Control of Volume and Turgor in Stomatal Guard Cells

Water loss from plants is determined by the aperture of stomatal pores in the leaf epidermis, set by the level of vacuolar accumulation of potassium salt, and hence volume and turgor, of a pair of guard cells. Regulation of ion fluxes across the tonoplast, the key to regulation of stomatal aperture, can only be studied by tracer flux measurements. There are two transport systems in the tonoplast. The first is a Ca²⁺-activated channel, inhibited by phenylarsine oxide (PAO), responsible for the release of vacuolar K⁺(Rb⁺) in response to the “drought” hormone, abscisic acid (ABA). This channel is sensitive to pressure, down-regulated at low turgor and up-regulated at high turgor, providing a system for turgor regulation. ABA induces a transient stimulation of vacuolar ion efflux, during which the flux tracks the ion content (volume, turgor), suggesting ABA reduces the set-point of a control system. The second system, which is PAO-insensitive, is responsible for an ion flux from vacuole to cytoplasm associated with inward water flow following a hypo-osmotic transfer. It is suggested that this involves an aquaporin as sensor, and perhaps also as responder; deformation of the aquaporin may render it ion-permeable, or, alternatively, the deformed aquaporin may signal to an associated ion channel, activating it. Treatment with inhibitors of aquaporins, HgCl₂ or silver sulfadiazine, produces a large transient increase in ion release from the vacuole, also PAO-insensitive. It is suggested that this involves the same aquaporin, either rendered directly ion-permeable, or signalling to activate an associated ion channel.     

Osmotic Adjustment and Stomatal Response to Water Deficits in Maize

A pot experiment was carried out using five maize {Zea maysL.) cultivars under three soil moisture levels (MPa 0 to –0.05,–0.3 to –0.9 and –1.2 to –1.5) to investigatethe effects of water deficits on osmotic adjustment and stomatalconductance. The degree of leaf rolling and the sugar and nutrientconcentrations in leaf cell sap were measured. Leaf water potential and osmotic potential decreased and stomatalconductance decreased with increasing water deficits. Stomatalconductance correlated positively with leaf water potentialand osmotic potential. Degree of leaf rolling was lower in cultivarswhich maintained higher turgor. Osmotic adjustment of 0.08 to0.43 MPa was found under the lowest soil moisture level in fivecultivars used. Sugar and K were the major osmotic substancesin the maize plant. Sugar, K and Mg concentrations increasedunder water deficit, and correlated negatively with a decreasein osmotic potential. Key words: Zea mays L., leaf water relations, leaf rolling, osmotic adjustment, stomatal conductance, water deficit     

Turgor Pressure, Volumetric Elastic Modulus, Osmotic Volume and Ultrastructure of Chlorella emersonii Grown at High and Low External NaCl

Chlorella emersonii (211/11n) was grown at external NaCl concentrationsranging between 1.0 and 335 mM (0.08–1.64 MPa). Previousstudies showed that there was no significant change in the internalconcentrations of Na⁺ or Cl^– over this range, the concentrationsremaining below 35 mM. Relative growth rates of C. emersoniiwere 30–45% lower in 335 mM NaCl than in 1.0 mM NaCl.Turgor pressure varied with the osmotic pressure of the growthmedium. Plots of cell volume versus (external osmotic pressure)^–1indicated that cells grown in 1.0 mM NaCl (0.08 MPa) had turgorpressures ranging from 0.5 to 0.8 MPa, while cells in 335 mMNaCl (1.64 MPa) had turgor pressures of 0.0–0.14 MPa.Estimates of turgor pressure derived from the osmotic pressureof cell sap had a mean value of 0.6 MPa for cells in 1.0 mMNaCl, and 0.3 MPa for cells in 335 mM NaCl. The volumetric elasticmodulus () depended on the osmotic pressure of the growth medium: was 8.5 ± 1.7 MPa for cells grown in 1.0 mM NaCl, and0.9 ± 0.6 for cells in 335 mM NaCl. was measured bychanging turgor pressures over the range 0.0–0.5 MPa,and was found to be independent of turgor. Electron micrographsshowed that the walls of cells grown in 335 mM NaCl were 70%thicker than those grown in 1.0 mM NaCl. Other changes in cellularstructure were small, however, the area occupied by vacuolesincreased from 7% in cells grown in 1.0 mM NaCl to 14% in cellsin 335 mM. The percent osmotic volume of cells grown in 1.0–335mM NaCl (61 ± 17%, v/v) was similar to the percent watercontent (59 ± 13%, w/w). Key words: Chlorella emersonii, Sodium chloride, Osmotic volume, Turgor, Volumetric-elastic-modulus     

Turgor, Growth and Rheological Gradients of Wheat Roots Following Osmotic Stress

The growth rate of hydroponically grown wheat roots was reducedby mannitol solutions of various osmotic pressures. For example,following 24 h exposure to 0·96 MPa mannitol root elongationwas reduced from 1· mm h^–1 to 0·1 mm h^–1 Mature cell length was reduced from 290 µm in unstressedroots to 100 µm in 0·96 MPa mannitol. This indicatesa reduction in cell production rate from about 4 per h in theunstressed roots to 1 per h in the highest stress treatment. The growing zone extended over the apical 4·5 mm in unstressedroots but became shorter as growth ceased in the proximal regionsat higher levels of osmotic stress. The turgor pressure along the apical 5·0 mm of unstressedroots was between 0·5 and 0·6 MPa but declinedto 0·41 MPa over the next 50 mm. Following 24 h in 0·48(200 mol m^–3) or 0·72 MPa (300 mol m^–) mannitol,turgor along the apical 50 mm was indistinguishable from thatof unstressed roots but turgor declined more steeply in theregion 5·10 mm from the tip. At the highest level ofstress (0·96 MPa or 400 mol m^–3 mannitol) turgordeclined steeply within the apical 20 mm. Key words: Growth, turgor pressure, wall rheology, osmotic stress, osmotic adjustment     

Variation in Selection for. Leaf Water Conductance in Relation to Growth and Stomatal Dimensions in Lolium perenne L.

A simple, rapid technique for direct selection for leaf waterconductance (LWC) in two populations of Lolium perenne L. isdescribed. Measurements were made with a diffusion porometerin growth rooms on the youngest fully expanded leaf of eachtiller. Considerable variation in LWC was found between 100genotypes of each population. Most of the variation in totalLWC was attributed to variation in adaxial LWC and it was shownthat ignoring abaxial LWC gave very similar ranking of genotypesto those using total LWC. Selection criteria were then establishedwhich maximised the repeatability of LWC measurements and allowedselection of groups of plants significantly different from oneanother in LWC. The variation in LWC was not related eitherto stomatal length or number, or to growth rate. Lolium perenne L., perennial ryegrass, abaxial and adaxial, leaf water conductance, selection, diurnal rhythms, ontogenetic changes, growth, stomatal dimensions     

Osmotic Behavior of Oat Coleoptile Tissue in Relation to Growth

Efforts were made to estimate the water potential difference that is required, between rapidly growing oat coleoptile cylinders and dilute medium, to support the rate of water uptake involved in elongation, (a) by the traditional method of determining the concentration of mannitol in which the tissue neither gains nor loses water, and (b) by measuring the rates of osmotic exchanges induced by treating the tissue with different hypotonic mannitol concentrations. Both methods indicated large water potential differences (3 to 10 atm), in some cases approaching the osmotic pressure of the cells. However, indication was obtained that the rates of osmotic exchanges induced by mannitol solutions, and presumably also the equilibrium response sought in (a), are governed by the rate of diffusional exchange of mannitol with the free space rather than by the permeability of the tissue to water. Osmotic swelling of the tissue measured by immersing it in water after its turgor pressure had been reduced by evaporation, was at least two to four times more rapid than when mannitol was involved. The permeability to water estimated by the evaporation-immersion method indicated that rapidly elongating cylinders have water potentials between -0.8 and -2.5 atm, or between 10 and 25 per cent of their osmotic pressure.     

Enhanced Stomatal Conductance by a Spontaneous Arabidopsis Tetraploid,Me-0, Results from Increased Stomatal Size and Greater Stomatal Aperture

The rate of gas exchange in plants is regulated mainly by stomatal size and density. Generally, higher densities of smaller stomata are advantageous for gas exchange; however, it is unclear what the effect of an extraordinary change in stomatal size might have on a plant’s gas-exchange capacity. We investigated the stomatal responses to CO₂ concentration changes among 374 Arabidopsis (Arabidopsis thaliana) ecotypes and discovered that Mechtshausen (Me-0), a natural tetraploid ecotype, has significantly larger stomata and can achieve a high stomatal conductance. We surmised that the cause of the increased stomatal conductance is tetraploidization; however, the stomatal conductance of another tetraploid accession, tetraploid Columbia (Col), was not as high as that in Me-0. One difference between these two accessions was the size of their stomatal apertures. Analyses of abscisic acid sensitivity, ion balance, and gene expression profiles suggested that physiological or genetic factors restrict the stomatal opening in tetraploid Col but not in Me-0. Our results show that Me-0 overcomes the handicap of stomatal opening that is typical for tetraploids and achieves higher stomatal conductance compared with the closely related tetraploid Col on account of larger stomatal apertures. This study provides evidence for whether larger stomatal size in tetraploids of higher plants can improve stomatal conductance.Gas exchange is a vital activity for higher plants that take up atmospheric CO₂ and release oxygen and water vapor through epidermal stomatal pores. Gas exchange affects CO₂ uptake, photosynthesis, and biomass production (Horie et al., 2006; Evans et al., 2009; Tanaka et al., 2014). Stomatal conductance (gs) is used as an indicator of gas-exchange capacity (Franks and Farquhar, 2007). Maximum stomatal conductance (gsmax) is controlled mainly by stomatal size and density, two parameters that change with environmental conditions and are negatively correlated with each other (Franks et al., 2009).Given a constant total stomatal pore area, large stomata are generally disadvantageous for gas exchange compared with smaller stomata, because the greater pore depth in larger stomata increases the distance that gas molecules diffuse through. This increased distance is inversely proportional to g_smax (Franks and Beerling, 2009). The fossil record indicates that ancient plants had small numbers of large stomata when atmospheric CO₂ levels were high, and falling atmospheric [CO₂] induced a decrease in stomatal size and an increase in stomatal density to increase g_s for maximum carbon gain (Franks and Beerling, 2009). The positive relationship between a high g_s and numerous small stomata also holds true among plants living today under various environmental conditions (Woodward et al., 2002; Galmés et al., 2007; Franks et al., 2009). Additionally, the large stomata of several plant species (e.g. Vicia faba and Arabidopsis [Arabidopsis thaliana]) are often not effective for achieving rapid changes in g_s, due to slower solute transport to drive movement caused by their lower membrane surface area-to-volume ratios (Lawson and Blatt, 2014).Stomatal size is strongly and positively correlated with genome size (Beaulieu et al., 2008; Franks et al., 2012; Lomax et al., 2014). Notably, polyploidization causes dramatic increases in nucleus size and stomatal size (Masterson, 1994; Kondorosi et al., 2000). In addition to the negative effects of large stomata on gas exchange (Franks et al., 2009), polyploids may have another disadvantage; del Pozo and Ramirez-Parra (2014) showed that artificially induced tetraploids of Arabidopsis have a reduced stomatal density (stomatal number per unit of leaf area) and a lower stomatal index (stomatal number per epidermal cell number). Moreover, tetraploids of Rangpur lime (Citrus limonia) and Arabidopsis have lower transpiration rates and changes in the expression of genes involved in abscisic acid (ABA), a phytohormone that induces stomatal closure (Allario et al., 2011; del Pozo and Ramirez-Parra, 2014). On the other hand, an increase in the ploidy level of Festuca arundinacea results in an increase in the CO₂-exchange rate (Byrne et al., 1981); hence, polyploids may not necessarily have a reduced gas-exchange capacity.Natural accessions provide a wide range of information about mechanisms for adaptation, regulation, and responses to various environmental conditions (Bouchabke et al., 2008; Brosché et al., 2010). Arabidopsis, which is distributed widely throughout the Northern Hemisphere, has great natural variation in stomatal anatomy (Woodward et al., 2002; Delgado et al., 2011). Recently, we investigated leaf temperature changes in response to [CO₂] in a large number of Arabidopsis ecotypes (374 ecotypes; Takahashi et al., 2015) and identified the Mechtshausen (Me-0) ecotype among ecotypes with low CO₂ responsiveness; Me-0 had a comparatively low leaf temperature, implying a high transpiration rate. In this study, we revealed that Me-0 had a higher g_s than the standard ecotype Columbia (Col), despite having tetraploid-dependent larger stomata. Notably, the g_s of Me-0 was also higher than that of tetraploid Col, which has stomata as large as those of Me-0. This finding resulted from Me-0 having a higher g_s-to-g_smax ratio due to more opened stomata than tetraploid Col. In addition, there were differences in ABA responsiveness, ion homeostasis, and gene expression profiles in guard cells between Me-0 and tetraploid Col, which may influence their stomatal opening. Despite the common trend of smaller stomata with higher gas-exchange capacity, the results with Me-0 confirm the theoretical possibility that larger stomata can also achieve higher stomatal conductance if pore area increases sufficiently.     

A Comparison of the Growth, Photosynthesis, Stomatal Conductance and Water Use Efficiency of Moricandia and Brassica Species

When grown in pots and well-watered, the relative growth ratesof the above ground parts of two species of Moricandia (M. arvensis,an intermediate C₃–C₄ species, and M. moricandioides,a C₃ species) were inferior to those of two cultivated Brassicaspecies (B. campestris and B. napus). The Moricandia specieshad thicker leaves (greater d.wt per unit leaf area) with morechlorophyll than the Brassica species and had slightly greaterrates of photosynthesis per unit leaf area at an irradiance(400–700 nm) of 2000 µmol quanta m^–2 s ^–1.Leaves of M. arvensis, known to have a CO₂ compensation pointbetween that of C₃ and C₄ species, had a lower ratio of theintercellular to atmospheric partial pressure of CO₂ (C₁/C_a)and a greater instantaneous water use efficiency (WUE) thanthose of M. moricandioides and the Brassica species. Carbon isotope discrimination (     

Enhanced Photosynthesis and Growth in atquac1 Knockout Mutants Are Due to Altered Organic Acid Accumulation and an Increase in Both Stomatal and Mesophyll Conductance

Stomata control the exchange of CO₂ and water vapor in land plants. Thus, whereas a constant supply of CO₂ is required to maintain adequate rates of photosynthesis, the accompanying water losses must be tightly regulated to prevent dehydration and undesired metabolic changes. Accordingly, the uptake or release of ions and metabolites from guard cells is necessary to achieve normal stomatal function. The AtQUAC1, an R-type anion channel responsible for the release of malate from guard cells, is essential for efficient stomatal closure. Here, we demonstrate that mutant plants lacking AtQUAC1 accumulated higher levels of malate and fumarate. These mutant plants not only display slower stomatal closure in response to increased CO₂ concentration and dark but are also characterized by improved mesophyll conductance. These responses were accompanied by increases in both photosynthesis and respiration rates, without affecting the activity of photosynthetic and respiratory enzymes and the expression of other transporter genes in guard cells, which ultimately led to improved growth. Collectively, our results highlight that the transport of organic acids plays a key role in plant cell metabolism and demonstrate that AtQUAC1 reduce diffusive limitations to photosynthesis, which, at least partially, explain the observed increments in growth under well-watered conditions.Stomata are functionally specialized microscopic pores that control the essential exchange of CO₂ and H₂O with the environment in land plants. Stomata are found on the surfaces of the majority of the aerial parts of plants, rendering them as the main control point regulating the flow of gases between plants and their surrounding atmosphere. Accordingly, the majority of water loss from plants occurs through stomatal pores, allowing plant transpiration and CO₂ absorption for the photosynthetic process (Bergmann and Sack, 2007; Kim et al., 2010). The maintenance of an adequate water balance through stomatal control is crucial to plants because cell expansion and growth require tissues to remain turgid (Sablowski and Carnier Dornelas, 2014), and minor reductions in cell water volume and turgor pressure will therefore compromise both processes (Thompson, 2005). As a result, the high sensitivity of plant tissues to turgor has prompted the use of reverse genetic studies in attempt to engineer plants with improved performance (Cowan and Troughton, 1971; Xiong et al., 2009; Borland et al., 2014; Franks et al., 2015).In most land plants, not only redox signals invoked by shifts in light quality (Busch, 2014) but also the transport of inorganic ions (e.g. K⁺, Cl⁻, and NO⁻₃) as well as metabolites such as the phytohormone abscisic acid (ABA), Suc, and malate, are important players controlling stomatal movements (Hetherington, 2001; Roelfsema and Hedrich, 2005; Pandey et al., 2007; Blatt et al., 2014; Kollist et al., 2014). In this context, although organic acids in plants is known to support numerous and diverse functions both within and beyond cellular metabolism, only recently have we obtained genetic evidence to support that modulation of guard cell malate and fumarate concentration can greatly influence stomatal movements (Nunes-Nesi et al., 2007; Araújo et al., 2011b; Penfield et al., 2012; Medeiros et al., 2015). Notably malate, in particular, has been considered as a key metabolite and one of the most important organic metabolites involved in guard cell movements (Hedrich and Marten, 1993; Fernie and Martinoia, 2009; Meyer et al., 2010). During stomatal aperture, the flux of malate into guard cells coupled with hexoses generated on starch breakdown lead to decreases in the water potential, and consequently, water uptake by the guard cells ultimately opens the stomata pore (Roelfsema and Hedrich, 2005; Vavasseur and Raghavendra, 2005; Lee et al., 2008). On the other hand, during stomatal closure, malate is believed to be converted into starch, which has no osmotic activity (Penfield et al., 2012) or, alternatively, is released from the guard cells to the surrounding apoplastic space (Lee et al., 2008; Negi et al., 2008; Vahisalu et al., 2008; Meyer et al., 2010).The role of organic acids on the stomatal movements has been largely demonstrated by studies related to malate transport (Lee et al., 2008; Meyer et al., 2010; Sasaki et al., 2010). In the last decade, two protein families were identified and functionally characterized to be directly involved with organic acid transport at the guard cell plasma membrane and to be required for stomatal functioning (Lee et al., 2008; Meyer et al., 2010; Sasaki et al., 2010). In summary, AtABCB14, a member of the ABC (ATP binding cassette) family, which is involved in malate transport from apoplast to guard cells, was described as a negative modulator of stomatal closure induced by high CO₂ concentration; notably, exogenous application of malate minimizes this response (Lee et al., 2008). In addition, members of a small gene family, which encode the anion channels SLAC1 (slow anion channel 1) and four SLAC1-homologs (SLAHs) in Arabidopsis (Arabidopsis thaliana), have been described to be involved in stomatal movements. SLAC1 is a well-documented S-type anion channel that preferentially transports chloride and nitrate as opposed to malate (Vahisalu et al., 2008, 2010; Geiger et al., 2010; Du et al., 2011; Brandt et al., 2012; Kusumi et al., 2012). Lack of SLAC1 in Arabidopsis and rice (Oryza sativa) culminated in a failure in stomatal closure in response to high CO₂ levels, low relative humidity, and dark conditions (Negi et al., 2008; Vahisalu et al., 2008; Kusumi et al., 2012). Although mutations in AtSLAC1 impair S-type anion channel functions as a whole, the R-type anion channel remained functional (Vahisalu et al., 2008). Indeed, a member of the aluminum-activated malate transporter (ALMT) family, AtALMT12, an R-type anion channel, has been demonstrated to be involved in malate transport, particularly at the plasma membrane of guard cells (Meyer et al., 2010; Sasaki et al., 2010). Although AtALMT12 is a member of ALMT family, it is not activated by aluminum, and therefore Meyer et al. (2010) proposed to rename it as AtQUAC1 (quick-activating anion channel 1; Imes et al., 2013; Mumm et al., 2013). Hereafter, we will follow this nomenclature. Deficiency of a functional AtQUAC1 has been documented to lead to changes in stomatal closure in response to high levels of CO₂, dark, and ABA (Meyer et al., 2010). Taken together, these studies have clearly demonstrated that both S- and R-type anion channels are key modulators of stomatal movements in response to several environmental factors.Despite a vast number of studies involving the above-mentioned anion channels, little information concerning the metabolic changes caused by their impairment is currently available. Such information is important to understand stomatal movements, mainly considering that organic acids, especially the levels of malate in apoplastic/mesophyll cells, have been highlighted as of key importance in leaf metabolism (Fernie and Martinoia, 2009; Araújo et al., 2011a, 2011b; Lawson et al., 2014; Medeiros et al., 2015). Here, we demonstrate that a disruption in the expression of AtQUAC1, which leads to impaired stomatal closure (Meyer et al., 2010), was accompanied by increases in mesophyll conductance (g_m), which is defined as the conductance for the transfer of CO₂ from the intercellular airspaces (C_i) to the sites of carboxylation in the chloroplastic stroma (C_c). By further characterization of atquac1 knockout plants, we demonstrated that reduced diffusive limitations resulted in higher photosynthetic rates and altered respiration that, in turn, led to enhanced biomass accumulation. Overall, the results obtained are discussed both in terms of the importance of organic acid transport in plant cell metabolism and with regard to the contribution that it plays in the regulation of both stomatal function and growth.     

Responses of Photosynthetic Rate and Stomatal Conductance to Water Stress in Soybean Leaves

Responses of photosynthetic rate and stomatal conductance to water stress as weI1 as the relationship between photosynthetic rate and stomatal conductance were investigated with soybean cultivars “Ludou No. 4” and “7605”. The former was a high yield cultivars widely used in Shandong province, and the latter was a small grain soybean line bred by Shandong Academy of Agricultural science. Soil water stress decreased leaf apparent photosynthetic rate and stomatal conductance of two soybean cultivars, and “Ludou No. 4” decreased more than “7605”. At the same value of water potential, photosynthetic rate and stomatal conductance of “7605” were higher than those of “Ludou No,4”,but the rate of stomatal closure for “7605” was higher than “Ludou No. 4”. Decreasing of stomatal conductance caused rising of leaf temperature of two soybean cultivars, and the rising of “7605” was more rapid than “Ludou No. 4”, but at the same treatment of water stress, leaf temperature of “Ludou No. 4” was higher than “7605”. Leaf water use efficiecy (WUE) of two soybean cultivars were decreased under water stress, and the rate of decreasing in “Ludou No.4” was more rapid than in “7605”. These results showed that “7605” was more resistant to water:stress than “Ludou No. 4”.     

Responses of Soybean Leaf Angle, Photosynthesis and Stomatal Conductance to Leaf and Soil Water Potential

The hypothesis that soil water potential (_s) is better correlatedto heliotropic leaf orientation, photosaturated photosyntheticCO₂ assimilation and stomatal conductance during periods oflimited water availability than is bulk leaf water potential(₁) was examined in greenhouse-grown soybean (Glycine max) plants,submitted to a progressive drought. Paired plants were exposedto either 1000 or 100 µmol m^–2 s^–1 photonflux densities (PFD) for 45–60 mins. The higher irradianceinduced short-term decreases in ₁, due to increased transpiration,while _l in the plant exposed to low PFD did not decrease. Thesechanges in ₁ occurred independently of changes in soil waterstatus. Concurrent to the light treatments, a single attachedleaf from each of the two plants was isolated from the restof the plant by shading, and the pulvinus of its terminal leafletwas exposed to a perpendicular PFD of 500 µmol m–²S^–1. Leaf movement of this leaflet was recorded in responseto this light, until a stable leaflet angle was achieved. Valuesof _s and _l (before and after light treatment), and photosaturatedrates of photosynthesis and stomatal conductance, were thenmeasured on these leaves. Leaflet angle and gas exchange werebetter correlated with _s (r² = 0.50, 0.50 and 0.57 for angle,photosynthesis and conductance, respectively) than with _l especiallywhen _l was the result of short-term, high-light induced changesin leaf water status (r² = 0.36, 0.32 and 0.49, for the sameparameters). Leaflet angle was also correlated with stomatalconductance (r² = 0.61) and photosynthetic rate (r² = 0.60),suggesting a close association between leaf orientation, leafmetabolism and soil water availability. Glycine max (L.) Merr. cv. Essex, soybean, heliotropism, water potential, photosynthesis, stomatal conductance, solar tracking     

Rapid Response of the Yield Threshold and Turgor Regulation during Adjustment of Root Growth to Water Stress in Zea mays

Responses of cortical cell turgor (P) following rapid changes in osmotic pressure ([pi]m) were measured throughout the elongation zone of maize (Zea mays L.) roots using a cell pressure probe and compared with simultaneously measured root elongation to evaluate: yield threshold (Y) (minimum P for growth), wall extensibility, growth-zone radial hydraulic conductivity (K), and turgor recovery rate. Small increases in [pi]m (0.1 MPa) temporarily decreased P and growth, which recovered fully in 5 to 10 min. Under stronger [pi]m (up to 0.6 MPa), elongation stopped for up to 30 min and then resumed at lower rates. Recoveries in P through solute accumulation and lowering of Y enabled growth under water stress. P recovery was as much as 0.3 MPa at [pi]m = 0.6 MPa, but recovery rate declined as water stress increased, suggesting turgor-sensitive solute transport into the growth zone. Under strong [pi]m, P did not recover in the basal part of the growth zone, in conjunction with a 30% shortening of the growth zone. Time courses showed Y beginning to decrease within several minutes after stress imposition, from about 0.65 MPa to a minimum of about 0.3 MPa in about 15 min. The data concerning Y were not confounded significantly by elastic shrinkage. K was high (1.3 x 10-10 m2 s-1 MPa-1), suggesting very small growth-induced water potential gradients.     

Root Osmotic Adjustment and Stomatal Control of Leaf Gas Exchange are Dependent on Citrus Rootstocks Under Water Deficit

Journal of Plant Growth Regulation - Drought tolerance is defined by several morpho-physiological mechanisms that together improve plant development under water-limiting conditions. Previously, we...     

适应不同光环境小蜡叶片气孔导度、气孔开度和气孔密度的时空变化

对生长在强光环境和弱光环境小蜡叶片的气孔参数测定发现 :气孔导度和气孔开度在 4个取样部位存在异质性分布。气孔导度和气孔开度经回归分析呈线性、指数或多项式分布。统计分析表明 :强光下的叶片气孔导度和气孔开度的相关性明显高于弱光环境叶片的数值。无论强光环境还是弱光环境下的叶片 ,在取样部位编号为1和 3,其气孔导度和气孔开度均存在显著的线性关系。弱光环境下叶片的气孔密度要远低于强光环境下的叶片。强光环境下叶片对变化环境的敏感性要大于弱光环境下的叶片 ,这可能与强光环境叶片具有较高的气孔密度有关。     

适应不同光环境小蜡叶片气孔导度、气孔开度和气孔密度的时空变化

对生长在强光环境和弱光环境小蜡叶片的气孔参数测定发现:气孔导度和气孔开度在4个取样部位存在异质性分布.气孔导度和气孔开度经回归分析呈线性、指数或多项式分布.统计分析表明:强光下的叶片气孔导度和气孔开度的相关性明显高于弱光环境叶片的数值.无论强光环境还是弱光环境下的叶片,在取样部位编号为1和3,其气孔导度和气孔开度均存在显著的线性关系.弱光环境下叶片的气孔密度要远低于强光环境下的叶片.强光环境下叶片对变化环境的敏感性要大于弱光环境下的叶片,这可能与强光环境叶片具有较高的气孔密度有关.     

Methanol Emission from Leaves (Enzymatic Detection of Gas-Phase Methanol and Relation of Methanol Fluxes to Stomatal Conductance and Leaf Development)

We recently reported the detection of methanol emissions from leaves (R. MacDonald, R. Fall [1993] Atmos Environ 27A: 1709-1713). This could represent a substantial flux of methanol to the atmosphere. Leaf methanol production and emission have not been investigated in detail, in part because of difficulties in sampling and analyzing methanol. In this study we used an enzymatic method to convert methanol to a fluorescent product and verified that leaves from several species emit methanol. Methanol was emitted almost exclusively from the abaxial surfaces of hypostomatous leaves but from both surfaces of amphistomatous leaves, suggesting that methanol exits leaves via stomates. The role of stomatal conductance was verified in experiments in which stomates were induced to close, resulting in reduced methanol. Free methanol was detected in bean leaf extracts, ranging from 26.8 [mu]g g-1 fresh weight in young leaves to 10.0 [mu]g g-1 fresh weight in older leaves. Methanol emission was related to leaf development, generally declining with increasing leaf age after leaf expansion; this is consistent with volatilization from a cellular pool that declines in older leaves. It is possible that leaf emission could be a major source of methanol found in the atmosphere of forests.     

小麦和玉米叶片两面气孔导性的差异及其与环境因素的关系

气孔是作物与大气进行水汽和CO_2交换的通道,当环境条件发生变化时,作物通过调节气孔的开度来控制叶片内部和外部的气体交换速率。作物群体中叶片两面的小生境有明显差异,在长期进化和适应中形成了两面气孔特性     

Growth, Turgor, Water Potential, and Young's Modulus in Pea Internodes

The relations between longitudinal growth, Young's modulus, turgor, water potential, and tissue tensions have been studied on growing internodes of etiolated pea seedlings in an attempt to apply some physical concepts to the growth of a well-known plant material. The modulus has been determined by the resonance frequency method and expressed as E_tissue It increases nearly proportional to the turgor pressure and is at water saturation more than 50 times higher than at plasmolysis. E_tissue is higher in the epidermis than in the ground parenchyma. Indoleacetic acid causes a decrease in Etissue Other properties have been studied on intact and split segments of internodes in solutions of graded mannitol additions. — The following tentative picture of the normal course of the growth has been obtained. Auxin induces growth both in the periphery (epidermis) and in the central core (parenchyma) under a decrease in E_tissue This is followed by an increase of E_tissue which is independent of auxin but depending upon the turgor pressure. It is assumed to involve internal structural changes of the cell walls of the type of creep. The rapid growth takes place in a dynamic system with a low water potential despite favourable water conditions. Epidermis and parenchyma grow equally rapid without tissue tensions. — Such can be produced artificially by splitting of segments and water uptake. The parenchyma thereby loses its sensitivity to auxin. This is the background of the split stem test for auxin. — E_tissue increases when growth is slowing down, probably owing to both synthesis of wall substance and structural changes within the wall. The cells attain a more static condition with E_tissue higher in epidermis than in parenchyma. This leads to the normal tissue tensions. — The result agrees with growth according to the multi-net-principle. The cause of the low water potential and low turgor is discussed with reference to the dynamic nature of both growth and water transport and a probably low matric potential of the streaming water. The decrease in E_tissue following auxin addition is small but is the net difference between an auxin-induced decrease and an increase through the assumed creep.