The Relation between Photosynthesis and Stomatal Resistance of Each Leaf Surface in Cotton Leaves

The relation between photosynthesis and stomatal resistance of each leaf surface in cotton leaves (Gossypium hirsutum) was studied during ageing and with increase in light intensity. During ageing of leaves the stomatal resistance of the upper surface increases before that of the lower surface. This observation would suggest that the stomata of the upper leaf surface commence to age before those of the lower leaf surface. The earlier commencement of the increase in upper stomatal resistance results in the decline in photosynthesis at an early stage of ageing being negatively correlated with the upper stomatal resistance and not related to the lower stomatal resistance. The results indicate that in this initial phase of ageing the decrease in photosynthesis and increase in stomatal resistance of the upper surface are not causally related but occur simultaneously. At a more advanced stage of ageing the lower stomatal resistance also increases and then the decline in photosynthesis becomes related to the increase in stomatal resistance of both leaf surfaces. When illumination of leaves is increased stepwise, the resultant increase in photosynthesis is related to the reduction in stomatal resistance of each leaf surface. At low light intensities the stomatal resistance of the lower surface is low while that of the upper surface is relatively high. The difference between them decreases with increase in light intensity. This would indicate that the upper stomata require a higher light intensity to open than the lower stomata. The stormatal resistance of the two leaf surfaces is not related to the stomatal frequencey.     

The effect of light intensity on stomatal frequency in leaves ofIris hollandica hort., var. wedgwood

Stomatal frequency in leaves of plants of Iris hollandica, cultivated under four light intensities, about 12%, 37%, 75% and 100% of natural daylight, was studied. The stomatal frequency decreases with lower light intensities. The gradients of stomatal frequency of successive leaves show an increase up to the third or fourth leaf with a subsequent decrease of values up to the highest leaf. The total number of stomata per leaf is not constant; differences exist in successive leaves of a shoot as well as among the leaves of plants growing under various light intensities. The character of gradients was changed at lower light intensities. The largest reaction to the light intensity appeared in leaves with higher stomatal frequency. The gradients in individual leaves depend on the insertion of the leaf. Both the shape and the slope of these gradients are influenced by the light intensity.     

Stomatal development in new leaves is related to the stomatal conductance of mature leaves in poplar (Populus trichocarpaxP. deltoides)

In general, stomatal density (SD) decreases when plants are grown at high CO2 concentrations. Recent studies suggest that signals produced from mature leaves regulate the SD of expanding leaves. To determine the underlying driver of these signals in poplar (Populus trichocarpaxP. deltoides) saplings, a cuvette system was used whereby the environment around mature (lower) leaves could be controlled independently of that around developing (upper) leaves. A series of experiments were performed in which the CO2 concentration, vapour pressure deficit (D), and irradiance (Q) around the lower leaves were varied while the (ambient) conditions around the upper leaves were unchanged. The overall objective was to break the nexus between leaf stomatal conductance and transpiration and photosynthesis rates of lower leaves and determine which, if any, of these parameters regulate stomatal development in the upper expanding leaves. SD, stomatal index (SI), and epidermal cell density (ED) were measured on the adaxial and abaxial surfaces of fully expanded upper leaves. SD and SI decreased with increasing lower leaf CO2 concentration (150-780 ppm) at both ambient (1.3-1.6 kPa) and low (0.7-1.0 kPa) D. SD and SI at low D were generally higher than at ambient D. By contrast, ED was relatively insensitive to both vapour pressure and CO2 concentration. When lower leaves were shaded, upper leaf SD, SI, and ED decreased but did not change with varying CO2 concentration. These results suggest that epidermal cell development and stomatal development are regulated by different physiological mechanisms. SI of the upper leaves was positively and highly correlated (r2>0.84) with the stomatal conductance of the lower leaves independent of their net photosynthesis and transpiration rates, suggesting that the stomatal conductance of mature leaves has a regulatory effect on the stomatal development of expanding leaves.     

Stomatal numbers of soybean and response to water stress

The relationship among stomatal density, photosynthetic rate, leaf conductance, plant growth, bean yield and kaempferol triglucoside (K9) in the leaves of soybean (Glycine max (L.) Merr.) was examined in two field tests. K9 in the leaves was associated with reduced stomatal density, reduced photosynthetic rate, reduced stomatal conductance, reduced plant weight and lower bean yield. Plants with high stomatal frequency (lacking K9) were better able to take advantage of increased water supply by increasing stomatal conductance (upper surface), transpiration and bean yield. Plants with low stomatal frequency (with K9) were unresponsive to irrigation and in this sense were more tolerant of water stress, but their overall yield was low.     

青藏高原不同海拔3种菊科植物叶片结构变化及其生态适应性

采用常规石蜡制片技术和显微观察方法,对分布于青藏高原祁连山东部山地冷龙岭3600—4400 m不同海拔高度的蒲公英(Taraxacum mongolicum Hand-Mazz)、火绒草(Leontopodium leontopodioides(Willd.)Beauv)和美丽风毛菊(Saussurea superba Anthony)的叶片形态解剖结构进行了研究,并探讨了其对海拔高度的响应及生态适应性。结果表明:随海拔高度的升高,3种植物气孔器外拱盖内缘、角质层纹饰、气孔与表皮细胞的位置关系以及上、下表皮气孔器内缘呈现不同的变化趋势;火绒草上、下表皮气孔密度随海拔升高而增加,而蒲公英和美丽风毛菊的气孔密度则降低;3种植物上、下表皮气孔指数随海拔高度的增加均未出现规律性变化;3种植物叶片厚度、上下表皮厚度、上下角质层厚度、栅栏细胞系数均随海拔升高而增加;解剖学指标之间大多呈明显的协同进化;叶片结构,尤其是气孔密度对海拔高度变化表现出较大的可塑性。研究表明3种植物采取不同的响应机制来适应海拔高度的变化,植物对高原环境变化的适应具有多样性。     

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.     

冠层部位对金叶女贞叶片结构的影响

金叶女贞是一种在园林绿化上广泛应用的彩叶植物,上层叶金黄色,下层叶深绿色。本文利用常规植物制片技术,研究了叶绿素缺少的金叶女贞上层金叶和下层绿叶的解剖结构。结果表明:(1)金叶女贞上层金叶和下层绿叶的上表皮无显著差异,但是下层绿叶的下表皮细胞较小,气孔密度较低;(2)下层绿叶的叶肉组织较薄,细胞较大,排列疏松;(3)在下层绿叶中叶绿体类囊体片层排列密集,叶肉组织内单位体积叶绿体数量较少;(4)冠层部位对光合色素的积累有明显影响。叶绿素缺少的金叶女贞叶片结构显著受到冠层部位的影响,光合色素的变化幅度较一般植物更大,不同冠层部位叶片叶色有明显差异,可以作为研究植物冠层部位对叶形态、结构和生理生化影响的好材料。     

Stomatal activity within the crowns of tall deciduous trees under forest conditions

The variation in stomatal activity within the crowns ofAcer campestre, Carpinus betulus andQuercus cerris was measured by vapour exchange porometer on several summer days in an oak-hornbeam forest, in SW Slovakia, Czechoslovakia. Variation resulted from crown position in the forest stand and from leaf position within the canopy. The highest stomatal conductance was in sunlit sun leaves in the upper part of the canopy. Stomatal conductance decreased with increasing depth in the canopy. The steepest decrease was in the upper canopy, in the intermediate zone between fully sunlit and fully shaded leaves, and was caused by the decline in leaf irradiance and in stomatal density. In codominant trees, the conductance in shade leaves at the base of the crown was significantly lower than in the sun leaves at the top of the crown. In a dominant tree,Q. cerris, the differences in stomatal conductance were small and most frequently insignificant. Variation in incident light also determined the diurnal variation of stomatal conductance with respect to crown aspect. Differences between sun leaves on the east and west facing aspects of the overstory crown ofQ. cerris were demonstrated for several days.     

不同R∶FR值对菊花叶片气孔特征和气孔导度的影响

以切花菊品种"神马(Jinba)"为试材,2010年10月至2011年2月间在南京信息工程大学试验温室采用不同Red (660±10) nm: Far-red (730±10) nm值的LED光源短日处理,研究了温室切花菊叶片气孔特征和气孔导度对不同R∶FR值的响应。结果表明:不同R∶FR值短日处理35d菊花叶片的上表皮和下表皮的气孔直径分别以R∶FR值4.5和6.5处理最大,均以R∶FR值2.5处理最小,气孔密度和气孔开度均以R∶FR=2.5处理最高,以R∶FR值6.5处理最低,下表皮的气孔密度、气孔开度明显高于上表皮;不同R∶FR值处理叶片的气孔开张比和气孔指数差异不显著;在相同光强下,菊花叶片气孔导度和光合速率由大到小的R∶FR值顺序依次为2.5、4.5、0.5、6.5。叶片气孔导度与气孔指数、气孔密度、气孔开张比和气孔开度成正相关,与气孔长度和气孔宽度呈负相关;R∶FR值在2.5-6.5范围内,随光质中红光成分增加,叶片气孔密度、气孔指数、气孔开度、气孔开张比和气孔导度显著降低。     

Photocontrol of the Functional Coupling between Photosynthesis and Stomatal Conductance in the Intact Leaf : Blue Light and Par-Dependent Photosystems in Guard Cells

The photocontrol of the functional coupling between photosynthesis and stomatal conductance in the leaf was investigated in gas exchange experiments using monochromatic light provided by lasers. Net photosynthesis and stomatal conductance were measured in attached leaves of Malva parviflora L. as a function of photon irradiance at 457.9 and 640.0 nanometers.

Photosynthetic rates and quantum yields of photosynthesis were higher under red light than under blue, on an absorbed or incident basis.

Stomatal conductance was higher under blue than under red light at all intensities. Based on a calculated apparent photon efficiency of conductance, blue and red light had similar effects on conductance at intensities higher than 0.02 millimoles per square meter per second, but blue light was several-fold more efficient at very low photon irradiances. Red light had no effect on conductance at photon irradiances below 0.02 millimoles per square meter per second. These observations support the hypothesis that stomatal conductance is modulated by two photosystems: a blue light-dependent one, driving stomatal opening at low light intensities and a photosynthetically active radiation (PAR)-dependent one operating at higher irradiances.

When low intensity blue light was used to illuminate a leaf already irradiated with high intensity, 640 nanometers light, the leaf exhibited substantial increases in stomatal conductance. Net photosynthesis changed only slightly. Additional far-red light increased net photosynthesis without affecting stomatal conductance. These observations indicate that under conditions where the PAR-dependent system is driven by high intensity red light, the blue light-dependent system has an additive effect on stomatal conductance.

The wavelength dependence of photosynthesis and stomatal conductance demonstrates that these processes are not obligatorily coupled and can be controlled by light, independent of prevailing levels of intercellular CO₂. The blue light-dependent system in the guard cells may function as a specific light sensor while the PAR-dependent system supplies a CO₂-modulated energy source providing functional coupling between the guard cells and the photosynthesizing mesophyll.

     

Leaf photosynthetic parameters of two maize (Zea mays) cultivars in response to various patterns of chilling temperatures and photon flux densities

Chilling‐induced photosynthetic impairment was examined in leaves of maize (Zea mays L.) seedlings of two cultivars, one adapted to western Europe and one adapted to Mexican highlands. Three experiments were performed in a controlled environment. The effects of chilling night temperatures, of chilling at high light intensity and of variable chilling day temperatures on photosynthetic parameters, were evaluated. Chilling in the dark period resulted in stomatal limitation of net photosynthesis. Chilling at moderate to high light intensities caused chilling‐dependent photoinhibition of CO₂ uptake. Photobleached maize leaves did not resume normal photosynthetic function. Maize cv. Batan 8686 from the highlands of Mexico was less susceptible to photosynthetic damage than maize cv. Bastion adapted for cultivation in W. Europe, when exposed to chilling night temperatures, or to mild chilling photoinhibitory conditions.     

适应不同光环境小蜡叶片气孔导度、气孔开度和气孔密度的时空变化

对生长在强光环境和弱光环境小蜡叶片的气孔参数测定发现:气孔导度和气孔开度在4个取样部位存在异质性分布.气孔导度和气孔开度经回归分析呈线性、指数或多项式分布.统计分析表明:强光下的叶片气孔导度和气孔开度的相关性明显高于弱光环境叶片的数值.无论强光环境还是弱光环境下的叶片,在取样部位编号为1和3,其气孔导度和气孔开度均存在显著的线性关系.弱光环境下叶片的气孔密度要远低于强光环境下的叶片.强光环境下叶片对变化环境的敏感性要大于弱光环境下的叶片,这可能与强光环境叶片具有较高的气孔密度有关.     

Polar current patterns in the leaves of the aquatic angiospermElodea Canadensis

Summary The pattern and density of ionic currents near the surface ofElodea canadensis leaves were investigated using a 3-d recording vibrating probe. Positive current always entered the adaxial (upper) surface, and left the abaxial (lower) surface of mature leaves. Current densities ranged from 1 to 15Acm^–2 in leaves irradiated with white light submerged in artificial pond water of pH 8. When the pH of the medium was lowered from 8 to 6, a rapid, approximate 40% reduction in current density on both sides of the leaf occured. Within 15 min of turning the light off, the current at the surface almost vanished. After re-introducing light, current began to rise after 10 to 15 min.Immature leaves exhibited no polarity in the current pattern, i.e., inward and outward current was found on both surfaces. Immature leaves also produced less current than mature ones, and responded less strongly to darkness. The results support the conclusion thatElodea leaves pump H⁺ out at the lower surface, and leak OH^– at the upper surface. The H⁺ current at the lower surface presumably facilitates the intake of carbon in form of CO₂.Abbrevations APW artificial pond water - Tris Tris (hydoxymethyl) amino methane - MES Morpholinoethane sulfonic acid - SD standard deviation - SEM standard error of the mean Dedicated to the memory of Professor Oswald Kiermayer     

Variations in stomatal density and index: implications for palaeoclimatic reconstructions

The variation in stomatal characters in leaves from one Alnus glutinosa (L.) Gaertn. tree is analysed. Measurements were taken from over 70 sites on the abaxial surfaces of representative ‘sun’ and ‘shade’ leaves having the same insertion point. The mean values of stomatal density and index in the shade leaf were significantly lower (71 and 93%, respectively) than those for the sun leaf. Within leaves, up to 2.5-fold differences in stomatal density values were observed. Contour maps derived from the data reveal non-random trends over the leaf surface. Correlations between stomatal density, epidermal cell density and stomatal index indicate that the variation in stomatal density within leaves arose primarily from local differences in stomatal differentiation, rather than from local differences in leaf expansion. This research demonstrates that a high level of variation in stomatal characters occurs both within and between leaves. We conclude that a well-defined sampling strategy should be used when estimating stomatal characters for (tree) leaves. Furthermore, the leaf's insertion point and situation within the tree crown should be taken into account. We discuss the implications of these findings for palaeoclimatic interpretations and emphasize the need for great caution when drawing conclusions based solely on stomatal characters.     

Effect of varying co2 and light levels on growth of hedyotis and sugarcane shoot cultures

Summary Shoot cultures of Hedyotis corymbosa, a C3 species, and sugarcane, a C4 species, were used to examine the effects of various CO₂ concentrations and two light intensities on growth and photosynthetic rates. The fresh and dry weights of new growth of Hedyotis shoots were higher when grown under the higher light intensity, while differences among shoots grown under different CO₂ levels were marginal. After 14 d of growth in various CO₂ concentrations, no significant differences could be observed in the newly produced leaves of Hedyotis with respect to stomatal distribution and number of mesophyll cell layers. Shoots grown under high light intensity did not show higher rates of photosynthesis than those grown under low light intensity. Also, sugarcane shoots grown in a CO₂-enriched environment did not have higher photosynthetic rates, perhaps because the C4 pathway is less sensitive to the ambient CO₂ concentration. The quantum yield of Hedyotis shoots grown on medium with 20 g l⁻¹ sucrose was lower than that of shoots on lower sucrose concentrations, supporting the view that photosynthesis is inhibited by high levels of sucrose. Our results suggest that Hedyotis shoots in culture exhibit some form of acclimation to high CO₂. so that there is no net gain in productivity by photosynthesis.     

Photosynthesis in an Australian rainforest tree,Argyrodendron peralatum,during the rapid development and relief of water deficits in the dry season

Summary Rates of apparent photosynthesis were measured in situ at five positions between the upper crown and a lower branch of a 34 m tall Argyrodendron peralatum (F.M. Bailey) H.L. Edlin ex I.H. Boas tree, and on an understorey sapling of the same species growing in a northern Australian rainforest. At the end of the dry season, rapid reductions in photosynthetic rates occurred in the upper crown within three days after a rain event, but changes in the lower crown and the sapling were less marked. Complete recovery of photosynthesis followed a second rain event. At high photon flux densities, stomatal conductance to water vapour decreased in a curvilinear fashion as the vapour pressure difference between leaf and air increased. Apparent photosynthesis was linearly related to stomatal conductance on the first clear day after each rain event, but there was no relationship between these parameters at the end of a brief natural drying cycle. Under conditions of adequate water supply, stomatal conductances of both upper crown and understorey leaves increased linearly with increasing photon flux density up to about 300 mol m^-2 s^-1. During water deficits, stomatal conductances in leaves from the understorey increased much more rapidly at very low photon flux densities than did conductances in leaves from the upper canopy.     

遮阴对高山杜鹃叶片解剖和光合特性的影响

为了解高山杜鹃对光能的需求和适应性,该研究以盆栽3 a生高山杜鹃品种cv. Furnivall's Daughter为材料,探讨了遮阴对高山杜鹃叶片解剖结构和光合特性的影响。结果表明:光照强度对高山杜鹃品种cv. Furnivall's Daughter叶片的气孔密度没有显著影响,其气孔密度范围在299.70~327.22个·mm~(-2)之间,但光照对气孔开度和单个气孔器的面积影响显著,100%全光照和30%全光照处理植株分别具有最小和最大的叶片气孔开度。在处理的光强范围内,随着光强减弱,叶片厚度、栅栏组织厚度、海绵组织厚度以及上、下表皮厚度逐渐降低,有利于提高叶片的光能利用效率。100%全光照处理下,高山杜鹃叶片的光饱和点(LSP)、净光合速率(P_n)、饱和光合速率(P_(max))、气孔导度(Gs)、蒸腾速率(Tr)均较低,遮阴处理有效提高了Pn、P_(max)、G_s、T_r和光能利用效率(LUE),且30%全光照处理植株的叶片光补偿点(LCP)、暗呼吸速率(Rd)最低,而LSP、P_n、P_(max)、G_s、T_r和LUE最高。这表明高山杜鹃在云南昆明地区的最适光照条件是30%左右的全光照,在高山杜鹃的栽培及应用中,应采取适当的遮阴措施以满足其生长的最佳光照条件。     

Selective Formation of Photosystem 1 under Illumination at Low Intensity

Analyses of chlorophylls a and b and P700 in the wheat leaves grown for 8 days under illumination with white light at different intensities suggested selective formation of photosystem 1 of the photosynthesis at low light intensities. This was confirmed for the two types of chloroplasts isolated from leaves grown at light intensities of 1.1 and 240 μ W/cm², respectively, by measuring their pigment compositions, activities of photosystems 1 and 2, and absorption and fluorescence spectra. The chloroplasts developed at the low intensity showed properties only of photosystem 1 while those developed at the high intensity showed properties of both photosystems 1 and 2. Only photosystem 1 particles were obtained by fractionation of low intensity chloroplasts by treatment with digitonin followed by centrifugation, while high intensity chloroplasts could be fractionated into photosystem-1 and photosystem-2 particles. When the leaves grown at low light intensity were illuminated with strong light, photosystem 2 was developed. The fluorescence emission spectrum of low intensity chloroplasts at 77°K showed two peaks at 685 and 734 nm, and the spectrum of high intensity chloroplasts showed three peaks at 685, 697 and 740 nm.     

Dynamic acclimation to sunlight in an alpine plant,Soldanella alpina L.

In the French Alps, Soldanella alpina (S. alpina) grow under shade and sun conditions during the vegetation period. This species was investigated as a model for the dynamic acclimation of shade leaves to the sun under natural alpine conditions, in terms of photosynthesis and leaf anatomy. Photosynthetic activity in sun leaves was only slightly higher than in shade leaves. The leaf thickness, the stomatal density and the epidermal flavonoid content were markedly higher, and the chlorophyll/flavonoid ratio was significantly lower in sun than in shade leaves. Sun leaves also had a more oxidised plastoquinone pool, their PSII efficiency in light was higher and their non-photochemical quenching (NPQ) capacity was higher than that of shade leaves. Shade-sun transferred leaves increased their leaf thickness, stomatal density and epidermal flavonoid content, while their photosynthetic activity and chlorophyll/flavonoid ratio declined compared to shade leaves. Parameters indicating protection against high light and oxidative stress, such as NPQ and ascorbate peroxidase, increased in shade-sun transferred leaves and leaf mortality increased. We conclude that the dynamic acclimation of S. alpina leaves to high light under alpine conditions mainly concerns anatomical features and epidermal flavonoid acclimation, as well as an increase in antioxidative protection. However, this increase is not large enough to prevent damage under stress conditions and to replace damaged leaves.     

LIGHT,TEMPERATURE AND SALINITY EFFECTS ON GROWTH,LEAF ANATOMY AND PHOTOSYNTHESIS OF DISTICHLIS SPICATA (L.) GREENE

The effects of light, temperature, and salinity on growth, net CO₂ exchange and leaf anatomy of Distichlis spicata were investigated in controlled environment chambers. When plants were grown at low light, growth rates were significantly reduced by high substrate salinity or low temperature. However, when plants were grown at high light, growth rates were not significantly affected by temperature or salinity. The capacity for high light to overcome depressed growth at high salinity cannot be explained completely by rates of net photosynthesis, since high salinity caused decreases in net photosynthesis at all environmental conditions. This salinity-induced decrease in net photosynthesis was caused largely by stomatal closure, although plants grown at low temperature and low light showed significant increases in internal leaf resistance to CO₂ exchange. Increased salinity resulted in generally thicker leaves with lower stomatal density but no significant differences in the ratio of mesophyll cell surface area to leaf area. Salinity and light during growth did not significantly affect rates of dark respiration. The mechanisms by which Distichlis spicata tolerates salt appear to be closely coulpled to the utilization of light energy. Salt-induced leaf succulence is of questionable importance to gas exchange at high salinity in this C₄ species.