Effects of Age and Water Stress on Endogenous Levels of Plant Growth Regulators in Euphorbia lathyrus L.

The effect of droughting and waterlogging on changes in endogenouslevels of hormones have been studied in Euphorbia lathyrus L.with particular reference to plant age and leaf age. Younger tissues (i.e. immature leaves and young plants) synthesizedsignificantly greater amounts of ABA per unit fresh weight thandid corresponding older tissue. The increased ABA levels correlatedwell with corresponding increases in water saturation deficit(WSD) and stomatal resistance in a given treatment but the relationshipbetween the absolute values of these three parameters variedgreatly between seedlings of different ages. Endogenous ethylenelevels in leaves and stems were unaffected by droughting irrespectiveof age. Imposition of waterlogging led to marked increases in ABA levelsin young plants though this was not strictly related to changesin WSD or stomatal resistance. Endogenous levels of ethyleneincreased in response to waterlogging, this being marked inthe older seedlings which also had a higher incidence of senescenceand abscission. The more complex effects of waterlogging, the significance ofage of tissue on hormonal responses to stress, and the adaptivevalue of such responses to younger tissues are discussed.     

Water balance of tissue segments from leaves of different insertion levels

Actual heterogeneity of indicators of water balance was studied on tissue segments from leaves ofBrassica oleracea L. v.acephala andNicotiana tabacum L. under controlled conditions. Maximal values of transpiration rate, water absorption rate, water transport rate and low values of WSD at the same time were observed at an initial WSD of 0% in segments from middle leaves and at an initial WSD of 20% mostly in segments from upper leaves. Although the properties of leaf tissue itself were probably the basic factors determining the actual heterogeneity of indicators of water balance, without connection with all other factors (e. g. various microclimates, supply of water) heterogeneity did not appear to a full degree.     

Experimental Studies of the Factors Controlling Transpiration: II. THE RELATION BETWEEN TRANSPIRATION RATE AND LEAF WATER CONTENT

Data are presented which show, when stomatal control is eliminated,that wheat leaves may lose 5–6 per cent. and Pelargoniumleaves 10–12 per cent. of their water without any reductionin the transpiration rate. Experiments in which Pelargonium and wheat leaves, with stomatalcontrol present, were submitted to cycles of changing watercontent also failed to establish any direct relation betweentranspiration rate and leaf water content. It is concluded that leaf water content over the range of 70–100per cent. of that present in the turgid state has no significanteffect in determining the rate of water loss from leaves. A repetition of Knight's experiment showed that stomata openedin still air and closed in moving air. This was not recordedby Knight, who used a porometer cup permanently attached tothe leaf. It is concluded that the higher transpiration raterecorded by Knight after a period of still air was due to widerstomatal aperture and not to the higher leaf water content assuggested by him.     

彰武松、樟子松光合生产与蒸腾耗水特性

本文采用Li-6400光合测定系统对性成熟(18a)阶段彰武松(Pinus densiflora var.zhangwuensis)和樟子松(Pinus sylvestris var.mongolica)光合及蒸腾指标不同季节日变化进行了测定,并采用切枝蒸腾法对两个树种叶片气孔蒸腾和角质层蒸腾进行对比测定,评价了气孔开闭敏感性,探讨了两个树种光合生产与蒸腾耗水特性。结果表明：在同样生境条件下,彰武松比樟子松有较大的光合速率(Pn)和较小的蒸腾速率(Tr)。在5月和7月,彰武松的Pn和Tr日变化呈现明显双峰型,其Pn和Tr“午休”现象均主要受气孔限制;在10月呈单峰型。樟子松的Pn和Tr日变化在整个生长季均呈单峰型,而且,彰武松日光合量(DAP)均高于樟子松,是樟子松的163.4%(5月)、211.1%(7月)和183.6%(10月)。光响应曲线参数表明：在不同月份,彰武松最大光合速率(Pmax)均大于樟子松,且光饱和点(LSP)较高,光补偿点(LCP)较低。在任意被测时刻,彰武松气孔导度(Gs)和Tr都小于樟子松。彰武松具有较小气孔和角质层蒸腾速度,并且在同样干旱条件下,彰武松气孔下陷,其气孔的开闭反应更加敏感。彰武松水分利用效率(WUE)较高,约是樟子松的2.29倍。这些结果暗示,彰武松以其高的光合速率和低的蒸腾耗水特性,提高水分利用效率,以其敏感的气孔开闭机制和旱生叶片结构进而实现在干旱半干旱地区的速生特性。     

Effect of ouabain on stomatal movements and transpiration rate of Secale cereale

Young rye seedlings (Secale cereale cv. Petkus) were grown for 12 or 18 din aerated nutrient solution. CO2-free air in light or in darkness induced stomatal opening. When light and CO2-free air were applied together, maximum stomatal opening was observed. Na+ channel inhibitor [3H]ouabain was taken up from nutrient medium and translocated into the leaves via the xylem sap. The presence of 10-5 M ouabain in the root medium for 20 h increased stomatal opening and transpiration rate. On the other hand, stomatal closing was not affected by the presence of ouabain. These results suggested that Na+ might play a role in stomatal movements. This revised version was published online in July 2006 with corrections to the Cover Date.     

Leaf age and salinity influence water relations of pepper leaves

Plant growth is reduced under saline conditions even when turgor in mature leaves is maintained by osmotic adjustment. The objective of this study was to determine if young leaves from salt-affected plants were also osmotically adjusted. Pepper plants (Capsicum annuum L. cv. California Wonder) were grown in several levels of solution osmotic potential and various components of the plants' water relations were measured to determine if young, rapidly growing leaves could accumulate solutes rapidly enough to maintain turgor for normal cell enlargement. Psychrometric measurements indicated that osmotic adjustment is similar for both young and mature leaves although osmotic potential is slightly lower for young leaves. Total water potential is also lower for young leaves, particularly at dawn for the saline treatments. The result is reduced turgor under saline conditions at dawn for young but not mature leaves. This reduced turgor at dawn, and presumably low night value, is possibly a cause of reduced growth under saline conditions. No differences in leaf turgor occur at midday. Porometer measurements indicated that young leaves at a given salinity level have a higher stomatal conductance than mature leaves, regardless of the time of day. The result of stomatal closure is a linear reduction of transpiration.     

不同生长时期多伞阿魏土壤养分及光合特性比较研究

以多伞阿魏(Ferula ferulaeoides)为研究对象,分别对其展叶期、成熟期、果期土壤含水量、pH、有机质、养分和叶片的净光合速率、蒸腾速率、水分利用率等光合指标的日变化进行比较研究。结果表明:土壤含水量、有机质、养分随着多伞阿魏生长呈下降趋势,而pH却有所上升,但变化幅度不大。在不同土壤深度上,土壤含水量、pH随着土层深度的加深逐渐升高,而土壤有机质、养分含量却随土层的加深逐渐减少。多伞阿魏3个时期的净光合速率日变化均为双峰曲线,在14:00出现明显的“光合午休”现象,日平均大小为展叶期>成熟期>果期;蒸腾速率日变化趋势表现为单峰曲线,在中午14:00达到最高值,日平均大小为展叶期>果期>成熟期;水分利用率日变化总体呈上升趋势,在14:00出现最低值,日平均大小为成熟期>果期>展叶期。通过对多伞阿魏净光合速率和蒸腾速率与影响因子的相关分析显示:在整个生长期气孔导度,光合有效辐射与净光合速率呈显著正相关性;光照强度与蒸腾速率有明显正相关性。     

Retarded Stomatal Closure by Phenylmercuric Acetate

Excised leaves of Nerium oleander, which were treated with phenylmercuric acetate (PMA) 1_1/2 h before excising, transpired faster than untreated excised leaves. Similarly, PMA-treated oleander plants transpired more than untreated plants in the dark. These effects were due to retarded stomatal closure caused by PMA. Measurements of stomatal apertures on disks of Vicia faba leaves kept in the dark, and of diffusive resistance to water vapor from Phaseolus vulgaris leaves, confirmed that PMA retards stomatal closing as well as stomatal opening. However, day-time reductions in transpiration by PMA greatly exceed night-time increases in water loss. The mechanisms of stomatal movement, as affected by PMA, are discussed. PMA may conceivably decrease the permeability of guard cell membranes to solutes, thereby retarding all stomatal movements that are osmotically induced.     

H_2O_2作为根源信号介导盐胁迫诱导的蚕豆气孔关闭反应

H2O2作为信号分子可被多种胁迫诱导产生并在细胞内积累,进而参与调节植物的抗逆反应。文章通过远红外热成像观察等实验发现,根部NaCl胁迫可诱导蚕豆气孔关闭,叶片温度上升,叶片内Na＋和H2O2含量增加,蒸腾流汁液中H2O2浓度升高。另外,NaCl可直接诱导离体蚕豆根产生H2O2,却不能影响表皮条内H2O2含量。NaCl胁迫条件下产生的蒸腾流汁液可直接诱导表皮条气孔关闭,该过程可被抗氧化剂抗坏血酸（AsA）所逆转。这些结果表明,H2O2作为盐胁迫的根源信号,可能通过维管系统运输参与调节蚕豆气孔的关闭反应。     

Localization of mechanisms involved in hydropassive and hydroactive stomatal responses of Sambucus nigra to dry air

The response of stomata to a reduction of air humidity is composed of a hydropassive opening followed by active closure. Whereas the mechanisms behind the hydropassive opening are largely understood, the location and physiological basis of the sensing mechanisms leading to active closure are not yet known. This study attempts to evaluate the importance of a single pore's transpiration on its own response and that of adjacent pores. Selected stomata on attached intact leaves of Sambucus nigra were sealed with mineral oil and the response to a reduction of humidity was continuously observed in situ. Blocking a pore's transpiration had no appreciable effect on hydropassive opening and subsequent stomatal closure. If the adjacent stomata were additionally sealed, the closing response was reduced, but not the hydropassive opening. On the other hand, sealing the entire leaf surface, except a small area including the observed stomata, also reduced stomatal closure. These results indicate that strictly local processes triggered by a pore's own transpiration are not required to induce stomatal closure. To describe the effect of one pore's transpiration on the hydropassive and hydroactive responses of neighboring stomata, a simple spatial model was constructed. It suggests that 90% of the closing effect covers an area of approximately 0.5 mm2, whereas the effect on hydropassive opening affects an area of approximately 1 mm2. This divergence may suggest mechanisms other than or in addition to those involving changes of local leaf water potential.     

Effect of Potassium Levels on the Stomatal Behavior of the Hemi-Parasite Striga hermonthica

The hemi-parasite Striga hermonthica, exhibits an anomalous pattern of stomatal response, stomata remaining open in darkness and when subjected to water stress. This suggests irregularity in stomatal response due to malfunction of the stomatal mechanism. To test this suggestion guard cells were isolated from the effects of surrounding cells, by incubating epidermal strips at low pH. These stomata responded rapidly to low CO₂ concentrations, darkness, and ABA. Thus, a paradox exists between stomatal behavior observed in whole leaves and that in isolated guard cells. However, when incubated in the presence of high potassium concentrations (>200 millimolar KCl) stomatal responses in epidermal strips resembled those found in whole leaves, with enhanced opening and reduced closing responses. It is suggested that the anomalous behavior of stomata in Striga and other leafy hemiparasites can be explained by the modulatory effects of high potassium concentrations which accumulate in the leaves as a consequence of high transpiration rates and the lack of a retranslocation system.     

Cadmium effects on leaf transpiration of sugar beet (Beta vulgaris)

Seedlings of sugar beet ( Beta vulgaris L. cv Monohill) were cultivated for 4 weeks in nutrient solution containing different concentrations of CdCl₂ (0 to 10 μ M ). The effects of Cd on appearance and function of stomata and leaf cuticle were investigated by water loss measurements and microscopy. The leaf transpiration rate increased with increasing Cd concentrations while the sum total of stomatal aperture area per unit leaf area decreased. Already at low Cd levels. an increase of defective and undeveloped stomata was found in Cd treated plants. These stomata are closed or have small apertures and probably lack a functional closing mechanism. The number of intact stomata per unit leaf area was lower in leaves of Cd treated plants than in controls, and Cd induced closure of intact stomata. The total number of stomata per leaf area slightly increases with increasing Cd concentration. as does the percentage of small stomata. Furthermore. specific leaf area increased, while the density of leaf structure was decreased by Cd. From this observation we conclude that the increase in transpiration rate caused by Cd is primarily due to effects on the permeability of the leaf cuticle to water.     

Stomatal oscillations at small apertures: indications for a fundamental insufficiency of stomatal feedback-control inherent in the stomatal turgor mechanism.

Continuous measurements of stomatal aperture simultaneously with gas exchange during periods of stomatal oscillations are reported for the first time. Measurements were performed in the field on attached leaves of undisturbed Sambucus nigra L. plants which were subjected to step-wise increases of PPFD. Oscillations only occurred when stomatal apertures were small under high water vapour mole fraction difference between leaf and atmosphere (DeltaW). They consisted of periodically repeated opening movements transiently leading to very small apertures. Measurements of the area of the stomatal complex in parallel to the determination of aperture were used to record volume changes of guard cells even if stomata were closed. Stomatal opening upon a light stimulus required an antecedent guard cell swelling before a slit occurred. After opening of the slit the guard cells again began to shrink which, with some delay, led to complete closure. Opening and closing were rhythmically repeated. The time-lag until initial opening was different for each individual stoma. This led to counteracting movements of closely adjacent stomata. The tendency to oscillate at small apertures is interpreted as being a failure of smoothly damped feedback regulation at the point of stomatal opening: Volume changes are ineffective for transpiration if stomata are still closed; however, at the point of initial opening transpiration rate rises steeply. This discontinuity together with the rather long time constants inherent in the stomatal turgor mechanism makes oscillatory overshooting responses likely if at high DeltaW the 'nominal value' of gas exchange demands a small aperture.     

Movement of water from old to young leaves in three species of succulents

A hypothetical adaptive response of succulent plants to drought-stress is the redistribution of water from old to young leaves. We examined the effects of possible movement of water from old to young leaves in three succulent species, Carpobrotus edulis (weak CAM-inducible), Kalanchoe tubiflora (CAM) and Sedum spectabile (possibly a CAM-cycler or CAM-inducible). Old leaves were removed from plants, and photosynthesis, transpiration, f. wt : d. wt ratios, diurnal acid fluctuations, stomatal conductance and internal CO2 concentrations of the remaining young leaves were measured during drought-stress. Comparison was made with plants retaining old leaves. There was no evidence that water moved from old to young leaves during drought-stress as previously hypothesized. Only in drought-stressed plants of K. tubiflora, were photosynthetic and transpiration rates of young leaves greater on shoots with old leaves removed compared with attached. There was a trend in all species for greater fluctuations in acidity in young leaves on shoots that lacked older leaves. For two of the three species studied, the f. wt : d. wt ratios of young leaves were greater under drought-stress, on shoots with old leaves removed than with them attached. Absence of old leaves may reduce competition for water with young leaves, which consequently have higher water content and greater photosynthetic rates.     

Stomatal malfunctioning under low VPD conditions: induced by alterations in stomatal morphology and leaf anatomy or in the ABA signaling?

Exposing plants to low VPD reduces leaf capacity to maintain adequate water status thereafter. To find the impact of VPD on functioning of stomata, stomatal morphology and leaf anatomy, fava bean plants were grown at low (L, 0.23 kPa) or moderate (M, 1.17 kPa) VPDs and some plants that developed their leaves at moderate VPD were then transferred for 4 days to low VPD (M→L). Part of the M→L‐plants were sprayed with ABA (abscisic acid) during exposure to L. L‐plants showed bigger stomata, larger pore area, thinner leaves and less spongy cells compared with M‐plants. Stomatal morphology (except aperture) and leaf anatomy of the M→L‐plants were almost similar to the M‐plants, while their transpiration rate and stomatal conductance were identical to that of L‐plants. The stomatal response to ABA was lost in L‐plants, but also after 1‐day exposure of M‐plants to low VPD. The level of foliar ABA sharply decreased within 1‐day exposure to L, while the level of ABA‐GE (ABA‐glucose ester) was not affected. Spraying ABA during the exposure to L prevented loss of stomatal closing response thereafter. The effect of low VPD was largely depending on exposure time: the stomatal responsiveness to ABA was lost after 1‐day exposure to low VPD, while the responsiveness to desiccation was gradually lost during 4‐day exposure to low VPD. Leaf anatomical and stomatal morphological alterations due to low VPD were not the main cause of loss of stomatal closure response to closing stimuli.     

In Situ Measurement of Epidermal Cell Turgor, Leaf Water Potential, and Gas Exchange in Tradescantia virginiana L

A combined system has been developed in which epidermal cell turgor, leaf water potential, and gas exchange were determined for transpiring leaves of Tradescantia virginiana L. Uniform and stable values of turgor were observed in epidermal cells (stomatal complex cells were not studied) under stable environmental conditions for both upper and lower epidermises. The changes in epidermal cell turgor that were associated with changes in leaf transpiration were larger than the changes in leaf water potential, indicating the presence of transpirationally induced within-leaf water potential gradients. Estimates of 3 to 5 millimoles per square meter per second per megapascal were obtained for the value of within-leaf hydraulic conductivity. Step changes in atmospheric humidity caused rapid changes in epidermal cell turgor with little or no initial change in stomatal conductance, indicating little direct relation between stomatal humidity response and epidermal water status. The significance of within-leaf water potential gradients to measurements of plant water potential and to current hypotheses regarding stomatal response to humidity is discussed.     

Specification of adaxial and abaxial stomata, epidermal structure and photosynthesis to CO2 enrichment in maize leaves

Acclimation to CO2 enrichment was studied in maize plants grown to maturity in either 350 or 700 microl l-1 CO2. Plants grown with CO2 enrichment were significantly taller than those grown at 350 microl l-1 CO2 but they had the same number of leaves. High CO2 concentration led to a marked decrease in whole leaf chlorophyll and protein. The ratio of stomata on the adaxial and abaxial leaf surfaces was similar in all growth conditions, but the stomatal index was considerably increased in plants grown at 700 microl l-1 CO2. Doubling the atmospheric CO2 content altered epidermal cell size leading to fewer, much larger cells on both leaf surfaces. The photosynthesis and transpiration rates were always higher on the abaxial surface than the adaxial surface. CO2 uptake rates increased as atmospheric CO2 was increased up to the growth concentrations on both leaf surfaces. Above these values, CO2 uptake on the abaxial surface was either stable or increased as CO2 concentration increased. In marked contrast, CO2 uptake rates on the adaxial surface were progressively inhibited at concentrations above the growth CO2 value, whether light was supplied directly to this or the abaxial surface. These results show that maize leaves adjust their stomatal densities through changes in epidermal cell numbers rather than stomatal numbers. Moreover, the CO2-response curve of photosynthesis on the adaxial surface is specifically determined by growth CO2 abundance and tracks transpiration. Conversely, photosynthesis on the abaxial surface is largely independent of CO2 concentration and rather independent of stomatal function.     

The effect of root cooling on hormone content, leaf conductance and root hydraulic conductivity of durum wheat seedlings (Triticum durum L.)

Root cooling of 7-day-old wheat seedlings decreased root hydraulic conductivity causing a gradual loss of relative water content during 45 min (RWC). Subsequently (in 60 min), RWC became partially restored due to a decrease in transpiration linked to lower stomatal conductivity. The decrease in stomatal conductivity cannot be attributed to ABA-induced stomatal closure, since no increase in ABA content in the leaves or in the concentration in xylem sap or delivery of ABA from roots was found. However, decreased stomatal conductance was associated with a sharp decline in the content of cytokinins in shoots that was registered shortly after the start of root cooling and linked to increases in the activity of cytokinin-oxidase. This decrease in shoot cytokinin content may have been responsible for closing stomata, since this hormone is known to maintain stomatal opening when applied to plants. In support of this, pre-treatment with synthetic cytokinin benzyladenine was found to increase transpiration of wheat seedlings with cooled roots and bring about visible loss of turgor and wilting.     

Transpiration, CO2 assimilation, WUE, and stomatal aperture in leaves of Viscum album (L.): Effect of abscisic acid (ABA) in the xylem sap of its host (Populus x euamericana).

Leaves of the mistletoe Viscum album (L.) show a high rate of transpiration, even when the host is under severe drought stress. The hypothesis that a strong control of ABA influx from the xylem sap of the host into the mistletoe prevents stomatal closure in mistletoe leaves was tested under the following conditions: sections of poplar twigs carrying a mistletoe were perfused with artificial xylem sap that contained different ABA concentrations and both transpiration and ABA levels were analysed in mistletoe leaves. Despite variation by a factor of 10(4), the ABA content of the host xylem did not affect ABA levels, leaf transpiration, CO(2) assimilation, WUE, or the degree of stomatal aperture in mistletoe leaves. These observations support the hypothesis of a strong control of ABA influx from the host of the xylem into the mistletoe, although degradation of ABA before it enters the mistletoe leaves cannot be excluded. This mechanism may ensure a water and nutritional status favourable for the mistletoe, even if the water status of the host is impaired. Despite the lack of short-term sensitivity of ABA levels in mistletoe leaves to even strong changes of ABA levels in the xylem sap of the host, ABA levels in mistletoe leaves were relatively high compared to ABA levels in the leaves of several tree species including poplar. Since significant transpiration of the mistletoe leaves was observed despite high ABA levels, a diminished sensitivity of the stomata of mistletoe leaves to ABA has to be concluded. The stomatal density of adaxial Viscum leaves of 89+/-23 stomata per mm is lower than those reported in a study performed at the end of the 19th century.     

Stomatal characteristics and leaf anatomy of potato plantlets cultured in vitro under photoautotrophic and photomixotrophic conditions

Summary Potato plantlets (Solanum tuberosum L. cv. Benimaru) were cultured under photoautotrophic (without any sucrose in the nutrient medium and with enriched CO₂ and high photosynthetic photon flux) and photomixotrophic conditions (20 g 1⁻¹ sucrose in the medium). Leaf anatomy and stomatal characteristics of the leaves were studied in relation to stomatal size and density. Leaf diffusive resistance, transpiration rate, and wax content of the leaves were also investigated. In the photoautotrophic treatment, stomata behaved normally by closing in the dark and opening in the light. The stomatal density increased twofold compared to that of the photomixotrophic treatment. Relatively thick leaves and an organized palisade layer were observed and the epicuticulal wax content was remarkably higher in this treatment, i.e., seven times greater than that of photomixotrophic treatment. In general, higher diffusive resistance of the leaves was observed than under photomixotrophic conditions; also the resistance increased in darkness and decreased in the light. All these characteristics led the plantlets to have a normal and controlled transpiration rate, which was exceptionally high in the photomixotrophic treatment throughout the light and the dark period.