Cell wall water content has a direct effect on extensibility in growing hypocotyls of sunflower (Helianthus annuus L.)

It has been proposed that spacing between cellulose microfibrils within plant cell walls may be an important determinant of their mechanical properties. A consequence of this hypothesis is that the water content of cell walls may alter their extensibility and that low water potentials may directly reduce growth rates by reducing cell wall spacing. This paper describes a number of experiments in which the water potential of frozen and thawed growing hypocotyls of sunflower (Helianthus annuus L.) were altered using solutions of high molecular weight polyethylene glycol (PEG) or Dextran while their extension under constant stress was monitored using a creep extensiometer (frozen and thawed tissue was used to avoid confounding effects of turgor or active responses to the treatments). Clear reductions in extensibility were observed using both PEG and Dextran, with effects observed in hypocotyl segments treated with PEG 35 000 solutions with osmotic pressures of > or =0.21 MPa suggesting that the relatively mild stresses required to reduce water potentials of plants in vivo by 0.21 MPa may be sufficient to reduce growth rates via a direct effect on wall extensibility. It is noted, therefore, that the water binding capacity of plant cell walls may be of ecophysiological importance. Measurements of cell walls of sunflower hypocotyls using scanning electron microscopy confirmed that treatment of hypocotyls with PEG solutions reduced wall thickness, supporting the hypothesis that the spatial constraint of movement of cellulose microfibrils affects the mechanical properties of the cell wall.     

Hydrogen peroxide-mediated cell-wall stiffening in vitro in maize coleoptiles

It has recently been proposed that H₂O₂-dependent peroxidative formation of phenolic cross-links between cell-wall polymers serves as a mechanism for fixing the viscoelastically extended wall structure and thus confers irreversibility to wall extension during cell growth (M. Hohl et al. 1995, Physiol. Plant. 94: 491–498). In the present paper the isolated cell wall (operationally, frozen/thawed maize coleoptile segments) was used as an experimental system to investigate H₂O₂-dependent cell-wall stiffening in vitro. Hydrogen peroxide inhibited elongation growth (in vivo) and decreased cell-wall extensibility (in vitro) in the concentration range of 10–10000 mol·1^–1. In rheological measurements with a constant-load extensiometer the stiffening effect of H₂O₂ could be observed with both relaxed and stressed cell walls. In-vitro cell-wall stiffening was a time-dependent reaction that lasted about 60 min in the presence of saturating concentrations of H₂O₂. The presence of peroxidase in the growth-limiting outer epidermal wall of the coleoptile was shown by histochemical assays. Peroxidase inhibitors (azide, ascorbate) suppressed the wall-stiffening reaction by H₂O₂ in vitro. Hydrogen peroxide induced the accumulation of a fluorescent, insoluble material in the cell walls of living coleoptile segments. These results demonstrate that primary cell walls of a growing plant organ contain all ingredients for the mechanical fortification of the wall structure by H₂O₂-inducible phenolic cross-linking.Supported by Deutsche Forschungsgemeinschaft. I thank Ms. Bärbel Huvermann for expert technical assistance.     

The partitioning of water uptake between growth forms in a Neotropical savanna: do herbs exploit a third water source niche?

In addition to trees and grasses, the savannas of central Brazil are characterised by a diverse herbaceous dicot flora. Here we tested whether the coexistence of a highly diversified assemblage of species resulted in stratification or strong overlap in the use of soil water resources. We measured oxygen and hydrogen isotope ratios of stem water from herbs, grasses and trees growing side by side, as well as the isotopic composition of water in soil profile, groundwater and rainfall, and predawn (Ψ_pd) and midday (Ψ_md) leaf water potentials. We used a stable isotope mixing model to estimate vertical partitioning of soil water by the three growth forms. Grasses relied on shallow soil water (5–50 cm) and were strongly anisohydric. Ψ_pd and Ψ_md decreased significantly from the wet to the dry season. Trees extracted water from deeper regions of the soil profile (60–120 cm) and were isohydric. Ψ_pd and Ψ_md did not change from the wet to the dry season. Herbs overlapped with grasses in patterns of water extraction in the dry season (between 10 and 40 cm), but they took up water at soil depths intermediate (70–100 cm) to those of trees and grasses during the wet season. They showed seasonal changes in Ψ_pd but not in Ψ_md. We conclude that vertical partitioning of soil water may have contributed to coexistence of these three growth forms and resulted in a more complex pattern of soil water extraction than the two‐compartment model of soil water uptake currently used to explain the structure and function of tropical savanna ecosystems.     

Rapid and reversible modifications of extension capacity of cell walls in elongating maize leaf tissues responding to root addition and removal of NaCl

A creep extensiometer technique was used to provide direct evidence that short (20 min) and long-term (3d) exposures of roots to growth inhibitory levels of salinity (100mol m^-3 NaCl) induce reductions in the irreversible extension capacity of cell walls in the leaf elongation zone of intact maize seedlings (Zea mays L.). The long-term inhibition of cell wall extension capacity was reversed within 20 min of salt withdrawal from the root medium. Inhibited elongation of leaf epidermal tissues was also reversed after salt removal. The salt-induced changes in wall extension capacity were detected using in vivo and in vitro assays (shortly after localized freeze/thaw treatment of the basal elongation zone). The rapid reversal of the inhibition of wall extensibility and leaf growth after salt removal from root medium of long-term salinized plants, suggested that neither deficiencies in growth essential mineral nutrients nor toxic effects of NaCl on plasmamembrane viability were directly involved in the inhibition of leaf growth. There was consistent agreement between the scale, direction and timing of salinity-induced changes in leaf elongation growth and wall extension capacity. Rapid metabolically regulated changes in the physical properties of growing cell walls, caused by osmotic (or other) effects, appear to be a factor regulating maize leaf growth responses to root salinization.     

Temperature responses of photosynthesis and leaf hydraulic conductance in rice and wheat

Studies on the temperature (T) responses of photosynthesis and leaf hydraulic conductance (K_leaf) are important to plant gas exchange. In this study, the temperature responses of photosynthesis and K_leaf were studied in Shanyou 63 (Oryza sativa) and Yannong 19 (Triticum aestivum). Leaf water potential (Ψ_leaf) was insensitive to T in Shanyou 63, while it significantly decreased with T in Yannong 19. The differential Ψ_leaf − T relationship partially accounted for the differing g_m–T relationships, where g_m was less sensitive to T in Yannong 19 than in Shanyou 63. With different g_m–T and Ψ_leaf–T relationships, the temperature responses of photosynthetic limitations were surprisingly similar between the two lines, and the photosynthetic rate was highly correlated with g_m. With the increasing T, K_leaf increased in Shanyou 63 while it decreased in Yannong 19. The different K_leaf–T relationships were related to different Ψ_leaf–T relationships. When excluding the effects of water viscosity and Ψ_leaf, K_leaf was insensitive to T in both lines. g_m and K_leaf were generally not coordinated across different temperatures. This study highlights the importance of Ψ_leaf on leaf carbon and water exchanges, and the mechanisms for the g_m–T and K_leaf–T relationships were discussed.     

Rheological analysis of viscoelastic cell wall changes in maize coleoptiles as affected by auxin and osmotic stress

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 looseining). 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 rheological 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.     

The ecosystem wilting point defines drought response and recovery of a Quercus-Carya forest

Soil and atmospheric droughts increasingly threaten plant survival and productivity around the world. Yet, conceptual gaps constrain our ability to predict ecosystem-scale drought impacts under climate change. Here, we introduce the ecosystem wilting point (Ψ_EWP), a property that integrates the drought response of an ecosystem's plant community across the soil–plant–atmosphere continuum. Specifically, Ψ_EWP defines a threshold below which the capacity of the root system to extract soil water and the ability of the leaves to maintain stomatal function are strongly diminished. We combined ecosystem flux and leaf water potential measurements to derive the Ψ_EWP of a Quercus-Carya forest from an “ecosystem pressure–volume (PV) curve,” which is analogous to the tissue-level technique. When community predawn leaf water potential (Ψ_pd) was above Ψ_EWP (=−2.0 MPa), the forest was highly responsive to environmental dynamics. When Ψ_pd fell below Ψ_EWP, the forest became insensitive to environmental variation and was a net source of carbon dioxide for nearly 2 months. Thus, Ψ_EWP is a threshold defining marked shifts in ecosystem functional state. Though there was rainfall-induced recovery of ecosystem gas exchange following soaking rains, a legacy of structural and physiological damage inhibited canopy photosynthetic capacity. Although over 16 growing seasons, only 10% of Ψ_pd observations fell below Ψ_EWP, the forest is commonly only 2–4 weeks of intense drought away from reaching Ψ_EWP, and thus highly reliant on frequent rainfall to replenish the soil water supply. We propose, based on a bottom-up analysis of root density profiles and soil moisture characteristic curves, that soil water acquisition capacity is the major determinant of Ψ_EWP, and species in an ecosystem require compatible leaf-level traits such as turgor loss point so that leaf wilting is coordinated with the inability to extract further water from the soil.     

Water potential in soil and Atriplex nummularia (phytoremediator halophyte) under drought and salt stresses

Atriplex nummularia is a halophyte widely employed to recover saline soils and was used as a model to evaluate the water potentials in the soil-plant system under drought and salt stresses. Potted plants grown under 70 and 37% of field capacity irrigated with solutions of NaCl and of a mixture of NaCl, KCl, MgCl₂ and CaCl₂ reproducing six electrical conductivity (EC): 0, 5, 10, 20, 30, and 40 dS m^?1. After 100 days, total water (Ψ_{w, plant}) and osmotic (Ψ_{o, plant}) potentials at predawn and midday and Ψ_{o, soil}, matric potential (Ψ_{m, soil}) and Ψ_{w, soil} were determined. The type of ion in the irrigation water did not influence the soil potential, but was altered by EC. The soil Ψ_o component was the largest contributor to Ψ_{w, soil}. Atriplex is surviving ECs close to 40 dS m^?1 due to the decrease in the Ψ_w. The plants reached a Ψ_w of approximately ?8 MPa. The water potentials determined for different moisture levels, EC levels and salt types showed huge importance for the management of this species in semiarid regions and can be used to recover salt affected soils.     

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° and 30°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.Abbreviations DTT dithiothreitol - EDTA ethylenediaminetetraacetic acid - EGTA ethyleneglycol-bis-(-aminoethylether)-N,N,N,N-tetraacetic acid - Hepes N-2-hydroxyethylpiperazine-N-2-ethansulfonic acid     

Physical extensibility of maize coleoptile cell walls: apparent plastic extensibility is due to elastic hysteresis

Segments of maize (Zea mays L.) coleoptiles demonstrate plastic cell-wall extensibility (E_pl) as operationally defined by the amount of irreversible strain elicited by stretching living or frozen-thawed tissue under constant load in an extensiometer (creep test). Changes of E_pl are correlated with auxin- and abscisic-acid-dependent growth responses and have therefore been causally related to hormone-controlled cell-wall loosening. Auxin induces an increase of E_pl specifically in the outer epidermal wall of maize coleoptiles which is considered as the growth-limiting wall of the organ. However, detailed kinetic measurements of load-induced extension of frozen-thawed coleoptile segments necessitates a revision of the view that E_pl represents a true plastic (irreversible) wall deformation. Segments demonstrate no significant irreversible extension when completely unloaded between loading cycles. Moreover, E_pl can be demonstrated repeatedly if the same segment is subjected to repeated loading cycles in the extensiometer. It is shown that these phenomena result from the hysteresis behaviour of the cell wall. Stress-strain curves for loading and unloading form a closed hysteresis loop, the width of which represents E_pl at a particular load. Auxin-treatment of segments leads to a deformation of the hysteresis loop, thereby giving rise to an increase of E_pl. These results show that the creep test estimates the viscoelastic (retarded elastic) properties rather than the plastic properties of the wall.Abbreviations E_tot, E_el, E_pl total, elastic, and plastic cell-wall extensibility as defined by the standard creep test - L load Supported by Deutsche Forschungsgemeinschaft (SFB 206).     

Cooperation of epidermis and inner tissues in auxin-mediated growth of maize coleoptiles

The function of the epidermis in auxinmediated elongation growth of maize (Zea mays L.) coleoptile segments was investigated. The following results were obtained: i) In the intact organ, there is a strong tissue tension produced by the expanding force of the inner tissues which is balanced by the contracting force of the outer epidermal wall. The compression imposed by the stretched outer epidermal wall upon the inner tissues gives rise to a wall-pressure difference which can be transformed into a water-potential difference between inner tissues and external medium (water) by removal of the outer epidermal wall. ii) Peeled segments fail to respond to auxin with normal growth. The plastic extensibility of the inner-tissue cell walls (measured with a constant-load extensiometer using living segments) is not influenced by auxin (or abscisic acid) in peeled or nonpeeled segments. It is concluded that auxin induces (and abscisic acid inhibits) elongation of the intact segment by increasing (decreasing) the extensibility specifically in the outer epidermal wall. In addition, tissue tension (and therewith the pressure acting on the outer epidermal wall) is maintained at a constant level over several hours of auxin-mediated growth, indicating that the inner cells also contribute actively to organ elongation. However, this contribution does not involve an increase of cell-wall extensibility, but a continuous shifting of the potential extension threshold (i.e., the length to which the inner tissues would extend by water uptake after peeling) ahead of the actual segment length. Thus, steady growth involves the coordinated action of wall loosening in the epidermis and regeneration of tissue tension by the inner tissues. iii) Electron micrographs show the accumulation of striking osmiophilic material (particles of approx. 0.3 m diameter) specifically at the plasma membrane/cell-wall interface of the outer epidermal wall of auxin-treated segments. iv) Peeled segments fail to respond to auxin with proton excretion. This is in contrast to fusicoccin-induced proton excretion and growth which can also be readily demonstrated in the absence of the epidermis. However, peeled and nonpeeled segments show the same sensitivity to protons with regard to the induction of acid-mediated in-vivo elongation and cell-wall extensibility. The observed threshold at pH 4.5–5.0 is too low to be compatible with a second messenger function of protons also in the growth response of the inner tissues. Organ growth is described in terms of a physical model which takes into account tissue tension and extensibility of the outer epidermal wall as the decisive growth parameters. This model states that the wall pressure increment, produced by tissue tension in the outer epidermal wall, rather than the pressure acting on the inner-tissue walls, is the driving force of growth.Abbreviations and symbols E _el, E _pl elastic and plastic in-vitro cell-wall extensibility, respectively - E _tot E _el+E _pl - FC fusicoccin - IAA indole-3-acetic acid - IT inner tissue - ITW inner-tissue walls - OEW outer epidermal wall - osmotic pressure - P wall pressure - water potential     

Hydroxyl radical-induced cell-wall loosening in vitro and in vivo: implications for the control of elongation growth

Hydroxyl radicals (OH) are capable of unspecifically cleaving cell-wall polysaccharides in a site-specific reaction. I investigated the hypothesis that cell-wall loosening underlying the elongation growth of plant organs is controlled by apoplastically produced OH attacking load-bearing cell-wall matrix polymers. Isolated cell walls (operationally, frozen/thawed, abraded segments from coleoptiles or hypocotyls, respectively) from maize, cucumber, soybean, sunflower or Scots pine seedlings were pre-loaded with catalytic Cu or Fe ions and then incubated in a mixture of ascorbate + H2O2 for generating OH in the walls. This treatment induced irreversible wall extension (creep) in walls stretched in an extensiometer. The reaction could be promoted by acid pH and inhibited by several OH scavengers. Generation of OH by the same reaction in living coleoptile or hypocotyl segments caused elongation growth. Auxin-induced elongation growth of maize coleoptiles could be inhibited by OH scavengers. Auxin promoted the production of superoxide radicals (O2(-)), an OH precursor, in the growth-controlling outer epidermis of maize coleoptiles. It is concluded that OH fulfils basic criteria for a wall-loosening factor acting in auxin-mediated elongation growth of plant species with widely differing cell-wall polysaccharide compositions.     

Stomatal closure of Pelargonium × hortorum in response to soil water deficit is associated with decreased leaf water potential only under rapid soil drying

Soil water deficits applied at different rates and for different durations can decrease both stomatal conductance (g_s) and leaf water potential (Ψ_leaf). Understanding the physiological mechanisms regulating these responses is important in sustainable irrigation scheduling. Glasshouse‐grown, containerized Pelargonium × hortorum BullsEye plants were irrigated either daily at various fractions of plant evapotranspiration (100, 75 and 50% ET) for 20 days or irrigation was withheld for 4 days. Xylem sap was collected and g_s and Ψ_leaf were measured on days 15 and 20, and on days 16–19 for the respective treatments. Xylem sap pH and NO₃^? and Ca²⁺ concentrations did not differ between irrigation treatments. Xylem abscisic acid (ABA) concentrations ([ABA]_xyl) increased within 24 h of irrigation being withheld whilst g_s and Ψ_leaf decreased. Supplying irrigation at a fraction of daily ET produced a similar relationship between [ABA]_xyl and g_s, but did not change Ψ_leaf. Treatment differences occurred independently of whether Ψ_leaf was measured in whole leaves with a pressure chamber, or in the lamina with a thermocouple psychrometer. Plants that were irrigated daily showed lower [ABA]_xyl than plants from which irrigation was withheld, even at comparable soil moisture content. This implies that regular re‐watering attenuates ABA signaling due to maintenance of soil moisture in the upper soil levels. Crucially, detached leaves supplied with synthetic ABA showed a similar relationship between [ABA]_xyl and g_s as intact plants, suggesting that stomatal closure of P. hortorum in response to soil water deficit is primarily an ABA‐induced response, independent of changes in Ψ_leaf.     

Response of eucalypt species to drought

The response of three eucalypt species (Eucalyptus pulchella, Eucalyptus coccifera and Eucalyptus delegatensis) to a severe drought in the summer of 1982/83 was examined at Snug Plains, south-eastern Tasmania. Few large differences in leaf water potential (Ψ_l) or stomatal conductance (gs) were apparent even at the height of the drought when both Ψ_l and soil water potentials (Ψ_s) reached ca. — 4.5 MPa. However, E. pulchella maintained a higher relative water content (RWC) in its leaves than E. coccifera and E. delegatensis, and showed less severe crown damage. After the first light rains substantial interspecific differences in Ψ_l and gs occurred. Eucalyptus pulchella restored normal Ψ_l, gs and RWC more rapidly than the other two species and, even for severely droughted trees, crown growth commenced via epicormic buds near the ends of its branches while for E. delegatensis and E. coccifera crown regeneration was via epicormic buds arising from stems and larger branches. This resulted in a change in dominance in certain stands and showed that E. pulchella was more drought-resistant than E. coccifera, which was in turn more resistant than E. delegatensis. This conclusion was confirmed during competition experiments using potted seedlings. However, potted seedlings differed from mature field trees by maintaining moderate gs at high vapour pressure deficits and closing stomata at Ψ_l below ca. — 2.0 MPa. Substantial variation in the severity of drought symptoms was observed over short distances. This variation appeared to be determined by the moisture-holding capacity of the soil and the biomass of the stand. Although differences in the rooting patterns of seedlings were evident, field measurements of Ψ_l and Ψ_s suggested that all three species were exploiting the same water resource. In contrast to previous studies, the results suggest that large interspecific differences in tissue hydration and crown damage may be present, even though differences in Ψ_l, gs and characteristics of the root system may appear small.     

Osmoregulation and membrane-mediated responses to altered water potential in plant cells

Plant cells respond to short-term stress dehydration by modification of internal Ψ_π such that an inward gradient of Ψ_ω is maintained. In response to lowered Ψ_ω, increases in internal Ψ_π are created by alteration of cell inorganic ions and small organic solute content. Passive movement of water follows, changing cell hydration and forcing the plasma membrane against the elastic cell wall. The stretched cell wall presses against the cell contents, creating a hydrostatic pressure, Ψ_π, which tends to force water out of the cell. The resulting hydrostatic pressure eventually comes into equilibrium with forces bringing water into the cell, largely Ψ_π, and the net flow of water ceases.The mechanism for sensing cell Ψ_ω changes is unknown but the initial event must be physical, not biochemical. The method of translation of such physical events into biochemical actions is also unknown but the Zimmermann model provides a means of signal transduction and amplification, through the alteration of membrane parameters, which could account for the observed changes. As for animal cells, cell levels of Ca²⁺ are important for their regulation of membrane P_j in these responses but unlike osmoregulation in higher animals, the involvement of plant hormones in these responses have not been clearly established. However, the important role of plant cell limiting membranes in plant cell osmoregulation responses seems obvious.     

In-vivo measurement of cell-wall extensibility in maize coleoptiles: Effects of auxin and abscisic acid

Plastic and elastic in-vivo extensibilities (E_pl and E_el, respectively) of cell walls of growing maize (Zea mays L.) coleoptile segments were measured by stretching living tissue at constant force (creep test) in an extensiometer. The linear displacement transducer used as a measuring device permits the determination of load-induced extensions in the range of 0–1% of the segment's length, leading to a minimal disturbance of the hydraulic parameters of the tissue and allowing the measurement of unidirectional cell-wall creep at virtually unchanged turgor and metabolic activity. A rein-vestigation of the time-course of indole-3-acetic acid-promoted and abscisic acid-inhibited wall loo-sening revealed that the in-vivo creep test yields results very similar to those obtained previously with the in-vitro creep test [Kutschera and Schopfer, 1986, Planta 167, 527–535]. The hormones affect elongation rate and E_pl in a closely correlated manner both in step-up as well as step-down growth changes whereas E_el remains unaltered. It is argued that both hormones influence growth by modifying E_pl of the outer epidermis and that this effect can be quantitatively measured, in relative units, by either the in-vivo or the in-vitro creep test.Abbreviations ABA ±abscisic acid - E_el, E_pl elastic and plastic in-vivo cell-wall extensibility, respectively - E_tot E_el+E_pl - IAA indole-3-acetic acid; m, cell-wall yielding coefficient     

The mechanical behavior of isolated Avena coleoptile walls subjected to constant stress: properties and relation to cell elongation

In order to assess the role of the mechanical properties of the wall in auxin-induced cell elongation, a study has been made of the ability of isolated Avena coleoptile walls to extend (creep) when subjected to a constant applied stress. Creep occurs as a viscoelastic extension which has the following characteristics: the extension is proportional to log time and is partly reversible, and the extension rate has a Q₁₀ of about 1.05 and is markedly greater in auxin-pretreated walls. In nonconditioned walls the extension rate is proportional to applied stress, but pre-extension causes the appearance of an apparent yield strain. The similarity of creep and instantaneous plastic deformation in response to temperature or to pretreatment with auxin or KCN suggests that the instantaneous deformation is simply the viscoelastic extension which occurs at very short times. A comparison of these viscoelastic properties with the properties of auxin-induced cell elongation indicates that cell elongation requires more than just a physical extension of the wall. It is suggested that elongation occurs as a series of extension steps, each of which involves a viscoelastic extension preceded or accompanied by an auxin-dependent biochemical change in the wall properties.     

Stomatal control by both [ABA] in the xylem sap and leaf water status: a test of a model for draughted or ABA-fed field-grown maize

A model of maize stomatal behaviour has been developed, in which stomatal conductance is linked to the concentration of abscisic acid ([ABA]) in the xylem sap, with a sensitivity dependent upon the leaf water potential (Ψ₁). It was tested against two alternative hypotheses, namely that stomatal sensitivity to xylem [ABA] would be linked to the leaf-to-air vapour pressure difference (VPD), or to the flux of ABA into the leaf. Stomatal conductance (g_s) was studied: (1) in field-grown plants whose xylem [ABA] and Ψ₁ depended on soil water status and evaporative demand; (2) in field-grown plants fed with ABA solutions such that xylem [ABA] was artificially raised, thereby decreasing g_s and increasing Ψ₁ and leaf-to-air VPD; and (3) in ABA-fed detached leaves exposed to varying evaporative demands, but with a constant and high Ψ₁. The same relationships between g_s, xylem [ABA] and Ψ₁, showing lower stomatal sensitivity to [ABA] at high Ψ₁, applied whether variations in xylem [ABA] were due to natural increase or to feeding, and whether variations in Ψ₁, were due to changes in evaporative demand or to the increased Ψ₁ observed in ABA-fed plants. Conversely, neither the leaf-to-air VPD nor the ABA flux into the leaf accounted for the observed changes in stomatal sensitivity to xylem [ABA]. The model, using parameters calculated from previous field data and the detached-leaf data, was tested against the observations of both ABA-fed and droughted plants in the field. It accounted with reasonable accuracy for changes in g_s (r² ranging from 0.77 to 0.81). These results support the view that modelling of stomatal behaviour requires consideration of both chemical and hydraulic aspects of root-to-shoot communication.     

Temperature and water content effects on the viscoelastic behavior of<Emphasis Type="Italic"> Tilia americana</Emphasis> (Tiliaceae) sapwood

The effects of temperature and water content on the viscoelasticity of living and dehydrated Tilia americana sapwood were examined using transient creep (time- and load-dependent deformation) tests under sustained bending loads. Creep tests were performed at 21.1°C and –20.5°C to determine the magnitudes and types of strains in living and dehydrated samples. Temperature had no effect on the creep rate of living sapwood. However, the creep rate of dehydrated samples at –20.5°C was significantly faster than that at 21.1°C. Regardless of temperature, sapwood had a faster creep rate than dehydrated samples. With small bending loads, the residual strains in sapwood were larger at 21.1°C compared to –20.5°C. Temperature did not significantly affect the residual strains in dehydrated samples. For small bending loads, frozen sapwood recovered all residual creep strains when thawed. With larger loads, residual and plastic (permanent) strains increased. We speculate that ice formation in cell lumens partially dehydrates (and thus stiffens and strengthens) cell wall materials and prevents cell wall buckling and elastic restoration after unloading. However, when thawed, sapwood can elastically restore its original configuration, provided it is not excessively bent (by ice or snow accumulations) when frozen.     

Water movement through Quercus rubra I. Leaf water potential and conductance during polycyclic growth

Two experiments examined simultaneous changes in leaf area (A_L), root length (L_r), stomatal conductance (g_s), leaf water potential (Ψ_L), transpiration and hydraulic plant conductance per unit leaf area (G) during the first three shoot cycles of northern red oak (Quercus rubra L.) grown under favourable and controlled conditions. Each shoot cycle consisted of bud swell, stem elongation, leaf expansion and rest; roots grew almost continuously. The g_s of all leaves decreased substantially while leaves of the newest flush were expanding and increased modestly when seedling leaf area remained constant. Overall, g_s decreased. The Ψ_L of mature leaves decreased during leaf expansion and increased by an equivalent amount during intervening periods. Possible explanations for the paired changes in g_s and Ψ_L are considered. Changes in G closely paralleled those of canopy g_s. These parallel changes during polycyclic seedling growth should act to keep seedling Ψ_L relatively constant as plant size increases and thereby help prevent Ψ_L from dropping to levels that would cause runaway embolism.