Sensitivity of water relations and photosynthesis to summer precipitation pulses for <Emphasis Type="Italic">Artemisia tridentata</Emphasis> and <Emphasis Type="Italic">Purshia tridentata</Emphasis>

For much of the western USA, precipitation occurs in pulses, the nature of which determine soil water potential and plant physiological performance. This research utilized three experiments to examine the sensitivity of photosynthesis and water relations for two widespread Great Basin Desert shrub species, Artemisia tridentata (which has both deep and shallow roots) and Purshia tridentata (which reportedly has only deep roots), to (1) variation in pulse magnitude size, (2) the kinetics of responses to pulses, and (3) the relationship between pulse-size and antecedent soil water content. At the study site in the southwestern Great Basin Desert, USA, summer rainfall exhibits a greater frequency of larger-sized events, and longer inter-pulse intervals, compared to annual patterns. Compared to pre-watering values, stem water potential initially increased by about 2.00 MPa for A. tridentata and 1.00 MPa for P. tridentata following watering to simulate an 11.5 mm rainfall pulse. For the same water addition, stomatal conductance increased by 0.3 mol m⁻² s⁻¹ and photosynthetic CO₂ assimilation increased 8-fold for A. tridentata and 6-fold for P. tridentata. Water potential and photosynthetic gas exchange were maximal for both species 2–3 days following a pulse addition. In comparison to P. tridentata, the increase in photosynthesis for A. tridentata was more pronounced for plants treated incrementally with several small pulses compared to plants treated with one pulse of an equivalent total volume. The results indicate that both species can respond to a range of summer rainfall pulse magnitudes within about 2 days, with A. tridentata generally exhibiting larger responses in comparison to the co-dominant shrub species P. tridentata, which at this study site does indeed have shallow roots. In a future climate, the timing and magnitude of summer rainfall pulses will determine the extent to which these two species undergo changes in water status and photosynthetic carbon uptake, with implications for their fitness.     

Comparative water relations of four Mediterranean oak species

The water relations and responses of two evergreen (Quercus ilex L. and Q. suber L.) and two deciduous (S. afares Pomel. and Q. faginea Will.) Quercus species were studied under experimental conditions. Two-year old seedlings grown in 30 l. pots were subjected to a drying period during which stomatal conductance, pre-dawn potential and minimum foliar potential were measured.The results shows that, for all species, the daily course of stomatal conductance agrees with the patterns proposed by Hinckley et al. (1978 & 1983). Concurrent with the species responses to short-term variation in water availability, it was found that pre-dawn leaf water potential controlled the maximum daily leaf conductance. There was a strong correlation between pre-dawn leaf potential and maximum daily conductance as described by the reciprocal function g_{srmax for}=(-0.47+2.61._p)^-1 the evergreen oaks and g_{srmax for}=(-1.94+7.39._p)^-1 for the deciduous species. These differences between the two groups may partialy explain their geograhic distributions, and suggest general questions concerning the mechanisms which optimize water-use efficiency in Mediterranean oak species.     

Can water responses in Stipa tenacissima L. during the summer season be promoted by non-rainfall water gains in soil?

This article reports on quantified soil water gains and their possible effects on summer water relationships in a semiarid Stipa tenacissima L. grasslands located in SE Spain. We believe that the net soil water gains detected using minilysimeters could be from soil water vapour adsorption (WVA). Our study of high water-stress showed stomatal conductance (21.8–43.1 mmol H₂O m⁻² s⁻¹) in S. tenacissima leaves unusual for the summer season, and the evapotranspiration from S. tenacissima grassland, estimated by a multi-source sparse evapotranspiration model, closely corresponding to total WVA. This highlights the importance of summer soil WVA to stomatal conductance and vital transpiration in S. tenacissima. This study measured pre-dawn leaf water potential (ψ) response to sporadic light rainfall, finding that a light summer rainfall (1.59 mm day⁻¹) was sufficient to vary ψ in S. tenacissima from −3.8 (close to the turgour loss point) to −2.7 MPa. We hypothesize that soil WVA can supply vegetation with water vital to its survival in seasons with a severe water deficit, giving rise to a close relationship between soil water dynamics and plant water response.     

Indications of Hydraulic Lift by Pinus halepensis and Its Effects on the Water Relations of Neighbour Shrubs

We measured the stable deuterium isotopic composition of xylem sap, the shoot predawn and midday water potentials, and the leaf δ¹³C of Mediterranean shrubs Pistacia lentiscus, Globularia alypum and Rosmarinus officinalis in a south-oriented transect from a large (12 m tall) Aleppo pine tree, Pinus halepensis. We aimed to study the possibility of hydraulic lift from the deep roots of this pine tree to the shallow soil layers and its influence on these neighbour shrubs. These same traits were also studied in several individuals of the shrub Pistacia lentiscus growing with different types of neighbours: just shrubs, a small (3 – 4 m) pine tree, and the above mentioned large pine tree. The greater the distance from P. halepensis the plants grew, the higher xylem water δD, the lower the soil water content, and, the lower the predawn and midday water potentials were found. These results suggest the existence of an hydraulic lift from deep roots to shallow soil in this big tree. Further indication of this existence is provided by the improved water status of P. lentiscus (higher water potentials and δD, and lower δ¹³C and, therefore, lower water use efficiencies) when growing close to the big pine in comparison with the same shrub species growing close to small pines or just surrounded by other shrubs. Moreover, all these trends occurred in the dry summer season, but disappeared in the wet spring season. This revised version was published online in July 2006 with corrections to the Cover Date.     

Photosynthetic responses of Mojave Desert shrubs to free air CO2 enrichment are greatest during wet years

It has been suggested that desert vegetation will show the strongest response to rising atmospheric carbon dioxide due to strong water limitations in these systems that may be ameliorated by both photosynthetic enhancements and reductions in stomatal conductance. Here, we report the long‐term effect of 55 Pa atmospheric CO₂ on photosynthesis and stomatal conductance for three Mojave Desert shrubs of differing leaf phenology (Ambrosia dumosa—drought‐deciduous, Krameria erecta—winter‐deciduous, Larrea tridentata—evergreen). The shrubs were growing in an undisturbed ecosystem fumigated using FACE technology and were measured over a four‐year period that included both above and below‐average precipitation. Daily integrated photosynthesis (A_day) was significantly enhanced by elevated CO₂ for all three species, although Krameria erecta showed the greatest enhancements (63% vs. 32% for the other species) enhancements were constant throughout the entire measurement period. Only one species, Larrea tridentata, decreased stomatal conductance by 25–50% in response to elevated CO₂, and then only at the onset of the summer dry season and following late summer convective precipitation. Similarly, reductions in the maximum carboxylation rate of Rubisco were limited to Larrea during spring. These results suggest that the elevated CO₂ response of desert vegetation is a function of complex interactions between species functional types and prevailing environmental conditions. Elevated CO₂ did not extend the active growing season into the summer dry season because of overall negligible stomatal conductance responses that did not result in significant water conservation. Overall, we expect the greatest response of desert vegetation during years with above‐average precipitation when the active growing season is not limited to ～ 2 months and, consequently, the effects of increased photosynthesis can accumulate over a biologically significant time period.     

Effects of plant size on photosynthesis and water relations in the desert shrub Prosopis glandulosa (Fabaceae)

The Jornada del Muerto basin of the Chihuahuan Desert of southern New Mexico, USA, has undergone a marked transition of plant communities. Shrubs such as mesquite (Prosopis glandulosa) have greatly increased or now dominate in areas that were previously dominated by perennial grasses. The replacement of grasses by shrubs requires an establishment phase where small shrubs must compete directly with similar-sized grass plants. This is followed by a phase in which large, established shrubs sequester nutrients and water within their biomass and alter soil resources directly under their canopy, creating “islands” of fertility. We hypothesized that these two phases were associated with shrubs having different physiological response capacities related to their age or size and the resource structure of the environment. As a corollary, we hypothesized that responses of small shrubs would be more tightly coupled to variation in soil moisture availability compared to large shrubs. To test these hypotheses, we studied gas exchange and water relations of small (establishing) and large (established) shrubs growing in the Jornada del Muerto as a function of varying soil moisture during the season. The small shrubs had greater net assimilation, stomatal conductance, transpiration, and xylem water potential than large shrubs following high summer rainfall in July, and highest seasonal soil moisture at 0.3 m. High rates of carbon assimilation and water use would be an advantage for small shrubs competing with grasses when shallow soil moisture was plentiful. Large shrubs had greater net assimilation and water-use efficiency, and lower xylem water potential than small shrubs following a dry period in September, when soil moisture at 0.3 m was lowest. Low xylem water potentials and high water-use efficiency would allow large shrubs to continue acquiring and conserving water as soil moisture is depleted. Although the study provides evidence of differences in physiological responses of different-sized shrubs, there was not support for the hypothesis that small shrubs are more closely coupled to variation in soil moisture availability than large shrubs. Small shrubs may actually be less coupled to soil moisture than large shrubs, and thus avoid conditions when continued transpiration could not be matched by equivalent water uptake.     

PHYSIOLOGICAL CONTROL OF CHAMISE SHOOT GROWTH AFTER FIRE

Five controlled burns (ca. 2 ha each) were conducted in the Coast Range of northern California near Hopland, California, between November 18, 1975 and September 15, 1976 to determine the effect of shrub phenology on the sprouting response of chamise (Adenostoma fasciculatum H&A) following fire. The time of burn had little effect on the amount of shoot growth that occurred after fire, although the pattern of growth was altered. Shrubs burned on June 22 or August 2, 1976 grew continually until August 1977, while unburned shrubs or those burned at other times ceased seasonal growth during the first summer after fire. Neither the amount nor the pattern of shoot growth was influenced by the shrub water status before the fire occurred. The relationship between chamise shoot growth after fire, ¹⁴CO₂ fixation, xylem sap tension, and air temperature was also studied. The growth rate of regrowing shrubs was highly correlated with ¹⁴CO₂ fixation prior to the growth measurement. Water status (pre-dawn xylem sap tension) had a negative correlation with ¹⁴CO₂ fixation. The abundance of carbohydrate at the shoot apex is believed to influence the seasonal pattern and rate of chamise shoot growth following fire.     

The Influence of Spatial Patterns of Soil Moisture on the Grass and Shrub Responses to a Summer Rainstorm in a Chihuahuan Desert Ecotone

The cycling of surface water, energy, nutrients, and carbon is different between semiarid grassland and shrubland ecosystems. Although differences are evident when grasslands are compared to shrublands, the processes that contribute to this transition are more challenging to document. We evaluate how surface redistribution of precipitation and plant responses to the resulting infiltration patterns could contribute to the changes that occur during the transition from grassland to shrubland. We measured soil water potential under grasses (Bouteloua eriopoda), shrubs (Larrea tridentata) and bare soil and changes in plant water relations and gas exchange following a 15 mm summer storm in the grassland–shrubland ecotone at the Sevilleta National Wildlife Refuge in central New Mexico USA. Following the storm, soil water potential (Ψ_s) increased to 30 cm depth beneath both grass and shrub canopies, with the greatest change observed in the top 15 cm of the soil. The increase in Ψ_s was greater beneath grass canopies than beneath shrub canopies. Ψ_s under bare soil increased only to 5 cm depth. The substantial redistribution of rainfall and different rooting depths of the vegetation resulted in high Ψ_s throughout most of the rooting volume of the grasses whereas soil moisture was unchanged throughout a large portion of the shrub rooting volume. Consistent with this pattern, predawn water potential (Ψ_PD) of grasses increased more than 5 MPa to greater than −1 MPa whereas Ψ_PD of shrubs increased to −2.5 MPa, a change of less than 2 MPa. Transpiration increased roughly linearly with Ψ_PD in both grasses and shrubs. In grasses, assimilation was strongly correlated with Ψ_PD whereas there was no relationship in shrubs where assimilation showed no significant response to the pulse of soil moisture following the storm. These data show that preferential redistribution of water to grass canopies enhances transpiration and assimilation by grasses following large summer storms. This process may inhibit shrubland expansion at the ecotone during periods without extreme drought.     

Effects of manipulation of water and nitrogen regime on the water relations of the desert shrub Larrea tridentata

Summary Water and nitrogen regimes of Larrea tridentata shrubs growing in the field were manipulated during an annual cycle. Patterns of leaf water status, leaf water relations characteristics, and stomatal behavior were followed concurrently. Large variations in leaf water status in both irrigated and nonirrigated individuals were observed. Predawn and midday leaf water potentials of nonirrigated shrubs were lowest except when measurements had been preceded by significant rainfall. Despite the large seasonal variation in leaf water status, reasonably constant, high levels of turgor were maintained. Pressure-volume curve analysis suggested that changes in the bulk leaf osmotic potential at full turgor were small and that nearly all of the turgor adjustment was due to tissue elastic adjustment. The increase in tissue elasticity with increasing water deficit manifested itself as a decrease in the relative water content at zero turgor and as a decrease in the tissue bulk elastic modulus. Because of large hydration-induced displacement in the osmotic potential and relative water content at zero turgor, it was necessary to use shoots in their natural state of hydration for pressure-volume curve determinations. Large diurnal and seasonal differences in maximum stomatal conductance were observed, but could not easily be attributed to variations in leaf water potential or leaf water relations characteristics such as the turgor loss point. The single factor which seemed to account for most of the diurnal and seasonal differences in maximum stomatal conductance between individual shrubs was an index of soil/root/ shoot hydraulic resistance. Daily maximum stomatal conductance was found to decrease with increasing soil/root/ shoot hydraulic resistance. This pattern was most consistent if the hydraulic resistance calculation was based on an estimate of total canopy transpiration rather than the more commonly used transpiration per unit leaf area. The reasons for this are discussed. It is suggested that while stomatal aperture necessarily represents a major physical resistance controlling transpiration, plant hydraulic resistance may represent the functional resistance through its effects on stomatal aperture.     

Gas exchange and water relations of two Rocky Mountain shrub species exposed to a climate change manipulation

Gas exchange and water relations responses to warming were compared for two shrub species, Artemisia tridentata spp. vaseyana (Asteraceae), a widely distributed evergreen species of the Great Basin and the western slope of the Rocky Mountains, and Pentaphylloides floribunda (Rosaceae), a deciduous shrub limited in distribution to moist, high-elevation meadows. Plants were exposed to an in situ infrared (IR) climate change manipulation at the Rocky Mountain Biological Laboratory, near Crested Butte, CO. Measurements of gas exchange and water relations were made on the two species in July and August, 1993 from plants growing in situ in infrared-heated and control plots. Carbon dioxide uptake, water loss, leaf temperature, water use efficiency, and water potential were compared to test the hypothesis that leaf and soil responses to IR will cause leaf level changes in photosynthesis. Photosynthetic CO₂ uptake and water use efficiency increased for A. tridentata (2.9 vs. 1.9 mol m^–2 s^–1 and 1.2 vs. 0.7 mmol C/mol H₂O) in the heated plots compared to the controls, while water potential was significantly lower in the heated plots (–1.1 vs. –0.5 MPa). The heating treatment decreased rates of photosynthesis for P. floribunda, but not significantly so. For A. tridentata, the results are consistent with the community-level changes observed with heating. Taken together, the evidence suggests that global warming is likely to result in increasing dominance of A. tridentata in subalpine meadow habitat now dominated by forbs.     

Water relations and growth of shrubs before and after fire in a semi-arid woodland

Summary Plant water relations and shoot growth rate of shrubs resprouting after fire or unburnt were measured in a semi-arid poplar box (Eucalyptus populnea) shrub woodland of eastern Australia. In vegetation unburnt for about 60 years, the dawn xylem water potential (_x) of the dominant shrub species was about-1.0 MPa when the soil was wet and-8.0 MPa when the soil was very dry. At any one time, the dominant shrub species,Eremophila mitchellii, E. sturtii, Geijera parviflora andCassia nemophila, were similar in _x butAcacia aneura andDodonaea viscosa were consistently higher in _x than this group when the soil was moist and lower when the soil was dry. The dominant tree species,Eucalyptus populnea andE. intertexta, appeared to have access to additional water beneath the hardpan which is located 60–80 cm below the surface. When shrubs were under extreme water stress (_x of-8 MPa), the trees had a _x of-3 to-3.6 MPa. Following a fire, both _x and leaf stomatal conductance (g _s) of resprouting shrubs were higher for about 5 years than comparable-aged unburnt vegetation, with relative differences in _x increasing with drought stress. Elongation rate of resprouts was positively linked to prefire shrub height in 3 of 4 species. However, shrubs resprouting after high intensity fires had substantially higher rates of shoot elongation than after low intensity fires which were in turn higher than for foliar expansion of unburnt shrubs. It is concluded that the growth rate of resprouting shrubs is primarily determined by physiological/ morphological factors associated with plant size but is also assisted by greater availability of water and possibly nutrients for a period after fire.     

Detecting the Differences in Responses of Stomatal Conductance to Moisture Stresses between Deciduous Shrubs and Artemisia Subshrubs

Shrubs and subshrubs can tolerate wider ranges of moisture stresses in both soil and air than other plant life forms, and thus represent greater nonlinearity and uncertainty in ecosystem physiology. The objectives of this paper are to model shrub/subshrub stomatal conductance by synthesizing the field leaf gas exchanges data of 24 species in China, in order to detect the differences between deciduous shrubs and Artemisia subshrubs in their responses of stomatal conductance to changes in the moisture stresses. We revised a model of stomatal conductance by incorporating the tradeoff between xylem hydraulic efficiency and cavitation loss risk. We then fit the model at the three hierarchical levels: global (pooling all data as a single group), three functional groups (deciduous non-legume shrubs, deciduous legume shrubs, and subshrubs in Artemisia genus), and individual observations (species × sites). Bayesian inference with Markov Chain Monte Carlo method was applied to obtain the model parameters at the three levels. We found that the model at the level of functional groups is a significant improvement over that at the global level, indicating the significant differences in the stomatal behavior among the three functional groups. The differences in tolerance and sensitivities to changes in moisture stresses are the most evident between the shrubs and the subshrubs: The two shrub groups can tolerate much higher soil water stress than the subshrubs. The analysis at the observation level is also a significant improvement over that at the functional group level, indicating great variations within each group. Our analysis offered a clue for the equivocal issue of shrub encroachment into grasslands: While the invasion by the shrubs may be irreversible, the dominance of subshrubs, due to their lower resistance and tolerance to moisture stresses, may be put down by appropriate grassland management.     

Water relations of crok-oak (Quercus suber L.) under natural conditions

Daily and annual courses of leaf transpiration, stomatal conductance and shoot water potential of four Quercus suber individuals were compared in a semi-natural stand in southwest Portugal, from spring 1989 to early summer 1990.The trees investigated showed annual patterns typical of evergreen sclerophyllous species but varied in their range of stomatal operation. This appeared to be related to differences in hydraulic conductivity in the root-to-leaf pathway.Maximum stomatal conductance and transpiration rates occurred from March to June.Water stress was found to be moderate and winter cold stress due to low air and soil temperatures appeared to have an influence on plant water balance through their effects on flow resistances.Abbreviations g_sw stomatal conductance - g_max maximum stomatal conductance - PAR photosynthetically active radiation - RH relative humidity of the air - T leaf transpiration - Ta air temperature - TL leaf temperature - T_max maximum leaf transpiration - W air-to-leaf vapor pressure difference - shoot water potential - PD predawn shoot water potential - MIN minimum shoot water potential     

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.     

Gas exchange of a desert shrub (Zygophyllum dumosum Boiss.) under different soil moisture regimes during summer drought

The effects of soil water potential on photosynthesis and transpiration of whole Zygophyllum dumosum Boiss. shrubs were examined with a field IRGA system during a rainless summer. Daily photosynthesis and transpiration activities were not notably different on a unit phyllode area basis among shrubs at naturally differing soil water potentials. Irrigation of shrubs caused phyllodes to increase significantly in water content and new leaflets to appear. Leaflets had three times as many stomata per unit area (23000 stomata cm^-2) as phyllodes (7100 stomata cm^-2) but photosynthesis and transpiration rates were not measurably different between irrigated and non-irrigated shrubs on a unit area basis. This finding suggests that sufficient soil moisture will lead to increased carbon uptake of the entire shrub simply because the total area of photosynthesizing tissue increases. Gas exchange rates appear to be controlled solely by atmospheric conditions under the stresses of summer.     

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 Reactions of Two Different Maize Lines to Osmotically Induced Drought Stress

Two maize lines differing in drought resistance were grown at different drought stress induced by polyethylene glycol (PEG 10 000) solutions with osmotic potentials of –0.20, –0.40 and –0.80 MPa in the semipermeable membrane system. During the five days soil water content decreased (from 0.43 to 0.29, 0.25 and 0.23 g cm^–3 for three PEG solutions, respectively) as well as leaf water potentials (_w; from – 0.54 to –0.76, –1.06 and –1.46 MPa). These values were not significantly different between the investigated lines, indicating that a controlled and consistent soil moisture stress was achieved. Soil drying induced an increase in the ABA content of leaves and xylem of both lines and the effects on stomatal conductance were greater in drought susceptible line (B-432) compared to drought resistant line (ZPBL-1304). To test possible difference in stomatal sensitivity to xylem ABA between lines and to assess any ABA vs. _w interaction, roots were fed with 10, 50 and 100 mmol m^–3 ABA solutions in another set of experiments. These results showed that manipulation of xylem ABA affected stomata of both lines similarly. Comparison of stomatal sensitivity to drought-induced and applied ABA demonstrated that drought treatment affected stomata of investigated lines by differentially increasing their sensitivity to xylem ABA, thus confirming an interaction between chemical signalling and hydraulic signalling.     

Growth response of barley and tomato to nitrogen stress and its control by abscisic acid,water relations and photosynthesis

Barley (Hordeum vulgare L.) and tomato Lycopersicon esculentum Mill.) were grown hydroponically and examined 2, 5, and 10 d after being deprived of nitrogen (N) supply. Leaf elongation rate declined in both species in response to N stress before there was any reduction in rate of dryweight accumulation. Changes in water transport to the shoot could not explain reduced leaf elongation in tomato because leaf water content and water potential were unaffected by N stress at the time leaf elongation began to decline. Tomato maintained its shoot water status in N-stressed plants, despite reduced water absorption per gram root, because the decline in root hydraulic conductance with N stress was matched by a decline in stomatal conductance. In barley the decline in leaf elongation coincided with a small (8%) decline in water content per unit area of young leaves; this decline occurred because root hydraulic conductance was reduced more strongly by N stress than was stomatal conductance. Nitrogen stress caused a rapid decline in tissue NO ₃ ^- pools and in NO ₃ ^- flux to the xylem, particularly in tomato which had smaller tissue NO ₃ ^- reserves. Even in barley, tissue NO ₃ ^- reserves were too small and were mobilized too slowly (60% in 2 d) to support maximal growth for more than a few hours. Organic N mobilized from old leaves provided an additional N source to support continued growth of N-stressed plants. Abscisic acid (ABA) levels increased in leaves of both species within 2 d in response to N stress. Addition of ABA to roots caused an increase in volume of xylem exudate but had no effect upon NO ₃ ^- flux to the xylem. After leaf-elongation rate had been reduced by N stress, photosynthesis declined in both barley and tomato. This decline was associated with increased leaf ABA content, reduced stomatal conductance and a decrease in organic N content. We suggest that N stress reduces growth by several mechanisms operating on different time scales: (1) increased leaf ABA content causing reduced cell-wall extensibility and leaf elongation and (2) a more gradual decline in photosynthesis caused by ABA-induced stomatal closure and by a decrease in leaf organic N.Abbreviation and symbols ABA abscisic acid - c_i leaf internal CO₂ concentration - L_p root hydraulic conductance     

Ecophysiological studies of Sonoran Desert plants

Summary The gas exchange of two Sonoran Desert plants was measured near optimum soil water conditions occurring in the summer and winter. Photosynthesis and stomatal conductance rates of a drought-deciduous shrub, Ambrosia deltoidea, and an evergreen non-riparian tree, Olneya tesota, are mainly affected by plant water potential. During such periods the diurnal gas exchange patterns are characterized by maximum rates of photosynthesis and stomatal conductance occurring early in the day, which decrease progressively thereafter. The effect of plant water potential on gas exchange is both direct and indirect. Decreasing plant water potential indirectly affects ¹⁴CO₂ assimilation by closure of stomata, and the effect is similar in both species. However, the direct effects of decreasing plant water potential are dissimilar in the two species. Plants of the shrub species have a higher potential maximum photosynthesis, but are more sensitive to plant water stress than are plants of the tree species. Both species respond to favorable growth conditions in the summer and winter, and have the potential for rapid carbon input into the Sonoran Desert ecosystem.This research was supported by National Science Foundation Grant BMS 74-02671-A04, through the U.S./I.B.P. Desert Biome     

Above- and below-ground environmental influences on leaf conductance ofCeanothus thyrsiflorus growing in a chaparral environment: drought response and the role of abscisic acid

Small shrubs ofCeanothus thyrsiflorus were grown in 19-1 pots irrigated under natural conditions in a chaparral region of Southern California and then subjected to soil drying. Characteristics of leaf gas exchange, leaf water potential, and concentrations of the stress hormone abscisic acid in the xylem sap, ABA_xyl, were determined at various stages of drought. Diurnal changes in conductance were strongly correlated with leaf net photosynthesis rate, which provides an effective, integrative predictor of above-ground climate effects on conductance. In drought conditions, ABA_xyl concentration increased. Increases in the concentration range of 50–500 nmol/l appeared to induce stomatal closure, restricting water loss and carbon dioxide uptake. When the momentary water potential is related to ABA_xyl, ABA appeared to increase significantly only after a threshold of approximately –1.5 MPa was exceeded. At less negative water potentials, large variation in ABA_xyl in the 50–1000 nmol/l range occurred for all water-potential values, because ABA_xyl remains relatively constant over diurnal courses as water potentials decrease and then recover. When the water potential became more negative than –1.5 MPa, ABA_xyl concentrations occurred between approximately 500 and 10 000 nmol/l and even greater in isolated cases. An approximately linear relationship is recognizable between ABA_xyl and momentary water potential in this range because in plants under drought conditions, ABA_xyl increases during the course of the day as water potential decreases. Increases in ABA_xyl in the high concentration range were associated with relatively minor additional restrictions in gas exchange, but they might contribute to improved water use efficiency and explain diurnal changes in the potential for stomatal opening that have been observed in Mediterranean sclerophyllous species. When we examined long-term seasonal change in the response of irrigated plants, changes in average daily temperature greater than 10°C occurred (also associated with shifts in relative humidity and radiation input), which apparently led to small changes in predawn water potential in the –0.1 to –0.7 MPa range. Increases in ABA_xyl occurred that were in turn negatively correlated with daily maximum leaf conductance. Thus, chaparral shrubs under non-drought conditions seem to sense even small changes in environmental conditions, in our opinion most probably due to initial drying of the uppermost soil and synthesis of ABA in the shallow roots. The results support the hypothesis that information of photosynthesis rate and predawn water potential may be used as primary variables to predict canopy conductance of Mediterranean sclerophyll shrub vegetation.