Comparative water-use efficiency of Sporobolus arabicus and Leptochloa fusca and its relation with carbon-isotope discrimination under semi-arid conditions

Water-use efficiency (WUE) of Leptochloa fusca (L.) Kunth (Kallar grass) and Sporobolus arabicus Boiss. was determined under different soil moisture regimes. Plants grown in lysimeters were subjected to three soil moisture regimes, viz. well-watered (100%), medium-watered (75%), and low-watered (50%) of total available water (TAW). The soil moisture was restored on alternate days by adding the required volume of water on the basis of neutron moisture meter readings taken from neutron access tubes installed in each lysimeter. The grasses were harvested after suitable intervals (4 months) to obtain maximum biomass. Leaf samples collected at each harvest were analyzed for carbon-isotope discrimination (¹³C) with an isotope ratio (¹³C/¹²C) mass spectrometer. Results indicated significant differences in WUE of both grasses subjected to different water regimes. Sporobolus arabicus showed higher WUE than Kallar grass. However, Kallar grass showed better value of yield response factor (k _y = 0.649) compared with Sporobolus (k _y = 1.06) over the entire season. The data confirm that these grasses can be grown successfully in water-limited environments by selecting an optimum soil moisture level for maximum biomass production. The mean carbon-isotope discrimination (¹³C) of Kallar grass (–14.4) and Sporobolus (–12.8) confirm that both are C₄ plants. The carbon-isotope discrimination () was significantly and negatively correlated with WUE of the two species studied. The results of the present study confirm that ¹³C or of leaves can be used as good predictor of WUE in some C₄ plants.     

Traits of the invasive Centaurea maculosa and two native grasses: effect of N supply

The Eurasian forb Centaurea maculosa (Lam.; spotted knapweed) has invaded millions of hectares of semi-arid grasslands in western North America. It readily colonizes disturbed areas, but also invades pristine grasslands. C. maculosa's success could be attributed to greater use, or more efficient use, of available soil nitrogen (N). Soil N often limits growth on semi-arid grasslands. Greater or more efficient use of soil N by C. maculosa, if this occurred, may inhibit establishment, survival, or reproduction of native grasses. In a glasshouse, C. maculosa and two native grasses, Pseudoroegneria spicata [Scribn. and Smith] A. Love and Pascopyrum smithii [Rybd.] A. Love, were grown in mixed- and monoculture for 8 weeks to determine growth response to two soil N supplies, which mimicked low and high N mineralization rates in semi-arid grasslands. At the end of the 8 weeks, plants were exposed to ¹⁵N-labeled nitrate for 24 h, and harvested to compare uptake of NO₃ ^–. C. maculosa's growth response to N indicated that it was more competitive for N than the tussock grass P. spicata, but less competitive than the rhizomatous grass P. smithii. C. maculosa used nitrogen less efficiently than both of these native grasses. C. maculosa roots took up more ¹⁵N per unit root mass than the grasses, but acquired less N than P. smithii because P. smithii had greater root mass than C. maculosa. Total biomass and ¹⁵N uptake of C. maculosa varied depending on which species it was growing with. C. maculosa's success cannot be explained wholly by greater or more efficient use of soil N than that of the native grasses with which it competes.     

The effect of nitrogen supply on growth and water-use efficiency of xylem-tapping mistletoes

Xylem-tapping mistletoes are known to have normally a higher rate of transpiration and lower water-use efficiency than their hosts. The relationships between water relations, nutrients and growth were investigated for Phoradendron juniperinum growing on Juniperus osteosperma (a non-nitrogen-fixing tree) and for Phoradendron californicum growing on Acacia greggii (a nitrogen-fixing tree). Xylem sap nitrogen contents were approximately 3.5 times higher in the nitrogen-fixing host than in the non-nitrogen-fixing host. The results of the present study show that mistletoe growth rates were sevenfold greater on a nitrogen-fixing host. At the same time, however, the differences in water-use efficiency between mistletoes and their hosts, which were observed on the non-nitrogen-fixing host did not exist when mistletoes were grown on hosts with higher nitrogen contents in their xylem sap. Growth rates and the accumulation of N, P, K, and Ca as well as values for carbon-isotope ratios of mistletoe tissues support the hypothesis that the higher transpiration rates of mistletoes represent a nitrogen-gathering mechanism.Abbreviation ¹³C carbon-isotope ratio Dedicated to Professor Dr. Hubert Ziegler on the occasion of his 60th birthday     

Water relations of native and introduced C4 grasses in a neotropical savanna

Introduced African grasses are invading Neotropical savannas and displacing the native herbaceous community. This work, which is part of a program to understand the success of the African grasses, specifically investigates whether introduced and native grasses differ in their water relations. The water relations of the native Trachypogon plumosus and the successful invader Hyparrhenia rufa were studied in the field during two consecutive years in the seasonal savannas of Venezuela. The two C₄ grasses differed clearly in their responses to water stress. H. rufa consistently had higher stomatal conductance, transpiration rate, leaf water and osmotic potential and osmotic adjustment than the native T. plumosus. Also, leaf senescence occurred much earlier during the dry season in H. rufa. Both grasses showed a combination of water stress evasion and tolerance mechanisms such as stomatal sensitivity to atmospheric or soil water stress, decreased transpiring area and osmotic adjustment. Evasion mechanisms are more conspicuous in H. rufa whereas T. plumosus is more drought tolerant and uses water more conservatively. The evasion mechanisms and oportunistic use of water by H. rufa, characteristic of invading species, contribute to, but only partially explain, the success of this grass in the Neotropical savannas where it displaces native plants from sites with better water and nutrient status. Conversely, the higher water stress tolerance of t. plumosus is consistent with its capacity to resist invasion by alien grasses on shallow soils and sites with poorer nutrient and water status.     

DIFFERING SELECTION ON PLANT PHYSIOLOGICAL TRAITS IN RESPONSE TO ENVIRONMENTAL WATER AVAILABILITY: A TEST OF ADAPTIVE HYPOTHESES

I used phenotypic selection analysis to test the prediction from functional and comparative studies of plants that smaller leaves and more efficient water use are adaptive in drier environments. I measured selection gradients on leaf size and instantaneous water-use efficiency (a measure of carbon gain per unit water loss) in experimental populations of Cakile edentula var. lacustris placed into wet and dry environments in the field. Linear and nonlinear selection differed significantly between the two environments as predicted. Water-use efficiency was selected to be higher, and leaf area was selected toward a small intermediate optimum, in the dry environment. There was also significant positive correlational selection on water-use efficiency and leaf size, suggesting that the optimum leaf size in the dry environment is greater for plants with higher water-use efficiency. In contrast, neither leaf size nor water-use efficiency were selected in the wet environment, though larger leaves resulted in greater vegetative biomass. Path analysis of the linear selection gradients found that water-use efficiency affected plant fitness primarily because it increased vegetative biomass, as suggested by the hypotheses about the function of physiological traits. These results were not only consistent with the functional hypotheses but also with the observed genetic differentiation in water-use efficiency and leaf size between wet and dry site populations.     

Response of the invasive Centaurea maculosa and two native grasses to N-pulses

Nitrogen is often a limiting resource on semi-arid grasslands. During the growing season, N is often only available during short-term pulses associated with wetting events. The Eurasian forb Centaurea maculosa Lam. has invaded millions of hectares of semi-arid grasslands in western North America. C. maculosa's success could be attributed to greater use of N-pulses, or more efficient use of N supplied in those pulses compared with native grasses. In a glasshouse, C. maculosa and two native grasses, the caespitose Pseudoroegneria spicata [Scribn. and Smith] A. Love and the rhizomatous Pascopyrum smithii [Rybd.] A. Love, were established in mixed- and monoculture combinations, and then conditioned to weekly N-pulses of 8, 24, or 72 h for 8 weeks. These pulse durations are typical on semi-arid grasslands. At the end of the 8 weeks, plants were exposed to ¹⁵N-labeled nitrate (¹⁵NO₃ ^–) for 8 h and harvested 16 h later to compare short-term root uptake of ¹⁵NO₃ ^–. C. maculosa did not have greater enrichment (atom % ¹⁵N), rate of ¹⁵N-uptake (mol g^–1 h^–1), or ¹⁵N acquired (relative to ¹⁵N applied) than the grasses. C. maculosa's ¹⁵N-uptake per unit mass was relatively consistent across pulse durations, whereas ¹⁵N-uptake was lower at the longer pulse durations for the grasses. In general, C. maculosa acquired more of the applied ¹⁵N than P. spicata but less than P. smithii. ¹⁵N acquired was often influenced by the neighbour's identity. Regarding growth responses, C. maculosa produced more total biomass than the grasses, except for P. smithii plants growing with C. maculosa conditioned to 72 h pulses of N. Root mass ratios varied depending on the neighbor. Overall, C. maculosa used nitrogen less efficiently than the grasses. C. maculosa's success as an invasive species cannot be explained wholly by a greater response to N-pulses or more efficient use of N-pulses compared with native grasses with which it competes.     

Response of net ecosystem gas exchange to a simulated precipitation pulse in a semi-arid grassland: the role of native versus non-native grasses and soil texture

Physiological activity and structural dynamics in arid and semi-arid ecosystems are driven by discrete inputs or pulses of growing season precipitation. Here we describe the short-term dynamics of ecosystem physiology in experimental stands of native (Heteropogon contortus) and invasive (Eragrostis lehmanniana) grasses to an irrigation pulse across two geomorphic surfaces with distinctly different soils: a Pleistocene-aged surface with high clay content in a strongly horizonated soil, and a Holocene-aged surface with low clay content in homogenously structured soils. We evaluated whole-ecosystem and leaf-level CO₂ and H₂O exchange, soil CO₂ efflux, along with plant and soil water status to understand potential constraints on whole-ecosystem carbon exchange during the initiation of the summer monsoon season. Prior to the irrigation pulse, both invasive and native grasses had less negative pre-dawn water potentials ( _pd), greater leaf photosynthetic rates (A _net) and stomatal conductance (g _s), and greater rates of net ecosystem carbon exchange (NEE) on the Pleistocene surface than on the Holocene. Twenty-four hours following the experimental application of a 39 mm irrigation pulse, soil CO₂ efflux increased leading to all plots losing CO₂ to the atmosphere over the course of a day. Invasive species stands had greater evapotranspiration rates (ET) immediately following the precipitation pulse than did native stands, while maximum instantaneous NEE increased for both species and surfaces at roughly the same rate. The differential ET patterns through time were correlated with an earlier decline in NEE in the invasive species as compared to the native species plots. Plots with invasive species accumulated between 5% and 33% of the carbon that plots with the native species accumulated over the 15-day pulse period. Taken together, these results indicate that system CO₂ efflux (both the physical displacement of soil CO₂ by water along with plant and microbial respiration) strongly controls whole-ecosystem carbon exchange during precipitation pulses. Since CO₂ and H₂O loss to the atmosphere was partially driven by species effects on soil microclimate, understanding the mechanistic relationships between the soil characteristics, plant ecophysiological responses, and canopy structural dynamics will be important for understanding the effects of shifting precipitation and vegetation patterns in semi-arid environments.     

Water-use strategy of three central Asian desert shrubs and their responses to rain pulse events

Plant water-use strategy is considered to be a function of the complex interactions between species of different functional types and the prevailing environmental conditions. The functional type of a plant’s root system is fundamental in determining the water-use strategy of desert shrubs and the physiological responses of the plant to an occasional rainfall event, or rain pulse. In this current study of Tamarix ramosissima Ledeb. Fl.Alt., Haloxylon ammodendron (C.A.Mey.) Bunge and Reaumuria soongorica (Pall.) Maxim., three dominant shrub species in the Gurbantonggut Desert (Central Asia), plant root systems were excavated in their native habitat to investigate their functional types and water-use strategies. We monitored leaf water potential, photosynthesis and transpiration rate during a 39-day interval between successive precipitation events during which time the upper soil water changed markedly. Plant apparent hydraulic conductance and water-use efficiency were calculated for the varying soil water conditions. Our results show that: 1) The three species of shrub belong to two functional groups: phreatophyte and non-phreatophyte; 2) The photosynthetic capacity and leaf-specific apparent hydraulic conductance of the three species was stable during the time that the water condition in the upper soil changed; 3) Transpiration, leaf water potential and water-use efficiency in Tamarix ramosissima Ledeb. Fl.Alt. were stable during the period of observation, but varied significantly for the other two species. Tamarix ramosissima Ledeb. Fl.Alt., as a phreatophyte, relies mostly on groundwater for survival; its physiological activity is not inhibited in any way by the deficiency in upper soil water. Non-phreatophyte Haloxylon ammodendron (C.A.Mey.) Bunge and Reaumuria soongorica (Pall.) Maxim. use precipitation-derived upper soil water for survival, and thus respond clearly to rain pulse events in terms of leaf water potential and transpiration. The observed similarity in leaf-specific photosynthesis capacity among all three species indicates that the two non-phreatophyte species are able to maintain normal photosynthesis within a wide range of plant water status. The observed stability in leaf-specific apparent hydraulic conductance indicates that the two non-phreatophyte species are able to maintain sufficient water supply to their foliage via, mostly likely, effective morphological adjustment at the scale of the individual plant.Section editor: H. Lambers     

Differences in water-use strategies along an aridity gradient between two coexisting desert shrubs (<Emphasis Type="Italic">Reaumuria soongorica</Emphasis> and <Emphasis Type="Italic">Nitraria sphaerocarpa</Emphasis>): isotopic approaches with physiological evidence

Background and aims

Understanding the responses of different plant species to changes in available water sources is critical for accurately modeling and predicting species dynamics. Our study aimed to explore whether there were differences in water-use strategies between the two coexisting shrubs (Reaumuria soongorica and Nitraria sphaerocarpa) in response to different amounts of summer precipitation.

Methods

We conducted 3 years of field observations at three sites along an aridity gradient from the middle to lower reaches of the Heihe River basin, northwestern China. Stable oxygen composition (δ¹⁸O) in plant xylem water, soil water and groundwater were analyzed concurrently with ecophysiological measurements at monthly intervals during the growing seasons.

Results

Water source for coexisting R. soongorica and N. sphaerocarpa did not differ at the sites with high precipitation, but significantly differed in more arid locations. The N. sphaerocarpa was more sensitive to summer precipitation than R. soongorica in terms of predawn water potential, stomatal conductance and foliage carbon-isotope discrimination.

Conclusions

The plants relying on groundwater maintained consistent water use strategies, but not plants that took up precipitation-derived water. We also found that the difference in water source uptake between the coexisting species was more apparent in more arid locations.

     

Comparisons of carbon isotope discrimination in populations of aridland plant species differing in lifespan

Summary Carbon isotope discrimination () was compared between populations of dominant perennial plant species, differing in life expectancy, in two deserts with contrasting vegetation types. In both deserts, plants of the shorter-lived species showed significantly higher and greater intrapopulation variance in this character compared to the long-lived species. These results indicate underlying differences in gas-exchange physiology, and suggest a positive correlation between water-use efficiency and lifespan in desert plants. Differences in variance for this character may reflect greater microenvironmental variation experienced by shorter-lived plants and/or different forms of selection acting on water-use traits. Spatial distributions were significantly clustered for the shorter-lived species and significantly uniform for the long-lived species, indicating that competition has been important in the development of the long-lived populations. The long-lived Larrea tridentata showed a significant, negative correlation between and Thiessen polygon area, suggesting a positive relationship between water-use efficiency and longevity within this species. This relationship was weakly supported in the other warm desert species, Encelia farinosa, but was not observed within populations of the cold desert species, Gutierrezia microcephala and Coleogyne ramosissima. These results suggest that reflects key aspects of plant metabolism related to lifespan; these differences may ultimately influence interactions among desert plants and the structure of desert plant communities.     

The importance of factors controlling species abundance and distribution varies in native and non‐native ranges

How variation in factors controlling species abundance and distribution between native and non‐native ranges compares to that within ranges remains poorly understood. We used a globally distributed ruderal, Centaurea solstitialis (Centaurea), to explore the possibility that the importance of those factors exhibits great variation between and within ranges. To test our hypothesis, we established seed addition experiments with soil disturbance (turnover and control) and biocide (fungicides, insecticide, and control) treatments in two regions within native (the Caucasus and south‐western Turkey) and non‐native (the western United States – US – and central Argentina) distributions. Also, we estimated the rate of vegetation recovery after disturbance (resilience) and related it to Centaurea density in experimental plots. Disturbance strongly increased Centaurea density in all regions. Density was similar between the native Caucasus and non‐native Argentina and much greater in those regions than in the native Turkey and non‐native US in biocide‐free plots. Fungicides had positive effects on density in the US and negative ones in the Caucasus and Argentina, resulting in no differences between those three regions and greater density in the US than Turkey. Insecticide applications promoted Centaurea density in Turkey and Argentina, but inter‐regional comparisons of density in treated plots were comparable to those in biocide‐free plots. Overall, plants were smaller and less fecund in Turkey than the other regions, except the US. The greatest fungal attack was documented in Turkey, and herbivory was stronger there and in Argentina than in the Caucasus and US. The resilience of the local community explained a large proportion of variation in Centaurea density. These results support our hypothesis, and reveal that the speed at which competition is re‐gained after disturbance may influence global variation in Centaurea abundance. Because many ruderals exhibit native and non‐native distributions, our results are likely to be generalized to other systems.     

Stress responses of native and exotic grasses in a Neotropical savanna predict impacts of global change on invasion spread

Communities subject to stress, including those with low invasibility, may be dominated by exotic generalist species. African grasses are aggressive invasive species in Neotropical savannas, where their response to abiotic stress remains unknown. We assessed the role of waterlogging and canopy closure on the presence, abundance and reproductive tillering of African and native grasses in a Neotropical savanna in southeastern Brazil. We obtained abundance and reproductive tillering data of exotic (Melinis minutiflora, Melinis repens and Urochloa decumbens) and common native grasses in 20 sites. We also determined the groundwater depth, soil surface water potential and canopy cover at these sites. The grass species generally had a low frequency and performed poorly where soil remained waterlogged throughout the year, except for two native species. Most native species were exclusive to either well‐drained savannas or better drained wet grasslands. However, two species (Loudetiopsis chrysothrix and Trachypogon spicatus) occurred in both vegetation types. Two exotic species (M. minutiflora and M. repens) were less common but demonstrated reasonable performance in wet grasslands, possibly due to their root system plasticity. Furthermore, U. decumbens had a lower occurrence, density and reproductive tillering at these sites, but was successful at sites where the groundwater level was slightly deeper. Although the favourable water regime in the savannas increases their invasibility in general, resistance to invasion by African grasses may be greater at microsites with high canopy closure, where these species showed lower performance and did not affect the abundance of co‐occurring native grasses. In summary, the Brazilian savanna becomes more susceptible to the spread of African grasses when disturbances decrease canopy closure or lower rainfall associated with climate change reduces the average groundwater depth and consequently releases invasive species from soil waterlogging in grasslands.     

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.     

Effects of N and water supply on water use-efficiency of a semiarid grassland in Inner Mongolia

Productivity of semiarid grasslands is primarily limited by seasonal rainfall amount and becomes increasingly limited by nutrient availability under wet conditions. Interactive effects of water and N availability on grassland productivity and parameters related to water use were studied on a grassland site in Inner Mongolia, China, in a 2-factorial experiment with two levels of water (rainfed: 158 mm; irrigated: 839 (N0) and 972 (N1) mm) and N supply (0 or 180 kg N ha^?1). RUE was calculated from ANPP and cumulative water supply. Bare soil evaporation (E) was calculated from climatic data and leaf area dynamics, and percolation (D) and transpiration (T) were estimated with HYDRUS-1D. Water-use efficiency (WUE, ANPP / (T + D)) and transpiration efficiency (TE, ANPP / T) were calculated. Resource availability had pronounced effects on the water-use efficiency of semiarid grassland. RUE, WUE, and TE all decreased under irrigated compared to rainfed conditions and were significantly increased with N fertilizer application at both levels of water supply. While the irrigation effect on parameters of water-use efficiency were accordingly reflected in stable carbon isotope signatures, N supply resulted in significantly less negative δ¹³C-values under rainfed but not irrigated conditions. It is concluded, that spatial or temporal gradients in resource availability have pronounced effects on the water-use efficiency of semiarid grassland. The decrease of water use-efficiency under high water supply was related to differences in TE and not to a relative increase of unproductive water loss. Carbon isotope discrimination was highly correlated with WUE and TE, but can be a poor predictor of RUE.     

Water resource partitioning,stem xylem hydraulic properties,and plant water use strategies in a seasonally dry riparian tropical rainforest

This study investigated seasonal variation in the origin of water used by plants in a riparian tropical rainforest community and explored linkages between plant water source, plant xylem hydraulic conductivity and response to the onset of dry conditions. The study focused on five co-dominant canopy species, comprising three tree species (Doryphora aromatica, Argyrodendron trifoliolatum, Castanospora alphandii) and two climbing palms (Calamus australis and Calamus caryotoides). Stable isotope ratios of oxygen in water (¹⁸O) from soil, groundwater, stream water and plant xylem measured in the wet season and the subsequent dry season revealed water resource partitioning between species in the dry season. Measurement of stem-area-specific hydraulic conductivity (K_S) in the wet season and subsequent dry season showed a significant dry-season loss of K_S in three of the five species (Castanospora alphandii, Calamus australis and C. caryotoides) and a decrease in mean K_S for all species. This loss of hydraulic conductivity was positively correlated with the difference between wet-season and dry-season midday leaf water potentials and with leaf carbon isotope discrimination, indicating that plants that were less susceptible to loss of conductivity had greater control over transpiration rate and were more water-use efficient.     

Biotic constraints on the invasion of diffuse knapweed (Centaurea diffusa) in North American grasslands

Knapweeds (Centaurea spp.) are among the most invasive of non-indigenous plant species that have colonized western North America over the last century. We conducted a 4-year experiment in a reconstructed grassland to test hypotheses related to the ability of grasslands to resist the invasion of diffuse knapweed (C. diffusa). We experimentally invaded C. diffusa and three native species into areas where we manipulated soil nitrogen (N) and phosphorus (P) availability and removed extant grasses to reduce competition. We evaluated the growth response of these species to these resources and competitive manipulations. Of the native species that were experimentally added, only one species, Ratibida pinnata (prairie coneflower), established in any numbers. Establishment values in intact vegetation were low for both species, but establishment by C. diffusa (0.02%) clearly outperformed that of R. pinnata (0.001%). Under reduced grass competition, establishment was enhanced, but the values for C. diffusa (0.68%) were not statistically different from those of R. pinnata (0.57%). Neither species performed better under higher soil nutrients in the presence of competing grasses. In plots with both species, biomass of the two planted species was positively correlated, but the biomass of both species was negatively correlated with non-added weedy species. Subsequent harvests of C. diffusa indicated that establishment was enhanced in treatments with higher soil nutrients but that the biomass of these plants could only be enhanced when plant competition was also reduced. These results indicate that C. diffusa can establish in intact grasslands at rates higher than natives, but opportunism rather than competitive ability best describes the invasiveness of C. diffusa. Thus, the mechanisms contributing to the establishment of this knapweed species are different from factors identified as contributing to the dominance of this invader.     

Exotic plant communities shift water-use timing in a shrub-steppe ecosystem

Semiarid areas in the US have realized extensive and persistent exotic plant invasions. Exotics may succeed in arid regions by extracting soil water at different times or from different depths than native plants, but little data is available to test this hypothesis. Using estimates of root mass, gravimetric soil water, soil-water potential, and stable isotope ratios in soil and plant tissues, we determined water-use patterns of exotic and native plant species in exotic- and native-dominated communities in Washington State, USA. Exotic and native communities both extracted 12 ± 2 cm of water from the top 120 cm of soil during the growing season. Exotic communities, however, shifted the timing of water use by extracting surface (0–15 cm) soil water early in the growing season (i.e., April to May) before native plants were active, and by extracting deep (0–120 cm) soil water late in the growing season (i.e., June to July) after natives had undergone seasonal senescence. We found that δ ¹⁸O values of water in exotic annuals (e.g., −11.8 ± 0.4 ‰ for Bromus tectorum L.) were similar to δ ¹⁸O values of surface soil water (e.g., −13.3 ± 1.4 ‰ at −15 cm) suggesting that transpiration by these species explained early season, surface water use in exotic communities. We also found that δ ¹⁸O values of water in taprooted exotics (e.g., −17.4 ± 0.3 ‰ for Centaurea diffusa Lam.) were similar to δ ¹⁸O values of deep soil water (e.g., −18.4 ± 0.1 ‰ at −120 cm) suggesting that transpiration by these species explained late season, deep water use. The combination of early-season, shallow water-use by exotic winter-actives and late-season, deep water-use by taprooted perennials potentially explains how exotic communities resist establishment of native species that largely extracted soil water only in the middle of the growing season (i.e., May to June). Early season irrigation or the planting of natives with established root systems may allow native plant restoration.     

Soil water exploitation after fire: competition between Bromus tectorum (cheatgrass) and two native species

Summary Causes for the widespread abundance of the alien grass Bromus tectorum (cheatgrass) after fire in semiarid areas of western North America may include: (1) utilization of resources freed by the removal of fireintolerant plants; and (2) successful competition between B. tectorum and individual plants that survive fire. On a site in northwestern Nevada (USA), measurements of soil water content, plant water potential, aboveground biomass production, water use efficiency, and B. tectorum tiller density were used to determine if B. tectorum competes with either of two native species (Stipa comata and Chrysothamnus viscidiflorus) or simply uses unclaimed resources. Soil water content around native species occurring with B. tectorum was significantly lower (P<0.05) than around individuals without B. tectorum nearby. Native species had significantly more negative plant water potential when they occurred with B. tectorum. Aboveground biomass was significantly higher for native species without B. tectorum. However, the carbon isotope ratio of leaves for native species with B. tectorum was not significantly different from individuals without B. tectorum. Thus, B. tectorum competes with native species for soil water and negatively affects their wate status and productivity, but the competition for water does not affect water use efficiency of the native species. These adverse effects of B. tectorum competition on the productivity and water status of native species are also evident at 12 years after a fire. This competitive ability of B. tectorum greatly enhances its capability to exploit soil resources after fire and to enhance its status in the community.     

Effects of environmental change, including drought, on water use by competing Calluna vulgaris (heather) and Pteridium aquilinum (bracken)

1. Competition for water between Calluna vulgaris (heather) and Pteridium aquilinum (bracken) was studied in conditions of increased temperature, drought and increased nitrogen supply. All these factors increased the intensity of competition for water, with the combination of drought and increased nitrogen having the greatest effect on water use.
2. Both species increased water-use efficiency in response to increased nitrogen and drought. The effects of temperature were however, equivocal. Calluna had a greater water demand than Pteridium but acclimated to water stress more readily.
3. Calluna was the superior competitor for water; its water-use efficiency was reduced as a consequence of its roots depleting water from the Pteridium rooting zone. Pteridium , the poorer competitor, increased water-use efficiency to cope with reduced water availability owing to competition.
4. There was a strong relationship between carbon isotope discrimination (Δ) and instantaneous water-use efficiency for both species, but discrimination provided a more sensitive measure of seasonal water-use efficiency. Reconstruction of the plant's history of water-use efficiency by retrospective measurement of Δ proved a useful technique for Calluna leaves but was inappropriate for Pteridium rhizome.     

Carbon isotope composition of N2-fixing and N-fertilized legumes along an elevational gradient

Dinitrogen-fixing legumes are frequently assumed to be less water-use efficient than plants utilizing soil mineral N, because of the high respiratory requirements for driving N₂ fixation. However, since respiration is assumed not to discriminate against ¹³C, any differences in water-use efficiency exclusively due to respiration should not be apparent in carbon isotope discrimination () values. Our objective was to determine if the source of N (N₂ fixation versus soil N) had any effect on of field-grown grain legumes grown at different elevations. Four legume species, Glycine max, Phaseolus lunatus, P. vulgaris, and Vigna unguiculata, were grown on five field sites spanning a 633 m elevational gradient on the island of Maui, Hawaii. The legumes were either inoculated with a mixture of three effective strains of rhizobia or fertilized weekly with urea at 100 kg N ha^-1 in an attempt to completely suppress symbiotic N₂-fixing activity. In 14 of 20 analyses of stover and 12 of 15 analyses of seed values were significantly higher (p=0.10) in the inoculated plants than the N-fertilized plants. Nitrogen concentrations were generally higher in the fertilized treatments than the inoculated treatments. The different values obtained depending on N-source may have implications in using as an indicator of water-use efficiency or yield potential of legumes.