Interpretation of an empirical model for stomatal conductance in terms of guard cell function

An empirical model for stomatal conductance (g), proposed by Leuning (1995, this issue) as a modification of Ball, Woodrow & Berry's (1987) model, is interpreted in terms of a simple, steady-state model of guard cell function. In this model, stomatal aperture is a function of the relative turgor between guard cells and epidermal cells. The correlation between g and leaf surface vapour pressure deficit in Leuning's model is interpreted in terms of stomatal sensing of the transpiration rate, via changes in the gradient of total water potential between guard cells and epidermal cells. The correlation between g, CO₂ assimilation rate and leaf surface CO₂ concentration in Leuning's model is interpreted as a relationship between the corresponding osmotic gradient, irradiance, temperature, intercellular CO₂ concentration and stomatal aperture itself. The explicit relationship between osmotic gradient and stomatal aperture (possibly describing the effect of changes in guard cell volume on the membrane permeability for ion transport) results in a decrease in the transpiration rate in sufficiently dry air. Possible extension of the guard cell model to include stomatal responses to soil water status is discussed.     

The Effect of Phosphorus and Potassium Deficiencies on Transpiration in Tea (Camellia sinensis)

The effect of phosphorus and potassium deficiencies on transpiration in tea (Camellia sinensis L. Clone DT 1) was studied. The plants were grown in sand culture, and measurements were made after the plants showed phosphorus or potassium deficiency symptoms. The overall growth of plants was reduced by potassium deficiency but not by phosphorus deficiency. Both deficiencies reduced stomatal aperture and increased leaf water potential. Stomatal density decreased in phosphorus deficient leaves and it increased in potassium deficient leaves. The transpiration of whole plants was reduced by both deficiencies. The relative sensitivity of transpiration to water stress was increased by potassium deficiency but not by phosphorus deficiency.     

Does transpiration control stomatal responses to water vapour pressure deficit?

Three types of observations were used to test the hypothesis that the response of stomatal conductance to a change in vapour pressure deficit is controlled by whole-leaf transpiration rate or by feedback from leaf water potential. Varying the leaf water potential of a measured leaf by controlling the transpiration rate of other leaves on the plant did not affect the response of stomatal conductance to vapour pressure deficit in Glycine max. In three species, stomatal sensitivity to vapour pressure deficit was eliminated when measurements were made at near-zero carbon dioxide concentrations, despite the much higher transpiration rates of leaves at low carbon dioxide. In Abutilon theophrasti, increasing vapour pressure deficit sometimes resulted in both decreased stomatal conductance and a lower transpiration rate even though the response of assimilation rate to the calculated substomatal carbon dioxide concentration indicated that there was no ‘patchy’ stomatal closure at high vapour pressure deficit in this case. These results are not consistent with stomatal closure at high vapour pressure deficit caused by increased whole-leaf transpiration rate or by lower leaf water potential. The lack of response of conductance to vapour pressure deficit in carbon dioxide-free air suggests that abscisic acid may mediate the response.     

ABA-deficient (aba1) and ABA-insensitive (abi1-1, abi2-1) mutants of Arabidopsis have a wild-type stomatal response to humidity

In most plant species, a decrease in atmospheric humidity at the leaf surface triggers a decrease in stomatal conductance. While guard cells appear to respond to humidity‐induced changes in transpiration rate, as opposed to relative humidity or vapour pressure difference, the underlying cellular mechanisms for this response remain unknown. In the present set of experiments, abscisic acid (ABA)‐deficient (aba1) and ABA‐insensitive (abi1‐1 and abi2‐1) mutants of Arabidopsis thaliana were used to test the hypothesis that the humidity signal is transduced by changes in the flux or concentration of ABA delivered to the stomatal complex in the transpiration stream. In gas exchange experiments, stomatal conductance was as sensitive to changes in vapour pressure difference in aba1, abi1‐1 and abi2‐1 mutant plants as in wild‐type plants. These experiments appear to rule out an obligate role for either the concentration or flux of ABA or ABA conjugates as mediators of the guard cell response to atmospheric water potential. The results stand in contrast to the well‐established role of ABA in mediating guard cell responses to decreases in soil water potential.     

Temperature-dependent stomatal movement in tulip petals controls water transpiration during flower opening and closing

Temperature‐dependent tulip petal opening and closing movement was previously suggested to be regulated by reversible phosphorylation of a plasma membrane aquaporin ( Azad et al., 2004a ). Stomatal apertures of petals were investigated during petal opening at 20°C and closing at 5°C. In completely open petals, the proportion of open stomata in outer and inner surfaces of the same petal was 27 ± 6% and 65 ± 3%, respectively. During the course of petal closing, stomatal apertures in both surfaces reversed, and in completely closed petals, the proportion of open stomata in outer and inner surfaces of the same petal was 74 ± 3% and 29 ± 6%, respectively, indicating an inverse relationship between stomatal aperture in outer and inner surfaces of the petal during petal opening and closing. Both petal opening and stomatal closure in the outer surface of the petal was inhibited by a Ca²⁺ channel blocker and a Ca²⁺ chelator, whereas the inner surface stomata remained unaffected. On the other hand, sodium nitroprusside, a nitric oxide donor, had no effect on stomatal aperture of the outer surface but influenced the inner surface stomatal aperture during petal opening and closing, suggesting different signalling pathways for regulation of temperature‐dependent stomatal changes in the two surfaces of tulip petals. Stomata were found to be differentially distributed in the bottom, middle and upper parts of tulip petals. During petal closing, water transpiration was observed by measuring the loss of ³H₂O. Transpiration of ³H₂O by petals was fivefold greater in the first 10 min than that found after 30 min, and the transpiration rate was shown to be associated with stomatal distribution and aperture. Thus, the stomata of outer and inner surfaces of the petal are involved in the accumulation and transpiration of water during petal opening.     

Direct Observation of Reversible and Irreversible Stomatal Responses of Attached Sunflower Leaves to SO(2)

The effects of SO₂ on stomatal aperture of attached sunflower leaves were observed with a remote-control light microscope system that permitted continuous observation of stomatal responses over periods of several hours. The relationship between actual stomatal aperture and stomatal conductance, measured with a porometer, also was examined on leaves before and after exposure to SO₂.

A distinction between uninjured and injured regions was clearly visible on leaves after exposure to 1.5 microliters per liter SO₂ for less than an hour. During the exposure, the mean value of apertures for many stomata, which indicates stomatal conductance and transpiration rate, tended to decrease simultaneously in the uninjured and injured regions. However, the rate of decrease in the injured region was slower than that in the uninjured region because of a transient opening induced by water-soaking in the injured region. The transient opening was less common in stomata near veins and veinlets.

There was a good correlation between pore width and stomatal conductance measured with a porometer before exposure to SO₂. This correlation continued in leaves exposed to SO₂ until visible, irreversible injury occurred, but then it disappeared.

The results of these experiments indicate the necessity of continuous observation of individual stomata under the microscope to understand the effects of air pollutants such as SO₂ on stomatal behavior.

     

Changes in Transpiration, Net Carbon Dioxide Assimilation and Leaf Water Potential Resulting from Application of Hydrostatic Pressure to Roots of Intact Pepper Plants

Hydrostatic pressures varying from 0 to 6.0 bar were applied to roots of intact Capsicum annuum L. cv. California Wonder plants growing in nutrient solution and the rates of transpiration, and net CO₂ assimilation, apparent compensation point and leaf water potential measured. Increasing the pressure on the roots of plants with roots in solution with either -0.5 or -5.0 bar osmotic potential with 1 bar increments resulted in a decrease in transpiration. With the application of 1 or 2 bar pressure the rate of transpiration returned to near or above the original rate. An application of 3 or 4 bar pressure reduced the rate of transpiration of all plants. The transpiration of plants with roots in solution with -0.5 bar osmotic potential remained at the reduced rate for as long as these pressures were maintained. The transpiration of plants with roots in solution with -5.0 bar was only temporarily suppressed at these pressures. Changing the applied pressure from 3 or 4 bar to 0 resulted in a rapid increase in transpiration which lasted approximately 15 minutes. This was followed by a decrease in transpiration to a rate lower than before the pressure was applied. The pattern of response was similar for plants at low or high light intensity or at normal or low CO₂ concentrations. When leaf diffusive resistance was 6.0 s cm^?1 or greater, changes in net CO₂ assimilation were similar to those of transpiration. The apparent CO₂ compensation point increased as pressure was applied and decreased with a release in pressure. Leaf water potential increased with an increase in pressure and decreased with a decrease in pressure. The changes in leaf water potential were frequently but not always proportional to changes in pressure. It is postulated that the respouses noted were due to changes in resistance to flow of water from xylem terminals through the mesophyll cells and stomatal cavities to the atmosphere.     

Signalling mechanisms involved in the response of two varieties of Humulus lupulus L. to soil drying: I. changes in xylem sap pH and the concentrations of abscisic acid and anions

Background and aims

Soil drying leads to the generation of chemical signals in plants that regulate water use via control of the stomatal aperture. The aim of our work was to identify the presence and identity of potential chemical signals, their dynamics, and their relationship with transpiration rate during soil drying in hop (Humulus lupulus (L.)) plants.

Methods

We used pressure chamber technique for measurement of shoot water potential and collection of shoot xylem sap. We analyzed concentrations of abscisic acid (ABA), nitrate, phosphate, sulphate and malate in sap and also the rate of whole plant transpiration.

Results

Transpiration rate decreased prior to changes in shoot water potential. The concentration of ABA in xylem sap continuously increased from early to later stages of water stress, whereas in leaves it increased only at later stages. Shoot sap pH increased simultaneously with the decrease of transpiration rate. Xylem sap alkalization was in some cases accompanied by a decrease in nitrate concentration and an increase in malate concentration. Concentration of sulphate increased in xylem sap during drying and sulphate in combination with a higher ABA concentration enhanced stomatal closure.

Conclusions

Several early chemical signals appear in sap of hop plants during soil drying and their impact on transpiration may vary according to the stage of soil drying.     

樟子松幼苗生长及光合特性对强风沙流吹袭的响应

为了解樟子松幼苗对不同时间强风沙流吹袭的生理生态响应,2013年春季在内蒙古科尔沁沙地研究了8级大风风沙流(风速18m·s-1,风沙流强度173g·cm-1·min-1)吹袭10、20和30min下樟子松幼苗生长与光合特性的变化。结果显示:(1)随着风吹时间的增加,樟子松的株高生长量减少、茎粗生长加快,落叶数量增加,其中30min处理与CK相比的株高生长量下降52.63%,茎粗生长量增加233.30%,落叶指数增加466.70%。(2)风沙流吹袭没有改变樟子松幼苗的净光合速率、蒸腾速率、气孔导度和胞间CO2浓度的日变化规律,但日光合峰值下降,日最大蒸腾速率增加;与CK相比,30min处理的日最大光合速率下降22.69%,日最大蒸腾速率增加11.89%。(3)随风吹时间增加,其叶片温度、叶片相对含水量、日均光合速率、水分利用效率下降,30min处理较CK依次下降0.60%、4.37%、28.57%和31.58%,且日均蒸腾速率、气孔导度和胞间CO2浓度增加,30min处理较CK依次增加6.25%、6.67%和12.60%。研究表明,在持续风沙流胁迫下,樟子松幼苗光合作用受到抑制,蒸腾耗水增加,水分利用效率降低;樟子松幼苗生长速率降低主要源于光合面积减少和光合作用受到抑制,而其光合速率下降主要因幼苗叶片叶温和叶片含水量下降所致,蒸腾速率的增加主要源于气孔导度的增加;为了适应风沙流持续吹袭的胁迫,樟子松幼苗采取了降低株高生长速率,加快茎粗生长速率的适应策略。     

Transpiration, potassium uptake and flow in tobacco as affected by nitrogen forms and nutrient levels

BACKGROUND AND AIMS: Ammonium can result in toxicity symptoms in many plants when it is supplied as the sole source of N. In this work, influences of different nitrogen forms at two levels (2 and 15 mm N) on growth, water relations and uptake and flow of potassium were studied in plants of Nicotiana tabacum 'K 326'. METHODS: Xylem sap from different leaves was collected from 106-d-old tobacco plants cultured in quartz sand by application of pressure to the root system. Whole-shoot transpiration for each of the treatments was measured on a daily basis by weight determination. KEY RESULTS: Total replacement of NO(3)(-)N by NH(4)(+)-N caused a substantial decrease in dry weight gain, even when plants grew under nutrient deficiency. Increasing nutrient concentration resulted in a greater net dry weight gain when nitrogen was supplied as NO(3)(-) or NH(4)NO(3), but resulted in little change when nitrogen was supplied as NH(4)(+). NH(4)(+)-N as the sole N-source also caused reduction in transpiration rate, changes in plant WUE (which depended on the nutrient levels) and a decrease in potassium uptake. However, the amount of xylem-transported potassium in the plants fed with NH(4)(+) was not reduced: it was 457 % or 596 % of the potassium currently taken up at low or high nutrient level, respectively, indicating a massive export from leaves and cycling of potassium in the phloem. CONCLUSIONS: Ammonium reduces leaf stomatal conductance of tobacco plants. The flow and partitioning of potassium in tobacco plants can be changed, depending on the nitrogen forms and nutrient levels.     

Ecophysiology of Salt Excretion in Aeluropus litoralis (Graminae)

Various aspects of salt excretion from leaves of Aeluropus were investigated. Salt excretion exhibited an optimum-type of curve when measured against external salt concentration, while sodium content of the leaves increased linearly. The ‘relative excretion’, i.e. rate of excreted ions: change in leaf ion content, was maximal in the low salt concentration range, and decreased when external sodium chloride concentration increased. Concentration of the excreted droplets was higher than the external concentration when the leaves were exposed to low salt concentrations in the medium, but the reverse occurred when the external salt concentrations were high. The excretion process was sensitive to water-stress conditions, caused either by high external salt concentrations or by exposure to dry atmosphere. A considerable fraction of the leaf sodium content in salt-treated leaves was only slightly available for excretion. Salt excretion in Aeluropus was enhanced by light. Such enhancement was indirect and is attributed to the increase of salt transport via transpiration stream. Selectivity of the salt-excretion mechanism is in favour of sodium and against potassium. On the other hand, potassium has a high affinity for the accumulation systems within the leaves. The ecological significance of the results is discussed.     

Optimization of potassium for proper growth and physiological response of Houttuynia cordata Thunb.

Houttuynia cordata Thunb. is an edible herb with a variety of pharmacological activities, but only limited information is available about its response towards potassium supplementation. Sterile plantlets were cultured in media with different potassium levels, and parameters related to growth, foliar potassium, water and chlorophyll contents, photosynthesis, transpiration, H₂O₂ contents and antioxidative enzyme activities were determined after a month. Results showed that 1.28 mM potassium was the optimum for H. cordata as highest values of dry weight, shoot height, root length and number were obtained at this concentration. The optimum potassium concentration resulted in the maximum net photosynthetic rate which could be associated with the highest chlorophyll content rather than limited stomatal conductance. The supply of surplus potassium resulted in higher content of foliar potassium, but negatively correlated with the biomass. Both potassium starvation (0 mM) and high potassium (>1.28 mM) could lead to water loss through high transpiration rate and low water absorption, respectively, and resulted in H₂O₂ accumulation and increased activities of catalase and peroxidase, which suggested induction of oxidative stress. Moreover, H. cordata showed the minimum of H₂O₂ content and the maximum of superoxide dismutase activity on 1.28 mM potassium, implying its role in inducing tolerance against oxidative stress.     

采煤塌陷影响下土壤含水量变化对柠条气孔导度、蒸腾与光合作用速率的影响

采煤塌陷引起的土壤环境因子的变化对矿区植物生长的影响越来越受到人们的关注,气孔导度、蒸腾与光合作用作为环境变化响应的敏感因子,研究植物气孔导度、蒸腾与光合作用的变化是揭示荒漠矿区自然环境变化及其规律的重要手段之一。研究采煤塌陷条件下植物光合生理的变化是探究煤炭开采对植物叶片水分蒸腾散失和CO_2同化速率影响的关键环节,是探讨采煤塌陷影响下植物能量与水分交换动态的基础,而采煤矿区植物叶片气孔导度、蒸腾与光合作用速率对采煤塌陷影响下土壤含水量变化的响应如何尚不清楚。选取神东煤田大柳塔矿区52302工作面为实验场地,以生态修复物种柠条为研究对象,对采煤塌陷区和对照区柠条叶片气孔导度、蒸腾和光合作用速率以及土壤体积含水量进行监测,分析了采煤塌陷条件下土壤含水量的变化以及其对柠条叶片气孔导度、蒸腾与光合作用速率的影响。结果显示:(1)煤炭井工开采在地表形成大量裂缝,破坏了土体结构,潜水位埋深降低,土壤含水量均低于沉陷初期,相对于对照区,硬梁和风沙塌陷区土壤含水量分别降低了18.61%、21.12%;(2)柠条叶片气孔导度、蒸腾和光合作用速率均与土壤含水量呈正相关关系;煤炭开采沉陷增加了地表水分散失,加剧了土壤水分胁迫程度,为了减少蒸腾导致的水分散失,柠条叶片气孔阻力增加,从而气孔导度降低,阻碍了光合作用CO_2的供应,从而导致柠条叶片光合作用速率的降低,蒸腾速率也显著降低。     

不同强度净风频繁吹袭对樟子松(Pinus sylvestris var.mongolica)幼苗光合蒸腾特征的影响

为了解不同强度净风频繁吹袭对樟子松幼苗光合蒸腾特性的影响,2013年春季在内蒙古科尔沁沙地研究了0(对照)、6、9、12、15、18 m/s等6个风速处理(分别相当于0、4、5、6、7、8级风)4次吹袭下樟子松幼苗光合速率、蒸腾速率、水分利用效率等指标的变化。结果表明,净风频繁吹袭没有改变樟子松幼苗的光合速率和蒸腾速率的日变化规律,但可使其光合蒸腾的"午休"时间加长、"休眠"程度加深;随着风吹强度的增加,其日均光合能力和蒸腾速率显著降低,其中18 m/s处理较对照分别下降27.6%和22.3%;随着风吹强度增加,气孔导度、胞间CO_2浓度均先下降后回升,除18 m/s处理胞间CO_2浓度显著高于CK外,其他处理均显著低于CK;随着风吹强度增加,水分利用效率和光能利用效率均先增加后下降,其中除18 m/s处理的水分利用效率显著低于CK,6 m/s处理的光能利用效率高于CK外,其他处理的水分利用效率均高于CK,光能利用效率均低于CK;日均光合蒸腾速率的下降主要源于气孔导度的降低,而水分利用效率和光能利用效率的变化均受制于光合速率和蒸腾速率的变化。     

Photosynthesis and Transpiration as a Function of Gaseous Diffusive Resistances in Orange Leaves

The CO₂ and H₂O vapour exchange of single attached orange, Citrus sinensis (L.), leaves was measured under laboratory conditions using infrared gas analysis. Gaseous diffusive resistances were derived from measurements at a saturating irradiance and at a leaf temperature optimum for photosynthesis. Variation in leaf resistance (within the range 1.6 to 60 s cm^-1) induced by moisture status, or by cyclic oscillations in stomatal aperture, was associated with changes in both photosynthesis and transpiration. At low leaf resistance (ri less than 10 s cm^-1) the ratio of transpiration to photosynthesis declined with reduced stomatal aperture, indicating a tighter stomatal control over H₂O vapour loss than over CO₂ assimilation. At higher leaf resistance (ri greater than 10 s cm^-1) changes in transpiration and photosynthesis were linearly related, but leaf resistance and mesophyll resistance were also positively correlated, so that strictly stomatal control of photosynthesis became more apparent than real. This evidence, combined with direct measurements of CO₂ diffusive resistances (in a -O₂ gas stream) emphasised the presence of a significant mesophyll resistance; i.e., an additional and rate limiting resistance to CO₂ assimilation over and above that encountered by H₂O vapour escaping from the leaf.     

Active chloride transport in the leaf epidermis of Commelina communis in relation to stomatal activity

Summary A chloride selective micro-electrode has been used to determine vacuolar chloride concentrations in individual cells of the leaf epidermis of Commelina communis. When the stomata were open a gradient in chloride concentration across the stomatal complex was observed with the highest concentration in the guard cells. On stomatal closure the chloride gradient was reversed. Calculation of the driving forces on chloride indicated that active transport of chloride was occurring during both stomatal opening and closure. This transport appeared to be energetically independent of the transport of potassium. These results are discussed in relation to the behaviour of other anions during stomatal movements.     

Dynamics of stomatal water relations during the humidity response: implications of two hypothetical mechanisms

The feasibility of two hypothetical mechanisms for the stomatal response to humidity was evaluated by identifying theoretical constraints on these mechanisms and by analysing timecourses of stomatal aperture following a step change in humidity. The two hypothetical mechanisms, which allow guard cell turgor pressure to overcome the epidermal mechanical advantage, are: (1) active regulation of guard cell osmotic pressure, requiring no hydraulic disequilibrium between guard and epidermal cells, and (2) a substantial hydraulic resistance between guard and epidermal cells, resulting in hydraulic disequilibrium between them. Numerical simulations of the system are made possible by recently published empirical relationships between guard cell pressure and volume and between stomatal aperture, guard cell turgor pressure, and epidermal cell turgor pressure; these data allow the hypothetical control variables to be inferred from stomatal aperture and evaporative demand, given physical assumptions that characterize either hypothesis. We show that hypothesis (1) predicts that steady‐state π_g is monotonically related to transpiration rate, whereas hypothesis (2) suggests that the relationship between transpiration rate and the steady‐state guard to epidermal cell hydraulic resistance may be either positive or negative, and that this resistance must change substantially during the transient phase of the stomatal response to humidity.     

Changes in the Water Balance and Photosynthesis of Onion, Bean and Cotton Plants under Saline Conditions

The osmotic concentration (osmotic potential) of onion leaf sap did not adjust to chloride salinity, and consequently water potential, turgor, stomatal aperture and transpiration were reduced. Although osmotic concentration of bean and cotton leaf sap did adjust to a saline root medium and turgor was no less in the salinized plants than in the controls, stomata of the salinized plants remained only partly open and transpiration was reduced. Net photosynthesis of onion plants was reduced by salinity (this effect being much enhanced in a hot dry atmosphere) but it could be rapidly raised to the level of the controls by inducing elevated leaf turgor. Stomatal closure was initially responsible for most of the ～30 % reduction in photosynthesis of salinized beans. This was due to interference with CO₂ diffusion and could be overcome by raising the CO₂ concentration in the air. At a later stage of growth, salinity affected the light reaction of bean photosynthesis, and elevation of the air CO₂ had little effect. Closure of stomata of salinized cotton plants had only a relatively small effect on net photosynthesis. Light intensity and CO₂ concentration experiments showed that salinity was reducing the photosynthesis of cotton leaves mainly by affecting the light reaction of photosynthesis. It is concluded that chloride salinity does affect the water balance and rate of photosynthesis of plants and that the nature and degree of the effects will depend upon climatic conditions and may be very different between plant species and in the same species at different periods of growth.     

Transpiration of spring barley under nitrogen and phosphorus deficiency during leaf wilting

In an open gas exchange system with a thermocouple psychrometer the transpiration rate of the first leaf in 8-day plants of spring barley was measured in dependence on the water saturation deficit (ΔW _sat). The plants were cultivated in Richter’s nutrient solution, either complete, or deficient in nitrogen or phosphorus. The cuticular transpiration (as measured in the dark) was unaffected by N and P deficiency. The N deficiency reduced the transpiration rate by increasing stomatal resistance since full water saturation of the leaf (67% rate of the control variant) up to stomatal closing at Δ W_sat = 14%. The P deficiency does not affect the transpiration rate at initial phases of wilting, but the stomata close only at a higher Δ W_sat (25%) than those in the control.     

盐胁迫对沙拐枣光合生理特性的影响

以采自甘肃民勤一年生的沙拐枣幼苗为试材,对不同NaCl浓度(0、50、100、200、300mmol·L~(-1))处理下沙拐枣光合生理特性进行分析,并对各生理指标与地上生物量进行灰色关联度分析,以探讨荒漠植物沙拐枣的抗盐机理,为沙拐枣的保护及其恢复荒漠生态系统稳定提供理论依据。结果显示:随着NaCl浓度的升高,沙拐枣同化枝内脯氨酸含量逐渐增大,而其可溶性糖含量逐渐减小;在低浓度NaCl(50mmol·L~(-1) NaCl)处理下,同化枝光合参数均增加,且净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)均达到最大值,比对照分别显著增加了33.3%、68.0%、60.8%;与50mmol·L~(-1) NaCl相比,处理浓度超过50mmol·L~(-1) NaCl时,Pn、Gs、Tr均降低;同化枝叶绿素b含量随着NaCl浓度的增加而降低,而叶绿素a和总叶绿素含量均呈先增加后降低的趋势。灰色关联度分析发现,同化枝的Tr、Gs、Ci以及叶绿素b与地上生物量的关联度较大。研究表明,低盐浓度NaCl激活了沙拐枣的某些生理机制,有利于植株的光合作用和生长,而植物在高盐浓度胁迫时能通过调节脯氨酸和可溶性糖的含量,减少叶绿素含量、Pn和Tr等维持自身的生长。