Effects of drought stress on photosynthetic gas exchange, chlorophyll fluorescence and stem diameter of soybean plants

Changes in plant growth, photosynthetic gas exchange, chlorophyll fluorescence and stem diameter of soybean [Glycine max (L.) Merr.] plants under drought stress were studied. Total plant dry mass was reduced by 30 % compared to well-watered control plants. Leaf water potential was slightly decreased by water stress. Water stress induced daytime shrinkage and reduced night-time expansion of stem. Photosynthetic rate, stomatal conductance and transpiration rate were significantly declined by water stress, while the intercellular CO₂ concentration was changed only slightly at the initiation of stress treatment. The maximum photochemical efficiency of photosystem 2 and apparent photosynthetic electron transport rate were not changed by water 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。     

Diurnal courses and factorial dependencies of leaf conductance and transpiration of differently potassium fertilized and watered field grown barley plants

During the grain filling period we followed diurnal courses in leaf water potential (ψ₁), leaf osmotic potential (ψ_π), transpiration (E), leaf conductance to water vapour transfer (g) and microclimatic parameters in field-grown spring barley (Hordeum distichum L. cv. Gunnar). The barley crop was grown on a coarse textured sandy soil at low (50 kg ha⁻¹) or high (200 kg ha⁻¹) levels of potassium applied as KCl. The investigation was undertaken at full irrigation or under drought. Drought was imposed at the beginning of the grain filling period. Leaf conductance and rate of transpiration were higher in the flag leaf than in the leaves of lower insertion. The rate of transpiration of the awns on a dry weight basis was of similar magnitude to that of the flag leaves. On clear days the rate of transpiration of fully watered barley plants was at a high level during most part of the day. The transpiration only decreased at low light intensities. The rate of transpiration was high despite leaf water potentials falling to rather low values due to high evaporative demands. In water stressed plants transpiration decreased and midday depression of transpiration occurred. Normally, daily accumulated transpirational water loss was lower in high K leaves than in low K leaves and generally the bulk water relations of the leaves were more favourable in high K plants than in low K plants. The factorial dependency of the flag leaf conductances on leaf water potential, light intensity, leaf temperature, and leaf-to-air water vapour concentration difference (ΔW) was analysed from a set of field data. From these data, similar sets of microclimatic conditions were classified, and dependencies of leaf conductance on the various environmental parameters were ascertained. The resulting mathematical functions were combined in an empirical simulation model. The results of the model were tested against other sets of measured data. Deviations between measured and predicted leaf conductance occurred at low light intensities. In the flag leaf, water potentials below-1.6 MPa reduced the stomatal apertures and determined the upper limit of leaf conductance. In leaves of lower insertion level conductances were reduced already at higher leaf water potentials. Leaf conductance was increased hyperbolically as photosynthetic active radiation (PAR) increased from darkness to full light. Leaf conductance as a function of leaf temperature followed an optimum curve which in the model was replaced by two linear regression lines intersecting at the optimum temperature of 23.4°C. Increasing leaf-to-air water vapour concentration difference caused a linear decrease in leaf conductance. Leaf conductances became slightly more reduced by lowered water potentials in the low K plants. Stomatal closure in response to a temperature change away from the optimum was more sensitive in high K plants, and also the decrease in leaf conductance under the influence of lowered ambient humidity proceeded with a higher sensitivity in high K plants. Thus, under conditions which favoured high conductances increase of evaporative demand caused an about 10% larger decrease in leaf conductance in the high K plants than in the low K plants. Stomatal sizes and density in the flag leaves differed between low and high K plants. In plants with partially open stomata, leaf conductance, calculated from stomatal pore dimensions, was up to 10% lower in the high K plants than in the low K plants. A similar reduction in leaf conductance in high K plants was measured porometrically. It was concluded that the beneficial effect of K supply on water use efficiency reported in former studies primarily resulted from altered stomatal sizes and densities.     

Cadmium-Induced Changes in Leaf Epidermes, Photosynthetic Rate and Pigment Concentrations in Cajanus Cajan

Application of different concentrations of cadmium [5, 10, 15, 25 and 50 g(CdCl₂) g^–1(soil d.m.)] markedly affected leaves of Cajanus cajan (Linn.) Huth. Due to increased Cd content in leaves, stomatal density and size on abaxial epidermis, and the size of stomatal aperture and length and density of trichomes on both leaf epidermes decreased significantly in the treated plants. Net photosynthetic rate and stomatal conductance were reduced significantly at each concentration of cadmium, whereas reduction in intercellular carbon dioxide concentration was significant at 10 g Cd onwards. The contents of chlorophyll a, chlorophyll b and carotenoids were relatively low during early stages of plant development under the effect of Cd. Nitrate content, nitrate reductase activity and protein content were also lower in treated plants, compared with control.     

Effects of zinc and soil moisture on photosynthetic rate and chlorophyll fluorescence parameters of maize

Effects of zinc [0 and 5.0 mg Zn kg⁻¹ (soil)] on photosynthetic rate (P_N), and chlorophyll fluorescence in leaves of maize (Zea mays L.) cv. Zhongdan 9409 seedlings grown under different soil moisture regimes (40–45 % and 70–75 % of soil saturated water content) were studied. Zn application did not enhance maize plant adaptation to drought stress. The relative water content and the water potential of leaves were not affected by Zn treatment. Moreover, The P_N of drought-stressed plants was not improved by Zn supply. The increases of plant biomass, stomatal conductance and quantum yield of photosystem 2 due to Zn addition were notable in well-watered plants.     

Stomatal function,density and pattern,and CO2 assimilation in Arabidopsis thaliana tmm1 and sdd1‐1 mutants

• Stomata modulate the exchange of water and CO₂ between plant and atmosphere. Although stomatal density is known to affect CO₂ diffusion into the leaf and thus photosynthetic rate, the effect of stomatal density and patterning on CO₂ assimilation is not fully understood.
• We used wild types Col‐0 and C24 and stomatal mutants sdd1‐1 and tmm1 of Arabidopsis thaliana, differing in stomatal density and pattern, to study the effects of these variations on both stomatal and mesophyll conductance and CO₂ assimilation rate. Anatomical parameters of stomata, leaf temperature and carbon isotope discrimination were also assessed.
• Our results indicate that increased stomatal density enhanced stomatal conductance in sdd1‐1 plants, with no effect on photosynthesis, due to both unchanged photosynthetic capacity and decreased mesophyll conductance. Clustering (abnormal patterning formed by clusters of two or more stomata) and a highly unequal distribution of stomata between the adaxial and abaxial leaf sides in tmm1 mutants also had no effect on photosynthesis.
• Except at very high stomatal densities, stomatal conductance and water loss were proportional to stomatal density. Stomatal formation in clusters reduced stomatal dynamics and their operational range as well as the efficiency of CO₂ transport.

     

Effects of water stress on the chlorophyll content,nitrogen level and photosynthesis of leaves of two maize genotypes

The dynamics of leaf chlorophyll level, nitrogen content, photosynthesis and stomatal conductance were followed in detail in two cultivars of maize (Zea mays) during a short period of water stress, applied at tasseling, and during the subsequent recovery phase. Plants used in the experiment were grown in sand-nutrient solution culture under field weather conditions. Water stress reduced chlorophyll levels, stomatal conductance and photosynthesis, but the nitrogen content of the leaves was not affected. It is concluded that the stress-induced loss of chlorophyll is not mediated by a lack of nitrogen. Considerable differences were observed between genotypes in the rate of post-stress recovery of chlorophyll level. Recovery, upon rewatering, of stomatal conductance and photosynthesis preceded that of chlorophyll level. Losses of up to 40% of leaf chlorophyll content were insufficient to affect rates of photosynthesis measured at mid-day.     

水分亏缺对蚕豆光合特性影响的初步研究

本文报道了蚕豆现蕾至饱荚期不同时间土壤水分亏缺情况下的光合特性、光合产量及蚕豆水分亏缺敏感期。蚕豆现蕾后给予土壤干旱处理,光合速率、叶绿素含量、叶面积、气孔开度、生物产量及籽粒产量下降,但气孔密度和呼吸速率增加。水分亏缺使叶片光饱和点由50kLx降至30kLx,气孔开度日变化呈单峰(9—11时)曲线。始荚至盛荚期对土壤干旱最敏感,此期是蚕豆灌水的关键时期。     

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.     

The effect of take-all disease on gas-exchange rates and biomass in two winter wheat lines with different drought response

There have been no studies of the effect of take-all on leaf gas-exchange rates, despite the fact that take-all severely restricts plant water and nutrient uptake, which results in significant biomass and grain yield reduction. Here we describe the effect of inoculation with Gaeumannomyces graminis (Sacc.) var. tritici (Ggt) on carbon assimilation rate (A) and biomass production of wheat plants grown under two water regimes. We show that the impact of Ggt inoculation on plant growth and leaf A may be through reduced photosynthetic capacity of the leaves and not water stress per se. The nature of this reduced photosynthetic capacity remains uncertain but may involve nutrient deficiency and different enzymes produced by the fungus. In each of the 3 years the experiment was conducted, Ggt significantly reduced A, i.e. at anthesis by 18% in 2000, 15% in 2001, and 12% in 2002. In agreement with other field studies, Ggt reduced tiller number and production of all plant components, mostly root dry mass and grain mass per plant. Highly significant negative correlations were found between disease rating and A in all years, showing that at disease ratings equal or higher than 3 (on a scale from 1 to 4) A could practically be zero. While A decreased, intercellular CO₂ concentration increased or did not change, and stomatal conductance was relatively high. In addition, A was more reduced under high than under low soil moisture content. These results support the idea that water stress per se did not contribute to the observed reduction of A. The mechanism of photosynthetic capacity reduction due to the Ggt root-rotting fungus is of interest as it may lead to the molecular mechanisms of plant resistance and ultimately to the development of take-all resistant plants.     

Growth and Gas Exchange of Three Sorghum Cultivars Under Drought Stress

A field study was conducted to evaluate the drought tolerance of three sorghum [Sorghum bicolor (L.) Moench] cultivars, Gadambalia, Arous elRimal and Tabat, and quantify the physiological bases for differences in their drought tolerance. Water stress reduced shoot dry mass of Gadambalia, Arous elRimal and Tabat by 43, 46 and 58 %, respectively. The respective reduction in leaf area of the three cultivars was 28, 54 and 63 %. The reduction in net photosynthetic rate, stomatal conductance and transpiration rate due to water stress was lowest in Gadambalia and highest in Tabat. The leaf water potentials and relative water contents of Gadambalia under wet and dry treatments were similar, while those of Tabat were significantly reduced by water stress. The lowest and highest liquid water flow conductance was displayed by Tabat and Gadambalia, respectively. Drought tolerance in Gadambalia is associated with its smaller leaf area, higher liquid water flow conductance, and ability to maintain high leaf water potential, relative water content, stomatal conductance, transpiration rate and photosynthetic rate under drought stress. This revised version was published online in July 2006 with corrections to the Cover Date.     

Stomatal and photosynthetic responses to shade in sorghum, soybean and eastern gamagrass

We studied photosynthetic and stomatal responses of grain sorghum ( Sorghum bicolor [L.] Moench cv. Pioneer 8500), soybean ( Glycine max L. cv. Flyer) and eastern gamagrass ( Tripsacum dactyloides L.) during experimental sun and shade periods simulating summer cloud cover. Leaf gas exchange measurements of field plants showed that short-term (5 min) shading of leaves to 300–400 μmol m⁻² s⁻¹ photosynthetic photon flux density reduced photosynthesis, leaf temperature, stomatal conductance, transpiration and water use efficiency and increased intercellular CO₂ partial pressure. In all species, photosynthetic recovery was delayed when leaves were reilluminated, apparently by stomatal closure. The strongest stomatal response was in soybean. Photosynthetic recovery was studied further with soybeans grown indoors (maximum photosynthetic photon flux density 1 200 μmol m⁻² s⁻¹). Plants grown indoors had responses to shade similar to those of field plants, except for brief nonstomatal limitation immediately after reillumination. These responses indicated the importance of the light environment during leaf development on assimilation responses to variable light, and suggested different limitations on carbon assimilation in different parts of the soybean canopy. Photosynthetic oxygen evolution recovered immediately upon reillumination, indicating that the light reactions did not limit soybean photosynthetic recovery. While shade periods caused stomatal closure and reduced carbon gain and water loss in all species, the consequences for carbon gain/water loss were greatest in soybean. The occurrence of stomatal closure in all three species may arise from their shared phenologies and herbaceous growth forms.     

Effect of leaf surface wetness and wettability on photosynthesis in bean and pea

Leaf surface wetness that occurs frequently in natural environments has a significant impact on leaf photosynthesis. However, the physiological mechanisms for the photosynthetic responses to wetness are not well understood. The responses of leaf CO₂ assimilation rate (A) to 72 h of artificial mist of a wettable (bean; Phaseolus vulgaris) and a non‐wettable species (pea; Pisum sativum) were compared. Stomatal and non‐stomatal limitations to A were investigated. A 28% inhibition of A was observed in the bean leaves as a result of a 16% decrease in stomatal conductance and a 55% reduction in the amount of Rubisco. The decrease of Rubisco was mainly due to its partial degradation. In contrast to the bean leaves, a 22% stimulation of A was obtained in the 72 h mist‐treated pea leaves. Mist treatment increased stomatal conductance by 12.5% and had no effect on the amount of Rubisco. These results indicated that a positive photosynthetic response to wetness occurred only in non‐wettable species and is due to the change in stomatal regulation.     

番茄叶片光合作用对快速水分胁迫的响应

采用聚乙二醇(PEG 6000)溶液控制番茄根际水势和叶片离体的方式设置了水分胁迫处理,测算了光合诱导过程中净光合速率、暗呼吸速率和CO₂补偿点等光合参数的变化.结果表明： 在1000 μmol·m^-2·s^-1光诱导下,水分胁迫处理的番茄叶片净光合速率(P_n)达到最大值所需时间缩短为对照的1/3,气孔导度(g_s)快速增大为对照的1.5倍.水分胁迫处理的番茄叶片光饱和点(LSP)比对照降低了65%～85%,而光补偿点(LCP)比对照增加了75%～100%,缩小了番茄叶片利用光能的有效范围.水分胁迫处理的番茄叶片最大光合能力(A_max)低于对照40%以上,暗呼吸速率(R_d)增大了约45%.可见,快速水分胁迫处理使番茄叶片气孔迅速开放,光合诱导初始阶段消失.水分胁迫导致植物利用光能的效率和潜力降低是植物生产力下降的重要原因,而气孔调节是番茄适应快速水分胁迫的重要生理机制.     

Co-ordination of hydraulic and stomatal conductances across light qualities in cucumber leaves

Long-term effects of light quality on leaf hydraulic conductance (K(leaf)) and stomatal conductance (g(s)) were studied in cucumber, and their joint impact on leaf photosynthesis in response to osmotic-induced water stress was assessed. Plants were grown under low intensity monochromatic red (R, 640 nm), blue (B, 420 nm) or combined red and blue (R:B, 70:30) light. K(leaf) and g(s) were much lower in leaves that developed without blue light. Differences in g(s) were caused by differences in stomatal aperture and stomatal density, of which the latter was largely due to differences in epidermal cell size and hardly due to stomatal development. Net photosynthesis (A(N)) was lowest in R-, intermediate in B-, and highest in RB- grown leaves. The low A(N) in R-grown leaves correlated with a low leaf internal CO(2) concentration and reduced PSII operating efficiency. In response to osmotic stress, all leaves showed similar degrees of stomatal closure, but the reduction in A(N) was larger in R- than in B- and RB-grown leaves. This was probably due to damage of the photosynthetic apparatus, which only occurred in R-grown leaves. The present study shows the co-ordination of K(leaf) and g(s) across different light qualities, while the presence of blue in the light spectrum seems to drive both K(leaf) and g(s) towards high, sun-type leaf values, as was previously reported for maximal photosynthetic capacity and leaf morphology. The present results suggest the involvement of blue light receptors in the usually harmonized development of leaf characteristics related to water relations and photosynthesis under different light environments.     

Stomatal and photosynthetic responses ofCichorium intybus leaves to sulfur dioxide treatment at different stages of plant development

Fifty-day-oldCichorium intybus Linn, plants were exposed to 1 ppm sulfur dioxide gas, 2 h per day for 7 consecutive days. Their leaves as well as those from the control plants were sampled at pre-flowering, flowering, and post-flowering stages to study their morphological, physiological, and biochemical responses to SO₂ stress. The number, dimensions, area, and biomass of leaves were less in the treated plants. Length and width of stomatal apertures on both epidermises were greater for leaves exposed to SO₂. The Stomata were longer on the adaxial epidermis, but shorter on the abaxial epidermis, except at the pre-flowering stage. Stomatal widths varied widely. Compared with the controls, the abaxial epidermis on treated leaves showed consistently lower stomatal densities as well as stomatal indices. This was also true for the adaxial epidermis during the post-flowering stage. The photosynthetic rate and stomatal conductance were reduced in the SO₂-exposed plants, but intercellular CO₂ concentrations increased at the pre-flowering stage and, subsequently, declined. Chlorophyll a, carotenoid, and total chlorophyll contents increased at the pre-flowering stage, and then decreased. The level of chlorophyllb was reduced throughout plant development compared with the untreated controls.     

Effects of water stress and high nocturnal temperature on photosynthesis and nitrogen level of a perennial grass Leymus chinensis

Water deficit and high temperature are important environmental factors restricting plant growth and photosynthesis. The two stresses often occur simultaneously, but their interactions on photosynthesis and nitrogen level have been less studied. In the present experiment, we measured photosynthetic parameters, stomatal density, and nitrogen levels, as well as soluble sugar content of leaves of a perennial grass, Leymus chinensis, experiencing two day/night temperature regimes of 30/20 °C and 30/25 °C, and five different soil moisture contents (the soil relative-water content ranged from 80% to 25%). Leaf relative water content, leaf biomass, whole plant biomass, the ratio between the leaf biomass and total plant biomass, and the photosynthetic rate, as well as water-use efficiency decreased at high night temperature, especially under severe water stress conditions. Stomatal index was also increased by soil water stress except very severe water stress, and high nocturnal temperature decreased the leaf stomatal index under soil water stress. Nocturnal warming decreased nitrogen concentration in the leaves and increased it in the roots, particularly when plants were subjected to severe water stress. There were significant positive correlations between the photosynthetic rate and both soluble sugar concentration and nitrogen concentration at low nocturnal temperature. It is suggested that nocturnal warming significantly exacerbates the adverse effects of soil water stress, and their synergistic interactions might reduce the plant productivity and constrain its distribution in the region dominated by L. chinensis, based on predictions of global climate change.     

Responses of Rice Cultivars to the Elevated CO2

The effect of CO₂ concentration elevated to 575 – 620 µmol mol^–1 on growth, tillering, grain yield, net photosynthetic rate, dark respiration rate, stomatal conductance, sugar content and protein profile of two rice (Oryza sativa L.) cultivars Pusa Basmati-1 and Pusa-677 at flowering stage was studied using open top chambers. The cultivar Pusa Basmati-1 responded more markedly for most of the growth and physiological parameters compared to Pusa-677. The increase in grain yield in Pusa Basmati-1 attributed largely to increased grain number. The increased net photosynthetic rate and greater accumulation of sugar contributed significantly to the accelerated development of leaves and tillers in both the cultivars. The reduction in the low molecular mass proteins including Rubisco and increase in high molecular mass photosystem 2 proteins was observed in both the cultivars. Additional sugars may possibly help in balancing the profile of photosynthetic proteins and sustain greater growth and productivity in rice cultivars.     

Effects of root flooding and stage of development on the growth and photosynthesis of field bean (Vicia faba L. minor)

Field bean plants were subjected to flooding stress for 7 days, during two stages of development: at the vegetative phase (4-week-old seedlings) and at the generative phase (8-week-old plants). The height of plants, total area of leaves, the number of undamaged leaves, dry plant matter, chlorophyll content, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) activity, the maximum quantum yield of PS2 photochemistry (Fv/Fm ratio), the photosynthesis rate (P _N) and stomatal conductance (g _s) were determined. A strong reduction in stem elongation and leaf area as well as in dry matter production was observed as a result of flooding. The responses from vegetative plants were greater than in generative plants. Waterlogging decreased chlorophyll a and b in leaves, notably at the vegetative stage, and persisted after cessation of flooding. After flooding, photosynthesis was strongly reduced and positively correlated with decreased stomatal conductance. Damage to the photosynthetic apparatus resulted in a lower Fv/Fm especially in young seedlings. In vegetative plants Fv/Fm quickly returned to the control levels after the soil was drained. The results show that an excess of water in the soil limits growth and injures the photosynthetic apparatus in field beans, but that the extent of the injury is strongly age dependent.     

高粱、紫苏叶脉密度与光合特性的关系

叶脉是植物叶片光合作用水分输送的重要结构。为阐述叶脉与光合特性之间的关系,以C4植物高粱(Sorghum bicolor)、C3植物紫苏(Perilla frutescens)为实验材料研究了叶脉密度和光合特性之间的关系。结果表明,与紫苏相比,高粱叶片叶脉密度大,导水能力强,蒸腾速率高,但气孔密度小。进一步分析表明,高粱叶片近轴侧气孔密度占总气孔的比例明显高于紫苏。叶脉密度大的高粱具有较高的净光合速率;而紫苏叶脉密度小,净光合速率也较低。由此表明,较高的叶脉密度有利于支持较高的光合速率,但研究表明叶脉密度和气孔密度可能不存在严格的协同变异关系。研究结果对理解植物光合作用适应有重要意义。