Water stress-induced modifications of leaf hydraulic architecture in sunflower: co-ordination with gas exchange

The hydraulic architecture, water relationships, and gas exchange of leaves of sunflower plants, grown under different levels of water stress, were measured. Plants were either irrigated with tap water (controls) or with PEG600 solutions with osmotic potential of -0.4 and -0.8 MPa (PEG04 and PEG08 plants, respectively). Mature leaves were measured for hydraulic resistance (R(leaf)) before and after making several cuts across minor veins, thus getting the hydraulic resistance of the venation system (R(venation)). R(leaf) was nearly the same in controls and PEG04 plants but it was reduced by about 30% in PEG08 plants. On the contrary, R(venation) was lowest in controls and increased in PEG04 and PEG08 plants as a likely result of reduction in the diameter of the veins' conduits. As a consequence, the contribution of R(venation) to the overall R(leaf) markedly increased from controls to PEG08 plants. Leaf conductance to water vapour (g(L)) was highest in controls and significantly lower in PEG04 and PEG08 plants. Moreover, g(L) was correlated to R(venation) and to leaf water potential (psi(leaf)) with highly significant linear relationships. It is concluded that water stress has an important effect on the hydraulic construction of leaves. This, in turn, might prove to be a crucial factor in plant-water relationships and gas exchange under water stress conditions.     

Diurnal and seasonal changes of leaf lamina hydraulic conductance in bur oak (Quercus macrocarpa) and trembling aspen (Populus tremuloides)

The aim of this study was to examine the diurnal and seasonal variations in the sensitivity of leaf lamina (K _lam) hydraulic conductance to irradiance in bur oak (Quercus macrocarpa Michx.) and trembling aspen (Populus tremuloides Michx.), which vary in their responses of K _lam to irradiance. K _lam was determined using the high-pressure method and the measurements were carried out in June, August and September. The irradiance response of K _lam in bur oak was present throughout the day and declined in senescing leaves. In trembling aspen, K _lam declined from morning to late afternoon and drastically decreased before the onset of leaf senescence, but it was not sensitive to irradiance. In both tree species, the capacity of the petioles to supply water to leaf lamina changed during the day in accordance with the ability of the leaf lamina to transport water. Petiole hydraulic conductivity (K _pet) declined during the season in bur oak leaves, while it tended to increase in trembling aspen leaves. There was no correlation between the K _lam values and air temperature or light intensity at the time of leaf collection. For trembling aspen, K _pet was negatively correlated with the air temperature suggesting sensitivity to drought. We conclude that the water transport properties of petioles and leaf lamina in the two studied tree species reflect their ecological adaptations. Trembling aspen leaves have high hydraulic conductivity and high stomatal conductance regardless of the irradiance level, consistent with the rapid growth and high demand for water. In contrast, the increased lamina hydraulic conductivity and stomatal conductance under high irradiance in bur oak trees reflect a water conservation strategy.     

Fluoride inhibits root water transport and affects leaf expansion and gas exchange in aspen (Populus tremuloides) seedlings

The effects of sodium fluoride (0.3, 5 and 10 m M NaF) on root hydraulic conductivity, and gas exchange processes were examined in aspen ( Populus tremuloides Michx.) seedlings grown in solution culture. A long-term exposure of roots to NaF significantly decreased root hydraulic conductivity ( L _p) and stomatal conductance ( g _s). Root absorbed NaF significantly affected electrolyte leakage in leaf tissues and substantially restricted leaf expansion. NaF did not significantly affect leaf chlorophyll contents but decreased net photosynthesis ( P _n). A short-term exposure of excised roots to 5 m M NaF and KF significantly decreased root water flow ( Q _v) with a concomitant decline in root respiration and reduced g _s when applied through intact roots or excised stems. The same molar concentration of NaCl also decreased Q _v and g _s in intact seedlings, but to a lesser extent than NaF or KF, and did not significantly affect root respiration. The results suggest that fluoride metabolically inhibited Q _v or L _p, probably by affecting water channel activity. We suggest that the metabolic inhibition of L _p by root-absorbed fluoride affected gas exchange and leaf expansion in aspen seedlings.     

Impact of epidermal leaf mining by the aspen leaf miner (Phyllocnistis populiella) on the growth, physiology, and leaf longevity of quaking aspen

The aspen leaf miner, Phyllocnistis populiella, feeds on the contents of epidermal cells on both top (adaxial) and bottom (abaxial) surfaces of quaking aspen leaves, leaving the photosynthetic tissue of the mesophyll intact. This type of feeding is taxonomically restricted to a small subset of leaf mining insects but can cause widespread plant damage during outbreaks. We studied the effect of epidermal mining on aspen growth and physiology during an outbreak of P. populiella in the boreal forest of interior Alaska. Experimental reduction of leaf miner density across two sites and 3 years significantly increased annual aspen growth rates relative to naturally mined controls. Leaf mining damage was negatively related to leaf longevity. Leaves with heavy mining damage abscised 4 weeks earlier, on average, than leaves with minimal mining damage. Mining damage to the top and bottom surfaces of leaves had different effects on physiology. Mining on the top surface of the leaf had no significant effect on photosynthesis or conductance and was unrelated to leaf stable C isotope ratio (delta(13)C). Mining damage to the bottom leaf surface, where stomata are located, had significant negative effects on net photosynthesis and water vapor conductance. Percent bottom mining was positively related to leaf delta(13)C. Taken together, the data suggest that the primary mechanism for the reduction of photosynthesis by epidermal leaf mining by P. populiella is the failure of stomata to open normally on bottom-mined leaves.     

Decline of leaf hydraulic conductance with dehydration: relationship to leaf size and venation architecture

Across plant species, leaves vary enormously in their size and their venation architecture, of which one major function is to replace water lost to transpiration. The leaf hydraulic conductance (K(leaf)) represents the capacity of the transport system to deliver water, allowing stomata to remain open for photosynthesis. Previous studies showed that K(leaf) relates to vein density (vein length per area). Additionally, venation architecture determines the sensitivity of K(leaf) to damage; severing the midrib caused K(leaf) and gas exchange to decline, with lesser impacts in leaves with higher major vein density that provided more numerous water flow pathways around the damaged vein. Because xylem embolism during dehydration also reduces K(leaf), we hypothesized that higher major vein density would also reduce hydraulic vulnerability. Smaller leaves, which generally have higher major vein density, would thus have lower hydraulic vulnerability. Tests using simulations with a spatially explicit model confirmed that smaller leaves with higher major vein density were more tolerant of major vein embolism. Additionally, for 10 species ranging strongly in drought tolerance, hydraulic vulnerability, determined as the leaf water potential at 50% and 80% loss of K(leaf), was lower with greater major vein density and smaller leaf size (|r| = 0.85-0.90; P < 0.01). These relationships were independent of other aspects of physiological and morphological drought tolerance. These findings point to a new functional role of venation architecture and small leaf size in drought tolerance, potentially contributing to well-known biogeographic trends in leaf size.     

Root hydraulic conductance,gas exchange and leaf water potential in seedlings of Pistacia lentiscus L. and Quercus suber L. grown under different fertilization and light regimes

Differences in morphology, biomass allocations and physiological responses were investigated in seedlings of Mastic tree (Pistacia lentiscus L.) and Cork oak (Quercus suber L.) submitted to contrasting fertilization and light regimes during early growth. These species are two evergreen sclerophyllous Mediterranean species frequently used in Mediterranean reforestation programmes. Fertilization was the treatment that affected most of the morphological and physiological variables evaluated in P. lentiscus and Q. suber seedlings. Leaf area and specific leaf area (SLA) were affected by shading treatment in both species, showing higher values in seedlings grown under shade. P. lentiscus seedlings showed a high capacity to modify root morphological variables and root hydraulic conductance (K_R) with the fertilization treatment. In contrast, Q. suber showed low to moderate root system changes with the treatments applied, although the fertilization level affected biomass allocation (i.e., root to shoot ratio) in both species. Under high water demand, P. lentiscus seedlings with high K_R allowed transpiration (E) to increase without increasing the water potential gradient between soil and leaves. In Q. suber, high fertilization induced significant increases in photosynthesis (A), as well as a tendency to increase E with significantly lower leaf water potential (ψ_L).     

The effect of root pressurization on water relations, shoot growth, and leaf gas exchange of peach (Prunus persica) trees on rootstocks with differing growth potential and hydraulic conductance

It is well known that rootstocks can have an effect on the vegetative growth and development of the tree; however, there has been no clear explanation about the physiological mechanism involved in this phenomenon. Evidence indicates that the rootstock effects on tree vegetative growth may be related to hydraulic limitations of the rootstock. The objective of these experiments was to investigate the shoot growth, water potential, and gas exchange of peach trees on different rootstocks in response to manipulations of water relations of trees on rootstocks that differ in root hydraulic conductance. Tree water relations were manipulated by applying different amounts of pneumatic pressure on the root system and then relative shoot extension growth rate, tree transpiration rate, leaf water potential, leaf conductance, leaf transpiration, and net CO(2) exchange rate responses were measured. Root pressurization increased leaf water potential, relative shoot extension growth rate, leaf conductance, leaf transpiration, and net CO(2) exchange rates of trees on both vigorous and dwarfing rootstocks. There was a significant positive linear correlation between applied pneumatic pressure and tree transpiration rate and leaf water potential. Leaf conductance, transpiration rate, and net CO(2) exchange rate as well as relative shoot extension growth rates were also positively correlated with the applied pneumatic pressure on the root system. These relationships were consistent across both vigorous and size-controlling rootstocks, indicating that rootstock hydraulic limitation may be directly involved in the vegetative growth control of peach trees.     

Effect of vacuum infiltration on photosynthetic gas exchange in leaf tissue

Using a manometric method, photosynthetic oxygen evolution and ¹⁴CO₂ fixation have been determined for leaf tissue of Triticum aestivum L., Hordeum vulgare L., Phaseolus vulgaris L., and Lemna minor L. Approximately similar values in the range 0.2 to 0.4 millimoles grams fresh weight⁻¹ hour⁻¹ were obtained for both gases. In tissue subjected to vacuum infiltration, O₂ evolution and ¹⁴CO₂ fixation were barely measurable. It is considered that the elimination of photosynthetic gas exchange results from a decreased supply of CO₂ to the chloroplasts. Chopping wheat laminae also leads to a reduction in photosynthetic gas exchange, slices 1 millimeter or less giving only 10 to 20% of the value for whole tissue. Respiration is unaffected by either treatment. Carbonic anhydrase did not improve photosynthetic gas exchange in infiltrated tissue. The use of sliced or vacuum-infiltrated leaf tissue in photosynthetic studies is discussed.     

Effects of mannose on photosynthetic gas exchange in spinach leaf discs

When mannose is provided in the transpiration stream to spinach (Spinacia oleracea) leaf discs, a series of specific and nonspecific changes occur in CO₂ and H₂O vapor exchange as a function of feeding time. The initial increases in apparent photosynthesis and transpiration are nonspecific effects due to osmotic changes leading to passive stomatal opening. The mannose-specific effects are: (a) time-dependent changes in the CO₂ concentration required for saturation; (b) complex kinetics of the inhibition of CO₂ assimilation dependent on CO₂ and O₂ concentrations and the duration of feeding (high CO₂ and low O₂ lead to rapid inhibitions of photosynthesis); (c) elimination of the capacity of 2% O₂ to stimulate photosynthesis; and (d) oscillations in the CO₂ exchange rate following transitions from 20% to 2% O₂. The mannose-specific effects are reversible by orthophosphate. The mannose-dependent changes in gas exchange are attributed to altered [ATP]/[ADP] ratios.     

Root water flow and leaf stomatal conductance in aspen (Populus tremuloides) seedlings treated with abscisic acid

Exogenous abscisic acid (ABA) applied to the roots and excised shoots of aspen (Populus tremuloides Michx.) inhibited stomatal conductance. However, the effect of ABA on stomatal conductance was more pronounced in the excised shoots compared with the intact seedlings. Approximately 10% of the ABA concentration applied to the roots was found in the xylem exudates of root systems exposed to a hydrostatic pressure of 0.3 MPa. A similar concentration of ABA applied to the excised shoots produced a faster and greater reduction of stomatal conductance. ABA applied to the roots had no effect on root steady-state flow rate over the 5-h experimental period. Moreover, pre-incubating root systems of intact seedlings for 12 h with 5 x 10(-5) M ABA did not significantly reduce volume flow density. Similarly, ABA had no effect on root hydraulic conductivity and the activation energy of root water flow rates.     

Kinetics of recovery of leaf hydraulic conductance and vein functionality from cavitation-induced embolism in sunflower

The kinetics of leaf vein recovery from cavitation-induced embolism was studied in plants of sunflower cv. Margot, together with the impact of vein embolism on the overall leaf hydraulic conductance (Kleaf). During the air-dehydration of leaves to leaf water potentials (Psi L) of -1.25 MPa, Kleaf was found to decrease by about 46% with respect to values recorded in well-hydrated leaves. When leaves, previously dehydrated to Psi L= -1.1 MPa (corresponding to the turgor loss point), were put in contact with water, Kleaf recovered completely in 10 min and so did leaf water potential. Functional vein density was estimated in both dehydrating and rehydrating leaves in terms of total length of red-stained veins infiltrated with a Phloxine B solution per unit leaf surface area. Veins were found to embolize (unstained) with kinetics showing a linear relationship with Kleaf so that about a 70% loss of functional veins corresponded with a Kleaf loss of 46%. Cavitated veins recovered from embolism within 10 min from the beginning of leaf rehydration. These data indicate that: (a) leaves of sunflower underwent substantial vein embolism during dehydration; (b) vein embolism and leaf hydraulic efficiency apparently recovered from dehydration completely and rapidly upon rehydration; (c) vein refilling occurred while conduits were still at more negative xylem pressures than those required for spontaneous bubble dissolution on the basis of Henry's law. The possible consistent contribution of vital mechanisms for vein refilling is discussed.     

Effect of fruit load and girdling on leaf photosynthesis in Mangifera indica L

Leaf nitrogen concentration (Nm), mass-to-area ratio, amount of nitrogen per unit leaf area (Na), non-structural carbohydrate concentration (TNCa), maximal rate of carboxylation (Vcmax), light-saturated rate of photosynthetic electron transport (Jmax), dark respiration (Rd), net photosynthetic assimilation (Anet), quantum yield of photosystem II (PhiPSII), and intercellular CO2 concentration (Ci) were measured in Mangifera indica L. leaves on three types of fruit-bearing branches (non-girdled, NG; girdled with high (HFL) and low (LFL) fruit load), experiencing similar light exposure. TNCa, Vcmax/Na, Jmax/Na, Rd/Na, Ci, and the initial quantum yield of photosynthetic electron transport (alpha) were similar in both HFL and NG treatments, but Nm, Na, and photosynthetic capacity parameters (Vcmax and Jmax) were lower in the HFL than in the NG treatment. The strong depressing effect of girdling on leaf nitrogen concentration cannot therefore be attributed to a change in TNCa. By contrast, Na and TNCa were lower and higher, respectively, in the LFL than in the HFL treatment, suggesting that carbohydrate content may become the driving force behind photosynthetic acclimation to changing source-sink relationships, like the ones resulting from the presence of developing fruits. Vcmax/Na and Jmax/Na were lower in the LFL than in the HFL treatment, while Rd/Na, Ci, and alpha were not affected by fruit load. It is concluded that girdling and high fruit load affect photosynthesis permanently by decreasing and increasing, respectively, leaf nitrogen concentration. Fruit load, moreover, may have an additional effect on photosynthetic capacity by affecting the relationship between Vcmax and Jmax, and Na.     

成熟马占相思水力导度对水分利用和光合响应的季节性差异

通过测定成熟马占相思叶片的水势、气孔导度、蒸腾速率(Tr)、叶面积指数、边材面积等参数,研究了湿季(5月)和干季(11月)叶片的水力导度(K1)、水分和光合特性的关系.结果表明:高大植株(平均树高20 m、胸径0.26 m)的边材面积与叶面积的比率(Asp/Acl)比较小植株(平均树高14.5 m、胸径0.19m)高8.5％,前者的木质部水分通量大于后者,以支持冠层叶片的水分利用.对木质部易损曲线进行分析,K1降低50％时,湿、干季的叶片水势(Ψ1)分别为-1.41和-1.55 MPa,且干季的木质部空穴化的易损性高于湿季.湿、干季的K1峰值分别为5.5和4.5 mmol·m-2·s-1·MPa-1,最大蒸腾速率(Trmax)分别为3.6和1.8 mmol·m-2·s-1,且湿季的K1和Trmax明显大于干季.一天中K1和Tr的多次波动反映了木质部空穴化和修复的往复循环,叶片气孔在K降低超过50％或Ψ1达到-1.6MPa时关闭,气孔导度在K1达到50％前仍保持较高水平.干季的水力导度与光合速率的相关性较湿季明显.季节更迭导致叶片水力导度损失是Tr和CO2交换下降的原因.     

Modelled influences of non-exchanging trichomes on leaf boundary layers and gas exchange

The two main resistances in the exchange of gases between plants and the atmosphere are stomatal and boundary layer resistances. We modeled boundary layer dynamics over glabrous and pubescent leaves (assuming non-exchanging trichomes) with leaf lengths varying from 0.01 to 0.2 m, and windspeeds of 0.1-5.0 m x s(-1). Results from theoretical and semi-empirical formulae were compared. As expected, boundary layer thickness decreased with decreasing leaf length and increasing windspeed. The presence of trichomes increased leaf surface roughness, resulting in lowered Reynolds numbers at which the boundary layer became turbulent. This effect is especially important at low windspeeds and over small leaves, where the Reynolds number over glabrous surfaces would be low. We derived a new simple dimensionless number, the trip factor, to distinguish field conditions that would lead to a turbulent boundary layer based on the influence of trichomes. Because modeled rates of CO2 and H2O(v) exchange over turbulent boundary layers are one or more orders of magnitude faster than over laminar boundary layers, a turbulent boundary layer may lead to increased carbon uptake by plants. The biological trade-off is potentially increased transpirational water loss. However, in understory habitats characterized by low windspeeds, even a few trichomes may increase turbulence in the boundary layer, thus facilitating photosynthetic gas exchange. Preliminary field data show that critical trip factors are exceeded for several plant species, both in understory and open habitats.     

Sap flow, gas exchange, and hydraulic conductance of young apricot trees growing under a shading net and different water supplies

The experiment was carried out in a research field near Murcia, Spain, over a 3-week period between September 26 and October 16, 2000. Sixteen trees were used in the experiment, eight of which were placed under a rectangular shading net, while the other eight were maintained in the open air. Trees were irrigated once per day and, after October 5th, water was witheld from eight trees (four shaded and four unprotected for 5 days). The leaf stomatal conductance and the photosynthesis rates were higher in the shaded trees than in the exposed plants, probably because the leaf water potential was lower in the unshaded plants. This higher leaf conductance partially compensated for the effect of low radiation on transpiration, and the reduction of daily sap flow registered in shaded trees was only around 10-20%. The net also affected trunk diameter changes, with the shaded trees showing lower values of maximum daily shrinkage. Soil water deficit and high radiation had a similar effect on plant water parameters, lowering leaf water potential, leaf stomatal conductance, and the photosynthesis rate. The effects of both conditions were accumulative and so the exposed water-stressed plants showed the lowest values of total hydraulic resistance and water use efficiency, while the shaded well-irrigated trees registered the highest values for both parameters. For this reason, we think that net shading could be extended to apricot culture in many areas in which irrigation water is scarce and insolation is high.     

Effects of soil flooding on leaf gas exchange of tomato plants

Carbon dioxide and water vapor exchange of tomato (Lycopersicon esculentum Mill. cv Rheinlands Ruhm) leaves were measured before and after 24 h of soil flooding to characterize both stomatal and nonstomatal responses to the stress. Leaf epidermal conductance to water vapor decreased by 47% after flooding, accompanied by an increase in the sensitivity of stomata to changes in CO₂ concentration. Assimilation rates under ambient conditions fell by 27%, and the inhibition could not be overcome by elevated CO₂ partial pressures. Stomatal conductance limited the assimilation rate to approximately the same degree both before and after flooding. The reduction in photosynthetic capacity was not due to a decrease in apparent quantum yield or to an increase in photorespiration. The results were analyzed according to a recent model of photosynthesis, and possible mechanisms underlying the flooding effect are discussed.     

Roles of leaf water potential and soil-to-leaf hydraulic conductance in water use by understorey woody plants

Diurnal changes of leaf water potential and stomatal conductance were measured for 12 deciduous shrubs and tree saplings in the understorey of a temperate forest. Sunflecks raised the leaf temperature by 4°C, and vapor pressure deficit to 2 kPa. Although the duration of the sunflecks was only 17% of daytime, the photon flux density (PFD) of sunflecks was 52% of total PFD on a sunny summer day. Leaf osmotic potential at full turgor decreased in summer, except in some species that have low osmotic potential in the spring. Plants that endured low leaf water potential had rigid cell walls and low osmotic potential at full turgor. These plants did not have lower relative water content and turgor potential than plants with higher leaf water potential. There were three different responses to an increase in transpiration rate: (i) plants had low leaf water potential and slightly increased soil-to-leaf hydraulic conductance; (ii) plants decreased leaf water potential and increased the hydraulic conductance; and (iii) plants had high leaf water potential and largely increased the hydraulic conductance.