Stomatal Opening Mechanism of CAM Plants

Stomata usually open when leaves are transferred from darkness to light. However, reverse-phase stomatal opening in succulent plants has been known. CAM plants such as cacti and Opuntia ficus–indica achieve their high water use efficiency by opening their stomata during the cool, desert nights and closing them during the hot, dry days. Signal transduction pathway for stomatal opening by blue light photoreceptors including phototropins and the carotenoid pigment zeaxanthin has been suggested. Blue light regulated signal transduction pathway on stomatal opening could not be applied to CAM plants, but the most possible theory for a nocturnal response of stomata in CAM plants is photoperiodic circadian rhythm.     

Stomatal responses to changes in temperature at increasing water stress

Summary The response of stomata to a gradual increase in temperature at increasing plant water stress was studied in a hot desert habitat (Negev, Israel) in the field, but under controlled temperature and humidity conditions. Four native species (Zygophyllum dumosum, Artemisia herba-alba, Hammada scoparia, Reaumuria negevensis) and one cultivated plant (Prunus armeniaca) were used in these studies. The stomatal response to temperature was compared with the response in well-irrigated plants of the same species.At low water stress, the diffusion resistance for water vapour decreased in response to a gradual increase in temperature. Transpiration increased accordingly. This response was reversible. All species responded in the same way. The opening of stomata with increasing temperature was apparently independent of the stomatal response regulated by atmospheric humidity. At high plant water stress, the stomatal response was reversed, i.e., the stomata closed when temperature was gradually increased. This stomatal closure was also independent of the closure regulated by atmospheric humidity. The plant water potential at which the stomatal response to temperature was reversed, differed among the species investigated.     

Plant transpiration at high elevations: Theory,field measurements,and comparisons with desert plants

Summary The influence of elevational changes on plant transpiration was evaluated using leaf energy balance equations and well-known elevational changes in the physical parameters that influence water vapor diffusion. Simulated transpirational fluxes for large leaves with low and high stomatal resistances to water vapor diffusion were compared to small leaves with identical stomatal resistances at elevations ranging from sea level to 4 km. The specific influence of various air temperature lapse rates was also tested. Validation of the simulated results was accomplished by comparing actual field measurements taken at a low elevation (300 m) desert site with similar measurements for a high elevation (2,560 m) mountain research site. Close agreement was observed between predicted and measured values of transpiration for the environmental and leaf parameters tested.Substantial increases in solar irradiation and the diffusion coefficient for water vapor in air (D _wv) occurred with increasing elevation, while air and leaf temperatures, the water vapor concentration difference between the leaf and air, longwave irradiation, and the thermal conductivity coefficient for heat in air decreased with increasing elevation. These changes resulted in temperatures for sunlit leaves that were further above air temperature at higher elevations, especially for large leaves. For large leaves with low stomatal resistances, transpirational fluxes for low-elevation desert plants were close to those predicted for high-elevation plants even though the sunlit leaf temperatures of these mountain plants were over 10°C cooler. Simulating conditions with a low air temperature lapse rate (0.003° C m^-1 and 0.004° C m^-1) resulted in predicted transpirational fluxes that were greater than those calculated for the desert site. Transpiration for smaller leaves decreased with elevation for all lapse rates tested (0.003° C m^-1 to 0.010° C m^-1). However, transpirational fluxes at higher elevations were considerably greater than expected for all leaves, especially larger leaves, due to the strong influence of increased solar heating and a greater D _wv. These results are discussed in terms of similarities in leaf structure and plant habit observed among low-elevation desert plants and high-elevation alpine and subalpine plants.     

Seasonal photosynthetic gas exchange and water-use efficiency in a constitutive CAM plant, the giant saguaro cactus (Carnegiea gigantea)

Crassulacean acid metabolism (CAM) and the capacity to store large quantities of water are thought to confer high water use efficiency (WUE) and survival of succulent plants in warm desert environments. Yet the highly variable precipitation, temperature and humidity conditions in these environments likely have unique impacts on underlying processes regulating photosynthetic gas exchange and WUE, limiting our ability to predict growth and survival responses of desert CAM plants to climate change. We monitored net CO₂ assimilation (A _net), stomatal conductance (g _s), and transpiration (E) rates periodically over 2 years in a natural population of the giant columnar cactus Carnegiea gigantea (saguaro) near Tucson, Arizona USA to investigate environmental and physiological controls over carbon gain and water loss in this ecologically important plant. We hypothesized that seasonal changes in daily integrated water use efficiency (WUE_day) in this constitutive CAM species would be driven largely by stomatal regulation of nighttime transpiration and CO₂ uptake responding to shifts in nighttime air temperature and humidity. The lowest WUE_day occurred during time periods with extreme high and low air vapor pressure deficit (D _a). The diurnal with the highest D _a had low WUE_day due to minimal net carbon gain across the 24 h period. Low WUE_day was also observed under conditions of low D _a; however, it was due to significant transpiration losses. Gas exchange measurements on potted saguaro plants exposed to experimental changes in D _a confirmed the relationship between D _a and g _s. Our results suggest that climatic changes involving shifts in air temperature and humidity will have large impacts on the water and carbon economy of the giant saguaro and potentially other succulent CAM plants of warm desert environments.     

Experimental removal of intraspecific competitors — effects on water relations and productivity of a desert bunchgrass,Hilaria rigida

Summary Intraspecific competition in the C₄ bunchgrass Hilaria rigida was examined on a Sonoran Desert site in southeastern California. Potential competition within monospecific stands was experimentally altered by removal of the aboveground portions of all plants within a 1.5 m radius of a monitored plant. Compared with unaltered plots, altered plots had less negative soil water potentials during periods of soil drying. Leaf blades on monitored plants of altered plots remained green longer and had greater stomatal conductances than those on monitored plants on unaltered plots. Production of new culms was twice as great on altered plots. Greater root biomass and root length were observed in altered plots, and root extension into soil areas formerly occupied by roots of neighboring plants occurred within one year after treatment. The results indicate that removal of the aboveground biomass of neighboring plants reduces the competition for limited available soil water in this desert environment.     

Stable isotope composition of water in desert plants

A survey of the stable isotope content of tissue waters of plants from the Negev desert was conducted. Large differences were observed in the extent of enrichment of the heavy isotopes in leaf water relative to local precipitation among different plants. This is apparently caused by the species-dependent stratagems adopted by the plants to cope with water stress, primarily by differences in the depth of water uptake in the soil and through the timing of stomatal openings during the daily cycle. Salt stressed plants showed extreme variability in the isotopic composition of leaf–water. The results show that plants with adaptation to arid conditions can avoid the transpiration regime, which would lead to the strong isotopic enrichment in their leaf water expected under arid conditions. This has implications for the use of stable isotopes in plants as indicators of either plant ecophysiology or paleoclimate. Responsible Editor: Hans Lambers. G. Goodfriend is deceased.     

Effect of drought stress on growth and water relations of the mycorrhizal association Helianthemum almeriense-Terfezia claveryi

 Plants of Helianthemum almeriense were micropropagated on MS medium and inoculated in vitro with Terfezia claveryi mycelium on MH medium and vermiculite. Mycorrhizal (M) and non-mycorrhizal (NM) plants were subjected to a drought stress period of 3 weeks in greenhouse conditions with the soil matric potential maintained at –0.5 MPa. Drought stress did not affect the amount of mycorrhizal colonization. The survival rate of M plants at the end of the drought stress period was higher than that of NM plants. The water potential was higher in M plants than in NM plants by 14% in well-watered and 26% in drought-stressed plants. Transpiration, stomatal conductance and net photosynthesis were higher in M plants than in NM plants. Transpiration was 92% higher in M plants than in NM plants under drought-stress conditions and 40% when irrigated. Stomatal conductance was 45% and 14% higher and net photosynthesis 88% and 54% higher, respectively, in M than in NM plants. Drought-stressed M plants accumulated more N, P and K than drought-stressed NM plants. Reduced negative effects of drought stress on H. almeriense by the desert truffle T. claveryi could be ascribed to specific physiological and nutritional mechanisms, suggesting that this mycorrhizal symbiosis aids adaptation to arid climates. Accepted: 7 July 2000     

Abscisic Acid and Desiccation Tolerance in Mosses

In higher plants the phytohormone ABA is involved in processes that are connected to water deficit, like stomatal closure or desiccation tolerance. In bryophytes, also containing ABA in their tissues, physiological functions remained uncertain for a long time. Quite recently, several papers have shown different effects of exogenously applied ABA: stomatal closure in Anthoceros, drought hardening in Funaria and production of the landform in Riccia. In all these cases the relevant conditions (water deficit) enhance the endogenous ABA level significantly. For induced desiccation tolerance, ABA serves as a mediator to induce specific proteins (dehydrins) strongly connected with this tolerance. Therefore, it can be concluded that in bryophytes ABA has the same function as in higher plants. It acts as a mediator in stress conditions.     

Stomatal cavity modulates the gas exchange of Sorghum bicolor (L.) Moench. grown under different water levels

This work aimed to evaluate the effects of lower water levels on leaf intercellular spaces and to assess their relations with the gas exchange, anatomy, and growth of Sorghum bicolor. Experiments were conducted in a greenhouse, in which plants were subjected to three water conditions (ten replicates, n = 30): well-irrigated, decreased irrigation, and limited irrigation. Lower water levels had no significant effect on the growth of S. bicolor but increased the biomass of the roots. Moreover, the number of leaves, leaf area, and leaf size as well as the chlorophyll content were not affected by lower water levels, and no significant changes were detected for whole plant photosynthesis, transpiration, or stomatal conductance. The water content of the plants and the water potential remained unchanged. However, compared with other treatments, the decreased irrigation decreased water loss and increased the water retention. Lower water levels increased the intercellular CO₂ percentage, mesophyll area, and proportion of stomatal cavities and promoted minor changes in leaf tissue and stomatal traits. The increased stomatal cavities provided higher CO₂ uptake and prevented excessive water loss. Thus, modifications to the intercellular spaces promoted conditions to avoid excessive water loss while concurrently improving CO₂ uptake, which are important traits for drought-tolerant plants.

     

Carbon assimilation, 𝛅13C and water relations of Elaeagnus angustifolia grown at two groundwater depths in the Minqin desert,China

Abstract

We investigated the physiological responses of Elaeagnus angustifolia to variation in groundwater depth. Elaeagnus angustifolia seedlings were grown in the Minqin desert in lysimeters supplied with underground water at the soil depth of 1.40 m and 3.40 m. Results showed that constant access to groundwater allowed plants supplied with water at the lower soil layer to meet their water requirement and, consequently, they were not affected by water stress. There were no differences in A _max (the net CO₂ assimilation rate under conditions of photosynthetically photon flux density and CO₂ saturation), J _max (maximum rate of electron transport) and stomatal conductance between the two underground water treatments. However, plants with deeper groundwater had a significantly higher V _cmax (i.e. a higher carboxylation efficiency of Rubisco) and mesophyll conductance resulting in increased photosynthesis measured at the CO₂ growth condition (A) and, consequently higher intrinsic water-use efficiency (WUE). However, respiration was also increased in plants grown with deeper groundwater. This may have offset the increased A and led to a similar long-term WUE, as expressed by carbon isotope discrimination (δ¹³C), between the two ground water treatments. In the present study, we also found a high foliage nitrogen concentration in the E. angustifolia plants (3.75% on average), that may be very significant ecologically in improving soil properties. The physiological traits of E. angustifolia found in this study confirm that the use of perennial phreatophytic, nitrogen fixing species has significant potential to positively impact soil fertility and carbon sequestration under environmental conditions found in the Minqin desert.     

Water use by the desert cucurbit Citrullus colocynthis (L.) Schrad.

Summary The rates of water use and leaf surface conductance of Citrullus colocynthis (Cucurbitacea) were evaluated from measurements of the surface temperature and microenvironment of leaves. At desert sites in Saudi Arabia the transpiration rates reached 0.13–0.17 g m^-2 s^-1 and the leaf temperatures were always close to air temperature. Leaf models (dry) placed in the canopy were considerably warmer than the air. To investigate responses over a wider range of conditions, plants were grown in a controlled environment room. It was found that when conditions were made hotter than those that occurred in the desert, the stomatal conductance increased greatly. Transpiration rate attained 0.6 g m^-2 s^-1 and the leaves were up to seven degrees cooler than the air. The results suggest a finely-tuned control mechanism working like a switch when the leaves experience extreme conditions, and enabling the plant to avoid lethal temperatures.     

Does engineering abscisic acid biosynthesis in Nicotiana plumbaginifolia modify stomatal response to drought?

The consequences of manipulating abscisic acid (ABA) biosynthesis rates on stomatal response to drought were analysed in wild‐type, a full‐deficient mutant and four under‐producing transgenic lines of N. plumbaginifolia. The roles of ABA, xylem sap pH and leaf water potential were investigated under four experimental conditions: feeding detached leaves with varying ABA concentration; injecting exogenous ABA into well‐watered plants; and withholding irrigation on pot‐grown plants, either intact or grafted onto tobacco. Changes in ABA synthesis abilities among lines did not affect stomatal sensitivity to ABA concentration in the leaf xylem sap ([ABA]_xyl), as evidenced with exogenous ABA supplies and natural increases of [ABA]_xyl in grafted plants subjected to drought. The ABA‐deficient mutant, which is uncultivable under normal evaporative demand, was grafted onto tobacco stock and then presented the same stomatal response to [ABA]_xyl as wild‐type and other lines. This reinforces the dominant role of ABA in controlling stomatal response to drought in N. plumbaginifolia whereas roles of leaf water potential and xylem sap pH were excluded under all studied conditions. However, when plants were submitted to soil drying onto their own roots, stomatal response to [ABA]_xyl slightly differed among lines. It is suggested, consistently with all the results, that an additional root signal of soil drying modulates stomatal response to [ABA]_xyl.     

Water relations and hydraulic control of stomatal behaviour in bell pepper plant in partial soil drying

Two experiments, a split-root experiment and a root pressurizing experiment, were performed to test whether hydraulic signalling of soil drying plays a dominant role in controlling stomatal closure in herbaceous bell pepper plants. In the split-root experiment, when both root parts were dried, synchronous decreases in stomatal conductance (g_s), leaf water potential (LWP) and stem sap flow (SF_stem) were observed. The value of g_s was found to be closely related to soil water potential (SWP) in both compartments. Tight relationships were observed between g_s and stem sap flow under all conditions of water stress, indicating a complete stomatal adjustment of transpiration. When the half-root system has been dried to the extent that its water uptake dropped to almost zero, declines in g_s of less than 20% were observed without obvious changes in LWP. The reduced plant hydraulic conductance resulting from decreased sap flow and unchanged LWP may be a hydraulic signal controlling stomatal closure; the results of root pressurizing supported this hypothesis. Both LWP and g_s in water-stressed plants recovered completely within 25 min of the application of root pressurizing, and decreased significantly within 40 min after pressure release, indicating the hydraulic control of stomatal closure. Our results are in contrast to those of other studies on other herbaceous species, which suggested that chemical messengers from the roots bring about stomatal closure when plants are in water stress.     

荒漠灌木与城市绿化灌木功能性状及其对夏季热浪响应机理的差异

为比较干旱荒漠区城市绿化灌木和荒漠乡土灌木在夏季热浪期的受损差异和其对高温和热浪冲击响应机理的不同,在新疆北疆2022年夏季热浪期末期,测定了9种荒漠乡土灌木和8种城市绿化灌木的4种水分相关功能性状(水力、叶片、光合和碳经济性状),同时调查植株受损程度。调查结果表明,荒漠乡土灌木接近74%的个体在夏季热浪中未受损伤,但有56%的城市绿化灌木个体受损;水力和叶片性状方面,荒漠灌木的枝叶水势、干物质含量和导管直径的平均值低于城市绿化植物,而枝比导率、准稳态导水率、胡伯尔值、比叶面积、导管密度呈相反态势。在碳经济和光合性状方面,荒漠乡土灌木的可溶性碳含量和水分利用效率高于城市绿化灌木(P<0.05),但气孔导度成相反模式(P <0.05),而淀粉和结构性碳含量、蒸腾速率无显著差异(P>0.05);荒漠乡土灌木的性状整合度(G=0.39)高于城市绿化植物(G=0.20),且前者的关键节点个数,以及其与其他性状显著关联的个数均高于后者。结果表明,2022年夏季热浪引起荒漠乡土灌木的受损程度低于城市绿化灌木;荒漠植物在水力性状和性状整合方面存在优势,其水分吸收、运输和减少蒸腾失水的能力要强于城市绿化灌木,热浪影响下更易存活;相对城市绿化灌木,荒漠灌木可以协调功能性状间的关系,采取更多样化的适应机理将降低夏季热浪的负面影响。本研究可为干旱地区城市绿化灌木筛选以及区域生态保护政策制定提供科学参考。     

Water relations and stomatal characteristics of Mediterranean plants with different growth forms and leaf habits: responses to water stress and recovery

The aim of this study was to extent the range of knowledge about water relations and stomatal responses to water stress to ten Mediterranean plants with different growth forms and leaf habits. Plants were subjected to different levels of water stress and a treatment of recovery. Stomatal attributes (stomatal density, StoD), stomatal conductance (g _s), stomatal responsiveness to water stress (SR), leaf water relations (pre-dawn and midday leaf water potential and relative water content), soil to leaf apparent hydraulic conductance (K _L) and bulk modulus of elasticity (ε) were determined. The observed wide range of water relations and stomatal characteristics was found to be partially depended on the growth form. Maximum g _s was related to StoD and the stomatal area index (SAI), while g _s evolution after water stress and recovery was highly correlated with K _L. Relationships between SR to water deficit and other morphological leaf traits, such as StoD, LMA or ε, provided no general correlations when including all species. It is concluded that a high variability is present among Mediterranean plants reflecting a continuum of leaf water relations and stomatal behaviour in response to water stress.     

Gas exchange and resource-use efficiency of Leymus cinereus (Poaceae): diurnal and seasonal responses to naturally declining soil moisture

We examined factors that limit diurnal and seasonal photosynthesis in Leymus cinereus, a robust tussock grass from shrub-steppes of western North America. Data from plants in a natural stand and in experimental field plots indicate that this bunchgrass has 1) a high photosynthetic capacity, 2) high leaf nitrogen content and high nitrogen-use efficiency, 3) a steep leaf-to-air diffusion gradient for carbon dioxide, which enhances intrinsic water-use efficiency, and 4) photosynthetic tissues that tolerate severe water stress and recover quickly from moderate water stress. Midday depressions of CO₂ assimilation (A) and stomatal conductance were slight in plants with plentiful water, but marked in plants subject to moderate water stress. Midday stomatal closure in moderately stressed plants reduced intercellular carbon dioxide concentration (c_i) by ≈40 μl liter^-1. The maximum rate of A achieved during the day for severely stressed plants (predawn water potential = -4 MPa) was one-third and daily carbon gain per unit leaf area was about one-fourth that of well-watered plants. For plants in the natural stand, CO₂-saturated photosynthesis declined almost linearly with decreasing soil water availability over the growing season, whereas there was little effect on A at CO₂ ambient levels or on carboxylation efficiency until predawn water potentials reached -1.8 MPa. Nitrogen-use efficiency declined with diminishing soil moisture, but there was no seasonal change in stomatal limitation or instantaneous water-use efficiency as estimated from A vs. c_i curves at optimal leaf temperature and moderate atmospheric evaporative demand. Thus, reduced stomatal conductance in response to increased evaporative demand may increase stomatal limitation diumally, but over the growing season, stomatal limitation estimated from A vs. c_i curves is relatively constant because maximum stomatal conductance is closely tuned to the CO₂ assimilatory capacity of the mesophyll.     

Water Relations,CO2 Exchange,Water-use Efficiency and Growth of Welwitschia mirabilis Hook. fil. in three Contrasting Habitats of the Namib Desert1

CO₂ exchange, transpiration and leaf water potential of Welwitschia mirabilis were measured in three contrasting habitats of the Namib desert. From these measurements stomatal conductance, internal CO₂concentration and WUE were calculated. In two of the three habitats photosynthetic CO₂ uptake decreased and transpiration increased with increasing leaf age while in the third habitat CO₂ uptake increased and transpiration decreased with leaf age. Except for the stomata of young leaf sections in this habitat, stomata closed with increasing δw leading to a pronounced midday depression of CO₂ uptake. The high stomatal limitation of photosynthetic CO₂ uptake of glasshouse-grown plants was verified in the natural habitat. Photosynthetic CO₂ uptake saturated between 800 and 1300 μmol photons m^?2 s^?1depending on leaf age and habitat. CO₂ uptake had a broad temperature optimum declining significantly beyond 32 °C. Predawn leaf water potential reflected water availability and atmospheric conditions in the three habitats and ranged from ? 2.5 to ? 6.2 MPa. There was a pronounced diurnal course of leaf water potential in all habitats. During the day a gradient in water potential developed along the leaf axis with the lowest potential at the leaf's tip. With respect to whole plant balances of CO₂ exchange and transpiration, there were marked differences between Welwitschias in the three habitats. Despite a negative CO₂ balance over a period of five months, leaves in the driest habitat grew constantly at the expense of carbon reserves in the plant. Only at the wettest site did carbon gain exceed carbon demand for growth. The WUE of whole plants was insignificant in all habitats. The results were as contrasting as the habitats and plants and did not allow generalisations about adaptational features of Welwitschia mirabilis.     

Environmental effects on circadian rhythms in photosynthesis and stomatal opening

Persistent circadian rhythms in photosynthesis and stomatal opening occurred in bean (Phaseolus vulgaris L.) plants transferred from a natural photoperiod to a variety of constant conditions. Photosynthesis, measured as carbon assimilation, and stomatal opening, as conductance to water vapor, oscillated with a freerunning period close to 24 h under constant moderate light, as well as under light-limiting and CO₂-limiting conditions. The rhythms damped under constant conditions conducive to high photosynthetic rates, as did rates of carbon assimilation and stomatal conductance, and this damping correlated with the accumulation of carbohydrate. No rhythm in respiration occurred in plants transferred to constant darkness, and the rhythm in stomatal opening damped rapidly in constant darkness. Damping of rhythms also occurred in leaflets exposed to constant light and CO₂-free air, demonstrating that active photosynthesis and not simply light was necessary for sustained expression of these rhythms. This is CIWDPB Publication No. 1142 This research was supported by National Science Foundation grant BSR 8717422 (C.B.F.) and a U.S. Department of Agriculture training grant to Stanford University (T.L.H.).     

Effects of evening and nighttime leaf wetting on stomatal behavior of Cryptomeria japonica growing in dry soil

To examine the hypothesis that stomatal behavior of plants in dry soil is influenced by a slow recovery from daytime water deficit, we studied the effect of repeated wetting of leaves during evening and night in Cryptomeria japonica seedlings grown in dry soil. After 7 and 10 days of leaf wetting treatment the midday leaf water potential decreased and the transpiration rate increased, respectively. Therefore, we suggest that rapid recovery from daytime water deficit could weaken the water conserving stomatal behavior that adapts to drought conditions in the roots.     

Relationships between gas-exchange characteristics and stomatal structural modifications in some desert grasses under high salinity

Two populations, one from lesser saline Derawar Fort (DF) and the other from highly saline Ladam Sir (LS) in the Cholistan desert, for each of the five grass species, Aeluropus lagopoides, Cymbopogon jwarancusa, Lasiurus scindicus, Ochthochloa compressa, and Sporobolus ioclados were examined to investigate the influence of salinity on structural and functional characteristics of stomata. Salinity tolerance in A. lagopoides mainly depended on controlled transpiration rate (E) and high water-use efficiency (WUE), which was found to be regulated by fewer and smaller stomata on both leaf surfaces as well as stomatal encryption by epidermal invaginations. C. jwarancusa had sunken stomata on the abaxial surface only, which largely reflected a reduced E, but less affected stomatal conductance (g _s) or WUE. L. scindicus had fewer but larger stomata along with hairs/trichomes which may function to avoid water loss through transpiration, and hence, to attain a high WUE. In O. compressa stomata were found only on the abaxial surface and these were completely encrypted by epidermal invaginations as well as a dense covering of microhairs, which was associated with a low E and high WUE under salinity stress. In S. ioclados, the traits of increased stomatal density and decreased stomatal area may be critical for stomatal regulation under salt-prone environments. High stomatal regulation depended largely on stomatal density, area, and degree of encryption under salinity, which is of great ecophysiological significance for plants growing under osmotic stresses.