Effects of long-term rainfall variability on evapotranspiration and soil water distribution in the Chihuahuan Desert: A modeling analysis

We used the patch arid land simulator (PALS-FT) – a simple, mechanistic ecosystem model – to explore long-term variation in evapotranspiration (ET) as a function of variability in rainfall and plant functional type (FT) at a warm desert site in southern New Mexico. PALS-FT predicts soil evaporation and plant transpiration of a canopy composed of five principal plant FTs: annuals, perennial forbs, C₄ grasses, sub-shrubs, and evergreen shrubs. For each FT, the fractional contribution to transpiration depends upon phenological activity and cover as well as daily leaf stomatal conductance, which is a function of plant water potential, calculated from root-weighted soil water potential in six soil layers. Simulations of water loss from two plant community types (grass- vs. shrub-dominated) were carried out for the Jornada Basin, New Mexico, using 100 years of daily precipitation data (1891–1990). In order to emphasize variability associated with rainfall and fundamental differences in FT composition between communities, the seasonal patterns cover of perennials were held constant from year to year. Because the relative amount of year to year cover of winter and summer annual species is highly variable in this ecosystem, we examined their influence on model predictions of ET by allowing their cover to be variable, fixed, or absent.Over the entire 100-yr period, total annual ET is highly correlated with total annual rainfall in both community types, although T and E alone are less strongly correlated with rainfall, and variation in transpiration is nearly 3 times greater than evaporation and 2 times greater than variation in rainfall (CV of rainfall = 35%). Water use shows a relatively high similarity between the grass- and shrub-dominated communities, with a 100-yr average T/ET of 34% for both communities. However, based on a year-by-year comparison between communities, T/ET was significantly greater in the grass-dominated community, reflecting the fact that over the long term more than half of the rain occurs in the summer and is used slightly more efficiently (T¿E) by the C₄-grass community than the shrub community, although we found some rainfall patterns that resulted in much greater T/ET in the shrub community in a given year. Percent of water lost as transpiration (T/ET) suggests that while there is a general trend toward increased T/ET with rainfall in both community types, T/ET is extremely variable over the 100-yr simulation, especially for normal and below normal amounts of rainfall (T/ET values range from 1 to 58% for the grass-dominated site and 6 to 60% for the shrub-dominated site).These predictions suggest that because of the relatively shallow distribution of soil water, there is little opportunity for vertical partitioning of the soil water resource by differential rooting depths of the plant FTs, in contrast to the two-layer hypothesis of Walter (1971). However, functional types may avoid competition by keying on particular `windows' of moisture availability via differences in phenologies. We found very little differences in average, long-term model predictions of T, E, and ET when annual plant cover was variable, fixed, or absent. The results of our simulations help reconcile some of the disparate conclusions drawn from experimental studies about the relative contribution of transpiration vs. evaporation to total evapotranspiration, primarily by revealing the great year-to-year variability that is possible.     

Plant Water Potential Gradients Measured in the Field by Freezing Point

A portable freezing point meter was used in the field to measure the water potential gradients in sunflower (Helianthus annuus), beans (Phaseolus vulgaris), corn (Zea mays), wheat (Triticum aestivum), pumpkin (Cucurbita pepo), potato (Solanum tuberosum), alfalfa (Medicago sativa), and sugarbeets (Beta vulgaris). The measurements were made between daybreak and sunrise, and again during the middle of the afternoon on days when the potential evapotranspiration varied between 6.5 and 8.0 mm of water. The gradients varied from a maximum of 0.2 bar per cm in a wheat, down to an undetectable value for pumpkin. Although most of the soil in the root zone was kept at potentials above –1 bar, the bulk of the root tissue had water potentials of –5 to –10 bars. Differences in water potential between shaded and unshaded leaves, and between leaf tissue and guttation fluid suggested a similar drop of several bars between xylem elements and the surrounding leaf tissue in some plant species. The implications of such drops are discussed with respect to plant water transport equations and pressure cell potential measurements.     

Growth of Alaskan tundra plants in relation to water potential

The relationship of growth to plant water potential was studied in several arctic tundra species by experimentally altering the water regimes on excised tussocks, by clipping leaf area, and by clipping roots. Most species, including evergreen shrubs, showed greater growth under waterlogged conditions. The reduction of leaf area did not affect plant water potentials or growth. Root pruning decreased both plant water potentials and growth. Growth was suppressed when plant water potentials were −12 to −15 bars.     

A comparison of two water potential predictors for use in plant and soil thermocouple psychrometry

Summary A linear and a non-linear equation for predicting soil and plant water potentials as measured by thermocouple psychrometric techniques were compared. The non-linear predictor was often just as accurate as the linear predictor. The non-linear predictor did not predict positive water potentials. The non-linear predictor was most accurate in the range of zero to 10 negative bars water potential range. The linear predictor was more accurate than the non-linear predictor when water potentials were larger than –30 bars.     

Effect of water potential on seed germination

The response of seed germination to substrate water potential was determined for several plant species of the arctic tundra. Seeds were collected from Cape Thompson and Eagle Summit, Alaska and germinated on dialysis membranes over water solutions of polyethylene glycol with osmotic potentials of 0 to −6 bars. Germination did not occur with potentials below −3 bars, except for three fellfield species. Germination was delayed at lower osmotic potentials. Because the response of most species was similar, substrate water potential is probably not a factor affecting the establishment of most tundra plant species from seeds.     

Components of water potential estimated from xylem pressure measurements in five tree species

Summary Pressure volume curves were measured with a pressure bomb in leaves collected in the field from Ilex opaca, Acer rubrum, Liquidambar styraciflua, Liriodendron tulipifera and Cornus florida. Water potential components were calculated from the curves. The species differed in the relationships measured. In all species the trends from summer to fall were toward lower (more negative) osmotic potentials, lower matric potentials more rapid loss of turgor with increasing leaf water deficit, and the occurrence of incipient plasmolysis at lower values of leaf water deficit. Initial osmotic potentials ranged from-14.8 to-19.8 bars, similar to values reported in the literature for other mesophytic plants. These values, however, were much higher than those reported for halophytes and xerophytes. The fraction of leaf water which contributes to the osmotic potential ranged from 0.74 to 0.98 in this study. Values reported for other mesophytes and for halophytes and xerophytes all fall well within this range. Patterns of component water potentials are discussed in relation to potential growth rates and water flow in the total plant system.     

Comparative resistance of the soil and the plant to water transport

The resistances to liquid water transport in the soil and plant were determined directly and simultaneously from measurements of soil, root, and leaf water potentials and the flux of water through the soil-plant system to the sites of evaporation in the leaf. For soybean (Merr.) transporting water at a steady rate, water potential differences between soil and root were smaller than between root and leaf over the range of soil water potentials from −0.2 to −11 bars. As soil water was depleted, water flow through the soil and plant decreased to one-tenth the maximum rate, but both the soil resistance and plant resistance increased. The plant resistance remained larger than the soil resistance over the entire range of soil water availability. Previous suggestions that the soil is the major resistance have ignored the increase in plant resistance and/or assumed root densities that were too low.     

The influence of soil moisture on water potential,transpiration and photosynthesis of conifer seedlings

Summary The influence of soil moisture content and soil water potential on plant water potential, transpiration and net-photosynthesis of potted larch (Larix decidua), spruce (Picea abies) and pine (Pinus cembra) was studied under constant and close to optimum conditions in a laboratory.The equilibrium plant water potential measured under non-transpiring conditions came close to soil water potential, but in moist soil the equilibrium potential was slightly lower, particularly in larch where transpiration was not fully arrested. In very dry soil, plants had higher water potential than soil, presumably due to roots exploiting the wettest points within the soil.Pine, spruce and larch utilised a large part of soil moisture (down to 25wt.% soil water content or –1.5 bars potential) while maintaining plant water potential near –8, –9.5 and –12.5 bars respectively. A similar pattern occurred in dry soil. The differences between species are explained by differing stomatal sensitivity to water potential.Pine began a gradual reduction in gas-exchange below a soil water potential of –0.4 bars. Larch showed no marked reduction until the soil potential fell to –3.5 bars but below this the shut-down in gas-exchange was rapid. Spruce lay in between.In spite of the early and sensitive gas-exchange reduction with decreasing soil moisture, pine maintained the highest net photosynthesis/transpiration ratio and thus used limited soil water more slowly and economically than the other species.Seedlings maintained a higher rate of gas-exchange in strong light than in weak light, especially at low soil water potentials.     

Response of the water status of soybean to changes in soil water potentials controlled by the water pressure in microporous tubes

Water transport through a microporous tube-soil-plant system was investigated by measuring the response of soil and plant water status to step change reductions in the water pressure within the tubes. Soybeans were germinated and grown in a porous ceramic 'soil' at a porous tube water pressure of -0.5 kpa for 28 d. During this time, the soil matric potential was nearly in equilibrium with tube water pressure. Water pressure in the porous tubes was then reduced to either -1.0, -1.5 or -2.0 kPa. Sap flow rates, leaf conductance and soil, root and leaf water potentials were measured before and after this change. A reduction in porous tube water pressure from -0.5 to -1.0 or -1.5 kPa did not result in any significant change in soil or plant water status. A reduction in porous tube water pressure to -2.0 kPa resulted in significant reductions in sap flow, leaf conductance, and soil, root and leaf water potentials. Hydraulic conductance, calculated as the transpiration rate/delta psi between two points in the water transport pathway, was used to analyse water transport through the tube-soil-plant continuum. At porous tube water pressures of -0.5 to-1.5 kPa soil moisture was readily available and hydraulic conductance of the plant limited water transport. At -2.0 kPa, hydraulic conductance of the bulk soil was the dominant factor in water movement.     

The effect of water potential on soil-borne diseases of wheat seedlings

Under glasshouse conditions, Fusarium culmorum caused more injury to wheat seedlings in soil at 33% saturation (approximately – 1 bar) than at 66% (–0·1 bar). The same effect occurred with inoculated seeds raised on soil moisture tension tables at –0·1 and –0·5 bar, at 20° and at 15°C. In controls, spontaneous infection by soil-borne F. culmorum was unaffected by soil water potential. Inoculated F. nivale, at 127deg; and at 15°C, showed a similar, but smaller, effect to that with inoculated F. culmorum. Water potential in this range did not affect disease incidence with inoculated Gaeumannomyces graminis, but disease was more severe in the drier soil. On tension tables, seedlings did not consistently develop faster under wetter conditions, and factors other than host growth rate probably played a part in limiting fungal attack.     

Drought adaptations in two Californian evergreen sclerophylls

Summary A field study was initiated to determine the patterns of water stress imposition and stomatal resistance to gas exchange in representative species of 2 evergreen sclerophyllous communities. In concurrent experiments plant water potential, temperature and vapor pressure gradient were varied to determine the relative importance of morphological and physiological parameters in delaying onset of water stress during drought periods.In general, stomatal and photosynthetic responses to water stress were similiar in both species. Both were able to fix carbon even when leaf water potentials dropped as low as-25 bars. Stomatal movements were positively correlated with soil water potential rather than to leaf water potential. However, water stress developed much more rapidly in Arbutus menziesii, a plant of more northerly distribution, than in Heteromeles arbutifolia where they occur on adjacent sites. Morphological parameters were primarily responsible for the very different patterns of water stress imposition. Consequently, Arbutus is limited to areas of shorter drought duration than is Heteromeles and this is reflected in their differing distributions.     

Effects of repeated application of water stress on water status and growth of wheat

The effects on water status and growth of controlled cycles of water stress applied at various stages of development were studied on a semi-dwarf spring wheat (Triticum aestivum L.). The plants were grown in controlled environment chambers of the Duke University Phytetron at 24/18°C with a 12-h photo-period at about 600 μE m^?2 s^?1. Groups of plants were subjected to severe water stress by withholding irrigation, beginning at the 7th leaf, early anthesis, or early dough stages of development. A second cycle started 9 to 13 days after termination of the first cycle and maintained until the flag leaf water potential reached –25 bars at each of the growth stages. The lower leaves showed sign of wilting as indicated by curling in the first drying cycle at –7 bars and in the second cycle at –9 bars of leaf water potential during all stages of growth. Although these leaves recovered completely upon rewatering, onset of senescence was accelerated by three days in stressed plants. A preliminary drying cycle did not increase the ability of the plants to withstand subsequent stress because of severity of stress. Water stress of –25 bars at all three stages of growth reduced seed yield. The reduction was greater when a second stress cycle was also applied. Stress applied during early anthesis stage produced the smallest and the least number of seeds. The lack of osmotic adjustment probably was due to very rapid and severe development of water stress.     

Influence of temperature gradients on leaf water potential

Water potential was monitored at nine locations along single maize (Zea mays L.) leaf blades with aluminum block in situ thermocouple hygrometers. Water potential showed a continuous decrease toward the tip, with a 2- to 4-bar difference between leaf base and tip under both moist and dry soil conditions. The water potential difference between the soil and the leaf base was about 4 bars. Water potentials decreased during the day and during a drying cycle, and increased at night and after irrigation. Heating a band of a leaf to 40 C or cooling it to 7 C had no influence on the water potential of the affected portion when this was corrected for hygrometer output over standard calibrating solutions at the respective temperatures. Heating or cooling a portion of a leaf had neither short nor long term effects on water potential of more distal leaf portions continuously monitored by hygrometers in dew point readout. Water potential fluctuated with an amplitude of about 1.5 bars and an irregular period of 10 to 30 minutes. Measurements with silver foil in situ psychrometers gave similar results.     

The Association between Soil and Xylem Water Potential, Leaf Resistance, and Abscisic Acid Content in Droughted Seedlings of Douglas-fir (Pseudotsuga menziesii)

During progressive stages of drought and subsequent recovery through rewatering, foliage of 2-year-old seedlings of Douglas-fir was sampled for water potential, stomatal resistance, and abscisic acid content (ABA). Simultaneous measurements of soil matric potential were also made. There was a rapid rise in stomatal resistance in the range of 1500–2000 ng ABA g^?1 dry weight of foliage corresponding with a plant water potential of –10 to –12 bars. Soil matric potential seemed more closely associated with ABA increase than did plant water potential. After rewatering, seedlings returned to normal ABA levels within 6 days.     

Effects of fire on water and salinity relations of riparian woody taxa

Water and salinity relations were evaluated in recovering burned individuals of the dominant woody taxa from low-elevation riparian plant communities of the southwestern U.S. Soil elemental analyses indicated that concentrations of most nutrients increased following fire, contributing to a potential nutrient abundance but also elevated alluvium salinity. Boron, to which naturalized Tamarix ramosissima is tolerant, was also elevated in soils following fire. Lower moisture in the upper 30 cm of burned site soil profiles was attributed to shifts in evapotranspiration following fire. Higher leaf stomatal conductance occurred in all taxa on burned sites. This is apparently due to higher photosynthetic photon flux density at the midcanopy level and may be partially mitigated by reduced unit growth in resprouting burned individuals. Predawn water potentials varied little among sites, as was expected for plants exhibiting largely phreatophytic water uptake. Midday water potentials in recovering Salix gooddingii growing in the Colorado River floodplain reached levels which are considered stressful. Decreased hydraulic efficiency was also indicated for this species by examining transpiration-water potential regressions. Recovering, burned Tamarix and Tessaria sericea had enriched leaf tissue ¹³C relative to unburned controls. Higher water use efficiency following fire in these taxa may be attributed to halophytic adaptations, and to elevated foliar nitrogen in Tessaria. Consequently, mechanisms are proposed which would facilitate increased community dominance of Tamarix and Tessaria in association with fire. The theory that whole ecosystem processes are altered by invading species may thus be extended to include those processes related to disturbance.     

WATER USE AND SALT BALANCE IN THREE SALT MARSH SUCCULENTS

Water-use characteristics and potential salt accumulation rates were studied in three halophytes, Salicornia virginica, Balis marítima and Borrichia frutescens, inhabiting a salinity gradient in the high marsh. Xylem pressure potential (ψ_ρ), leaf osmotic potential (ψ_π) and leaf relative water content were measured seasonally in the three species. Species growing on the high end of the salinity gradient developed more negative xylem pressure potentials compared to species growing at lower soil salinities. This trend was also observed for leaf osmotic potentials. Low mean leaf ψ_π (below –15 to –36 bars) and high ash contents (0.27–0.48 g NaCl/g DW) indicated salt accumulation in transpiring tissues. However, calculations of potential salt accumulation, based on rates of transpiration and substrate salinity, suggest that some mechanism of salt exclusion at the roots may be operating.     

Can complementarity in water use help to explain diversity–productivity relationships in experimental grassland plots?

Positive diversity–productivity relationships have repeatedly been found in experimental grassland plots, but mechanistic explanations are still under debate. We tested whether complementarity for the exploitation of the soil water resource helps to explain these relationships. In the dry summer of 2003, evapotranspiration (ET) was assessed at the Swedish BIODEPTH site using two different approaches: snapshot measurements of canopy surface temperature and simulation of time-accumulated ET by means of a soil water balance model. More diverse plots were characterized by lower surface temperatures and higher accumulated ET. Transgressive overyielding tests revealed that ET in polycultures was higher than in the best-performing monocultures, but this pattern was reversed at high degrees of water stress. Our results indicate that a more complete exploitation of soil water by more diverse grassland systems is on the one hand likely to be a driver for their increased biomass production, but on the other hand causes the more diverse communities to be affected earlier by drought. Nevertheless, the results also suggest that productivity may (at least partially) be maintained under dry conditions due to the higher likelihood of including drought-tolerant species in the more diverse communities. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

WATER POTENTIAL,TEMPERATURE, AND KINETIN EFFECTS ON SEED GERMINATION IN SOIL AND SOLUTE SYSTEMS

Germination of lettuce and wheat in soil is reduced by a decrease in water potential, but a significant temperature-water potential interaction exists for lettuce. At 35 C kinetin permits lettuce germination at 0 and —1.1 bars, and at 25 C and 15 C it enhances germination at lower water potentials, causing 30% germination at —8.0 bars. Wheat germinates well at —8.0 bars, but no germination occurs at —14.9 bars; temperature had little effect on wheat germination. Germination in soil and solute systems was compared to determine the usefulness of solute germination data for predicting germination in dry soil. Total germination of lettuce in polyethylene glycol-6000 may approximate total germination in soil at the same water potential, but germination rates differ widely for the two systems. Kinetin-treated lettuce seeds nearly completed germination in two days in polyethylene glycol solutions, but five days were required for similar germination percentages in the soil. Sucrose is not useful for simulating soil water stress; wheat seeds germinate at —14.9 bars in sucrose but fail to germinate in soil at the same potential, and germination is more rapid in sucrose than in the soil.     

Soil water dynamics,transpiration, and water losses in a crested wheatgrass and native shortgrass ecosystem

The status of water in soil and vegetation was monitored in a stand of crested wheatgrass (Agropyron cristatum) and a nearby shortgrass steppe during a growing season. This was done to determine if water use and losses were similar among two very different communities in a similar climate. Precipitation was similar throughout the study period for both the crested wheatgrass and native shortgrass communities. However, the native shortgrass community with greater root biomass had consistently greater soil water depletion in the deeper soil horizons than was found in the crested wheatgrass community. Greater depletion of soil water by native shortgrass species suggests that they might be more competitive than crested wheatgrass in a water-limited environment.Crested wheatgrass maintained high leaf water potential early in the season, but lower water potential during the latter part of the growing season as compared with the major species of the shortgrass steppe, blue grama (Bouteloua gracilis) and western wheatgrass (Agropyron smithii). Leaf conductance was lower for crested wheatgrass than for the native grasses during the later part of the growing season. Consequently, seasonal transpiration for crested wheatgrass was lower when compared with blue grama or western wheatgrass. Lower conductance allowed crested wheatgrass to maintain relatively high internal water potential and may have accounted for less soil water use at deeper soil depths during the latter part of the growing season.Water loss through transpiration was less for western wheatgrass than for either blue grama or crested wheatgrass because western wheatgrass had less leaf area. However, western wheatgrass was as efficient as the other species in its use of water. Crested wheatgrass transpired more water than blue grama early in the growing season, but less than either native species for the remainder of the growing season. Estimated seasonal transpiration loss was greater in the shortgrass ecosystem than in the established crested wheatgrass stand.     

Relationship of water potential to growth of leaves

A thermocouple psychrometer that measures water potentials of intact leaves was used to study the water potentials at which leaves grow. Water potentials and water uptake during recovery from water deficits were measured simultaneously with leaves of sunflower (Helianthus annuus L.), tomato (Lycopersicon esculentum Mill.), papaya (Carica papaya L.), and Abutilon striatum Dickson. Recovery occurred in 2 phases. The first was associated with elimination of water deficits; the second with cell enlargement. The second phase was characterized by a steady rate of water uptake and a relatively constant leaf water potential. Enlargement was 70% irreversible and could be inhibited by puromycin and actinomycin D. During this time, leaves growing with their petioles in contact with pure water remained at a water potential of —1.5 to —2.5 bars regardless of the length of the experiment. It was not possible to obtain growing leaf tissue with a water potential of zero. It was concluded that leaves are not in equilibrium with the potential of the water which is absorbed during growth. The nonequilibrium is brought about by a resistance to water flow which requires a potential difference of 1.5 to 2.5 bars in order to supply water at the rate necessary for maximum growth.

Leaf growth occurred in sunflower only when leaf water potentials were above —3.5 bars. Sunflower leaves therefore require a minimum turgor for enlargement, in this instance equivalent to a turgor of about 6.5 bars. The high water potentials required for growth favored rapid leaf growth at night and reduced growth during the day.