Stomatal Responses to Sulphur Dioxide and Vapour Pressure Deficit

Stomatal conductances (g_s) of plants of Vicia faba, Raphanussativus, Phaseolus vulgaris, Heilanthus annuus, and Nicotianatabacum were measured in chambers containing either clean airor air containing between 18 and 1000 parts 10^–9 SO₂ atwater vapour pressure deficits (vpd) ranging from 1·0to 1·8 kPa. When vpd was low (<1·3 kPa at 22 °C) and stomatawere open, exposure to SO₂ induced rapid and irreversible increasesin g_s in V. faba. This response persisted throughout the exposure(3 d). The increase in g_s, 20–30% compared with cleanair, was independent of SO₂ concentration up to 350 parts 10^–9Stomatal conductances of polluted plants at night were greaterthan controls. When stomata were closed before exposure to SO₂,there was no effect on g_s. When vpd was varied, g_s of unpolluted plants of P. vulgarisshowed no response, but that of R. sativus increased slightlywith increasing vpd. In both species exposure to SO₂ causedan increase in g_s at all vpd values. g_s of unpolluted plantsof V. faba, H. annuus, and N. tabacum decreased with increasingvpd. At low vpd values exposure to SO₂ in these species causedan increase in g_s, but, above a certain value of vpd, dependingon species, g_s decreased with exposure to SO₂. It is postulated that SO₂, once in the substomatal cavity, entersthe stomatal complex via adjacent epidermal cells and at lowconcentrations leads to a reduction in turgor in these cellsand consequently to stomatal opening. In vpd-sensitive species,increased transpiration from guard cells or epidermal cellsadjacent to the stomata induced by SO₂ may lead to stomatalclosure at large vpd levels. Stomatal sensitivity to vpd insuch cases may be enhanced because adjacent epidermal cell turgoris lowered by SO₂. At high SO₂ concentrations direct disruptionof guard cell structure may lead to a loss of turgor and stomatalclosure.     

The Effects of Low Concentrations of Sulphur Dioxide on Stomatal Conductance and Epidermal Cell Survival in Field Bean (Vicia faba L.)

Exposure of 3 week old field bean plants to concentrations ofSO₂ from 50–500 µg m^-3 induced comparable 20–25%increases in mean leaf diffusive conductance regardless of whetherthe diffusive conductances were obtained by porometric measurementor calculation from gas exchange data. The stomatal conductancesof the adaxial and abaxial leaf surfaces were both increasedby exposure to SO₂. Microscopic examination of epidermal strips from control andpolluted plants revealed that the stomatal opening observedin treated plants was associated with a sharp reduction in theproportion of living epidermal cells adjacent to the stomata.The proportion of surviving adjacent epidermal cells was invariablysmaller on the lower epidermis and appeared to decrease as theSO₂ concentration was raised from 50 to 500 µg m^–3.Although the guard cells appeared to be undamaged at concentrationsbelow 200 µg m^–3, structural disorganization ordeath of one or both guard cells was observed frequently ator above 500 µg m^–3. The results are discussed in relation to the controversy concerningthe effects of SO₂ on stomatal aperture.     

Stomatal Conductance and Sulfur Uptake of Five Clones of Populus tremuloides Exposed to Sulfur Dioxide

Plants of five clones of Populus tremuloides Michx. were exposed to 0, 0.2 or 0.5 microliter per liter SO₂ for 8 hours in controlled environment chambers. In the absence of the pollutant, two pollution-resistant clones maintained consistently lower daytime diffusive conductance (LDC) than did a highly susceptible clone or two moderately resistant clones. Differences in LDC among the latter three clones were not significant. At 0.2 microliter per liter SO₂, LDC decreased in the susceptible clone after 8 hours fumigation while the LDC of the other clones was not affected. Fumigation with 0.5 microliter per liter SO₂ decreased LDC of all five clones during the fumigation. Rates of recovery following fumigation varied with the clone, but the LDC of all clones had returned to control values by the beginning of the night following fumigation. Night LDC was higher in the susceptible clone than in the other clones. Fumigation for 16 hours (14 hours day + 2 hours night) with 0.4 microliter per liter SO₂ decreased night LDC by half. Sulfur uptake studies generally confirmed the results of the conductance measurements. The results show that stomatal conductance is important in determining relative susceptibility of the clones to pollution stress.     

The Effect of Reduced Hydraulic Conductance on Stomatal Conductance and Xylem Cavitation

The vulnerability of xylem conduits to cavitation theoreticallydetermines the maximum flow rate of water through plants, andhence maximum transpiration (E), stomatal conductance (g_s),and leaf area (A₁. Field-grown Betula occidentalis with a favourablewater supply exhibit midday xylem pressures (     

Relating Stomatal Conductance to Leaf Functional Traits

Leaf functional traits are important because they reflect physiological functions, such as transpiration and carbon assimilation. In particular, morphological leaf traits have the potential to summarize plants strategies in terms of water use efficiency, growth pattern and nutrient use. The leaf economics spectrum (LES) is a recognized framework in functional plant ecology and reflects a gradient of increasing specific leaf area (SLA), leaf nitrogen, phosphorus and cation content, and decreasing leaf dry matter content (LDMC) and carbon nitrogen ratio (CN). The LES describes different strategies ranging from that of short-lived leaves with high photosynthetic capacity per leaf mass to long-lived leaves with low mass-based carbon assimilation rates. However, traits that are not included in the LES might provide additional information on the species'' physiology, such as those related to stomatal control. Protocols are presented for a wide range of leaf functional traits, including traits of the LES, but also traits that are independent of the LES. In particular, a new method is introduced that relates the plants’ regulatory behavior in stomatal conductance to vapor pressure deficit. The resulting parameters of stomatal regulation can then be compared to the LES and other plant functional traits. The results show that functional leaf traits of the LES were also valid predictors for the parameters of stomatal regulation. For example, leaf carbon concentration was positively related to the vapor pressure deficit (vpd) at the point of inflection and the maximum of the conductance-vpd curve. However, traits that are not included in the LES added information in explaining parameters of stomatal control: the vpd at the point of inflection of the conductance-vpd curve was lower for species with higher stomatal density and higher stomatal index. Overall, stomata and vein traits were more powerful predictors for explaining stomatal regulation than traits used in the LES.     

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.     

Variation of Leaf Stomatal Conductance in Winter Wheat Canopy

Leaf stomatal conductance measured and analysed in the canopies of two winter wheat varieties in the field revealed that the probability of adaxial to abaxial conductance ratio followed an approximately normal distribution with a peake value of about 1.5. The ratio changed with the developmental stages being maximium at the heading stage. Leaf stomata in wheat of the upper part of the canopy were more active and showed more pronounced diurnal change of conductance than those of the lower part. Stomatal conductance decreased from top to bottom in canopy as a negative exponential function. By comparing adaxial and abaxial conductances in the apical, middle and basal parts of a leaf, the distribution of the stomatal conductances of a wheat leaf was as follows: a steady decrease from the basal part of adaxial, through the middle and apical parts of the adaxial surface turning to the apical part of abaxial, and then the middle and lastly, the basal part of abaxial. Based on values of the correlation coefficients among the various stomatal conductance and average stomatal conductance, the authors suggested that optimal apical measurement of stomatal conductance would be at the middle and apical parts and that of abaxial would be at middle and basal parts.     

落叶阔叶林内的光合作用和气孔传导率特征(英文)

本文根据Wang和BMdocchi(1989)最近提出的冠层辐射模型,进一步给出了一个模拟冠层光合作用速率和气孔传导率的模式.模式将冠层中每一层的叶面积分为向光叶、半影叶、和全遮荫叶三种,并分别计算其光合作用速率和气孔传导率。计算得到的光合速率廓线表明,在落叶阔叶林内,冠层下部的叶片常处于光照不足状态;半影效应使得透过林冠达于底部的辐射量增大,这对于林下植物的光合作用是有利的。 模式计算值与实测值之间的微弱差别应归因于纯辐射模型无法考虑湍流输送机制造成的CO_2传输和冠层底部耐荫性叶对于低光照的适应能力。     

Injury, Stomatal Conductance, and Abscisic Acid Levels of Pea Plants following Ozone plus Sulfur Dioxide Exposures at Different Times of the Day

Pea (Pisum sativum L. cv Alsweet) plants were exposed to mixtures of ozone plus sulfur dioxide at different times of the day. Injury, evaluated either as necrosis or chlorophyll, was greatest at midday when stomatal conductance was greatest. Abscisic acid levels were similar over the day, and showed no relation to stomatal conductance.     

Stomatal Conductance Is Essential for Higher Yield Potential of C3 Crops

The present review of gas-exchange studies conducted in rainfed or irrigated field conditions clearly indicates that stomatal conductance (g_s) of major crops has been inadvertently increased in the last 50 to 80 years as higher yields were being reached through conventional plant breeding. These findings suggest that high g_s rates are critical to optimum growth and yield of modern crops, in particular to crop growth rate (CGR) during seed formation and filling. Several hypotheses are presented that may account for this increase. This review also includes studies in which increases in g_s were documented for genetically-engineered plants. However, field evaluations of their gas-exchange performance remain sparse. The main finding of this review is that higher rates of g_s should be sought, as we forego the usual consideration of crop plants as single organisms losing water, but rather acknowledge that water does exit a crop canopy along a Soil Plant Atmosphere Continuum (SPAC). In non-limiting or moderately-limiting soil water conditions, high yields are generated by an active SPAC that requires sustained levels of g_s. In conditions of high evaporative demand, as in summer afternoons, sustained g_s allows for cooler canopies with no mid-day depression in photosynthesis as CO₂ remains able to diffuse in the leaf through stomata.     

Effect of Interruption of Flow Path on Stomatal Conductance of Abies amabilis

Wounding of root or stem water conduction systems or coolingof roots in Abies amabilis produced rapid stomatal closure independentof evaporative demand or leaf water potential. The responsealso occurred in a branch if its xylem was only partially cut,but did not occur if the branch was completely severed. Removingpart of the root system or cooling the roots produced the sameeffect as partial severing of the stem. The speed and uniformityof stomatal closure indicated that the stimulus was physical,linked to water flux in the xylem, and not caused by releaseof a chemical stimulus at the point of xylem flow disruption.The results suggested that stomatal closure could be rapidlyinduced with a change in the flux of water through the soil-plant-atmospherecontinuum. Key words: Capacitance, Stomata, Xylem water flux, Xylem wounding     

Modification of the Conductance and Gating Properties of Ryanodine Receptors by Suramin

Suramin, a polysulfonated napthylurea, increases the open probability and the single-channel conductance of rabbit skeletal and sheep cardiac ryanodine receptor channels. The main mechanism for the increase in P _o is an increase in the duration of open lifetimes. The effects on conduction and gating are completely reversible and involve an interaction with the cytosolic side of the channel. 10 mm dithiothreitol had no effect on the suramin-induced increase in conductance or P _o . Therefore oxidation of sulfhydryl groups on the channels does not appear to be involved. Suramin has been used as an antagonist of ATP at P₂ purinoceptors, however, we find that suramin does not antagonize the effect of ATP at skeletal or cardiac ryanodine receptor channels. The unusual gating kinetics induced by suramin suggest that it does not interact with the adenine nucleotide binding site on the ryanodine receptor but rather binds at a novel site(s). The suramin-induced changes to channel gating and conduction do not prevent the characteristic modification of single-channel properties by micromolar ryanodine. Received: 19 March 1996/Revised: 5 June 1996     

Effects of Sulphur Dioxide and Nitrogen Dioxide on Growth and Translocation in Winter Wheat

Gould, R. P. and Mansfield, T. A. 1988. Effects of sulphur dioxideand nitrogen dioxide on growth and translocation in winter wheat.—J. exp. Bot 39: 389–99 Winter wheat (Triticum aestivum L. cv. Avalon) was grown undersimulated autumn conditions for 4 weeks and exposed to a mixtureof SO₂ and NO₂. Biomass was measured after 2, 3 and 4 weeksand the flag leaves of sample plants were labelled with ¹⁴CO₂.Biomass yields revealed an increase in shoot-to-root ratiosunder polluted conditions. The labelling experiments showedthat less assimilate was transported to the roots, whilst morewas allocated to the younger components of the plant. It appearedthat NO₂ and SO₂ also caused labelled photosynthate to be retainedin the labelled leaf. Reducing the photon flux exacerbated theeffects of SO₂ and NO₂ as indicated by changes in biomass andby the distribution of ¹⁴C. Key words: Wheat, SO₂, NO₂, growth, translocation     

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”.     

Rapid and Specific Modulation of Stomatal Conductance by Blue Light in Ivy (Hedera helix) : An Approach to Assess the Stomatal Limitation of Carbon Assimilation

Low intensity (0.015 millimole per square meter per second) blue light applied to leaves of Hedera helix under a high intensity red light background (0.50 millimole per square meter per second red light) induced a specific stomatal opening response, with rapid kinetics comparable to those previously reported for stomata with `grass type' morphology. The response of stomatal conductance to blue light showed a transient `overshoot' behavior at high vapor pressure difference (2.25 ± 0.15 kiloPascals), but not at low vapor pressure difference (VPD) (0.90 ± 0.10 kilo-Pascal). The blue light-induced conductance increase was accompanied by an increase in net photosynthetic carbon assimilation, mediated by an increase in the intercellular concentration of carbon dioxide. Values of assimilation once the blue light-stimulated conductance increase reached steady state were less than those at the peak of the overshoot, but the ratios of assimilation to transpiration (A/E) and blue light-stimulated ΔA/ΔE were greater during the steady-state response than during the overshoot. These results indicate that significant stomatal limitation of assimilation can occur, but that this limitation may improve water use efficiency under high VPD conditions. Under high intensity red light, the decline in A/E associated with an increase in VPD was minimized when conductance was stimulated by additional low intensity blue light. This effect indicates that the blue light response of stomata may be important in H. helix for the optimization of water use efficiency under natural conditions of high irradiance and VPD.     

The Effects of Sulphur Dioxide on Phloem Transport in Two Cereals

Gould, R. P., Minchin, P. E. H. and Young, P. C. 1988. The effectsof sulphur dioxide on phloem transport in two cereals.—J.exp. BoL 39: 997–1007. In vivo investigations using ¹¹C-labelled photosynthate revealedthat there is a change in the patterns of tracer profiles whencereal leaves are exposed to SO₂. The change after exposureto SO₂ was interpreted in terms of a decrease in lateral waterflow into the sieve tubes brought about by reduced phloem loadingalong the length of a leaf. Analysis also revealed that thespeed of translocation was reduced, as expected by the Munchmodel of phloem transport. Key words: Sulphur dioxide, phloem transport, cereal leaves     

Stomatal Conductance, Photosynthetic Rate, and Pigment Content in Ruellia Tuberosa Leaves as Affected by Coal-Smoke Pollution

Study of the effects of air pollution caused by thermal power plant emissions on some foliar traits of Ruellia tuberosa L. has shown that length and width of stomata, length of stomatal pore, stomatal density, photosynthetic rate, stomatal conductance and chlerophyll content were reduced in the polluted plants in pre-flowering, flowering as well as post-flowering phases of plant growth. Intercellular carbon dioxide concentration in the palisade tissue was increased at each stage of plant development. Stomatal index remained almost unchanged at the polluted site, except on the adaxial surface during the preflowering stage where it was higher as compared to the non-polluted plants.     

Relative Sensitivity and Tolerance of some Gladiolus Cultivars to Sulphur Dioxide

Five Gladiolus cultivars, namely ‘Aldebaran’, ‘BrightEye’, ‘Illusion’, ‘Manisha’ and‘Manmohan’, were exposed to 1 and 2 µg l^–1sulphur dioxide to test their relative-sensitivity toleranceto the pollutant Plants were fumigated experimentally for 2h daily Foliar injury symptoms were observed first in ‘Manisha’followed by ‘Aldebaran’ and ‘Illusion’at the higher dose Photosynthetic pigments and leaf extractpH were significantly decreased, particularly in ‘Manisha’and ‘Illusion’ Overall disturbances in the plantmetabolism due to SO₂ treatment led to retarded growth of plants,as evident from decreased shoot length and phytomass valuesThe order of sensitivity of the five Gladiolus cultivars toSO₂ was as follows, with the greatest first Manisha, Illusion,Aldebaran, Bright Eye, Manmohan Cultivars, Gladiolus, sensitivity, sulphur dioxide, tolerance