water usemay have determined the magnitude of osmotic and elastic adjustmentin response to drought. In the early stages of soil drying,reductions in SGR and g were not accompanied by significantreductions in bulk leaf water status. This suggested that otherfactors, presumably signals originating from the roots, mayhave regulated SGR and g.     

Effects of Rapidly and Slowly Permeating Osmotica on Macromolecule and Sucrose Synthesis

Metabolic effects induced by low water potentials were separatedfrom those induce by low turgor and hydration, by comparingresponses of plant cells to rapidly and slowly permeating osmotica. Metabolic processes selected for study were those which areclaimed to be very sensitive to water deficits. ‘Glucose-stimulated’respiration, and sucrose and starch synthesis, were measuredin Chlorella pyrenoidosa and protein synthesis in vacuolatedand non-vacuolated segments of Zea mays roots. Mannitol, a slowlypermeating osmoticum, stimulated sucrose synthesis and reducedrates of other metabolic processes, including starch and proteinsynthesis. In contrast, metabolism changed little during exposureto low water potentials induced by ethylene glycol, which rapidlypermeates cells. The experiments demonstrated that water and solute potentialsas low as—20 atm had only very small effects on metabolism,strongly suggesting that water deficits act principally viadecreases in turgor and hydration.     

Water Relations of Growing Maize Coleoptiles : Comparison between Mannitol and Polyethylene Glycol 6000 as External Osmotica for Adjusting Turgor Pressure

Water relations of growing segments of maize (Zea mays L.) coleoptiles were investigated with osmotic methods using either mannitol (MAN) or polyethylene glycol 6000 (PEG) as external osmotica. Segments were incubated in MAN or PEG solutions at 0 to - 15 bar water potential (Ψ_o) and the effects were compared on elongation growth, osmotic shrinkage, cell sap osmolality (OC), and osmotic pressure (π_i). The nonpenetrating osmoticum PEG affects π_i in agreement with Boyle-Mariotte's law, i.e. the segments behave in principle as ideal osmometers. There is no osmotic adjustment in the Ψ_o range permitting growth (0 to −5 bar) nor in the Ψ_o range inducing osmotic shrinkage (−5 to −10 bar). Promoting growth by auxin (IAA) has no effect on the osmotic behavior of the tissue toward PEG. In contrast to PEG, MAN produces an apparent increase in π_i accompanied by anomalous effects on segment elongation and shrinkage leading to a lower value for Ψ_o which establishes a growth rate of zero and to an apparent recovery from osmotic shrinkage after 2 hours of incubation. These effects can be quantitatively attributed to uptake of MAN into the tissue. MAN is taken up into the apoplastic space and the symplast as revealed by a large temperature-dependent component of MAN uptake. It is concluded that MAN, in contrast to PEG, is unsuitable as an extemal osmoticum for the quantitative determination of water relations of growing maize coleoptiles.     

Effects of rapidly and slowly permeating osmotica on metabolism

Zea mays was exposed to solutions of low water potentials by addition of ethylene glycol or mannitol. Intact seedlings were treated for 1 hr at potentials between −10 and −20 atmospheres and then returned to high water potentials. Subsequent root extension was slow after mannitol treatment, but rapid when ethylene glycol had been used as the osmoticum. Cellular activity of excised roots was also affected much less by ethylene glycol than by mannitol. Processes studied included respiration, glucose uptake, and synthesis of methanol-insoluble compounds. These differences in response to various osmotica applied both during and after treatment at low water potentials.     

Osmotic relations of the drought-tolerant shrub Artemisia tridentata in response to water stress

Turgor maintenance, solute content and recovery from water stress were examined in the drought-tolerant shrub Artemisia tridentata. Predawn water potentials of shrubs receiving supplemental water remained above ?2 MPa throughout summer, while predawn water potentials of untreated shrubs decreased to ?5 MPa. Osmotic potentials decreased in conjunction with water potentials maintaining turgor pressures above 0 MPa. The decreases in osmotic potentials were not the result of osmotic adjustment (i.e. solute accumulation). Leaf solute contents decreased during drought, but leaf water volumes decreased more than 75% from spring to summer, thereby passively concentrating solutes within the leaves. The maintenance of positive turgor pressures despite decreases in leaf water volumes is consistent with other studies of species with elastic cell walls. Inorganic ion, organic acid, and carbohydrate contents of leaves declined during drought. The only solutes accumulating in leaves of A. tridentata with water stress were proline and a cyclitol, both considered compatible solutes. Total and osmotic potentials recovered rapidly following rewatering of shrubs; solute contents did not change except for a decrease in proline. Maintaining turgor through the passive concentration of solutes may be advantageous compared to synthesis of new solutes for osmotic adjustment in arid environments.     

Seasonal progressions of tissue and cell water relations parameters in evergreen and deciduous perennials

Abstract. Tissue and cell water relations parameters were followed for Heteromeles arbutifolia, Cercis occidentalis and Aesculus californica , in an environment exhibiting seasonally increasing drought. The extensive seasonal osmotic adjustment of evergreen H. arbutifolia and the moderate adjustment in C. occidentalis closely matched their respective seasonal decreases in minimum daily water potential. Summer deciduous A. californica exhibited only small drops in osmotic potential and water potential. Experiments with irrigated plants indicated that drought was not required for the osmotic adjustment of H. arbutifolia and C. occidentalis. However, in H. arbutifolia drought treatment enhanced osmotic adjustment. In irrigated H. arbutifolia , osmotic adjustment was mainly the result of an accumulation of osmotica. In drought-stressed plants, the same change in osmotic potential resulted from a combination of accumulation of osmotica and a decrease in symplast volume.     

Taxonomic and ecological aspects of the distribution of glycinebetaine and related compounds in plants

Summary The concentrations of the major inorganic ions and glycinebetaine, choline and proline and the osmotic pressure of extract sap have been determined in eight salt marsh species and four sand dune species from local habitats. These results together with those previously reported on hydroponically grown plants and data assembled from the literature show that glycinebetaine accumulation is a feature of members of the Chenopodiaceae, Amaranthaceae, many Gramineae and some members of the Solanaceae and Compositae, particularly when exposed to conditions of low soil water potential. It is suggested that in these families betaine is employed as a non-toxic cytoplasmic osmoticum when decreased osmotic potentials are required. In some other plant species proline may fulfil a similar function. Another quaternary ammonium compound may be accumulated in the Plumbaginaceae in addition to proline. Some evidence suggests that the differences in the organic osmoticum used may relate to the different inorganic ion contents of the plants. The accumulation of nitrogen dipoles as cytoplasmic osmotica may make heavy demands on the nitrogen economy of the plants and this problem is discussed briefly.     

Kinetics of Adaptation to Osmotic Stress in Lentil (Lens culinaris Med.) Roots

When intact roots of lentil (Lens culinaris Med.) are subjected to severe osmotic stress by treatment with a solution of low water potential, they immediately begin to shrink. Within 10 to 15 minutes, shrinkage ceases, and within 20 minutes, the roots resume growth. The time lag between application of osmoticum and resumption of growth varies from about 10 to 30 minutes over the range of external water potentials of −2 to −12.4 bars. For external water potentials as low as −8.7 bars the new steady rate of growth in the presence of osmoticum is approximately equal to that prevailing before application of osmoticum. For external water potentials between −8.7 and −13 bars growth resumes, but the new rate is less than that prior to addition of osmoticum. Measurements of changes in the internal solute content during adaptation show that the solute content of the root increases but that the magnitude of the increase is, by itself, insufficient to account for the resumption of rapid growth.     

The role of solute accumulation, osmotic adjustment and changes in cell wall elasticity in drought tolerance in Ziziphus mauritiana (Lamk.)

Ber (Ziziphus mauritiana Lamk.) is a major fruit tree crop of the north-west Indian arid zone. In a study of the physiological basis of drought tolerance in this species, two glasshouse experiments were conducted in which trees were droughted during single stress-cycles. In the first experiment, during a 13 d drying cycle, pre-dawn leaf water (leaf) and osmotic () potentials in droughted trees declined from -0.5 and -1.4 MPa to -1.7 and -2.2 MPa, respectively, for a decrease in relative water content () of 14%. During drought stress, changes in sugar metabolism were associated with significant increases in concentrations of hexose sugars (3.8-fold), cyclitol (scyllo-inositol; 1.5-fold), and proline (35-fold; expressed per unit dry weight), suggesting that altered solute partitioning may be an important factor in drought tolerance of Ziziphus. On rewatering pre-dawn leaf and recovered fully, but remained depressed by 0.4 MPa relative to control values, indicating that solute concentration per unit water content had changed during the drought cycle.Evidence for osmotic adjustment was provided from a second study during which a gradual drought was imposed. Pressure-volume analysis revealed a 0.7 MPa reduction in osmotic potential at full turgor, with leaf at turgor loss depressed by 1 MPa in drought-stressed leaves. Coupled with osmotic adjustment, during gradual drought, was a 65% increase in bulk tissue elastic modulus (wall rigidity) which resulted in turgor loss at the same in both stressed and unstressed leaves. The possible ecological significance of maintenance of turgor potential and cell volume at low water potentials for drought tolerance in Ziziphus is discussed.Keywords: Ziziphus mauritiana, drought, solute accumulation, osmotic adjustment, proline.      

Root Growth and Water Uptake by Maize Plants in Drying Soil

Sharp, R. E and Da vies, W. J. 1985. Root growth and water uptakeby maize plants in drying soil.— J. exp. Bot. 36: 1441–1456. The influence of soil drying on maize (Zea mays L.) root distributionand use of soil water was examined using plants growing in thegreenhouse in soil columns. The roots of plants which were wateredwell throughout the 18 d experimental period penetrated thesoil profile to a depth of 60 cm while the greatest percentageof total root length was between 20–40 cm. High soil waterdepletion rates corresponded with these high root densities.Withholding water greatly restricted root proliferation in theupper part of the profile, but resulted in deeper penetrationand higher soil water depletion rates at depth, compared withthe well watered columns. The deep roots of the unwatered plantsexhibited very high soil water depletion rates per unit rootlength. Key words: Maize, roots, water deficit, soil water depletion     

Effects of turgor potential on 1-aminocyclopropane-1-carboxylic acid uptake into the vacuolar compartment and ethylene production in tomato pericarp slices

While solute transport and ethylene production by plant tissue are sensitive to the osmotic concentration of the solution bathing the tissue, the influence of tissue water relations and specifically tissue turgor potential on the kinetics of 1-aminocyclopropane-1-carboxylic acid (ACC) uptake into the vacuolar compartment and ethylene production have not been examined. 1-Aminocyclopropane-1-carboxylic acid transport and ethylene production were examined in tomato (Lycopersicon esculentum Mill. cv. Liberty) pericarp slices incubated in solutions having a range of mannitol, polyethylene glycol 3350 and ethylene glycol concentrations known to affect tissue water relations. Tissue osmotic and turgor potentials were derived from osmolality measurements of cell saps recovered by freeze-thawing and corrected for the contribution of the free-space solution. When relatively nonpermeable (mannitol or polyethylene glycol 3350) osmotica were used, both ACC uptake and ethylene production were greatest at a solution osmolality of 230 milliosmolal where tissue turgor potential ranged between 120 and 140 kPa. At higher and lower turgor potentials, the high-affinity saturating component of ACC uptake and ethylene production were inhibited, and ACC efflux from the vacuolar compartment was increased. The inhibition of ACC uptake was evident as a decrease in V_max with no effect on K_m. Turgor potential changes caused by adjusting solution osmolality with mannitol or polyethylene glycol 3350 were accompanied by changes in the osmotic potential and water potential of the tissue. The effects of turgor potential vs the osmotic and water potentials of tomato pericarp slices were differentiated by comparing responses to nonpermeable osmotica and mixtures of nonpermeable and permeable osmotica. Ethylene glycol-mannitol mixtures had effects on the osmotic potential and water potential of the tissue similar to those of nonpermeable osmotica but had less effect on tissue turgor, ACC transport and ethylene production. Incubating tissue in solutions without nonpermeable osmotica osmotically shocked the tissue. Increasing solution osmolality with ethylene glycol in the absence of nonpermeable osmotica increased tissue turgor and ethylene production. The present study indicates that tissue turgor is an important factor affecting the kinetics of ACC uptake into the vacuolar compartment and ethylene production in tomato pericarp slices.     

The effect of osmotic potential on anther culture in spring wheat (Triticum aestivum)

This study was conducted to determine the effect of osmotic potential in a modified 85D12 medium on both callus induction and plant regeneration in the anther culture of two wheat genotypes, cv. Chris and cv. Pavon. Altering the medium osmotic potential by changing the carbohydrate source and concentration or by adding a non-metabolized osmoticum appeared to have the greatest potential for improving anther-derived green plant production. The medium osmotic potentials were varied (-0.67 to –2.30 MPa) by altering sucrose and PEG concentration. Both osmotica affected callus production, with –0.9 to –1.4 MPa media producing the most calluses. Callus production depended on genotype and osmoticum. Only PEG concentration affected green plant regeneration. The greatest number of green plants (11.5 plants per 100 anthers in cv. Chris) was obtained with 0.0125 M of PEG. This experiment suggested that a low level of PEG in the medium was beneficial for producing green plants from wheat anthers.     

Rapid osmotic adjustment by a succulent halophyte to saline shock

The objective of this research was to measure the short term osmotic adjustment of Salicornia europaea L. ssp. rubra (A. Nels) Breitung when suddenly exposed to 100 millimolar NaCl. Plants were grown hydroponically, shocked with 100 millimolar NaCl added to the culture solution, and stem tips analyzed for free inorganic ions and small organic molecules at intervals up to 72 hours. In the first 2 hours, the calculated leaf osmoticum showed a net increase of 158.8 millimolar most of which was free Mg²⁺ (+135.3 millimolar). Total sugars increased almost 5-fold by the 6th hour, enough to provide sufficient osmoticum for the cytoplasm if only partially confined there. By 24 hours, all measured osmotica had decreased except Na⁺, Mg²⁺, Cl⁻, and proline, with the net increase being 208 millimolar. By 72 hours, there was a net gain of 356 millimolar in osmotica of the stem tips, due to Na⁺ (+233.3 millimolar), Cl⁻ (+306.7 millimolar), and a small increase in sugar and proline (+3.5 millimolar), with all other osmotica decreasing in concentration. Compatible osmotica did not change sufficiently to account for osmotic balance between vacuole and cytoplasm; consequently, there must have been a reapportionment of osmotica within the cell in the short time duration of this experiment.     

Soil solute concentration and water uptake by single lupin and radish plant roots

Application of computer assisted tomography to gamma and X-ray attenuation measurements and Na⁺-LIX microelectrodes were used to determine the spatial distributions of soil water content and Na⁺ concentrations respectively near single roots of eighteen day old lupin and radish plants. These quantities were monitored at root depths of 3, 6 and 9 cm and at zero, 2, 4, 6, and 8 hour intervals from the diurnal commencement of transpiration. The plants were subjected to two levels of transpirational demand and five Na⁺ soil solution concentration levels. Water extraction rates for the lupin and radish roots increased continuously with time but were substantially reduced with increasing Na⁺ concentration in the treatment. Water uptake was uniform along the length of the essentially constant diameter lupin roots but decreased along the tapering radish roots as the diameter and hence the surface area per unit length of the roots decreased. The accumulation of Na⁺ at the root surfaces of both plants increased gradually with time in a near linear fashion and was slightly higher under the higher transpiration demand. These increases were not exponential as would be expected with non-absorption by the roots and this is considered to be due to back diffusion at the relatively high water contents used. At these water contents matric potentials had a much smaller influence on transpiration than osmotic potentials. The relationships between leaf water potentials (Ψ₁) and osmotic potentials at the root surfaces were linear with the decreases in Ψ₁ almost exactly reflecting the decreases in Ψ_π indicating rapid plant adjustment. Leaf water potentials decreased progressively with time and the relationships between leaf water potential and the transpiration rate were also linear supporting the suggestion of constant plant resistances at any given concentration.