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.     

Gradients of several leaf characteristics on stems of two forest herbs

The plants of two herbaceous species, namelyPulmonaria officinalis L. andSymphytum tuberosum L., growing in two deciduous forests in SW. Slovakia, were used for analysis of anatomicphysiological gradients along the insertion level of individual leaves. The gradients were established for six leaf characteristics. Specific leaf area and development of leaf area increased, while leaf tissue hydration, degrees of succulence and consistency decreased with height of insertion of leaves upon the stems. The gradients indicate the decreasing of the leaf water content and dry mass with height of the leaf on the stem. Stomatal resistance was usually the lowest in basal leaves and the highest in apical leaves of the plant. The anatomic-physiological heterogeneity of leaves of a plant is large and cannot be neglected in any anatomical or physiological investigations.     

Observations of Hydrogen and Oxygen Isotopes in Leaf Water Confirm the Craig-Gordon Model under Wide-Ranging Environmental Conditions

The Craig-Gordon evaporative enrichment model of the hydrogen (δD) and oxygen (δ¹⁸O) isotopes of water was tested in a controlled-environment gas exchange cuvette over a wide range (400‰ δD and 40‰ δ¹⁸O) of leaf waters. (Throughout this paper we use the term “leaf water” to describe the site of evaporation, which should not be confused with “bulk leaf water” a term used exclusively for uncorrected measurements obtained from whole leaf water extractions.) Regardless of how the isotopic composition of leaf water was achieved (i.e. by changes in source water, atmospheric vapor δD or δ¹⁸O, vapor pressure gradients, or combinations of all three), a modified version of the Craig-Gordon model was shown to be sound in its ability to predict the δD and δ¹⁸O values of water at the site of evaporation. The isotopic composition of atmospheric vapor was shown to have profound effects on the δD and δ¹⁸O of leaf water and its influence was dependent on vapor pressure gradients. These results have implications for conditions in which the isotopic composition of atmospheric vapor is not in equilibrium with source water, such as experimental systems that grow plants under isotopically enriched water regimes. The assumptions of steady state were also tested and found not to be a major limitation for the utilization of the leaf water model under relatively stable environmental conditions. After a major perturbation in the δD and δ¹⁸O of atmospheric vapor, the leaf reached steady state in approximately 2 h, depending on vapor pressure gradients. Following a step change in source water, the leaf achieved steady state in 24 h, with the vast majority of changes occurring in the first 3 h. Therefore, the Craig-Gordon model is a useful tool for understanding the environmental factors that influence the hydrogen and oxygen isotopic composition of leaf water as well as the organic matter derived from leaf water.     

Changing water use and adaptive strategies along rainfall gradients in Mediterranean lupins

• There is growing interest in harnessing the genetic and adaptive diversity of crop wild relatives to improve drought resilience of elite cultivars. Rainfall gradients exert strong selection pressure on both natural and agricultural ecosystems. Understanding plant responses to these facilitates crop improvement.
• Wild and domesticated narrow‐leafed lupin (NLL) collected along Mediterranean terminal drought stress gradients was evaluated under contrasting reproductive phase water supply in controlled field, glasshouse and cabinet studies. Plant phenology, growth and productivity, water use and stress responses were measured over time.
• There is an integrated suite of adaptive changes along rainfall gradients in NLL. Low rainfall ecotypes flower earlier, accumulate lower seed numbers, biomass and leaf area, and have larger root:shoot ratios than high rainfall ecotypes. Water‐use is lower and stress onset slower in low compared to high rainfall ecotypes. Water‐use rates and ecotypic differences in stress response (Ψleaf decline, leaf loss) are lower in NLL than yellow lupin (YL). To mitigate the effects of profligate water use, high rainfall YL ecotypes maintain higher leaf water content over declining leaf water potential than low rainfall ecotypes. There is no evidence for such specific adaptation in NLL.
• The data suggests that appropriate phenology is the key adaptive trait to rainfall gradients in NLL because of the flow‐on effects on biomass production, fitness, transpiration and stress onset, and the lack of physiological adaptations as in YL. Accordingly, it is essential to match phenology with target environment in order to minimize risk and maximize yield potential.

     

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

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

DIFFERING SELECTION ON PLANT PHYSIOLOGICAL TRAITS IN RESPONSE TO ENVIRONMENTAL WATER AVAILABILITY: A TEST OF ADAPTIVE HYPOTHESES

I used phenotypic selection analysis to test the prediction from functional and comparative studies of plants that smaller leaves and more efficient water use are adaptive in drier environments. I measured selection gradients on leaf size and instantaneous water-use efficiency (a measure of carbon gain per unit water loss) in experimental populations of Cakile edentula var. lacustris placed into wet and dry environments in the field. Linear and nonlinear selection differed significantly between the two environments as predicted. Water-use efficiency was selected to be higher, and leaf area was selected toward a small intermediate optimum, in the dry environment. There was also significant positive correlational selection on water-use efficiency and leaf size, suggesting that the optimum leaf size in the dry environment is greater for plants with higher water-use efficiency. In contrast, neither leaf size nor water-use efficiency were selected in the wet environment, though larger leaves resulted in greater vegetative biomass. Path analysis of the linear selection gradients found that water-use efficiency affected plant fitness primarily because it increased vegetative biomass, as suggested by the hypotheses about the function of physiological traits. These results were not only consistent with the functional hypotheses but also with the observed genetic differentiation in water-use efficiency and leaf size between wet and dry site populations.     

Hydraulic and chemical signalling in the control of stomatal conductance and transpiration.

Abscisic acid (ABA) transported in the xylem from root to shoot and perceived at the guard cell is now widely studied as an essential regulating factor in stomatal closure under drought stress. This provides the plant with a stomatal response mechanism in which water potential is perceived in the root as an indication of soil water status and available water resources. There is also ample evidence that stomata respond directly to some component of leaf water status. This provides additional information about water potential gradients developing between root and shoot as the result of water transport, allowing for a more stable regulation of shoot water status and better protection of the transport system itself. The precise location at which leaf water status is sensed, however, and the molecular events transducing this signal into a guard cell response are not yet known. Major questions therefore remain unanswered on how water stress signals perceived at root and leaf locations are integrated at the guard cell to control stomatal behaviour.     

Efficiency and regulation of water transport in some woody and herbaceous species

The efficiency with which plants transport water is related to the water potential differences required to drive water fluxes from the soil to the leaf. A comparative study of two woody and three herbaceous species (Citrus sinensis L. cv. Koethen, Pyrus kawakami L., Helianthus annuus L. cv. Mammoth Russian, Capsicum frutescens L. cv. Yolo Wonder, and Sesamum indicum L. cv. Glauca) indicated contrasts in water transport efficiency. Depression of leaf water potential in response to transpiration increases was found in the woody species; the herbaceous species, however, had more efficient water transport systems and presented no measurable response of leaf water potential to transpiration changes. Different maximum transpiration rates under the same climatic conditions were observed with different species and may be accounted for by stomatal response to humidity gradients between leaf and air. Leaf diffusion resistance in sesame increased markedly as the humidity gradient was increased, while leaf resistance of sunflower responded less to humidity. Stomata appeared to respond directly to the humidity gradient because changes in leaf water potential were not detected when leaf resistance increased or decreased.     

Leaf water 18O and 2H enrichment along vertical canopy profiles in a broadleaved and a conifer forest tree

Distinguishing meteorological and plant‐mediated drivers of leaf water isotopic enrichment is prerequisite for ecological interpretations of stable hydrogen and oxygen isotopes in plant tissue. We measured input and leaf water δ²H and δ¹⁸O as well as micrometeorological and leaf morpho‐physiological variables along a vertical gradient in a mature angiosperm (European beech) and gymnosperm (Douglas fir) tree. We used these variables and different enrichment models to quantify the influence of Péclet and non‐steady state effects and of the biophysical drivers on leaf water enrichment. The two‐pool model accurately described the diurnal variation of leaf water enrichment. The estimated unenriched water fraction was linked to leaf dry matter content across the canopy heights. Non‐steady state effects and reduced stomatal conductance caused a higher enrichment of Douglas fir compared to beech leaf water. A dynamic effect analyses revealed that the light‐induced vertical gradients of stomatal conductance and leaf temperature outbalanced each other in their effects on evaporative enrichment. We conclude that neither vertical canopy gradients nor the Péclet effect is important for estimates and interpretation of isotopic leaf water enrichment in hypostomatous trees. Contrarily, species‐specific non‐steady state effects and leaf temperatures as well as the water vapour isotope composition need careful consideration.     

Water-stressed maize, barley and rice seedlings show species diversity in mechanisms of leaf growth inhibition

The possible occurrence of species diversity in mechanisms underlying leaf-growth inhibition by water stress, was investigated in related cereal plants. Water stress was generated by additions of the osmoticum polyethylene glycol 6000 to the root medium. Effects of external water potentials ranging from 0 to -0.6MPa, on early growth parameters of emerging leaves were measured under controlled environment conditions, using pairs of maize, barley or rice genotypes with differing resistance to water stress under field conditions. Water potentials of -0.4 MPa for 24 h, similarly reduced leaf growth, comparative production rates of leaf epidermal cells and cell size in all genotypes. These reductions did not appear to be caused by reductions in the osmotic potential gradients between the expanding leaf cells and their external water source. However, growth inhibition in maize and barley, was accompanied by significant reductions in comparative leaf and cell wall extensibility. Moreover, regression plots revealed good linear correlations (r=0.83** for maize and r=0.77** for barley) between the reductions in leaf growth induced by a series of water potentials and associated reductions in leaf extensibility. In contrast, the reduction in growth of rice leaves, was not accompanied by any significant changes in leaf or cell wall extensibility. Similarly, regression plots revealed poor correlations between leaf growth and leaf extensibility in both paddy and upland rice (r=0.17 and r=0.07, respectively). Thus, despite numerous inter-species similarities, biophysical changes associated with stress-induced leaf growth inhibition in maize and barley, differed from those in rice.Key words: Cell walls, extensibility, water stress, cereal diversity, leaf growth.      

敦煌阳关湿地芦苇叶性状对土壤水分的响应

为了了解干旱区湿地不同水分梯度下芦苇叶性状的构建模式及对不同水分环境的适应策略差异性,以敦煌阳关渥洼池湿地植物芦苇(Phragmites australis)为研究对象,分析不同水分梯度下芦苇叶性状对土壤水分的响应。结果表明:(1)低水分梯度下的叶厚度(LT)和叶片磷含量(LPC)显著高于中、高水分梯度(P0.05),高水分梯度下的叶碳含量(LCC)也显著高于中、低水分梯度(P0.05)。(2)芦苇叶性状之间的关系也因水分的变化而有所不同。就整体而言,叶片碳含量与叶干物质含量(LDMC)极显著正相关(P0.01);叶厚度与叶片磷含量极显著正相关(P0.01),比叶面积(SLA)与叶氮含量(LNC)极显著正相关(P0.01)、与叶碳氮比(C/N)极显著负相关(P0.01)。(3)在水分作为限制因素的条件下,叶片碳含量与比叶面积对水分的响应最为突出。土壤含水量与叶厚度、叶碳含量和叶片磷含量均显著负相关(P0.05)。芦苇在低水分环境下采取高收入低投入的积极型生存策略,高水分下则采取低获取高消耗的保守型生存策略,说明了芦苇在适应异质生境的自我调节机制。     

Stomatal Response to Environment with Sesamum indicum. L

Leaf resistance of Sesamum indicum L. increased when the humidity gradient between leaf and air was increased, at moderate temperatures, even though calculated carbon dioxide concentrations within the leaf decreased slightly. Mesophyll resistance remained relatively constant when humidity gradients were changed, indicating that the increases in leaf resistance were mainly caused by reductions in stomatal aperture and that nonstomatal aspects of photosynthesis and respiration were not affected. Low carbon dioxide concentrations inside the leaf decreased but did not eliminate resistance response to the humidity gradient. Internal carbon dioxide concentrations had little effect on resistance in humid air but had moderate effects on resistance with large humidity gradients between leaf and air. Stomatal response to humidity was not present at high leaf temperatures. Effects of humidity gradients on photosynthetic and stomatal responses to temperature suggested that large humidity gradients may contribute to mid-day closure of stomata and depressions in photosynthesis.     

水分对敦煌阳关湿地芦苇叶片与土壤C、N、P生态化学计量特征的影响

土壤水分是影响干旱区植物养分吸收和利用策略的关键因子之一。研究不同水分梯度叶片与土壤生态化学计量特征,有助于揭示植物对环境变化的响应特征及生态适应性。通过野外调查与实验分析,对敦煌阳关不同水分梯度芦苇叶片与土壤碳(C)、氮(N)、磷(P)生态化学计量特征及其关系进行了研究。结果表明:(1)随土壤含水率升高,叶片C、N、P含量降低,叶片C/N、C/P、N/P升高。(2)随土壤含水率升高,土壤有机碳(OC)、总氮(TN)、总磷(TP)含量及土壤N/P升高,土壤C/N降低,土壤C/P先升后降。(3)低水分梯度叶片N、C/N与土壤N、C/N显著负相关(P<0.05),叶片C、P、C/P、N/P与土壤C、P、C/P、N/P无显著相关性(P>0.05);高、中水分梯度叶片C、N、P与土壤C、N、P化学计量特征相关性均不显著(P>0.05)。低水分梯度叶片受干旱胁迫和土壤养分制约,且能够保持较高的叶养分含量,体现了干旱区湿地植物异质生境下独特的养分调节机制。     

The gradient between pre-dawn rhizoplane and bulk soil matric potentials, and its relation to the pre-dawn root and leaf water potentials of four species

Uptake of soil water by plants may result in significant gradients between bulk soil and soil in the vicinity of roots. Few experimental studies of water potential gradients in close proximity to roots, and no studies on the relationship of water potential gradients to the root and leaf water potentials, have been conducted. The occurrence and importance of pre-dawn gradients in the soil and their relation to the pre-dawn root and leaf water potentials were investigated with seedlings of four species. Pre-germinated seeds were grown without watering for 7 and lid in a silt loam soil with initial soil matric potentials of -0.02, -0.1 and -0.22 MPa. Significant gradients, independent of the species, were observed only at pre-dawn soil matric potentials lower than -0.25 MPa; the initial soil matric potentials were -0.1 MPa. At an initial bulk soil matric potential of -0.22 MPa, a steep gradient between bulk and rhizoplane soil was observed after 7 d for maize (Zea mays L. cv. Issa) and sunflower (Helianthus annuus L. cv. Nanus), in contrast to barley (Hordeum vulgare L. cv. Athos) and wheat (Triticum aestivum L. cv. Kolibri). Pre-dawn root water potentials were usually about the same as the bulk soil matric potential and were higher than the rhizoplane soil matric potential. Pre-dawn root and leaf water potentials tended to be much higher than rhizoplane soil matric potentials when the latter were lower than -0.5 MPa. It is concluded that plants tend to become equilibrated overnight with the wetter bulk soil or with wetter zones in the bulk soil. Plants can thus circumvent negative effects of localized steep pre-dawn soil matric potential gradients. This may be of considerable importance for water uptake and growth in drying soil.     

Transpiration rate and intercellular diffusive resistance in the tobacco leaf

The increase in the measured transpiration rate in tobacco leaves due to the experimentally decreased humidity of the bulk air was found to be significantly lower than the theoretical value calculated from the change of water vapour concentration gradients. Boundary layer and stomatal diffusive resistances remained unchanged under experimental conditions with no change of net photosynthetic CO₂ uptake. This suggests an increase in intercellular diffusive resistance with an increase in water vapour concentration gradient which is the driving force of water vapour diffusive part of transpiration flux. The increase can be ascribed to the lengthening of intercellular diffusive pathway as steeper water vapour concentration gradient in intercellular spaces results in an increased evaporating surface of intercellular cells thus moving the effective plane of vaporization in leaf mesophyll further inwards. Due to different and independent changes of concentration gradients for water vapour and CO₂, different length of intercellular diffusive pathways for CO₂ and water vapour may be expected.     

Nuclear magnetic resonance study of spin relaxation and magnetic field gradients in maple leaves.

1H Nuclear magnetic resonance techniques were used to measure the distributions of spin-spin relaxation times, T2, and of magnetic field gradients in both the chloroplast and nonchloroplast water compartments of maple leaves (Acer platanoides). Results showed that encounters between water molecules and membranes inside chloroplasts provide an inefficient relaxation mechanism; i.e., chloroplast membranes interact weakly with water molecules. Gradient measurements indirectly measured the sizes of chloroplasts by showing that water in the chloroplasts is confined to small compartments a few microns in diameter. A comparison between measured gradients and gradients calculated for a model leaf indicated that chloroplasts are somewhat more likely to occupy positions along cell walls adjacent to air spaces, but also they may be found in the interiors of cells.     

Antitranspirant-induced increases in leaf water potential increase tuber calcium and decrease tuber necrosis in water-stressed potato plants

Experiments were undertaken with field-grown potato (Solanum tuberosum L.) plants to test the hypothesis that altering leaf:tuber water potential gradients within a plant subjected to low soil moisture will allow greater Ca accumulation in tubers and reverse Ca deficiency-related tuber necrosis. Antitranspirant formulations containing a wax emulsion and a spreader/sticker surfactant increased leaf water potential during a drought episode, significantly reducing the potential gradient that develops between leaf and tuber during a period of stress. Increased leaf water potential in treated plants was associated with decreased leaf Ca and increased tuber Ca. Tuber necrosis was found to be reduced in treated plants, thus increasing tuber quality.     

地形因子对黄土高原山杏叶片功能性状的影响

坡向和坡度是重要的地形因子,调控水、热组合可以影响植物生长及叶片功能性状,研究叶片功能性状对不同地形因子的响应,有助于了解植物对环境的适应策略.以阳坡(半阳坡)和阴坡(半阴坡)2个坡向,以及15°~20°、21°~25°和26°~30°3个坡度为环境梯度,对比研究了其对黄土高原主要造林树种山杏叶片功能性状(包括叶面积、比叶面积、叶干物质含量和叶片含水量)的影响,结果表明:1)坡向与坡度分别对所有叶片功能性状的影响均显著,坡向和坡度仅对叶片含水量具有交互作用.2)坡向梯度上,阳坡叶片的叶面积与阴坡相当,但叶干物质含量(0.24 g·g^-1)和叶片含水量(59.6%)均小于后者(0.27 g·g^-1和67.0%);阳坡叶片的比叶面积显著高于阴坡,其值分别为183.72和163.05 cm2·g^-1.3)坡度梯度上,叶片的比叶面积和叶面积均随坡度增大呈先减后增趋势,二者分别在15°~20°和26°~30°上达最大值,分别为184.04 cm^2·g^-1和21.14 cm2.4)除15°~20°与26°~30°的土壤水分无差异外,其余坡度以及坡向之间的土壤水分均存在显著差异,且土壤水分是造成叶片功能性状差异的主要原因之一,其中0~10 cm土层含水量对叶片功能性状的影响最大.5)叶片比叶面积与叶片含水量、叶干物质含量呈负相关,与叶面积呈正相关;叶干物质含量与叶片含水量呈正相关,与叶面积呈负相关;土壤含水量仅与叶片含水量呈正相关,与其他叶片功能性状均不相关.     

Seasonal and diurnal leaf movements of Rhododendron maximum L. in contrasting irradiance environments

Summary Leaf orientation (azimuth and angle) and leaf curling were measured seasonally and diurnally on Rhododendron maximum L. under an evergreen and a deciduous canopy. The microclimatic conditions under the evergreen canopy (mixed pine and hemlock) were characterized by lower irradiance but similar temperature, and vapor pressure deficit (vpd) to that under the deciduous canopy (mixed oak and maple). Under both canopies irradiance was more intense during winter months.On a seasonal basis leaf angle was closer to horizontal under the evergreen canopy but there was no difference between leaf curling in the two sites. Stomatal conductance was higher under the deciduous canopy but stomata were closed in the winter (following canopy abscission) under the evergreen and deciduous canopies even during warm winter days. Leaf water potentials were lower in the winter and Rhododendron maximum had higher leaf water potentials under the evergreen canopy.Significant association between mean leaf angle and curling index were found above a mean leaf angle of 70°. Leaf curling was highly associated with leaf temperature where 0° C was a critical value stimulating leaf curling. Leaf angle was linearly related to leaf temperatures above 0° C although this relationship was different under the two canopy types as a result of differing irradiance or differing water potential.     

Light adaptation and the role of autotrophic epiphytes in primary production of the temperate seagrass, Zostera marina L.

Photosynthetic features of Zostera marina L. and its autotrophic epiphyte community were investigated in a population inhabiting a shallow (1.3 m depth) water meadow in Great Harbor, Woods Hole, MA (U.S.A.). Photosynthesis versus irradiance (P-I) relationships were measured with respect to leaf age determined by the leaf position in the shoot bundle and by location of the tissue along the leaf axis. Therefore both age and light intensity gradients along the leaf axis were considered. The maximum photosynthesis (P_max) per dm² typically increased nearly two-fold along the leaf axis from leaf bases to apices. Photosynthetic rate on a chlorophyll (Chl) basis did not increase as dramatically along the leaf axis, and rates were usually lowest in tissues with the highest Chl content. The P-I relationships of leaves of different ages did not reveal photoinhibition even at light intensities > 1400 μE • m⁻² • s ⁻¹. Furthermore, no photoinhibition was observed in tissues from leaf blade bases, which never experienced high light levels (> 500 μE • m ⁻² • s⁻¹) in situ in Great Harbor. The initial slopes of the P-I curves and light compensation and saturation values varied along the leaf axis in relation to in situ light intensity gradients and in relation to leaf or tissue age. It appeared that leaf and/or tissue age was more important than light environment in determining P-I responses. The contribution of the autotrophic epiphyte community on Z. marina leaves to total photosynthesis per dm² was between 27 and 50%, and between 10 and 44% per mg chlorophyll. These levels of epiphyte photosynthesis can double the primary production of Z. marina leaves. No detrimental effects of epiphyte cover were realized in leaf maximal photosynthesis or P-I relationships. Non-epiphytized leaves and leaves from which epiphytes were removed showed essentially identical photosynthetic features. Light intensity and age gradients along the leaf axis control both the photosynthetic performance of the leaves and epiphyte biomass and photosynthesis.