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At least 19 records

Water uptake by growing cells: an assessment of the controlling roles of wall relaxation, solute uptake, and hydraulic conductance

Growing plant cells increase in volume principally by water uptake into the vacuole. There are only three general mechanisms by which a cell can modulate the process of water uptake: (a) by relaxing wall stress to reduce cell turgor pressure (thereby reducing cell water potential), (b) by modifying the solute content of the cell or its surroundings (likewise affecting water potential), and (c) by changing the hydraulic conductance of the water uptake pathway (this works only for cells remote from water potential equilibrium). Recent studies supporting each of these potential mechanisms are reviewed and critically assessed. The importance of solute uptake and hydraulic conductance is advocated by some recent studies, but the evidence is indirect and conclusions remain controversial. For most growing plant cells with substantial turgor pressure, it appears that reduction in cell turgor pressure, as a consequence of wall relaxation, serves as the major initiator and control point for plant cell enlargement. Two views of wall relaxation as a viscoelastic or a chemorheological process are compared and distinguished.

Non-NASA Center↗

Wall relaxation and the driving forces for cell expansive growth

When water uptake by growing cells is prevented, the turgor pressure and the tensile stress in the cell wall are reduced by continued wall loosening. This process, termed in vivo stress relaxation, provides a new way to study the dynamics of wall loosening and to measure the wall yield threshold and the physiological wall extensibility. Stress relaxation experiments indicate that wall stress supplies the mechanical driving force for wall yielding. Cell expansion also requires water absorption. The driving force for water uptake during growth is created by wall relaxation, which lowers the water potential of the expanding cells. New techniques for measuring this driving force show that it is smaller than believed previously; in elongating stems it is only 0.3 to 0.5 bar. This means that the hydraulic resistance of the water transport pathway is small and that rate of cell expansion is controlled primarily by wall loosening and yielding.

NASA Program Space Biology↗

Wall relaxation in growing stems: comparison of four species and assessment of measurement techniques

This study was carried out to develop improved methods for measuring in-vivo stress relaxation of growing tissues and to compare relaxation in the stems of four different species. When water uptake by growing tissue is prevented, in-vivo stress relaxation occurs because continued wall loosening reduces wall stress and cell turgor pressure. With this procedure one may measure the yield threshold for growth (Y), the turgor pressure in excess of the yield threshold (P-Y), and the physiological wall extensibility (phi). Three relaxation techniques proved useful: "turgor-relaxation", "balance-pressure" and "pressure-block". In the turgor-relaxation method, water is withheld from growing tissue and the reduction in turgor is measured directly with the pressure probe. This technique gives absolute values for P and Y, but requires tissue excision. In the balance-pressure technique, the excised growing region is sealed in a pressure chamber, and the subsequent reduction in water potential is measured as the applied pressure needed to return xylem sap to the cut surface. This method is simple, but only measures (P-Y), not the individual values of P and Y. In the pressure-block technique, the growing tissue is sealed into a pressure chamber, growth is monitored continuously, and just sufficient pressure is applied to the chamber to block growth. The method gives high-resolution kinetics of relaxation and does not require tissue excision, but only measures (P-Y). The three methods gave similar results when applied to the growing stems of pea (Pisum sativum L.), cucumber (Cucumis sativus L.), soybean (Glycine max (L.) Merr.) and zucchini (Curcubita pepo L.) seedlings. Values for (P-Y) averaged between 1.4 and 2.7 bar, depending on species. Yield thresholds averaged between 1.3 and 3.0 bar. Compared with the other methods, relaxation by pressure-block was faster and exhibited dynamic changes in wall-yielding properties. The two pressure-chamber methods were also used to measure the internal water-potential gradient (between the xylem and the epidermis) which drives water uptake for growth. For the four species it was small, between 0.3 and 0.6 bar, and so did not limit growth substantially.

NASA Program Space Biology↗

How do plant cell walls extend?

This article briefly summarizes recent work that identifies the biophysical and biochemical processes that give rise to the extension of plant cell walls. I begin with the biophysical notion of stress relaxation of the wall and follow with recent studies of wall enzymes thought to catalyze wall extension and relaxation. Readers should refer to detailed reviews for more comprehensive discussion of earlier literature (Taiz, 1984; Carpita and Gibeaut, 1993; Cosgrove, 1993).

Review, Tutorial↗

The growing world of expansins

Expansins are cell wall proteins that induce pH-dependent wall extension and stress relaxation in a characteristic and unique manner. Two families of expansins are known, named alpha- and beta-expansins, and they comprise large multigene families whose members show diverse organ-, tissue- and cell-specific expression patterns. Other genes that bear distant sequence similarity to expansins are also represented in the sequence databases, but their biological and biochemical functions have not yet been uncovered. Expansin appears to weaken glucan-glucan binding, but its detailed mechanism of action is not well established. The biological roles of expansins are diverse, but can be related to the action of expansins to loosen cell walls, for example during cell enlargement, fruit softening, pollen tube and root hair growth, and abscission. Expansin-like proteins have also been identified in bacteria and fungi, where they may aid microbial invasion of the plant body.

NASA Discipline Plant Biology↗

Elliptic Relaxation of a Tensor Representation for the Redistribution Terms in a Reynolds Stress Turbulence Model

A formulation to include the effects of wall proximity in a second-moment closure model that utilizes a tensor representation for the redistribution terms in the Reynolds stress equations is presented. The wall-proximity effects are modeled through an elliptic relaxation process of the tensor expansion coefficients that properly accounts for both correlation length and time scales as the wall is approached. Direct numerical simulation data and Reynolds stress solutions using a full differential approach are compared for the case of fully developed channel flow.

Carlson, J. R.↗

Photoinhibition of stem elongation by blue and red light: effects on hydraulic and cell wall properties

The underlying mechanism of photoinhibition of stem elongation by blue (BL) and red light (RL) was studied in etiolated seedlings of pea (Pisum sativum L. cv Alaska). Brief BL irradiations resulted in fast transient inhibition of elongation, while a delayed (lag approximately 60 minutes) but prolonged inhibition was observed after brief RL. Possible changes in the hydraulic and wall properties of the growing cells during photoinhibition were examined. Cell sap osmotic pressure was unaffected by BL and RL, but both irradiations increased turgor pressure by approximately 0.05 megapascal (pressure-probe technique). Cell wall yielding was analyzed by in vivo stress relaxation (pressure-block technique). BL and RL reduced the initial rate of relaxation by 38 and 54%, while the final amount of relaxation was decreased by 48 and 10%, respectively. These results indicate that RL inhibits elongation mainly by lowering the wall yield coefficient, while most of the inhibitory effect of BL was due to an increase of the yield threshold. Mechanical extensibility of cell walls (Instron technique) was decreased by BL and RL, mainly due to a reduction in the plastic component of extensibility. Thus, photoinhibitions of elongation by both BL and RL are achieved through changes in cell wall properties, and are not due to effects on the hydraulic properties of the cell.

NASA Program Space Biology↗

Growth inhibition, turgor maintenance, and changes in yield threshold after cessation of solute import in pea epicotyls

The dependence of stem elongation on solute import was investigated in etiolated pea seedlings (Pisum sativum L. var Alaska) by excising the cotyledons. Stem elongation was inhibited by 60% within 5 hours of excision. Dry weight accumulation into the growing region stopped and osmotic pressure of the cell sap declined by 0.14 megapascal over 5 hours. Attempts to assay phloem transport via ethylenediaminetetraacetate-enhanced exudation from cut stems revealed no effect of cotyledon excision, indicating that the technique measured artifactual leakage from cells. Despite the drop in cell osmotic pressure, turgor pressure (measured directly via a pressure probe) did not decline. Turgor maintenance is postulated to occur via uptake of solutes from the free space, thereby maintaining the osmotic pressure difference across the cell membrane. Cell wall properties were measured by the pressure-block stress relaxation technique. Results indicate that growth inhibition after cotyledon excision was mediated primarily via an increase in the wall yield threshold.

NASA Discipline Number 40-30↗

Assembly and enlargement of the primary cell wall in plants

Growing plant cells are shaped by an extensible wall that is a complex amalgam of cellulose microfibrils bonded noncovalently to a matrix of hemicelluloses, pectins, and structural proteins. Cellulose is synthesized by complexes in the plasma membrane and is extruded as a self-assembling microfibril, whereas the matrix polymers are secreted by the Golgi apparatus and become integrated into the wall network by poorly understood mechanisms. The growing wall is under high tensile stress from cell turgor and is able to enlarge by a combination of stress relaxation and polymer creep. A pH-dependent mechanism of wall loosening, known as acid growth, is characteristic of growing walls and is mediated by a group of unusual wall proteins called expansins. Expansins appear to disrupt the noncovalent bonding of matrix hemicelluloses to the microfibril, thereby allowing the wall to yield to the mechanical forces generated by cell turgor. Other wall enzymes, such as (1-->4) beta-glucanases and pectinases, may make the wall more responsive to expansin-mediated wall creep whereas pectin methylesterases and peroxidases may alter the wall so as to make it resistant to expansin-mediated creep.

NASA Discipline Plant Biology↗

Mechanism of rapid suppression of cell expansion in cucumber hypocotyls after blue-light irradiation

Rapid suppression of hypocotyl elongation by blue light in cucumber (Cucumis sativus L.) was studied to examine possible hydraulic and wall changes responsible for diminished growth. Cell-sap osmotic pressure, measured by vapor-pressure osmometry, was not decreased by blue light; turgor pressure, measured by the pressure-probe technique, remained constant during the growth inhibition; and stem hydraulic conductance, measured by dynamic and static methods, was likewise unaffected by blue light. Wall yielding properties were assessed by the pressure-block technique for in-vivo stress relaxation. Blue light reduced the initial rate of relaxation by 77%, but had little effect on the final amount of relaxation. The results demonstrate that blue irradiation acts to decrease the wall yielding coefficient, but not the yield threshold. Stress-strain (Instron) analysis showed that irradiation of the seedlings had little effect on the mechanical extensibilities of the isolated wall. The results indicate that blue light can reduce cell-wall loosening without affecting bulk viscoelastic properties, and indicate a chemorheological mechanism of cell-wall expansion.

NASA Discipline Number 40-30↗

Characterization of long-term extension of isolated cell walls from growing cucumber hypocotyls

Walls from frozen-thawed cucumber (Cucumis sativus L.) hypocotyls extend for many hours when placed in tension under acidic conditions. This study examined whether such "creep" is a purely physical process dependent on wall viscoelasticity alone or whether enzymatic activities are needed to maintain wall extension. Chemical denaturants inhibited wall creep, some acting reversibly and others irreversibly. Brief (15 s) boiling in water irreversibly inhibited creep, as did pre-incubation with proteases. Creep exhibited a high Q10 (3.8) between 20 degrees and 30 degrees C, with slow inactivation at higher temperatures, whereas the viscous flow of pectin solutions exhibited a much lower Q10 (1.35). On the basis of its temperature sensitivity, involvement of pectic gel-sol transitions was judged to be of little importance in creep. Pre-incubation of walls in neutral pH irreversibly inactivated their ability to creep, with a half-time of about 40 min. At 1 mM, Cu2+, Hg2+ and Al3+ were strongly inhibitory whereas most other cations, including Ca2+, had little effect. Sulfhydryl-reducing agents strongly stimulated creep, apparently by stabilizing wall enzyme(s). The physical effects of these treatments on polymer interactions were examined by Instron and stress-relaxation analyses. Some treatments, such as pH and Cu2+, had significant effects on wall viscoelasticity, but others had little or no apparent effect, thus implicating an enzymatic creep mechanism. The results indicate that creep depends on relatively rugged enzymes that are firmly attached to or entangled in the wall. The sensitivity of creep to SH-reducing agents indicates that thiol reduction of wall enzymes might provide a control mechanism for endogenous cell growth.

NASA Program Space Biology↗

Elliptic Relaxation of a Tensor Representation of the Pressure-Strain and Dissipation Rate

A formulation to include the effects of wall-proximity in a second moment closure model is presented that utilizes a tensor representation for the redistribution term in the Reynolds stress equations. The wall-proximity effects are modeled through an elliptic relaxation process of the tensor expansion coefficients that properly accounts for both correlation length and time scales as the wall is approached. DNS data and Reynolds stress solutions using a full differential approach at channel Reynolds number of 590 are compared to the new model.

Carlson, John R.↗

A Reynolds stress model for near-wall turbulence

The paper formulates a tensorially consistent near-wall second-order closure model. Redistributive terms in the Reynolds stress equations are modeled by an elliptic relaxation equation in order to represent strongly nonhomogeneous effects produced by the presence of walls; this replaces the quasi-homogeneous algebraic models that are usually employed, and avoids the need for ad hoc damping functions. The model is solved for channel flow and boundary layers with zero and adverse pressure gradients. Good predictions of Reynolds stress components, mean flow, skin friction, and displacement thickness are obtained in various comparisons to experimental and direct numerical simulation data. The model is also applied to a boundary layer flowing along a wall with a 90-deg, constant-radius, convex bend.

Durbin, P. A.↗

Assessment and application of Reynolds stress closure models to high-speed compressible flows

The paper presents results from the development of higher order closure models for the phenomological modeling of high-speed compressible flows. The work presented includes the introduction of an improved pressure-strain correlationi model applicable in both the low- and high-speed regime as well as modifications to the isotropic dissipation rate to account for dilatational effects. Finally, the question of stiffness commonly associated with the solution of two-equation and Reynolds stress transport equations in wall-bounded flows is examined and ways of relaxing these restrictions are discussed.

Gatski, T. B.↗

Mechanism of gibberellin-dependent stem elongation in peas

Stem elongation in peas (Pisum sativum L.) is under partial control by gibberellins, yet the mechanism of such control is uncertain. In this study, we examined the cellular and physical properties that govern stem elongation, to determine how gibberellins influence pea stem growth. Stem elongation of etiolated seedlings was retarded with uniconozol, a gibberellin synthesis inhibitor, and the growth retardation was reversed by exogenous gibberellin. Using the pressure probe and vapor pressure osmometry, we found little effect of uniconozol and gibberellin on cell turgor pressure or osmotic pressure. In contrast, these treatments had major effects on in vivo stress relaxation, measured by turgor relaxation and pressure-block techniques. Uniconozol-treated plants exhibited reduced wall relaxation (both initial rate and total amount). The results show that growth retardation is effected via a reduction in the wall yield coefficient and an increase in the yield threshold. These effects were largely reversed by exogenous gibberellin. When we measured the mechanical characteristics of the wall by stress/strain (Instron) analysis, we found only minor effects of uniconozol and gibberellin on the plastic compliance. This observation indicates that these agents did not alter wall expansion through effects on the mechanical (viscoelastic) properties of the wall. Our results suggest that wall expansion in peas is better viewed as a chemorheological, rather than a viscoelastic, process.

NASA Discipline Plant Biology↗