Relative growth rate,resource allocation and root morphology in the perennial legumes, <Emphasis Type="Italic">Medicago sativa</Emphasis>, <Emphasis Type="Italic">Dorycnium rectum</Emphasis> and <Emphasis Type="Italic">D. hirsutum</Emphasis> grown under controlled conditions

Perennial pastures are needed in farming systems in southern Australia to combat environmental problems such as dryland salinity. The mediterranean climate in southern Australia imposes constraints to the growth and survival of perennial plants. The aim of this study was to compare growth rates, resource allocation and root distribution in three perennial legumes, Medicago sativa L., Dorycnium hirsutum (L.) Ser. and Dorycnium rectum (L.) Ser., to identify different plant traits and their ecological and agronomic significance. Plants were grown in 1-m deep split tubes and destructive harvests were made every 2 weeks after plant emergence for 10 weeks. Leaf area and leaf, stem and root fresh and dry weights were measured. Maximum root depth and the root distribution were also determined. Seedlings of Dorycnium were slower to emerge and had a lower relative growth rate (RGR) than M. sativa. The slower RGR was associated with a lower specific leaf area (SLA) in D. hirsutum and a lower net assimilation rate (NAR) in D. rectum. Although all species allocated a similar proportion of biomass to roots, D. rectum had a shallower root distribution and took longer to produce deep roots. The slow growth rates of Dorycnium seedlings suggest that they are more prone to establishment problems due to competition from weeds or other pastures, and because they have less access to water at greater depth during summer drought. However, D. hirsutum displayed characteristics of a plant that is adapted to stressful environments and therefore may be able to grow in conditions where other perennial legumes cannot.     

Does transpiration control stomatal responses to water vapour pressure deficit?

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

Water relations and stomatal characteristics of Mediterranean plants with different growth forms and leaf habits: responses to water stress and recovery

The aim of this study was to extent the range of knowledge about water relations and stomatal responses to water stress to ten Mediterranean plants with different growth forms and leaf habits. Plants were subjected to different levels of water stress and a treatment of recovery. Stomatal attributes (stomatal density, StoD), stomatal conductance (g _s), stomatal responsiveness to water stress (SR), leaf water relations (pre-dawn and midday leaf water potential and relative water content), soil to leaf apparent hydraulic conductance (K _L) and bulk modulus of elasticity (ε) were determined. The observed wide range of water relations and stomatal characteristics was found to be partially depended on the growth form. Maximum g _s was related to StoD and the stomatal area index (SAI), while g _s evolution after water stress and recovery was highly correlated with K _L. Relationships between SR to water deficit and other morphological leaf traits, such as StoD, LMA or ε, provided no general correlations when including all species. It is concluded that a high variability is present among Mediterranean plants reflecting a continuum of leaf water relations and stomatal behaviour in response to water stress.     

Photosynthetic parameters and leaf water potential of five common bean genotypes under mild water deficit

The leaf water potential, gas exchange and chlorophyll fluorescence were evaluated in five common bean (Phaseolus vulgaris) genotypes A222, A320, BAT477, Carioca and Ouro Negro subjected to moderate water deficit. At the maximum water deficit (10 d of water withholding), the leaf water potential of genotypes A320 and A222 was higher (−0.35 and −0.50 MPa) when compared to the other genotypes (−0.67 to −0.77 MPa). The stomatal conductance and net photosynthetic rate were significantly reduced in all genotypes due to the water deficit. The greater reduction in stomatal conductance of A320 under drought resulted in high intrinsic water use efficiency. Mild water deficit affected the photochemical apparatus in bean genotypes probably by down-regulation since plants did not show photoinhibition. The photochemical apparatus of A222 and A320 genotypes was more sensitive to drought stress, showing reduced apparent electron transport even after the recovery of plant water status. On the other hand, even after 10 d of water withholding, the maximum efficiency of photosystem 2 was not affected, what suggest efficiency of the photoprotection mechanisms.     

Gas Exchange,Water Loss and Biomass Production in Rice and Wild Oryza Species in Well-Watered and Water-Limiting Growth Conditions

A major reason for the drought susceptibility of cultivated rice is the inability to regulate water loss as effectively as other cereals. We report studies of growth and gas exchange under well-watered and water-limiting conditions on selected species of Oryza and seek to relate this physiological information to the ecology of the genus. The immediate ancestor of the Oryza sativa cultigen is native to swamps and marshes, and wet habitats are typical for the genus as a whole. However, the ecological range of some species does extend to habitats that dry out seasonally. In all species studied growth was reduced by water deficit. In all species studied leaves had small absolute water contents and began to roll at a relative water content above 90%. However, there were species differences in leaf rolling and the response of stomatal conductance to an increase in vapour pressure deficit. Following re-watering, there were persistent reductions in stomatal conductance in most of the species tested, but the assimilation rate was not reduced in all of these cases. Where there was a persistant reduction in assimilation rate, there was also a reduction in carboxylation efficiency. It was a frequent observation that plants had a stomatal conductance greater than expected for their carbon assimilation rate; that is in the range where substantial changes in conductance have little effect upon photosynthesis. It is suggested that a reason for this may be the cooling of leaves which have a small thermal capacity in environments which often combine high temperature, humidity and irradiance. Large conductances combined with small water contents may be no disadvantage in the natural habitats of Oryza, but provide some reasons for the poor regulation of water loss in cultivated rice. Although there were significant differences in gas exchange amongst species the advantages that were observed over O. sativa were not of a magnitude likely to justify wide hybridisation. This implies that improvement in the drought resistance of rice is more likely to come from increasing water acquisition than from decreasing water loss.     

Tissue water relations in elfin cloud forest tree species of Serrania de Macuira,Guajira, Colombia

Summary Diurnal courses of stomatal conductance, leaf water potential, and the components of tissue water potential were measured in six canopy species in an elfin cloud forest. High values of stomatal conductance were measured on cloudy days and during early morning and late afternoon of sunny days. Decreases in stomatal conductance with increases in vapour pressure deficit may have been a response to avoid further water deficits and suggested a stomatal response to changes in relative humidity. Daily transpiration varied between 470 and 1014 g m^-2 day^-1 during cloudy days and between 532 and 944 g m^-2 day^-1 during clear days. Stomatal conductance may have also responded to changes in leaf water potential, which was minimum at noon. The minimum tissue water potential measured in the field was -1.8 MPa in Myrcianthes fragrans, and the minimum turgor pressure was 0.49 MPa also in M. fragrans. There was a correlation between the osmotic potential and the minimum tissue water potential, suggesting that osmotic potential plays a major role in the maintenance of turgor in these species, in spite of the great variability in the elastic properties of leaf tissues. Turgor pressure decreased during the day following the course of water potential but never approached the turgor loss point, as it has been measured in some lowland rain forest trees. This is a strong indication that elfin cloud forest trees do not suffer severe water deficits, and that small tree stature is not directly related to water shortage.     

The responses of stomata and leaf gas exchange to vapour pressure deficits and soil water content

Summary The responses of photosynthesis, transpiration and leaf conductance to changes in vapour pressure deficit were followed in well-watered plants of the herbaceous species, Helianthus annuus, Helianthus nuttallii, Pisum sativum and Vigna unguiculata, and in the woody species having either sclerophyllous leaves, Arbutus unedo, Nerium oleander and Pistacia vera, or mesomorphic leaves, Corylus avellana, Gossypium hirsutum and Prunus dulcis. When the vapour pressure deficit of the air around a single leaf in a cuvette was varied from 10 to 30 Pa kPa^-1 in 5 Pa kPa^-1 steps, while holding the remainder of the plant at a vapour presure deficit of 10 Pa kPa^-1, the leaf conductance and net photosynthetic rate of the leaf decreased in all species. The rate of transpiration increased initially with increase in vapour pressure deficit in all species, but in several species a maximum transpiration rate was observed at 20 to 25 Pa kPa^-1. Concurrent measurements of the leaf water potential by in situ psychrometry showed that an increase in the vapour pressure deficit decreased the leaf water potential in all species. The decrease was greatest in woody species, and least in herbaceous species. When the vapour pressure deficit around the remainder of the plant was increased while the leaf in the cuvette was exposed to a low and constant vapour pressure deficit, similar responses in both degree and magnitude in the rates of transpiration and leaf conductance were observed in the remainder of the plant as those occurring when the vapour pressure deficit around the single leaf was varied. Increasing the external vapour pressure deficit lowered the water potential of the leaf in the cuvette in the woody species and induced a decrease in leaf conductance in some, but not all, speies. The decrease in leaf conductance with decreasing water potential was greater in the woody species when the vapour pressure deficit was increased than when it remained low and constant, indicating that changing the leaf-to-air vapour pressure difference had a direct effect on the stomata in these species. The low hydraulic resistance and maintenance of a high leaf water potential precluded such an analysis in the herbaceous species. We conclude that at least in the woody species studied, an increase in the vapour pressure deficit around a leaf will decrease leaf gas exchange through a direct effect on the leaf epidermis and sometimes additionally through a lowering of the mesophyll water potential.     

Evaluation of the potential to measure photosynthetic rates in C3 plants (Flaveria pringlei and Oryza sativa) by combining chlorophyll fluorescence analysis and a stomatal conductance model

The response curves of leaf photosynthesis to varying light, temperature and leaf-to-air vapour pressure deficit were measured in the C₃ plants Flaveria pringlei and Oryza sativa in normal air with a computerized open infrared gas analysis (IRGA) system, and the photochemical efficiency of photosystem II, described as (1–F,/F′_m) after Genty. Briantais & Baker (1989, Biochimica et Biophysica Acta 990, 87–92), was simultaneously measured with a modulated fluorometer. A model was written for rates of CO₂ fixation as a function of the true rate of O₂ evolution measured by fluorescene analysis (Jo₂), mesophyll conductance and intercellular CO₂ partial pressure. A second model was developed for rates of CO₂ fixation as a function of Jo₂, mesophyll conductance and stomatal conductance. In the latter case, leaf stomatal conductance was simulated using the stomatal model proposed by Leuning (1995, Plant, Cell and Environment 18 , 339–355). The rates of CO₂ fixation predicted from the models were similar to rates measured by IRGA. The results indicate that there is potential to measure CO₂ fixation in C₃ plants by combining the non-invasive measurement of Jo₂ by chlorophyll fluorescence analysis with the stomatal conductance model.     

Variation in response to heavy metals during vegetative growth in <Emphasis Type="Italic">Dorycnium pentaphyllum</Emphasis> Scop.

Seeds, young plants and adult plants of the perennial Mediterranean leguminous shrub Dorycnium pentaphyllum Scop. were exposed to Cd (1–100 μM) or Zn (10–10,000 μM) on nutrient solution. This species is resistant to Cd and Zn at different phenological stages. The lowest doses of Zn and Cd improved seed germination and young seedling growth, while only the highest doses of both heavy metals inhibited germination and decreased growth. High doses of Cd reduced seed imbibition and young seedling water content, while Zn did not. Osmotic adjustment was more efficient in Zn-treated young plants than in Cd-treated ones, while chlorophyll concentrations decreased in the former but not in the latter. Those differences were not observed anymore in adult plants. Exclusion processes were more efficient at the adult stage than at the young seedling stage and were more marked in response to Zn than to Cd. It is concluded that D. pentaphyllum could be used for phytostabilization of heavy metal-contaminated areas. The physiological strategies of tolerance, however, differ according to the age of the plants and the nature of the metal.     

Stomatal Control of Water Loss in Siratro (Macroptilium atropurpureum (DC) Urb.), a Tropical Pasture Legume

Stomatal conductance of siratro declined linearly as leaf waterpotential fell until zero conductance was reached at –10bar. In a grass/legume pasture stomata of siratro respondedto humidity (saturation deficit), and to a lesser extent toleaf water potential, such that leaf water potential did notfall below –9 bar, whereas that of the grass continuedto decline for most of the day. The dual response of siratroto both humidity and leaf water potential suggests that thisspecies has an efficient two-stage stomatal control of waterloss which provides an explanation of its higher leaf waterpotential and greater drought avoidance compared with sown grassesin semi-arid areas of north-eastern Australia. Macroptilium atropurpureum (DC) Urb., siratro, Desmodium uncinatum, stomatal control, stomatal conductance, water loss, leaf water potential, drought avoidance, saturation deficit     

Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought

A comparative study on stomatal control under water deficit was conducted on grapevines of the cultivars Grenache, of Mediterranean origin, and Syrah of mesic origin, grown near Montpellier, France and Geisenheim, Germany. Syrah maintained similar maximum stomatal conductance (g_max) and maximum leaf photosynthesis (A_max) values than Grenache at lower predawn leaf water potentials, Ψ_leaf, throughout the season. The Ψ_leaf of Syrah decreased strongly during the day and was lower in stressed than in watered plants, showing anisohydric stomatal behaviour. In contrast, Grenache showed isohydric stomatal behaviour in which Ψ_leaf did not drop significantly below the minimum Ψ_leaf of watered plants. When g was plotted versus leaf specific hydraulic conductance, K_l, incorporating leaf transpiration rate and whole‐plant water potential gradients, previous differences between varieties disappeared both on a seasonal and diurnal scale. This suggested that isohydric and anisohydric behaviour could be regulated by hydraulic conductance. Pressure‐flow measurements on excised organs from plants not previously stressed revealed that Grenache had a two‐ to three‐fold larger hydraulic conductance per unit path length (K_h) and a four‐ to six‐fold larger leaf area specific conductivity (LSC) in leaf petioles than Syrah. Differences between internodes were only apparent for LSC and were much smaller. Cavitation detected as ultrasound acoustic emissions on air‐dried shoots showed higher rates for Grenache than Syrah during the early phases of the dry‐down. It is hypothesized that the differences in water‐conducting capacity of stems and especially petioles may be at the origin of the near‐isohydric and anisohydric behaviour of g.     

Stomatal control of transpiration in the canopy of a clonal Eucalyptus grandis plantation

 Predawn leaf water potential, stomatal conductance and microclimatic variables were measured on 13 sampling days from November 1995 through August 1996 to determine how environmental and physiological factors affect water use at the canopy scale in a plantation of mature clonal Eucalyptus grandis Hill ex-Maiden hybrids in the State of Espirito Santo, Brazil. The simple ”big leaf” Penman-Monteith model was used to estimate canopy transpiration. During the study period the predawn leaf water potential varied from –0.4 to –1.3 MPa, with the minimum values observed in the winter months (June and August 1996), while the average estimated values for canopy conductance and canopy transpiration fell from 17.3 to 5.8 mm s^–1 and from 0.54 to 0.18 mm h^–1, respectively. On the basis of all measurements, the average value of the decoupling coefficient was 0.25. During continuous soil water shortage a proportional reduction was observed in predawn leaf water potential and in daily maximum values of stomatal conductance, canopy transpiration and decoupling coefficient. The results showed that water vapour exchange in this canopy is strongly dominated by the regional vapour pressure deficit and that canopy transpiration is controlled mainly by stomatal conductance. On a seasonal basis, stomatal conductance and canopy transpiration were mainly related to predawn leaf water potential and, thus, to soil moisture and rainfall. Good results were obtained with a multiplicative empirical model that uses values of photosynthetically active radiation, vapour pressure deficit and predawn leaf water potential to estimate stomatal conductance. Received: 10 June 1998 / Accepted: 20 July 1998     

Field photosynthesis,microclimate and water relations of an exotic temperate liana,Pueraria lobata,kudzu

Summary Kudzu occurs in a variety of habitats in the southeastern United States. It is most common in exposed, forest edge sites and road cuts where it forms an extensive ground canopy as well as a canopy overtopping nearby trees, but it can also be found in completely open fields and deeply shaded sites within a forest. Microclimate, stomatal conductance, leaf water potential and photosynthetic responses to light, temperature and humidity were measured in two contrasting microhabitats on Pueraria lobata, kudzu. Midsummer leaf temperatures and leaf-to-air water vapor deficits for plants growing in an exposed site were significantly greater than for those in a shaded site, exceeding 35° C and 50 mmol mol^-1, respectively. Maximum stomatal conductance exceeded 400 mmol m^-2 s^-1 in exposed leaves during peak vegetative growth. Stomatal conductance in shaded leaves was approximately half the value measured in exposed leaves on any particular dya. Maximum photosynthetic carbon uptake was also higher in leaves growing in exposed sites compared to leaves in shaded sites, exceeding 18.7 and 14.0 mol m^-2 s^-1, respectively. Photosynthesis, stomatal conductance and intercellular CO₂ concentration decreased dramatically in response to increasing water vapor deficit for leaves from both sites. However, transpiration showed an initial increase at intermediate water vapor deficits, leveling off or even decreasing at higher values. Leaf water potential demonstrated marked diurnal variation, but remained constant over a wide range of transpirational water fluxes. This latter feature, combined with microenvironmental modification through rapid leaf orientation and pronounced stomatal responses to water vapor deficits may represent important adaptive responses in the exploitation of a diverse array of habitats by kudzu.     

Hydraulic Conductance and Stomatal Sensitivity to Changes of Leaf Water Status in Six Deciduous Tree Species

The relationship between shoot hydraulic conductance (L) and stomatal sensitivity to changes in leaf water status was studied in the saplings of six deciduous tree species. L increased significantly in sequence: Acer platanoides < Tilia Cordata < Padus avium = Quercus robur < Salix caprea = Populus tremula. L was higher in the trees grown in soil with a higher nitrogen content and lower in the trees grown under mild water stress or kept in darkness for several days. L was higher in July than in September in all the species. L correlated positively with maximum photosynthesis, stomatal conductance and stomatal sensitivity to an increase in leaf water potential, but negatively with stomatal sensitivity to a decrease in leaf water potential. The correlations between L and any other parameter were approximated by three different curves: data for water-stressed plants fit to the first, data for plants kept in darkness fit to the second and all the other data fit to the third curve. The reasons of the differences of shoot hydraulic conductance in the different experimental sets and the mechanisms which may cause the correlation between L and the other characteristics are discussed.     

S‐nitrosoglutathione spraying improves stomatal conductance,Rubisco activity and antioxidant defense in both leaves and roots of sugarcane plants under water deficit

Water deficit is a major environmental constraint on crop productivity and performance and nitric oxide (NO) is an important signaling molecule associated with many biochemical and physiological processes in plants under stressful conditions. This study aims to test the hypothesis that leaf spraying of S‐nitrosoglutathione (GSNO), an NO donor, improves the antioxidant defense in both roots and leaves of sugarcane plants under water deficit, with positive consequences for photosynthesis. In addition, the roles of key photosynthetic enzymes ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPC) in maintaining CO₂ assimilation of GSNO‐sprayed plants under water deficit were evaluated. Sugarcane plants were sprayed with water or GSNO 100 μM and subjected to water deficit, by adding polyethylene glycol (PEG‐8000) to the nutrient solution. Sugarcane plants supplied with GSNO presented increases in the activity of antioxidant enzymes such as superoxide dismutase in leaves and catalase in roots, indicating higher antioxidant capacity under water deficit. Such adjustments induced by GSNO were sufficient to prevent oxidative damage in both organs and were associated with better leaf water status. As a consequence, GSNO spraying alleviated the negative impact of water deficit on stomatal conductance and photosynthetic rates, with plants also showing increases in Rubisco activity under water deficit.     

Water use efficiency by alfalfa: Mechanisms involving anti-oxidation and osmotic adjustment under drought

Water use and mechanisms relating to osmotic adjustment and anti-oxidation were investigated in alfalfa (Medicago sativa L.) plants under reduced water availability. Water use efficiency (WUE), MDA and proline contents, and antioxidant enzyme activities were measured in three alfalfa cultivars under three levels of soil water availability in a greenhouse pot experiment. WUE was determined indirectly using discriminating carbon isotope composition. WUE increased with the severity of water deficit. Under all water regimes examined, cv. Longdong showed the greatest WUE values and the least reduction in biomass production under a 50% soil field water capacity. Stomatal density increased with increasing water deficit, but stomatal conductance decreased. This suggests that water stress can increase WUE by modifying stomatal regulation of the balance between the rates of CO₂ assimilation and water loss. The maintenance of leaf physiological function and leaf water status suggests that alfalfa has some mechanisms to maintain cell function when the plant is subjected to water deficit. The increase in the MDA content under drought conditions indicates that some degree of damage to cell membranes is unavoidable, whereas other results showing increases in the contents of proline and soluble sugars and activities of superoxide dismutase, peroxide dismutase, and catalase indicate how cell function may be to some extent maintained to result in the higher WUE. Alfalfa is shown to exhibit cultivar-specific differences in WUE with the maintenance of cell function under water deficit being related to anti-oxidation and osmotic adjustment.     

Effects of nursery preconditioning through mycorrhizal inoculation and drought in Arbutus unedo L. plants

The influence of a water deficit treatment and mycorrhizal inoculation with Pisolithus tinctorius (Pers.) Coker and Couch on the water relations, gas exchange, and plant growth in Arbutus unedo L. plants was studied in order to evaluate the hardening process during the nursery period. The ability to withstand the adverse conditions after transplantation was also studied. Mycorrhizal and non-mycorrhizal seedlings of A. unedo were pot-grown for 4 months in a greenhouse (nursery period), during which time two irrigation treatments, well watered (100% water holding capacity, leaching 20% of the applied water) and deficit irrigation (50% of the well watered), were applied. Subsequently, the plants were transplanted to the field and well irrigated (transplanting period), after which and until the end of the experiment they received no water (establishment period). At the end of the nursery period, both water deficit and mycorrhizae were seen to have altered the plant morphology. Mycorrhizal plants had lower leaf area and improved leaf color parameters, while the water deficit increased root dry weight and the root/shoot ratio. Mycorrhizal plants had higher leaf water potential values than non-inoculated plants. Mycorrhizae increased stomatal conductance and photosynthesis values, especially in stressed plants. Drought led to an osmotic adjustment and a decrease in the leaf water potential values at turgor loss point in the mycorrhizal plants. Cell wall rigidity, measured as increased bulk modulus of elasticity, was decreased by the mycorrhizae effect. After transplanting, no differences were found in the water relations or gas exchange values between treatments. During the establishment period, the plants that had been exposed to both drought and mycorrhizae showed a better water status (higher leaf water and turgor potential values) and higher gas exchange values. In conclusion, water deficit and mycorrhizal inoculation of A. unedo plants in nursery produced changes in tissue water relations, gas exchange, and growth, related with the acclimation process in the seedlings, which could provide better resistance to drought and stress conditions following planting.     

Responses of Ribulose-1,5-Bisphosphate Carboxylase,Protein Content,and Stomatal Conductance to Water Deficit in Maize,Tomato, and Bean

We compared responses of maize, tomato, and bean plants to water stress. Maize reached a severe water deficit (leaf water potential –1.90 MPa) in a longer period of time as compared with tomato and bean plants. Maize stomatal conductance (g _s) decreased at mild water deficit. g _s of tomato and bean decreased gradually and did not reach values as low as in maize. The protein content was maintained in maize and decreased at low water potential (_w); in tomato it fluctuated and also decreased at low _w; in bean it gradually decreased. Ribulose-1,5-bisphosphate carboxylase/oxygenase activity remained high at mild and moderate stress in maize and tomato plants; in bean it remained high only at mild stress.     

The relations of stomatal closure and reopening to xylem ABA concentration and leaf water potential during soil drying and rewatering

Two tropical tree species, Acacia confusa and Leucaena leucocephala, were used to study the relationships among stomatal conductance, xylem ABA concentration and leaf water potential during a soil drying and rewatering cycle. Stomatal conductance of both A. confusa and L. leucocephala steadily decreased with the decreases in soil water content and pre-dawn leaf water potential. Upon rewatering, soil water content and pre-dawn leaf water potential rapidly returned to the control levels, whereas the reopening of stomata showed an obvious lag time. The length of this lag time was highly dependent not only upon the degree of water stress but also on plant species. The more severe the water stress, the longer the lag time. When A. confusa and L. leucocephala plants were exposed to the same degree of water stress (around –2.0 MPa in pre-dawn leaf water potential), the stomata of A. confusa reopened to the control level 6 days after rewatering. However, it took L. leucocephala about 14 days to reopen fully. A very similar response of leaf photosynthesis to soil water deficit was also observed for both species. Soil drying resulted in a significant increase in leaf and xylem ABA concentrations in both species. The more severe the water stress, the higher the leaf and xylem ABA concentrations. Both leaf ABA and xylem ABA returned to the control level following relief from water deficit and preceded the full recovery of stomata, suggesting that the lag phase of stomatal reopening was not controlled by leaf and/or xylem ABA. In contrast to drying the whole root system, drying half of the root system did not change the leaf water relations, but caused a significant increase in xylem ABA concentration, which could fully explain the decrease of stomatal conductance. After rewatering, the stomatal conductance of plants in which half of the roots were dried recovered more rapidly than those of whole-root dried plants, indicating that the leaf water deficit that occurred during the drying period was related to the post-stress stomatal inhibition. These results indicated that the decrease in stomatal conductance caused by water deficit was closely related to the increase in xylem ABA, but xylem ABA could not fully explain the reopening of stomata after relief of water stress, neither did the leaf ABA. Some unknown physiological and/or morphological processes in the guard cells may be related to the recovery process.     

Stomatal conductance tracks soil-to-leaf hydraulic conductance in faba bean and maize during soil drying

Although regulation of stomatal conductance is widely assumed to be the most important plant response to soil drying, the picture is incomplete when hydraulic conductance from soil to the leaf, upstream of the stomata, is not considered. Here, we investigated to what extent soil drying reduces the conductance between soil and leaf, whether this reduction differs between species, how it affects stomatal regulation, and where in the hydraulic pathway it occurs. To this end, we noninvasively and continuously measured the total root water uptake rate, soil water potential, leaf water potential, and stomatal conductance of 4-week-old, pot-grown maize (Zea mays) and faba bean (Vicia faba) plants during 4 days of water restriction. In both species, the soil–plant conductance, excluding stomatal conductance, declined exponentially with soil drying and was reduced to 50% above a soil water potential of −0.1 MPa, which is far from the permanent wilting point. This loss of conductance has immediate consequences for leaf water potential and the associated stomatal regulation. Both stomatal conductance and soil–plant conductance declined at a higher rate in faba bean than in maize. Estimations of the water potential at the root surface and an incomplete recovery 22 h after rewatering indicate that the loss of conductance, at least partly, occurred inside the plants, for example, through root suberization or altered aquaporin gene expression. Our findings suggest that differences in the stomatal sensitivity among plant species are partly explained by the sensitivity of root hydraulic conductance to soil drying.

The hydraulic conductance between soil and leaf decreases exponentially with decreasing soil water potential, at a species-specific rate related to the decline rate of stomatal conductance.