No abstract

Plant metabolic activity in arid and semi-arid environments is largely tied to episodic precipitation events or "pulses". The ability of plants to take up and utilize rain pulses during the growing season in these water-limited ecosystems is determined in part by pulse timing, intensity and amount, and by hydrological properties of the soil that translate precipitation into plant-available soil moisture. We assessed the sensitivity of an invasive woody plant, velvet mesquite (Prosopis velutina Woot.), to large (35 mm) and small (10 mm) isotopically labeled irrigation pulses on two contrasting soil textures (sandy-loam vs. loamy-clay) in semi-desert grassland in southeastern Arizona, USA. Predawn leaf water potential (psi(pd)), the isotopic abundance of deuterium in stem water (deltaD), the abundance of 13C in soluble leaf sugar (delta13C), and percent volumetric soil water content (theta(v)) were measured prior to irrigation and repeatedly for 2 weeks following irrigation. Plant water potential and the percent of pulse water present in the stem xylem indicated that although mesquite trees on both coarse- and fine-textured soils quickly responded to the large irrigation pulse, the magnitude and duration of this response substantially differed between soil textures. After reaching a maximum 4 days after the irrigation, the fraction of pulse water in stem xylem decreased more rapidly on the loamy-clay soil than the sandy-loam soil. Similarly, on both soil textures mesquite significantly responded to the 10-mm pulse. However, the magnitude of this response was substantially greater for mesquite on the sandy-loam soil compared to loamy-clay soil. The relationship between psi(pd) and delta13C of leaf-soluble carbohydrates over the pulse period did not differ between plants at the two sites, indicating that differences in photosynthetic response of mesquite trees to the moisture pulses was a function of soil water availability within the rooting zone rather than differences in plant biochemical or physiological constraints. Patterns of resource acquisition by mesquite during the dynamic wetting-drying cycle following rainfall pulses is controlled by a complex interaction between pulse size and soil hydraulic properties. A better understanding of how this interaction affects plant water availability and photosynthetic response is needed to predict how grassland structure and function will respond to climate change.

Summary• Sorghum bicolor was exposed to free-air CO 2 enrichment (FACE) and drought at the Maricopa Agricultural Center, AZ, USA, in summer 1998. We predicted that bundle sheath leakiness ( Φ ) would be insensitive to FACE under well-irrigated (wet) conditions, but would be lower in FACE compared with control-CO 2 treatments when irrigation was withheld (dry).• Leaf and air δ 13 C values and leaf p i / p a from gas exchange were measured to estimate carbon isotope discrimination ( ∆ ) and Φ . Midday leaf water potential ( Ψ ) and photosynthetic rate were simultaneously measured to evaluate the influence of plant water status on Φ and the association between Φ and carbon gain.• Irrigation treatments affected Ψ , p i / p a , ∆ and Φ in control CO 2 and FACE rings. Differences in leaf ∆ between wet-and dry-treatment plots resulted from changes in Φ and to stomatal influences on p i / p a . FACE had very little effect on Ψ , ∆ and Φ in wet-treatment plots. However, Φ and ∆ in dry plots were higher in control than in FACE rings.• FACE ameliorated the effects of drought on bundle sheath leakiness and ∆ by reducing transpiration, prolonging soil water availability and enhancing plant water status. Direct effects of CO 2 enrichment on C 4 photosynthetic metabolism in Sorghum apparently are minimal and indirect effects depend on soil water supply.

We investigated hydraulic constraints on water uptake by velvet mesquite (Prosopis velutina Woot.) at a site with sandy-loam soil and at a site with loamy-clay soil in southeastern Arizona, USA. We predicted that trees on sandy-loam soil have less negative xylem and soil water potentials during drought and a lower resistance to xylem cavitation, and reach E(crit) (the maximum steady-state transpiration rate without hydraulic failure) at higher soil water potentials than trees on loamy-clay soil. However, minimum predawn leaf xylem water potentials measured during the height of summer drought were significantly lower at the sandy-loam site (-3.5 +/- 0.1 MPa; all errors are 95% confidence limits) than at the loamy-clay site (-2.9 +/- 0.1 MPa). Minimum midday xylem water potentials also were lower at the sandy-loam site (-4.5 +/- 0.1 MPa) than at the loamy-clay site (-4.0 +/- 0.1 MPa). Despite the differences in leaf water potentials, there were no significant differences in either root or stem xylem embolism, mean cavitation pressure or Psi(95) (xylem water potential causing 95% cavitation) between trees at the two sites. A soil-plant hydraulic model parameterized with the field data predicted that E(crit) approaches zero at a substantially higher bulk soil water potential (Psi(s)) on sandy-loam soil than on loamy-clay soil, because of limiting rhizosphere conductance. The model predicted that transpiration at the sandy-loam site is limited by E(crit) and is tightly coupled to Psi(s) over much of the growing season, suggesting that seasonal transpiration fluxes at the sandy-loam site are strongly linked to intra-annual precipitation pulses. Conversely, the model predicted that trees on loamy-clay soil operate below E(crit) throughout the growing season, suggesting that fluxes on fine-textured soils are closely coupled to inter-annual changes in precipitation. Information on the combined importance of xylem and rhizosphere constraints to leaf water supply across soil texture gradients provides insight into processes controlling plant water balance and larger scale hydrologic processes.

The changes in composition and productivity of semi-arid C(4) grassland, anticipated with rising atmospheric CO(2), will depend on soil water and nutrient availability. The interactive effects of soil resource limitation and elevated CO(2 )on these grasses, furthermore, may vary among C(4) biochemical subtypes (NADP-ME, NAD-ME, PCK) that differ in bundle sheath leakiness (Phi) responses to drought and nitrogen supply. To address C(4) subtype responses to soil resource gradients, the carbon isotope discrimination (Delta), bundle sheath leakiness (Phi), leaf gas exchange (A, g(s), c(i)/c(a)) and above-ground biomass accumulation were measured on three dominant grasses of semi-desert grassland in south-eastern Arizona. Bouteloua curtipendula (PCK), Aristida glabrata (NADP-ME) and the non-native Eragrostis lehmanniana (NAD-ME) were grown in controlled-environment chambers from seed under a complete, multi-factorial combination of present ambient (370 ppm) and elevated (690 ppm) CO(2) concentration and under high and low water and nitrogen supply. E. lehmanniana (NAD-ME) had the highest photosynthetic rate (A) and lowest Phi compared to the other two grasses when grown under low nitrogen and water availability. However, favourable water and nitrogen supply and elevated atmospheric CO(2) enhanced photosynthetic performance and above-ground biomass production of B. curtipendula (PCK) to a greater extent than in A. glabrata and E. lehmanniana. Contrary to pre vious studies, Phi and Delta in the NADP-ME subtype (A. glabrata) were most affected by changing environmental conditions compared to the other subtypes; deviations from the classic NADP-ME anatomy in Aristida could have accounted for this result. Overall, response of semi-arid grasslands to rising atmospheric CO(2) may depend more on species-specific responses to drought and nitrogen limitation than on general C(4) subtype responses.