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Turgor pressure provides the force needed to stress and deform the cell walls of plants, algae, and fungi during expansive growth. However, turgor pressure plays another subtle but equally important role in expansive growth of walled cells: it connects the two biophysical processes of water uptake and wall deformation to ensure that the volumetric rates of water uptake and enlargement of the cell wall chamber are equal. In this study, the role of turgor pressure as a 'connector' is investigated analytically by employing validated and established biophysical equations. The objective is to determine the effect of 'wall loosening' on the magnitude of turgor pressure. It is known that an increase or decrease in turgor pressure and/or wall loosening rate increases or decreases the expansive growth rate, respectively. Interestingly, it is shown that an increase in the wall loosening rate decreases the turgor pressure slightly, thus reducing the effect of wall loosening on increasing the expansive growth rate. Other analyses reveal that reducing the rate of water uptake results in a larger decrease in turgor pressure with the same increase in wall loosening rate, which further reduces the effect of wall loosening on increasing the expansive growth rate. [ABSTRACT FROM AUTHOR]

Succulence is an adaptation to low water availability characterised by the presence of water-storage tissues that alleviate water stress under low water availability. The succulent syndrome has evolved convergently in over 80 plant families and is associated with anatomical, physiological and biochemical traits. Despite the alleged importance of cell wall traits in drought responses, their significance in the succulent syndrome has long been overlooked. Here, by analyzing published pressure--volume curves, we show that elastic adjustment, whereby plants change cell wall elasticity, is uniquely beneficial to succulents for avoiding turgor loss. In addition, we used comprehensive microarray polymer profiling (CoMPP) to assess the biochemical composition of cell walls in leaves. Across phylogenetically diverse species, we uncover several differences in cell wall biochemistry between succulent and non-succulent leaves, pointing to the existence of a 'succulent glycome'. We also highlight the glycomic diversity among succulent plants, with some glycomic features being restricted to certain succulent lineages. In conclusion, we suggest that cell wall biomechanics and biochemistry should be considered among the characteristic traits that make up the succulent syndrome. [ABSTRACT FROM AUTHOR]

The stiffness of a plant cell in response to an applied force is determined not only by the elasticity of the cell wall but also by turgor pressure and cell geometry, which affect the tension of the cell wall. Although stiffness has been investigated using atomic force microscopy (AFM) and Young's modulus of the cell wall has occasionally been estimated using the contact-stress theory (Hertz theory), the existence of tension has made the study of stiffness more complex. Elastic shell theory has been proposed as an alternative method; however, the estimation of elasticity remains ambiguous. Here, we used finite element method simulations to verify the formula of the elastic shell theory for onion (Allium cepa) cells. We applied the formula and simulations to successfully quantify the turgor pressure and elasticity of a cell in the plane direction using the cell curvature and apparent stiffness measured by AFM. We conclude that tension resulting from turgor pressure regulates cell stiffness, which can be modified by a slight adjustment of turgor pressure in the order of 0.1 MPa. This theoretical analysis reveals a path for understanding forces inherent in plant cells. [ABSTRACT FROM AUTHOR]

The ability of plants to absorb CO2 for photosynthesis and transport water from root to shoot depends on the reversible swelling of guard cells that open stomatal pores in the epidermis. Despite decades of experimental and theoretical work, the biomechanical drivers of stomatal opening and closure are still not clearly defined. We combined mechanical principles with a growing body of knowledge concerning water flux across the plant cell membrane and the biomechanical properties of plant cell walls to quantitatively test the long-standing hypothesis that increasing turgor pressure resulting from water uptake drives guard cell expansion during stomatal opening. To test the alternative hypothesis that water influx is the main motive force underlying guard cell expansion, we developed a system dynamics model accounting for water influx. This approach connects stomatal kinetics to whole plant physiology by including values for water flux arising fromwater status in the plant. [ABSTRACT FROM AUTHOR]

Plant cell deformations are driven by cell pressurization and mechanical constraints imposed by the nanoscale architecture of the cell wall, but how these factors are controlled at the genetic and molecular levels to achieve different types of cell deformation is unclear. Here, we used stomatal guard cells to investigate the influences of wall mechanics and turgor pressure on cell deformation and demonstrate that the expression of the pectin-modifying gene PECTATE LYASE LIKE12 (PLL12) is required for normal stomatal dynamics in Arabidopsis thaliana. Using nanoindentation and finite element modeling to simultaneously measure wall modulus and turgor pressure, we found that both values undergo dynamic changes during induced stomatal opening and closure. PLL12 is required for guard cells to maintain normal wall modulus and turgor pressure during stomatal responses to light and to tune the levels of calcium crosslinked pectin in guard cell walls. Guard cell-specific knockdown of PLL12 caused defects in stomatal responses and reduced leaf growth, which were associated with lower cell proliferation but normal cell expansion. Together, these results force us to revise our view of how wall-modifying genes modulate wall mechanics and cell pressurization to accomplish the dynamic cellular deformations that underlie stomatal function and tissue growth in plants. [ABSTRACT FROM AUTHOR]

Xyloglucan is believed to play a significant role in cell wall mechanics of dicot plants. Surprisingly, Arabidopsis plants defective in xyloglucan biosynthesis exhibit nearly normal growth and development. We investigated a mutant line, cslc-Δ5 , lacking activity in all five Arabidopsis cellulose synthase like-C (CSLC) genes responsible for xyloglucan backbone biosynthesis. We observed that this xyloglucan-deficient line exhibited reduced cellulose crystallinity and increased pectin levels, suggesting the existence of feedback mechanisms that regulate wall composition to compensate for the absence of xyloglucan. These alterations in cell wall composition in the xyloglucan-absent plants were further linked to a decrease in cell wall elastic modulus and rupture stress, as observed through atomic force microscopy (AFM) and extensometer-based techniques. This raised questions about how plants with such modified cell wall properties can maintain normal growth. Our investigation revealed two key factors contributing to this phenomenon. First, measurements of turgor pressure, a primary driver of plant growth, revealed that cslc-Δ5 plants have reduced turgor, preventing the compromised walls from bursting while still allowing growth to occur. Second, we discovered the conservation of elastic asymmetry (ratio of axial to transverse wall elasticity) in the mutant, suggesting an additional mechanism contributing to the maintenance of normal growth. This novel feedback mechanism between cell wall composition and mechanical properties, coupled with turgor pressure regulation, plays a central role in the control of plant growth and is critical for seedling establishment in a mechanically challenging environment by affecting shoot emergence and root penetration. • Normal xyloglucan levels are important for seedling establishment • Changes in xyloglucan content affect turgor pressure • Reduced xyloglucan induces a reduction in cellulose crystallinity • Reduced xyloglucan leads to softer and weaker cell walls Bou Daher et al. show that Arabidopsis mutants lacking xyloglucan have reduced cellulose crystallinity, altered cell wall mechanical properties, and a reduction in turgor pressure. Bou Daher et al. also demonstrate the importance of xyloglucan in early seedling establishment. [ABSTRACT FROM AUTHOR]

How the plant cell wall expands and forms shapes is a long-standing mystery. Traditional thought is that turgor pressure drives these processes. However, a recent study by Haas and colleagues shows for the first time that the expansion of pectin homogalacturonan nanofilaments drives morphogenesis without turgor pressure in plant epidermal cells. [ABSTRACT FROM AUTHOR]

The shape and function of plant cells are often highly interdependent. The puzzle- shaped cells that appear in the epidermis of many plants are a striking example of a complex cell shape, however their functional benefit has remained elusive. We propose that these intricate forms provide an effective strategy to reduce mechanical stress in the cell wall of the epidermis. When tissue-level growth is isotropic, we hypothesize that lobes emerge at the cellular level to prevent formation of large isodiametric cells that would bulge under the stress produced by turgor pressure. Data from various plant organs and species support the relationship between lobes and growth isotropy, which we test with mutants where growth direction is perturbed. Using simulation models we show that a mechanism actively regulating cellular stress plausibly reproduces the development of epidermal cell shape. Together, our results suggest that mechanical stress is a key driver of cell-shape morphogenesis. [ABSTRACT FROM AUTHOR]

This paper provides a mini-review of evidence for negative turgor pressure in leaf cells starting with experimental evidence in the late 1950s and ending with biomechanical models published in 2014. In the present study, biomechanical models were used to predict how negative turgor pressure might be manifested in dead tissue, and experiments were conducted to test the predictions. The main findings were as follows: (i) Tissues killed by heating to 60 or 80 °C or by freezing in liquid nitrogen all became equally leaky to cell sap solutes and all seemed to pass freely through the cell walls. (ii) Once cell sap solutes could freely pass the cell walls, the shape of pressure-volume curves was dramatically altered between living and dead cells. (iii) Pressure-volume curves of dead tissue seem to measure negative turgor defined as negative when inside minus outside pressure is negative. (iv) Robinia pseudoacacia leaves with small palisade cells had more negative turgor than Metasequoia glyptostroboides with large cells. (v) The absolute difference in negative turgor between R. pseudoacacia and M. glyptostroboides approached as much as 1.0 MPa in some cases. The differences in the manifestation of negative turgor in living versus dead tissue are discussed. [ABSTRACT FROM AUTHOR]

While many molecular players involved in growth control have been identified in the past decades, it is often unknown how they mechanistically act to induce specific shape changes during development. Plant morphogenesis results from the turgor-induced yielding of the extracellular and load-bearing cell wall. Its mechanochemical equilibrium appears as a fundamental link between molecular growth regulation and the effective shape evolution of the tissue. We focus here on force-driven polymerization of the cell wall as a central process in growth control. We propose that mechanical forces facilitate the insertion of wall components, in particular pectins, a process that can be modulated through genetic regulation. We formalize this idea in a mathematical model, which we subsequently test with published experimental results. [ABSTRACT FROM AUTHOR]

During desiccation, plant cells are subjected to very low water potentials. Osmoregulation through increase of soluble materials ( e.g. soluble sugars, compatible inorganic ions) is a response to the decreasing turgor pressure in the cells. In bryophytes, sucrose acts as an osmolyte and also stabilizing membranes and proteins through vitrification. We used psychrometric measurements in the aquatic bryophyte Fontinalis antipyretica Hedw. to construct pressure–volume isotherms and determine the water relations parameters under fast and slow dehydration rates. Sucrose was also quantified. The starting hypothesis was that a slow dehydration rate would increase sucrose concentration, thereby decreasing the osmotic potential at turgor loss point, and would also increase cell wall elasticity, postponing turgor loss and allowing time for induction of molecular and structural acclimation mechanisms. In fact, we found that slowly dehydrated samples presented more elastic cell walls, allowing cells to shrink and maintain turgor, helping to better preserve their metabolic functions and therefore to induce desiccation tolerance (DT). On the other hand, in fast dehydrated samples the osmotic potential at turgor loss point decreased, indicating the activity of osmoregulation processes, possibly connected to the increase observed in sucrose content. Upon rehydration, fast dried samples lost 50% of the sucrose through leakage due to cell membrane rupture, while slow dehydrated leaves maintained their sucrose content constant. DT appears to be achieved through slow dehydration, meaning that a high sucrose content alone does not contribute to DT establishment. Moreover, in natural conditions external water can be maintained at very high values due to the life form of F. antipyretica , which grows in long and compact floating stems in streams, allowing a slow dehydration rate required for induction of other DT mechanisms. [ABSTRACT FROM AUTHOR]

One of the main differences between plant and animal cells are the walls surrounding plant cells providing structural support during development and protection like an adaptive armor against biotic and abiotic stress. During recent years it has become widely accepted that plant cells use a dedicated system to monitor and maintain the functional integrity of their walls. Maintenance of integrity is achieved by modifying the cell wall and cellular metabolism in order to permit tightly controlled changes in wall composition and structure. While a substantial amount of evidence supporting the existence of the mechanism has been reported, knowledge regarding its precise mode of action is still limited. The currently available evidence suggests similarities of the plant mechanism with respect to both design principles and molecular components involved to the very well characterized system active in the model organism Saccharomyces cerevisiae. There the system has been implicated in cell morphogenesis as well as response to abiotic stresses such as osmotic challenges. Here the currently available knowledge on the yeast system will be reviewed initially to provide a framework for the subsequent discussion of the plant cell wall integrity maintenance mechanism. The review will then end with a discussion on possible design principles for the cell wall integrity maintenance mechanism and the function of the plant turgor pressure in this context. [ABSTRACT FROM AUTHOR]

Hydrodynamics, cell wall and ion dynamics are all important properties that regulate pollen tube growth. Currently, the two main pollen tube growth models, the cell wall model and the hydrodynamic model do not appear to be reconcilable. Here we develop an integrative model for pollen tube growth and show that our model reproduces key experimental observations: (1) that the hypertonic condition leads to a much longer oscillatory period and that the hypotonic condition halves the oscillatory period; (2) that oscillations in turgor are experimentally undetectable; (3) that increasing the extracellular calcium concentration or decreasing the pH decreases the growth oscillatory amplitude; (4) that knockout of Raba4d, a member of the Rab family of small GTPase proteins, decreases pollen tube length after germination for 24 h. Using the model generated here, we reveal that (1) when cell wall extensibility is large, pollen tube may sustain growth at different volume changes and maintain relatively stable turgor; (2) turgor increases if cell wall extensibility decreases; (3) increasing turgor due to decrease in osmolarity in the media, although very small, increases volume change. However, increasing turgor due to decrease in cell wall extensibility decreases volume change. In this way regulation of pollen tube growth by turgor is context dependent. By changing the osmolarity in the media, the main regulatory points are extracellular osmolarity for water flow and turgor for the volume encompassed by the cell wall. However, if the viscosity of cell wall changes, the main regulatory points are turgor for water flow and wall extensibility for the volume encompassed by the cell wall. The novel methodology developed here reveals the underlying context-dependent regulatory principle of pollen tube growth. [ABSTRACT FROM AUTHOR]

Hydrodynamics, cell wall and ion dynamics are all important properties that regulate pollen tube growth. Currently, the two main pollen tube growth models, the cell wall model and the hydrodynamic model do not appear to be reconcilable. Here we develop an integrative model for pollen tube growth and show that our model reproduces key experimental observations: 1) that the hypertonic condition leads to a much longer oscillatory period and that the hypotonic condition halves the oscillatory period; 2) that oscillations in turgor are experimentally undetectable; 3) that increasing the extracellular calcium concentration or decreasing the pH decreases the growth oscillatory amplitude; 4) that knockout of Raba4d, a member of the Rab family of small GTPase proteins, decreases pollen tube length after germination for 24 hours. Using the model generated here, we reveal that 1) when cell wall extensibility is large, pollen tube may sustain growth at different volume changes and maintain relatively stable turgor; 2) turgor increases if cell wall extensibility decreases; 3) increasing turgor due to decrease in osmolarity in the media, although very small, increases volume change . However, increasing turgor due to decrease in cell wall extensibility decreases volume change. In this way regulation of pollen tube growth by turgor is context dependent. By changing the osmolarity in the media, the main regulatory points are extracellular osmolarity for water flow and turgor for the volume encompassed by the cell wall. However, if the viscosity of cell wall changes, the main regulatory points are turgor for water flow and wall extensibility for the volume encompassed by the cell wall. The novel methodology developed here reveals the underlying context-dependent regulatory principle of pollen tube growth. [ABSTRACT FROM AUTHOR]

Highlights: [•] A generalized form of a strain energy function is proposed. [•] A mathematical relation between turgor pressure and stretch ratio is developed. [•] The developed relation shows good fit with experimental data. [•] Effects of turgor pressure on cell walls of apple and potato tissues are discussed. [ABSTRACT FROM AUTHOR]

Focus has centered on C-partitioning in stems of sugarcane (Saccharum sp.) due to their high-sucrose accumulation features, relevance to other grasses, and rising economic value. Here we review how sugarcane balances between sucrose storage, respiration, and cell wall biosynthesis. The specific topics involve 1) accumulation of exceptionally high sucrose levels (up to over 500 mM), 2) a potential, turgor-sensitive system for partitioning sucrose between storage inside (cytosol and vacuole) and outside cells, 3) mechanisms to prevent back-flow of extracellular sucrose to xylem or phloem, 4) apparent roles of sucrose-P-synthase in fructose retrieval and sucrose re-synthesis, 5) enhanced importance of invertases, and 6) control of C-flux at key points in cell-wall biosynthesis (UDP-glucose dehydrogenase) and respiration (ATP- and pyrophosphate-dependent phospho-fructokinases). A combination of emerging technologies is rapidly enhancing our understanding of these points and our capacity to shift C-flux between sucrose, cell-wall polymers, or other C-sinks. [ABSTRACT FROM AUTHOR]

A cell-growth-based algorithm is presented based on the biomechanics of plant cells in tissues to help explain the typical differences in cellular architecture found between different pome fruit species, cultivars and tissues. The cell was considered as a closed thin-walled structure, maintained in tension by turgor pressure. The cell walls of adjacent cells were modelled as parallel and linearly elastic elements, which obeyed Hooke's law. A Voronoi tessellation was used to generate the initial topology of the cells. Cell expansion then resulted from turgor pressure acting on the yielding cell wall material. To find the sequence positions of each vertex of the cell walls, and thus, the shape of the cells with time, a system of differential equations for the positions and velocities of each vertex were established and solved using a Runge-Kutta fourth and fifth order (ODE45) method. The model was used to generate realistic 2D fruit tissue structures composed of cells of random shapes and sizes, cell walls and intercellular spaces. Comparison was made with fruit tissue micrographs. The virtual tissues can be used for numerical simulation of heat and mass transfer phenomena or mechanical deformation during controlled atmosphere storage of fresh pome fruit. [ABSTRACT FROM AUTHOR]

Plant cell expansion is controlled by a fine-tuned balance between intracellular turgor pressure, cell wall loosening and cell wall biosynthesis. To understand these processes, it is important to gain in-depth knowledge of cell wall mechanics. Pollen tubes are tip-growing cells that provide an ideal system to study mechanical properties at the single cell level. With the available approaches it was not easy to measure important mechanical parameters of pollen tubes, such as the elasticity of the cell wall. We used a cellular force microscope ( CFM) to measure the apparent stiffness of lily pollen tubes. In combination with a mechanical model based on the finite element method ( FEM), this allowed us to calculate turgor pressure and cell wall elasticity, which we found to be around 0.3 MPa and 20-90 MPa, respectively. Furthermore, and in contrast to previous reports, we showed that the difference in stiffness between the pollen tube tip and the shank can be explained solely by the geometry of the pollen tube. CFM, in combination with an FEM-based model, provides a powerful method to evaluate important mechanical parameters of single, growing cells. Our findings indicate that the cell wall of growing pollen tubes has mechanical properties similar to rubber. This suggests that a fully turgid pollen tube is a relatively stiff, yet flexible cell that can react very quickly to obstacles or attractants by adjusting the direction of growth on its way through the female transmitting tissue. [ABSTRACT FROM AUTHOR]

Turgor pressure in plant cells is involved in many important processes. Stable and normal turgor pressure is required for healthy growth of a plant, and changes in turgor pressure are indicative of changes taking place within the plant tissue. The ability to quantify the turgor pressure of plant cells in vivo would provide opportunities to understand better the process of pressure regulation within plants, especially when plant stress is considered, and to understand the role of turgor pressure in cellular signaling. Current experimental methods do not separate the influence of the turgor pressure from the effects associated with deformation of the cell wall when estimates of turgor pressure are made. In this paper, nanoindentation measurements are combined with finite element simulations to determine the turgor pressure of cells in vivo while explicitly separating the cell-wall properties from the turgor pressure effects. Quasi-static cyclic tests with variable depth form the basis of the measurements, while relaxation tests at low depth are used to determine the viscoelastic material properties of the cell wall. Turgor pressure is quantified using measurements on Arabidopsis thaliana under three pressure states (control, turgid and plasmolyzed) and at various stages of plant development. These measurements are performed on cells in vivo without causing damage to the cells, such that pressure changes may be studied for a variety of conditions to provide new insights into the biological response to plant stress conditions. [ABSTRACT FROM AUTHOR]

Significant controversy still swirls around the regulation of extension by tip-growing cells, particularly during stable, oscillatory growth of pollen tubes. One explanation proposes that turgor pressure is both the controlling and driving force. We refute this hypothesis on theoretical and evidentiary grounds. Direct measurement of intracellular pressure reveals constant turgor even as growth rates change. Measured ion fluxes, notably potassium, are insufficient to account for the requisite osmotic changes. Water movement, and hence pressure gradients, occur throughout the cell, unrestricted to local domains. Increases in hydrostatic pressure alone would force water out of the cell rather than cause increased growth. We have recently demonstrated concomitant changes in the apical cell wall that account fully for observed changes in growth rate. [Copyright &y& Elsevier]

Low concentrations of some trace metals markedly reduce root elongation rate and cause ruptures to root rhizodermal and outer cortical cells in the elongation zone. The interactions between the trace metals and plant components responsible for these effects are not well understood but may be linked to changes in water uptake, cell turgor and cell wall extensibility. An experiment was conducted to investigate the effects of Al, La, Cu, Gd, Sc and Ru on the saturated hydraulic conductivity of bacterial cellulose (BC)–pectin composites, used as plant cell wall analogs. Hydraulic conductivity was reduced to ≈30% of the initial flow rate by 39 µ M Al and 0.6 µ M Cu, ≈40% by 4.6 µ M La, 3 µ M Sc and 4.4 µ M Ru and ≈55% by 3.4 µ M Gd. Scanning electron microscopy (SEM) revealed changes in the ultrastructure of the composites. The results suggest that trace metal binding decreases the hydraulic conductivity through changes in pectin porosity. The experiment illustrates the importance of metal interactions with pectin, and the implications of such an interaction in plant metal toxicity and in normal cell wall processes. [ABSTRACT FROM AUTHOR]

Abstract: Cellular morphogenesis involves changes to cellular size and shape which in the case of walled cells implies the mechanical deformation of the extracellular matrix. So far, technical challenges have made quantitative mechanical measurements of this process at subcellular scale impossible. We used micro-indentation to investigate the dynamic changes in the cellular mechanical properties during the onset of spatially confined growth activities in plant cells. Pollen tubes are cellular protuberances that have a strictly unidirectional growth pattern. Micro-indentation of these cells revealed that the initial formation of a cylindrical protuberance is preceded by a local reduction in cellular stiffness. Similar cellular softening was observed before the onset of a rapid growth phase in cells with oscillating growth pattern. These findings provide the first quantitative cytomechanical data that confirm the important role of the mechanical properties of the cell wall for local cellular growth processes. They are consistent with a conceptual model that explains pollen tube oscillatory growth based on the relationship between turgor pressure and tensile resistance in the apical cell wall. To further confirm the significance of cell mechanics, we artificially manipulated the mechanical cell wall properties as well as the turgor pressure. We observed that these changes affected the oscillation profile and were able to induce oscillatory behavior in steadily growing tubes. [Copyright &y& Elsevier]

Hand sections of root and stem xylem of diverse species of Nymphaea (including Ondinea) were studied with SEM in order to explore the diversity of wall structure within the genus. Lateral walls of root tracheary elements are untextured at the magnifications used, but end wall structure suggests that lysis of pit membranes leaves a cellulosic network with a large reticulum. Stem tracheids have lateral walls that are untextured or have a few prominent fibrils, but end walls have a dense accretion of coarse fibrils. These fibrils form a spongiform or compressed network overlying the pit membranes. In addition, on the lumen side of pit membranes, coarse fibrils cross the pits in an axial direction. These coarse fibrils are connected to pit borders, and they may extend onto wall surfaces of the tracheary elements. These fibrillar accretions are considered here to fulfill criteria as secondary-wall components. Variation within the genus with respect to the fibrillar accretions may occur, but there may be more extensive diversity within any given sample than among species. Root tracheary elements could be considered to be vessel elements because of the extensive removal of wall material from the end-wall pit membranes. Stem tracheary elements qualify as tracheids because, rather than showing lysis of pit-membrane portions, end-wall pit membranes bear fibrillar accretions that are only sparsely porose. The absence of coarse fibrillar accretions on lateral walls of stem tracheids (on tracheid-to-parenchyma interfaces) provides difficulty for any hypothesis that would relate wall thickenings to turgor pressure. The patterns of fibrillar accretion observed in Nymphaea include all patterns thus far reported from stem tracheids in the other genera of the family, and they must be considered to be a characteristic of Nymphaeaceae. [ABSTRACT FROM AUTHOR]

Abstract: In CAM-plants rising levels of malic acid in the early morning cause elevated turgor pressures in leaf chlorenchyma cells. Under specific conditions this process is lethal for sensitive plants resulting in chlorenchyma cell burst while other species can cope with these high pressures and do not show cell burst under comparable conditions. The non-cellulosic polysaccharide composition of chlorenchyma cell walls was investigated and compared in three cultivars of Aechmea with high sensitivity for chlorenchyma cell burst and three cultivars with low sensitivity. Chlorenchyma layers were cut from the leaf and the non-cellulosic carbohydrate fraction of the cell wall fraction was analyzed by gas–liquid chromatography. Glucuronoarabinoxylans (GAXs) were the major non-cellulosic polysaccharides in Aechmea. The fine structure of these GAXs was strongly related to chlorenchyma wall strength. Chlorenchyma cell walls from cultivars with low sensitivity to cell burst were characterized by an A/X ratio of ca. 0.13 while those from cultivars with high sensitivity showed an A/X ratio of ca. 0.23. Xylose chains from cultivars with high cell burst sensitivity were ca. 40% more substituted with arabinose compared to cultivars with low sensitivity for cell burst. The results indicate a relationship in vivo between glucuronoarabinoxylan fine structure and chlorenchyma cell wall strength in Aechmea. The evidence obtained supports the hypothesis that GAXs with low degrees of substitution cross-link cellulose microfibrils, while GAXs with high degrees of substitution do not. A lower degree of arabinose substitution on the xylose backbone implies stronger cell walls and the possibility of withstanding higher internal turgor pressures without cell bursting. [Copyright &y& Elsevier]

In this article we investigate aspects of turgor-driven plant cell growth within the framework of a model derived from the Eulerian concept of instability. In particular we explore the relationship between cell geometry and cell turgor pressure by extending loss of stability theory to encompass cylindrical cells. Beginning with an analysis of the three-dimensional stress and strain of a cylindrical pressure vessel, we demonstrate that loss of stability is the inevitable result of gradually increasing intemal pressure in a cylindrical cell. The turgor pressure predictions based on this model differ from the more traditional viscoelastic or creep-based models in that they incorporate both cell geometry and wall mechanical properties in a single term. To confirm our predicted working turgor pressures, we obtained wall dimensions, elastic moduli, and turgor pressures of sequential internodal cells of intact Chant coral/ma plants by direct measurement. The results show that turgor pressure predictions based on loss of stability theory fall within the expected physiological range of turgor pressures for this plant. We also studied the effect of varying wall Poisson's ratio v on extension growth in living cells, showing that while increasing elastic modulus has an understandably negative effect on wall expansion, increasing Poisson's ratio would be expected to accelerate wall expansion. [ABSTRACT FROM AUTHOR]

From a mechanical point of view, plant and hyphal cells are more complex than their animal counterparts because the variety of structural components determining cellular architecture is broader. In addition to cytoskeletal elements and the plasma membrane, the cell wall and turgor pressure equip plant and hyphal cells with structures analogous to an exoskeleton and a hydroskeleton, respectively. To quantify the physical properties of plant and hyphal cells, researchers have developed a plethora of experimental methods. This review provides an overview of experimental approaches that have been used to measure turgor pressure and to determine the mechanical properties of the plant cell wall at the subcellular level. It is completed by a glimpse into the arsenal of techniques that has been used to characterize the physical properties of cytoskeletal elements. These have mostly been used on animal cells, but we hope they will find their way into plant cell research. Finally, assays and tests to measure the generation of forces by cells and subcellular structures are discussed. [ABSTRACT FROM AUTHOR]

The conventional method of measuring plant cell turgor pressure is the pressure probe but applying this method to single cells in suspension culture is technically difficult and requires puncture of the cell wall. Conversely, compression testing by micromanipulation is particularly suited to studies on single cells, and can be used to characterise cell wall mechanical properties, but has not been used to measure turgor pressure. In order to demonstrate that the micromanipulation method can do this, pressure measurements by both methods were compared on single suspension-cultured tomato ( Lycopersicon esculentum vf36) cells and generally were in good agreement. This validates further the micromanipulation method and demonstrates its capability to measure turgor pressure during water loss. It also suggests that it might eventually be used to estimate plant cell hydraulic conductivity. [ABSTRACT FROM AUTHOR]

• Background and Aims New wall deposition usually accompanies plant growth. External osmotica inhibit both processes but wall precursors continue to be synthesized, and exocytosis follows. Consequently, the osmotica appear to act outside of the plasma membrane. Because this implies an action of turgor pressure (P) on the periplasm by unknown mechanisms, the following study was undertaken to determine whether P could act in a way that altered wall deposition and assembly in the periplasm while the cells grow.• Methods Cells of Chara corallina were exposed to P slightly below normal by using a pressure probe while supplying inorganic carbon in light. After labelling, the walls were isolated and the amount of new wall was determined. Similar measurements were made after treatment with osmotica. Chlortetracycline-stimulated exocytosis was determined microscopically. Polysaccharide properties were determined by confocal microscopy and vapour pressure osmometry in an ‘artificial periplasm’ in isolated Chara cell walls, using labelled dextran as an anologue of hemicellulose, and polygalacturonate as pectin.• Key Results Rapid growth and wall deposition occurred at normal P of 0.5 MPa but both processes decreased when P was lowered 0.1 MPa. Inorganic carbon uptake and exocytosis were unaffected. In the artificial periplasm, normal P caused high polysaccharide concentrations and rapid polysaccharide entry into the wall, and gel formation in the pectin. Lowering P decreased entry and gel formation.• Conclusions This is the first indication that normal P of 0.5 MPa can concentrate periplasmic polysaccharides sufficiently to cause cross-linking and gel formation in pectins while simultaneously fostering the entry of large polysaccharides into small interstices in the existing wall. This P-action would thicken the primary wall and form a smooth transition between the new and old structure, suggesting a molecular mechanism of wall deposition and assembly while the wall extends. [ABSTRACT FROM AUTHOR]

Biomechanical studies aim at understanding the relationship between the mechanical properties of biological structures and their function. In cytomechanical investigations, this approach is brought down to the scale of cells and subcellular structures. In plant cells and the hyphae of fungi and water molds, interactions between turgor pressure, the cell wall, and the cytoskeleton are considered of primary importance. This review is an overview of how the mechanical properties of these individual features and their interactions have been measured and how the experimental data are used to produce theoretical mechanical models of cellular architecture and dynamics. Several models are discussed, and focusing on the example of tip-growing cells, various approaches to understanding the mechanical aspects of cellular morphogenesis are analyzed. [ABSTRACT FROM AUTHOR]

This study addresses the mechanism of wall stress relaxation in growing plant cells. The current viscoelastic model of cell wall relaxation, which dates from the work of Preston, Cleland, Lockhart, and others in the 1960s, has serious shortcomings. It has been shown however that the theory of loss of stability (LOS) can be applied to materials in tension, leading to the conclusion that the relaxation of stresses in the walls of any pressure vessel is rigorously modeled using LOS. We propose that LOS also provides a more appropriate and versatile model of stress relaxation in growing plant cells. We argue that when treated as a manifestation of LOS, the regulation of cell turgor has a rigorous and demonstrable basis in the geometrical and physical properties of the cell wall and the cell''s ability to import water. Thus plant cell growth can be regarded as an inherently self-limiting process, tunable by biochemical or structural means. Lastly, despite the current limitations of our model, we apply direct measurement of elastic modulus, wall thickness and cell radius obtained from cylindrical Chara corallina cells to generate an initial calculation of critical pressures in a hypothetical spherical cell with the same material properties. [Copyright &y& Elsevier]

Based on a single regular cell structural model, the effects of loading rate on the compressive behaviour of plant and vegetative tissues have been qualitatively investigated. The cell walls were treated as a polymeric composite material with microfibrils embedded in the highly structured cell wall matrix. The rubber elasticity, the turgor permeability and the loading rate were taken into account to qualitatively predict the tissue stiffness, cell wall stress, turgor pressure, cell debonding force, and the percentage of weight loss of the cell fluid. The predicted results are consistant with the related experimental phenomena. [ABSTRACT FROM AUTHOR]

Examines the factors affecting the tensile properties of Arabidopsis cell walls. Accounts on the consequence of turgor pressure within the cell; Measurement of the rate of plant cell expansion; Emphasis on the evidence for the independence of cellulose and pectin networks.

Experimental values of the stiffness of carrot tissue were compared with the predictions from models of fluid-filled closed cell foams. Compared to an experimental value of 7 MPa for fresh carrot, predictions gave moduli in the range 2 to 33 MPa based on a compressible fluid, an isotonic state turgor pressure of 0.8 MPa, a cell wall modulus of 100 MPa and cell lengths from 54 to 3 times the wall thickness. The modulus was predicted to increase linearly with turgor pressure in agreement with experiment for turgor pressures up to about 1 MPa, whereafter the experimental modulus increased more sharply, reaching a value of 14 MPa at a turgor pressure of 2.1 MPa, closer to the predicted rate of increase in an earlier shell model. Predictions based on an incompressible fluid gave initial and equilibrium moduli of 14 and 2 MPa, respectively, in agreement with the experimental values at high and low levels of turgor, again assuming a cell wall modulus of 100 MPa but requiring small cells of length ratio less than three times their wall thickness. [ABSTRACT FROM AUTHOR]

Two mechanisms have been proposed as the primary control of oscillating tip growth in Lilium longiflorum Thunb. pollen tubes: changes in cell wall strength (Holdaway-Clarke et al. 1997) or alternatively, changes in turgor pressure (Messerli et al. 2000). Here we have modified the ionic and osmotic concentrations of the growth medium to test predictions derived from both models. Raising the [Ca2+]o tenfold above normal reduced the amplitude of the [Ca2+]i oscillations and growth oscillations while it raised the basal [Ca2+]i and growth rate such that the average growth rate did not change. Raising the [H+] of the growth medium tenfold reversibly decreased and sometimes eliminated the [Ca2+]i and growth oscillations without changing the average growth rate. Lowering the [H+] tenfold led to irregular frequency and amplitude [Ca2+]i oscillations, reduced the average growth rate of tubes and led to cell bursting in 33% of tubes. Addition of 50 mM H+ buffer, MES, to prevent pH changes in the cell wall increased the period, amplitude and duration of both [Ca2+]i and growth oscillations. Changing the [K+]o did not markedly effect [Ca2+]i oscillations. Reducing the osmolarity of the medium led to transient large-amplitude [Ca2+]i and growth oscillations while reducing large-amplitude oscillations over long periods. In many different conditions under which growth still occurs, lily pollen tubes maintain growth oscillations, albeit with modified frequency, amplitude and duration. We conclude that modifications to both proposed models are necessary to explain oscillating growth in this system. [ABSTRACT FROM AUTHOR]

BSTRACT: Melon samples were equilibrated with solutions of increasing mannitol concentration in the range 0.00 to 0.80 M for the purpose of evaluating turgor pressure through volume change measurement. Melon compression and relaxation behaviors were evaluated with an Instron Testing machine, and structural characteristics were observed through microscopy. Immersion in mannitol solutions affected tissue firmness, reducing its value in all studied cases. After plasmolysis and plasmotypsis, samples became less elastic and more viscous, and firmness reduced more than 50%. Textural parameters obtained and microscopy results indicated that, in general, fracture might be associated with cell-wall integrity, and residual relaxation force might be associated with tissue turgor pressure. [ABSTRACT FROM AUTHOR]

ABSTRACTBoth turgor pressure and differences in membrane tension are capable of providing an energy input into exocytosis, the process of fusion of Golgi vesicles with the cell membrane in plants. It is shown that the contribution of turgor pressure is much larger than that of membrane tension, so that the exocytotic process is not likely on thermodynamic grounds to be reversible unless another source of energy is made available. However, recycling of membrane material as flattened, empty vesicles is energetically possible and is likely to be favoured when the magnitude of membrane tension in the cell membrane is low. Thus the outward flows of membrane and cell wall material are in principle linked to turgor, whereas membrane tension influences the inward flow of membrane material. [ABSTRACT FROM AUTHOR]

The pressure probe is a micro manometer for the simultaneous direct recording and manipulation of plant cell hydrostatic pressure. It is used to map in space and time the turgor pressures of individual cells within tissues and organs of intact plants. This is used to study the hydraulic architecture of tissues, tissue movement and the responses of tissues to water stress. The approach can be augmented by simultaneous measurement of individual cell osmotic pressure. This permits the hydraulic driving forces across selectively permeable membranes and walls to be assessed fully. By manipulating manually the pressure, cell wall elasticity and its properties can also be mapped. Under some conditions this can be extended to plastic behaviour. [ABSTRACT FROM AUTHOR]

Leaf extension was stimulated following exposure of three interamerican hybrid poplar clones (Populus trichocarpa × P. deltoides); 'Unal', 'Boelare', and 'Beaupre' and a euramerican clone 'Primo' (Populus nigra × P. deltoides) to elevated CO2 in controlled environment chambers. For all three interamerican clones the evidence suggests that this was the result of increased leaf cell expansion associated with enhanced cell wall extensibility (WEx), measured as tensiometric increases in cell wall plasticity. For the interamerican clone 'Boelare', there was also a significant increase in cell wall elasticity following exposure to elevated CO2 (P ≤ 0.001). The effect of elevated CO2 in stimulating cell wall extensibility was confirmed in a detailed spatial analysis of extensibility made across the lamina of expanding leaves of the clone 'Boelare. For two of the interamerican hybrids, 'Unal' and 'Beaupre', both leaf cell water potential (ψ) and turgor pressure (P) were lower in elevated than in ambient CO2. By contrast, no significant effects on the cell wall properties or leaf water relations for the euramerican hybrid 'Primo' were observed following exposure to elevated CO2, suggesting that the mechanism for increased leaf extension in elevated CO2 differed, depending on clone. The cumulative total length of leaves of 'Boelare' grown in elevated CO2 was significantly increased (P ≤ 0.05) and since leaf number was not significantly increased in any inter-american clone it is hypothesized that final leaf size was stimulated in elevated CO2 for these clones. By contrast, there was no significant effect of CO2 on cumulative total leaf length for the euramerican clone 'Primo', but leaf number was significantly increased by elevated CO2. [ABSTRACT FROM AUTHOR]

The effect of osmotic stress on the regulation of extension of roots of pine seedlings (Pinus pinaster Ait.) was studied. Extension rate (r) was measured as increase in root length, and as local growth rate (LGR) along the growing region. Turgor pressure (P) was measured over the same profile using the cell pressure probe. Correlation of LGR and P allowed changes in cell wall mechanical properties to be followed. In unstressed plants, differences in LGR along the root apex were owing to variations in the mechanical properties of the cell walls since P remained constant at 0.5 MPa along the first 10 mm of the root tip. Contrary to expectation, a moderate water stress of 0.15 MPa stimulated root growth despite a small reduction in P to 0.4 MPa. The walls of the expanding cells became looser, allowing enhanced growth in spite of reduced turgor. Examination of LGR showed that these changes occurred uniformly over the whole elongation zone. The advantage to the plant of this is discussed. Higher stresses (0.45 and 0.66 MPa) decreased cell turgor to a greater degree (⩽ 0.3 MPa) and decreased growth rate. At the highest stress (0.66 MPa), P was maximal in cells that were still elongating, resulting in a longitudinal gradient in turgor. In both treatments LGR remained unchanged at the apex of the growing zone. Wall loosening appeared to have occurred in this region. Pine root extension is therefore modulated by changes in cell wall properties, with cell wall loosening resulting in increased growth at moderate stress. It appears that growth under higher stresses is limited by inability to maintain turgor. [ABSTRACT FROM AUTHOR]

The expansion of roots is considered at the level of the single cell. The water relations of cell expansion are discussed. Water entry, solute import and cell wall properties are considered as possible regulatory points. It is argued that root cell expansion can be understood in terms of cell turgor pressure and the physical properties of the cell wall, provided solute supply is not limiting. Various measurements of cell wall properties in roots are presented and the assumptions underlying their measurements are presented. It is concluded that cell wall properties must he measured over short time periods to prevent alterations in wall properties during the experiment. The radial location of the load-hearing layers is discussed and it is concluded that, unlike aerial tissue, growth is limited by the properties of the inner layer of the root cortex. Evidence is presented to show that cell wall properties can change both during development and following turgor perturbation. In general, however, turgor itself is tightly regulated, particularly towards the root tip. A number of environmental situations are presented in which root growth is altered. The mechanism of the alteration is discussed at the single cell level. These 'stresses' include osmotic stress, low temperature and soil compaction. In many cases the alteration of root growth is consistent with changes in the cell wall properties of the growing cells. Severe stress. resulting in near cessation of root cell extension, can result in a change (usually an increase) in turgor pressure. The change in turgor pressure of the cells in the growing zone is smaller than that which would be expected from a continuation of an unstressed solute import rate. This exemplifies both the change in cell wall properties and the tight turgor homeostasis of root tips. The biochemical processes which underlie the modulation of cell wall properties are presented as they are currently understood in roots. [ABSTRACT FROM AUTHOR]

Expansive growth of plant cells is controlled principally by processes that loosen the wall and enable it to expand irreversibly. The central role of wall relaxation for cell expansion is reviewed. The most common methods for assessing the extension properties of plant cell walls ('wall extensibility') are described, categorized and assessed critically. What emerges are three fundamentally different approaches which test growing cells for their ability (a) to enlarge at different values of turgor, (b) to induce wall relaxation, and (c) to deform elastically or plastically in response to an applied tensile force. Analogous methods with isolated walls are similarly reviewed. The results of these different assays are related to the nature of plant cell growth and pertinent biophysical theory. I argue that the 'extensibilities' measured by these assays are fundamentally different from one another and that some are more pertinent to growth than others. [ABSTRACT FROM AUTHOR]

Leaf water (ψ) and solute (ψs) potential were measured in field sorghum and maize under well irrigated (I) and dryland (NI) conditions throughout a season. Despite decreases in ψ due to slow soil water depletion and to apparent increases in liquid phase plant resistance, midday leaf turgor (ψp) in the NI sorghum was maintained at similar levels as in the I treatment throughout the season due to concomitant decreases in ψs. Osmotic adjustment was also observed in maize, although ψp was significantly lower in the NI treatment as compared to I during the final stages of grain filling. A seasonal shift in the ψ vs. relative water content relation of NI sorghum leaves was observed, more water being retained by the older leaf at any particular ψ. The major factor for turgor maintenance was a net increase in solutes per unit of tissue. The role played by increases in the proportion of tissue volume occupied by cell wall was also evaluated. No stomatal closure due to water stress was found in NI sorghum even though leaf ψ reached -20 bars late in the season. Under similar conditions, stomata closed at - 14 to - 16 bars in younger plants where water stress was made to develop much faster. [ABSTRACT FROM AUTHOR]

Regulation of the in vivo yielding properties of the cell wall, namely cell wall extensibility φ and effective turgor (Pi-Y of Vigna hypocotyl segments during adaptive recovery of growth after osmotic stress was investigated by successive measurements using the pressure-jump method. In the absence of IAA, cell wall extensibility φ showed no significant changes under osmotic stress but effective turgor (Pi-Y reduced to near zero, resulting in no growth recovery. In the presence of 10 μM IAA, the cell wall extensibility was slightly decreased even when distinct recovery of growth took place in response to osmotic stress caused by 60 mM sorbitol. The effective turgor, however, showed a characteristic increase in contrast to the wall extensibility. It is not wall extensibility but effective turgor that regulates the cell wall yielding during the adaptive recovery of growth. In excised hypocotyl segments, however, reduced turgor would not be expected to recover, as has been observed in hollow cylinders (Nakahori et al. 1990). The results presented here suggest that the IAA-dependent adjustment of the yield threshold plays an indispensable role in wall-yielding for rapid adaptive growth recovery under osmotic stress. [ABSTRACT FROM AUTHOR]

Leaf growth of first year seedlings of silver birch (Betula pendula Roth,) and sycamore (Acer pseudoplatanus L.) was measured continuously using linear variable transducers (LVTDs) when shoots were enclosed in small Perspex chambers, in which temperature, vapour pressure deficit (VPD) and photon fluence rate were all controlled. A new method was used to measure the yield turgor (Y) of growing leaves. The VPD of air in the chamber was altered so that transpiration rate was changed. Turgor pressures, measured using thermocouple psychrometers, and growth rates varied as a result of this treatment, and the relationship between leaf growth and turgor pressure (P) and the turgor pressure at which growth ceased (F) could he measured within a very short time period using well-watered plants. The value of Y was constant for illuminated and darkened leaves of both species, but Y was higher for sycamore leaves (Y = 0.250 MPa) compared with the value for birch leaves (Y = 0.071 MPa). Cell wall extensibility (m, MPa[sup-1]h[sup-1]) was calculated from the slope of the line relating turgor pressure to growth. For leaves of birch, there was a significant increase in pu when leaves were illuminated (m, dark = 1.3 x 10[sup-2] MPa[sup-1]h[sup-1]; m, light = 4.6 x 10[sup-2] MPa[sup-1]), similar in magnitude to the increase in wall extensibility measured previously by mechanical analysis of killed leaf tissue (WEX, the plasticity of tissue subjected to a uni-axial stress). No change in was detected when darkened sycamore leaves were illuminated (mn, dark = 4.2 x l0[sup-2] MPa[sup-1] h[sup-1]; m, light = 4.4 x 10[sup-2] MPa[sup-1] h[sup-1]). The results confirm earlier work on the factors controlling the diurnal patterns of leaf growth of these two species and suggest that leaf growth of birch is primarily limited by cell wall loosening hut that the growth of sycamore leaves may be limited by leaf turgor. [ABSTRACT FROM AUTHOR]

The possible role in drought resistance played by leaf water potential, osmotic potential, weight-averaged turgor pressure and bulk modulus of elasticity is studied in mature and young leaves of Senecio candidus and S. ambiguus. Small water losses cause relatively large reductions in leaf water potential in S. candidus without greatly altering its osmotic potential. This occurs because S. candidus develops thicker leaf cell walls with high values of leaf bulk elastic modulus. Recovery from water stress is very rapid in this species. Leaf growth is influenced by the structural pattern of the cell walls in that S. candidus leaves complete their expansion within a shorter period of time than those of S. ambiguus. The possible adaptive significance of changing the leaf bulk modulus of elasticity in mature and young leaves of the two species is discussed. [ABSTRACT FROM AUTHOR]

Freshwater algae have an intracellular osmolarity in excess of that of the medium. The tendency for water to enter the cells is resisted, in cells with a mechanically functional cell wail, by the ability of the wall to withstand a turgor pressure equivalent to the intracellular osmolarity; in cells lacking such a mechanically functional wall the water which enters is expelled by a contractile vacuole (or its functional equivalent). A comparison of the energy costs of cell volume regulation by these two mechanisms has been carried out making energy costings based on the minimum thermodynamic requirement and on plausible mechanisms for the two processes (wall synthesis and contractile vacuole function). This analysis yielded the following conclusions. (1) Wall synthesis is a 'growth' process, while contractile vacuole operation is a 'maintenance' process; other things being equal, a cell wall is a preferable mechanism of volume regulation in non-growing cells. (2) For a 5 µm radius cell with a specific growth rate of 8 × 10-6 s-1 and a 10 µm radius cell with a rate of 4 × 10-6 s-1 the energy input rate for contractile vacuole `operation is considerably lower than for wall synthesis on the basis of minimum thermodynamic requirement; the plausible mechanism approach makes the two energy costs quite similar. Decreased growth rates favours the wall over the contractile vacuole mechanism. (3) Increased cell size (10 µm radius rather than 5 µm) has similar effects on the energy casts of both volume regulation mechanisms, provided a (cell organic weight)s-0.33 dependence of intrinsic growth rate is incorporated into the computations.(4) Increased intracellular osmolarity (at constant (at constant extracellular osmolarity) causes a directly proportional increase in energy input rate for the cell wall mechanism, and a more than proportional increase in energy cost for the contractile vacuole mechanism. (5) A non-spherical shape increases the energy cost (at constant cell volume) for both the cell wall and the contractile cacuole mechanisms. These conclusions may be used in the interpretation of such phenomena as the lower osmolarity of wall-less than walled cells and the tendency for the resting stages of normally wall-less and flagellate cells to have walls. It appears that the majority of freshwater algae have genetic capabilities to produce both walls and contractile vacuoles for cell volume regulation (e.g. the production of walled cysts by organisms which are normally flagellate, and the production of wall-less motile spores and gametes by normally walled organism). [ABSTRACT FROM AUTHOR]

Cambial derivatives differentiating as xylem fibres characteristically increase in length by intrusive growth. Radial expansion of contiguous cells is required for this intrusive growth which, in consequence, is restricted to the expansion zone of the newly formed wood. The amount of intrusion depends primarily on the position of the tip of a fibre in relation to the lines of junction of tangential and radial walls in the surrounding cells; these lines ot junction are potential sites for intrusive growth. In general, the greatest intrusion occurs where there are two potential sites for intrusion (usually one on either radial face of the fibre's tip); tips with one site show less intrusive growth and those with none show least of all. These arrangements may be modified by the configuration and activity of neighbouring cells. The hypothesis formulated is that the tip of a fibre secretes an enzyme which weakens the middle lamella between the cells adjacent to it; where this occurs, these cells, during their turgor expansion, round off from each other at the corners. The thin-walled tip of the fibre fills this space as it is created. The notion that the intruding tip pushes its way into the cellular matrix is contested. [ABSTRACT FROM AUTHOR]

The effects of auxin and osmotic stress on elongation growth of maize (Zea mays L.) coleoptile segments are accompanied by characteristic changes in the extensibility of the growth-limiting cell walls. At full turgor auxin causes growth by an increase in wall extensibility (wall loosening). Growth can be stopped by an osmotically produced step-down in turgor of 0.45 MPa. Under these conditions auxin causes the accumulation of a potential for future wall extension which is released after restoration of full turgor. Turgor reduction causes a reversible decrease in wall extensibility (wall stiffening) both in the presence and absence of auxin. These changes in vivo are correlated with corresponding changes in the theological properties of the cell walls in vitro which can be traced back to specific modifications in the shape of the hysteretic stress-strain relationship. The longitudinally load-bearing walls of the coleoptile demonstrate almost perfect viscoelasticity as documented by a nearly closed hysteresis loop. Auxin-mediated wall loosening causes an increase of loop width and thus affects primarily the amount of hysteresis in the isolated wall. In contrast, turgor reduction by osmotic stress reduces loop length and thus affects primarily the amount of viscoelastic wall extensibility. Pretreatment of segments with anoxia and H2O2 modify the hysteresis loop in agreement with the conclusion that the wall-stiffening reaction visualized under osmotic stress in vivo is an O2-dependent process in which O2 can be substituted by H2O2. Cycloheximide specifically inhibits auxin-mediated wall loosening without affecting wall stiffening, and this is mirrored in specific changes of the hysteresis loop. Corroborating a previous in vivo study (Hohl et al. 1995, Physiol. Plant. 94: 491–498) these results show that cell wall stiffening in vivo can also be demonstrated by theological measurements with the isolated cell wall and that this process can be separated from cell wall loosening by specific changes in the shape of the hysteresis loop. [ABSTRACT FROM AUTHOR]

Auxin-induced changes of wall-rheological properties during different growth rates of rye coleoptile segments (Secale cereale L.) were investigated. In addition, changes of osmotic concentration and turgor pressure were measured. Decrease of turgor and of osmotic concentration followed a synchronous time course. Auxin-incubated segments exhibited a faster decrease and eventually lower values of both parameters. Creep test extensibility measurements demonstrate that apparent plastic as well as elastic extensibility of distilled-water-incubated segments strongly decreased during 24 h. In auxin-incubated segments apparent plastic as well as elastic extensibilities were strongly increased, even in the absence of growth due to insufficient turgor pressure. The increasing effect of auxin on elastic wall properties is also reflected by an increase in relative reversible length (part of segment length by which segments shrink after freezing/thawing as referred to total length) and a complementary decrease of relative irreversible length (remaining length after turgor elimination as referred to turgid length); again the effects were independent of growth rate and turgor pressure. Cellulose synthesis inhibition of approx. 80% by dichlorobenzonitrile (DCB) had no significant effect either on growth or on wall-rheological properties. Independent of whether the changed rheological wall behaviour of auxin-incubated segments is causally related to the mechanism of auxin-induced wall loosening, it indicates changes of wall polymer properties and/or interactions which are conserved when no actual length increase occurs due to insufficient turgor pressure. The results suggest that IAA-induced wall loosening may be primarily mediated by cell wall changes other than cleavage of covalent, load-bearing bonds as hypothesized in various wall loosening models. [ABSTRACT FROM AUTHOR]

The dependence of elongation on the osmotic potential of the medium was investigated, using coleoptile segments (CS) of Triticim aestivum L. (cv. Hartri) and an optoelectronic device. The study aimed at separating the osmoelastic response from the irreversible growth response when an osmoticum (mannitol) was added, and to compare both processes in order to consider the possibility of growth‐induced reduction in turgor pressure. The prompt inhibition of elongation registered just after addition of 50 mM mannitol as well as the subsequent resumption of the original elongation rate could be quantitatively explained by the extent and the kinetics of the osmoelastic relaxation. An initial reduction in the irreversible elongation component by mild osmotic stress could not be demonstrated. Above a critical value, the irreversible growth was insensitive to a further increase in water potential. The minimum turgor pressure required to drive steady growth was not far from zero in both the presence and absence of auxin. The rate (r) of osmotically caused shortening per unit change of water potential r = 1/l x dl/dt x 1/Δψ was determined from the kinetics of CS shortening induced by addition of mannitol at nearly isotonic concentration (300 mM). This parameter relates a fractional change in length to the difference in water potential between inside and outside, and was assumed to depend largely on the hydraulic resistance of the tissue and cuticle. It was found to be independent of IAA. The relatively low value of Γ suggests significant reduction of turgor at high growth rates. In accordance with this conclusion, the extent of osmoelastic shortening after a transfer to 300 mM mannitol (dependent on wall strain) was significantly decreased in the presence of IAA. Addition of 100 μM IAA to CS growing at a constant rate induced pronounced oscillations in the rate of elongation, which may be connected with the change in elastic cell wall strain. Whereas the steady state growth rate before the addition of IAA was the same in the presence and in the absence of 50 mM mannitol, the maximum growth rate found after addition of IAA was substantially reduced in the mannitol variant. [ABSTRACT FROM AUTHOR]