Tissue water relations of four co-occurring chaparral shrubs

Summary Chaparral shrubs of California have a suite of morphological and physiological adaptations to withstand the prolonged summer droughts of a mediterranean climate. Not all species of chaparral have the same rooting depth and there is some evidence that those with shallow roots have tissue that is most tolerant to water stress. We tested this notion by comparing the tissue water relations of four co-occurring chaparral shrubs: Quercus durata, Heteromeles arbutifolia, Adenostoma fasciculatum, and Rhamnus californica. We used a pressure-volume technique and a dew-point hygrometer to metsure seasonal changes in osmotic potential when plant tissue was fully hydrated and osmotic potential at predawn, midday, and the turgor loss point. We also calculated seasonal changes in the minimum daily turgor potential, saturated weight/dry weight ratio of leaf tissue, and the bulk modulus of elasticity. We had information on the seasonal water use patterns and apparent rooting depths of these same four shrubs from a previous study (Davis and Mooney 1986). All evidence indicated that Rhamnus had shallow roots and Quercus deep roots. Our results indicated that the tissue water relations of our four co-occurring chaparral shrubs were not alike. Even though Rhamnus had shallow roots, it had the least xerophytic tissue. Seasonal osmotic potential and saturated weight/dry weight ratios were relatively high and bulk modulus of elasticity and minimum daily turgor potentials were low. Furthermore, even though Quercus had deep roots and experienced no seasonal water stress at our study site, its tissue water relations indicated relatively high tolerance to water stress. We conclude that seasonal drought tolerance of stem and leaf tissue of co-occurring chaparral shrubs does not necessarily correspond to rooting depth, to soil moisture resources available to the shrub, or to the degree of seasonal water stress experienced by the shrub.     

Recovery patterns of three chaparral shrub species after wildfire

Summary In a mature, even aged stand of mixed chaparral, Rhus laurina (facultative resprouter) had consistently higher water potentials and deeper roots than Ceanothus spinosus (facultative resprouter) and Ceanothus megacarpus (obligate seeder). For two years following a wildfire, the same stand of chaparral had resprouts with higher survivorships, predawn water potentials, stomatal conductances, photosynthetic rates and shoot elongation rates than seedlings. Supplemental irrigation of seedlings during summer months removed differences between resprouts and seedlings suggesting that the cause of such differences was limited water availability to the shoot tissues of seedlings. After two years of postfire regrowth, mean seedling survivorship for the obligate seeder (C. megacarpus) was 42%, whereas seedling survivorship for facultative resprouters was only 18% (C. spinosus) and 0.01% (R. laurina). Our results are consistent with the hypothesis that lack of resprouting ability among obligate seeders is offset by an enhanced ability to establish seedlings after wildfire, allowing obligate seeders to maintain themselves in mixed populations through many fire cycles.     

Comparative field water relations of four co-occurring chaparral shrub species

Summary The seasonal course of water relations was measured in the field in Adenostoma fasciculatum, Quercus dumosa, Ceanothus greggii, and Arctostaphylos glauca, four prominent members of the southern California chaparral vegetation. Ceanothus greggii and A. glauca developed similar seasonal patterns of minimum leaf water potentials, as estimated by xylem pressure measurements, which were much less negative than A. fasciculatum and Q. dumosa growing in close proximity on the same pole-facing slope site. Adenostoma fasciculatum on an adjacent equator-facing slope developed more negative water potentials than did A. fasciculatum on the pole-facing slope.Leaf conductance differed between species, and by leaf age class and slope exposure within a species. The greatest differences were measured between leaf age classes in A. fasciculatum on the pole-facing slope, with new leaves showing the greatest conductances early in the season. The same trend was measured in A. fasciculatum on the equator-facing slope, but the differences were less between leaf age classes and diminished earlier in the season than in A. fasciculatum on the pole-facing slope. The analysis of daily hysteresis in the leaf conductance-water potential relation suggests that early in the season when water is available, stomatal behavior is simultaneously governed by a complex of environmental factors, while late in the season stomatal behavior becomes increasingly dominated by tissue water status.     

Seasonal variation in the tissue water relations of Picea glauca

Seasonal variation in water relations of 3-yearold white spruce (Picea glauca (Moench) Voss) shoots, monitored with pressure-volume curves over 28 months, was closely related to shoot phenology and was sensitive to environmental fluctuations during both summer growth and winter dormancy. Turgor maintenance capacity was lowest during rapid shoot elongation from late May to early July; this was indicated by the lowest total turgor pressures, the highest (least negative) osmotic potentials at full turgor and the turgor loss point, the smallest differences between osmotic potentials at full turgor and the turgor loss point, the highest relative water contents at turgor loss and a linear decline in cell elasticity with decreasing turgor pressure. This suggests that the high susceptibility of white spruce seedlings to growth check after transplanting is largely attributable to the poor turgor maintenance capacity of this species in early summer.     

Differential survival of chaparral seedlings during the first summer drought after wildfire

Summary Big Pod Ceanothus (Ceanothus megacarpus) is an obligate seeder after fire; Laurel Sumac (Rhus laurina) is primarily a resprouter after fire. Both species commonly occur together in mixed stands and are dominant members of the coastal chaparral of southern California. We compared the mean survival of post-fire seedlings of each species during the first summer drought after fire and found C. megacarpus to have a mean survival of 54% while R. laurina had a mean survival of only 0.1%. Rooting dephs were similar between species but predawn water potentials and leaf temperatures were higher for R. laurina seedlings. Leaf temperatures for R. laurina reached a mean value of 46.8° C on hot, summer days, about 5° C higher than seedlings of C. megacarpus. By the end of the first growing season, 92% of all C. megacarpus seedlings had suffered herbivory compared to only 17% of all R. laurina seedlings. Herbivory did not appear to be the immediate cause of seedling mortality. Transect data indicated that full recovery of prefire species composition and density at our study site was likely but the mode of recovery was different for the species examined. R. laurina recovered primarily by sprouting, C. megacarpus totally by seedling establishment and a third species, Adenostoma fasciculatum (chamise), by a combination of sprouting and seedling establishment. We attribute the higher mortality of R. laurina seedlings to the greater sensitivity of its tissue to water stress. It may be that differential survival of shrub seedlings and differential modes of reestablishment after fire play an important role in maintaining species diversity in the chaparral communities of coastal, southern California.     

Field water relations of a wet-tropical forest tree species,Pentaclethra macroloba (Mimosaceae)

Summary The water relations of Pentaclethra macroloba (Willd.) Kuntze, a dominant, shade-tolerant, tree species in the Atlantic lowlands of Costa Rica, were examined within the forest canopy. Pressure-volume curves and diurnal courses of stomatal conductance and leaf water potential were measured in order to assess differences in water relations between understory, mid-canopy and canopy leaves. Leaves in the canopy had the smallest pinnules but the largest stomatal frequencies and stomatal conductances of the three forest levels. Osmotic potentials at full turgidity decreased with height in the forest; in the canopy and midcanopy they were reduced relative to those in the understory just enough to balance the gravitational component of water potential. Consequently, maximum turgor pressures were similar for leaves from all three canopy levels. Bulk tissue elastic modulus increased with height in the canopy. Leaf water potentials were lowest in the canopy and highest in the understory, even when the gravitational component was added to mid-canopy and canopy values. As a result, minimum turgor pressures were also lowest in the canopy compared to those at lesser heights, and approached zero in full sunlight on clear days.Osmotic potentials at each canopy level were similar for both wet and dry season samples dates suggesting that seasonal osmotic adjustment does not occur. Despite lowered predawn water potentials during the dry season, turgor was maintained in the understory by reduced stomatal conductances.     

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.     

Plant water relations and control of cell elongation at low water potentials

Recent developments in water status measurement techniques using the psychrometer, the pressure probe, the osmometer and pressure chamber are reviewed, and the process of cell elongation from the viewpoint of plant-water relations is discussed for plants subjected to various environmental stress conditions. Under water-deficient conditions, cell elongation of higher plants can be inhibited by interruption of water flow from the xylem to the surrounding elongating cells. The process of growth inhibition at low water potentials could be reversed by increasing the xylem water potential by means of pressure application in the root region, allowing water to flow from the xylem to the surrounding cells. This finding confirmed that a water potential field associated with growth process,i.e., the growth-induced water potential, is an important regulating factor for cell elongation other than metabolic factors. The concept of the growth-induced water potential was found to be applicable for growth retardation caused by cold stress, heat stress, nutrient deficiency and salinity stress conditions. In the present review, the fact that the cell elongation rate is primarily associated with how much water can be absorbed by elongating cells under water-deficiency, nutrient deficiency, salt stress, cold stress and heat stress conditions is suggested.     

Daily and seasonal variation in water relations of macchia shrubs and trees in France (Montpellier) and Turkey (Antalya)

Based upon two different research studies in the mediterranean regions of France and Turkey, drought resistance strategies were investigated in a broad group of species. The diurnal and seasonal patterns of the water relations of different lifeforms from the thermo-mediterranean to submediterranean lifezones were compared. Three sites near Montpellier, in Southern France, and five sites near Antalya, Turkey were used for this comparison. Xylem pressure potential and relative stomatal aperture were the key water relations parameters collected in France while these parameters as well as osmotic potential and leaf conductance were studied in Turkey.From the 26 different study species investigated in France, 7 distinct types of stomatal control were observed, with the deciduous lifeforms showing the least control, the sclerophyllous and coniferous evergreens the greatest control and the malacophyllous shrublets intermediate levels of control. Predawn water potential values provided a means of classifying species according to their temporal and spatial utilization of site water reserves. The comparison of turgor potentials (difference between water and osmotic potentials) gave an insight into leaf adaptations to site moisture. Species with high predawn water potentials generally maintain positive turgor even at midday during the summer, whereas species with low predawn values were frequently at zero turgor even at predawn. Phlomis grandiflora was the most extreme species with mid-summer predawns and midday water potentials of –6 MPa and osmotic potentials never more negative than –2.4 MPa.     

Water relation parameters of six Peltigera species correlate with their habitat preferences

Water relation parameters were measured in six congeneric lichen species with different requirements for water availability and with green algae (Peltigera aphthosa, Peltigera leucophlebia, Peltigera venosa) or cyanobacteria (Peltigera horizontalis, Peltigera praetextata, Peltigera rufescens) as main photobionts. Pressure–volume analysis was performed with a dewpoint hygrometer and integrated with anatomical analyses. The Peltigera species typical of arid environments were characterized by relatively lower osmotic potential (π₀) and turgor loss point (Ψ_TLP), and higher values of bulk modulus of elasticity (?). Both π₀ and Ψ_TLP were correlated with the size of medullary cells, while ? was negatively correlated with cell dimensions. The adaptive value of low Ψ_TLP might reside in the capability to maintain cell turgor for longer time intervals under dry conditions. High ? might allow xerophilous lichens to regain cell turgor more promptly even for small amounts of water uptake, thus enlarging the cumulative period of positive carbon balance in environments with fluctuating water availability. The influence of the photobiont type is also discussed.     

Nitrate increases in soil water following conversion of chaparral to grass

An Arizona watershed converted from chaparral to grass, released high concentrations of nitrate to stream water. The nitrate originated from the rooting zone of the decomposing shrubs. High nitrate concentrations (44–373 ppm) were found in soil solutions from 1.5-, 3.0-, and 4.6-m depths on the converted watershed as compared with low nitrate concentrations (0.2–6.2 ppm) found in an adjacent undisturbed area. Soil solution nitrate concentrations at the 0.3-m depth were generally low, especially in the untreated area. High nitrate concentrations were balanced mainly by relative decreases in bicarbonate anions in the soil solutions and in the stream water. Multiple stepwise regression analyses showed improvement in the regression of bicarbonate on nitrate when chloride and sulfate anions were entered as variables.     

Effects of water stress on the water relations of Phaseolus vulgaris and the drought resistant Phaseolus acutifolius

The tepary bean ( Phaseolus acutifolius Gray var. latifolius ), a drought resistant species, was compared under water stress conditions with the more drought susceptible P. vulgaris L. cvs Pinto and White Half Runner (WHR). In order to better understand the basis for the superior drought resistance of tepary, this study was designed to determine the relationships among leaf water potential, osmotic potential, turgor potential, and relative water content (RWC).
Plants were prestressed by withholding irrigation water. These stress pretreatments changed the relation between leaf water potential and relative water content of both species so that prestressed plants had lower water potentials than controls at the same leaf RWC. Tepary had lower water potentials at given RWC levels than Pinto or WHR; this can account for part of the superior resistance of tepary. In all genotypes, prestressed plants maintained osmotic potentials approximately 0.2 MPa lower than controls. Tepary reached osmotic potentials that were significantly lower (0.15 to 0.25 MPa) than Pinto or WHR. Both control and prestressed tepary plants had 0.05 to 0.25 MPa more turgor than Pinto or WHR at RWC values between 65 and 80%. Both prestressed and control tepary plants had greater elasticity (a lower elastic modulus) than Pinto or WHR. This greater turgor of tepary at low RWC values could be caused by several factors including greater tissue elasticity, active accumulation of solutes, or greater solute concentration.
Tepary had significantly lower osmotic potentials than the P. vulgaris cultivars, but there was little difference in osmotic potential between Pinto and WHR. Knowledge of differences in osmotic and turgor potentials among and within species could be useful in breeding for drought resistance in Phaseolus.     

Turgor and osmotic relations of the desert shrub Larrea tridentata

Abstract Leaf water relations characteristics of creosote bush, Larrea tridentata, were studied in view of previous reports that its leaves commonly experience zero or negative turgor under dry conditions. Leaf turgor loss point () was determined by a pressure-volume method for samples subjected to a hydration procedure and for untreated samples. Hydration caused to increase by as much as 3 M Pa. Hydration of samples also caused changes in other leaf water relations characteristics such as symplastic solute content, tissue elasticity and symplasmic water fraction, but total leaf solute content was unchanged. Comparison of our field plant water potential data with values of obtained by the two methods resulted in predictions of turgor loss during part or all of a diurnal cycle based on hydrated samples, and turgor maintenance (at least 0.3 MPa) based on untreated samples. Pooled data for obtained from both partially hydrated and untreated samples showed that L. tridentata maintains fairly constant levels of turgor over a wide range of leaf water potential. Dilution of cell contents by apoplastic water introduced significant errors in psychrometric determinations of osmotic potential in both frozen and thawed leaf tissue and expressed cell sap. Use of these values of osmotic potential resulted in predictions of zero turgor at all plant water potentials measured in the field.     

Leaf water relations characteristics of Lupinus angustifolius and L. cosentinii

Summary Lupins (Lupinus angustifolius and L. cosentinii) growing in 321 containers in a glasshouse were exposed to drought by withholding water. Leaf water potential (₁), and leaf osmotic potential (_s) were measured daily as soil water became depleted. Leaf water relations were further assessed by a pressure-volume technique and by measuring _s and relative water content of leaves after rehydration. Analysis by pressure-volume or cryoscopic techniques showed that leaf osmotic potential at saturation (_s100) decreased from -0.6 MPa in well watered to -0.9 MPa in severely droughted leaves, and leaf water potential at zero turgor (_zt) decreased from about -0.7 to -1.1 MPa in well watered and droughted plants, respectively. Relative water content at zero turgor (RWC_zt) was high (88%) and tended to be decreased by drought. The ratio of turgid leaf weight to dry weight was not influenced by drought and was high at about 8.0. The bulk elastic modulus () was approximately halved by drought when related to leaf turgor potential (_p) and probably mediated turgor maintenance during drought. The latter was found to be negatively influenced by rate of drought. Supplying the plants with high levels of K salts did not promote adjustment or turgor maintenance.     

Comparative physiology of burned and unburned Rhus laurina after chaparral wildfire

Summary Laurel Sumac (Rhus laurina) is a dominant member of the coastal chaparral community of southern California that survives periodic burning by wildfires by resprouting from a lignotuber (root crown). We investigated the physiological basis for resprouting by comparing shoot elongation, leaf nitrogen content, tissue water status, leaf conductance to water vapor diffusion, and photosynthetic rates of post-fire R. laurina to those of adjacent unburned shrubs. Resprouts had higher rates of shoot elongation, leaf conductance, and photosynthesis than mature, unburned shrubs. Leaf nitrogen contents were elevated in burned shrubs even though their leaves developed interveinal chlorosis. A comparison of soil water potential to predawn water potential indicated that roots of R. laurina remain active below 2 m during the first summer drought after wildfire. Our results support the hypothesis that lignotubers not only contain dormant buds that develop into aerial shoots after wildfire but they also supply nutrient resources that enhance shoot elongation. Because R. laurina is relatively sensitive to drought, yet very successful in its rapid recovery after fire, maintaining an active root system after shoot removal may be the primary function of the massive lignotuber formed by this species.     

Effect of ABA on the water relations of winter-wheat cultivars varying in drought resistance

Water and osmotic potentials were measured in leaves of a drought-sensitive ('Ponca') and a drought-resistant ('KanKing') cultivar of winter wheat ( Triticum aestivum L . em. Thell.) to determine if the potentials of the drought-sensitive cultivar could be made similar to those of the drought-resistant cultivar through application of abscisic acid (ABA). Stomatal resistance was also measured. Plants were sprayed with ABA and grown in soil, which was watered or allowed to dry. In well-watered plants, ABA closed the stomata of both cultivars. Stomatal resistance of plants grown without added water and with ABA was less than that of plants grown without added water and without ABA. Under ample water supply, ABA decreased water and osmotic potentials of the drought-sensitive cultivar (Ponca), but had no effect on these potentials in the drought-resistant cultivar (KanKing). Under water-deprived conditions, ABA increased water and osmotic potentials of Ponca, but did not change these potentials in KanKing. The overall effect of ABA was to decrease the differences in the water and osmotic potentials between the two cultivars.     

Comparative field water relations of three Mediterranean shrub species co-occurring at a natural CO(2) vent

Annual variations in the water relations and stomatal response of Erica arborea, Myrtus communis and Juniperus communis occurring at a natural CO(2) vent were analysed under Mediterranean field conditions. A distinct gradient of CO(2)concentration ([CO(2)]) exists between two sites near a natural CO(2)-emitting vent, with higher [CO(2)] (700 micromol mol(-1)) in the proximity of the CO(2) spring. Plants at the CO(2) spring site have been growing for generations at elevated [CO(2)]. At both sites, maximum leaf conductance was related to predawn shoot water potential. The effects of water deficits during the summer drought were severe. Leaf conductance and water potential recovered after major rainfalls in September to predrought values. Strong relationships between leaf conductance, predawn water potential, and leaf-specific hydraulic resistance are consistent with the role of stomata in regulating plant water status. Considerable between-species variation in sensitivity of water potentials and stomatal characters to elevated [CO(2)] were observed. Common to all the shrubs were a reduction in leaf conductance and an increase in water potentials in response to elevated [CO(2)]. Elevated [CO(2)] decreased the sensitivity of leaf conductance to vapour pressure deficit. Morphological characters (including stomatal density and degree of sclerophylly) showed site-dependent variations, but degree and sign of such changes varied with the species and/or the season. Measurements of discrimination against (13)C provided evidence for long-term decreases of water use efficiency in CO(2) spring plants. Analysis of C isotope composition suggested that a downward adjustment of photosynthetic capacity may have occurred under elevated [CO(2)]. Elevated [CO(2)] effects on water relations and leaf morphology persisted in the long term, but the three shrubs growing in the same environment showed species-specific responses.     

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.     

Apoplasmic water fractions and osmotic potentials at full turgidity of some Bryidae

The fractions of apoplasmic water of six moss species were estimated by comparison of the osmotic potentials of fully turgid living tissues (Ψ _π(s)) and of killed shoots (Ψ _π(k)). The values of Ψ _π(s) were determined by pressure-volume analysis using thermocouple hygrometry, whereas those of Ψ _π(k) were obtained by cryoscopy using extracts produced from dried materials and taking into consideration the original saturation water content of the shoots. Most of the tissues had Ψ _π(s) values around-1 to -1.5 MPa and non-osmotic water fractions of roughly 20% of the total water content at full turgor. Quantitative analysis of a number of osmotically active cell constituents showed that about one-third of the osmotic potential resulted from the accumulation of sugars. The total free amino acids accounted for about 15–20% of the osmolality of the cell solution. Malic- and citricacid contents varied much more with species but, in general, both carboxylic acids together contributed nearly 10–20% to the osmotic potential. The contents of inorganic anions such as free chloride and phosphate in mosses were low.     

Leaf water relations and anatomy of a tropical rainforest tree species vary with crown position

Summary Leaf water potentials, osmotic properties and structural characteristics were examined in the Australian tropical rainforest tree species, Castanospermum australe. These features were compared for individuals growing in the understorey and canopy of the undisturbed forest and in an open pasture from which the forest had been cleared. Leaf water potentials during the day declined to significantly lower values in the open-grown and canopy trees than in the understorey trees. During most of the day the opengrown tree experienced the lowest water potentials. These differences were paralleled by significant differences in tissue osmotic properties. The tissue osmotic potential at full hydration was lowest in the open-grown tree (-1.80 MPa), intermediate in the canopy trees (-1.38 MPa), and highest in the understorey trees (-0.80 MPa). As a result, the degree to which high and positive turgor pressures were maintained as water potentials declined was highest in the open-grown tree, intermediate in the canopy trees, and lowest in the understorey trees. The differences in tissue osmotic properties between individuals in the three crown positions were paralleled, in turn, by differences in leaf structual characteristics. Relative to leaves of the canopy and open-grown trees, leaves of the understorey trees had significantly larger epidermal cells with thinner cell walls, larger specific leaf areas and turgid weight: dry weight ratios, and a higher proportion of intercellular air space.Abbreviations ₁ Leaf tissue water potential - _min Lowest value of ₁ during the day ( noon) - _{P=0 1} zero turgor - R Relative water content - P Tissue turgor pressure - Tissue osmotic potential - ₀ at full hydration