Seasonal variations in moisture use in a piñon–juniper woodland

In water-limited environments of the intermountain region of North America, summer precipitation may play a role in the structure and function of aridland communities and ecosystems. This study examined the potential reliance on summer precipitation of two widespread, coexisting woody species in the southwestern United States, Pinus edulis Englmn. (Colorado piñon) and Juniperus osteosperma (Torr) Little (Utah juniper). The current distributions of P. edulis and J. osteosperma are highly suggestive of different dependencies on summer rainfall. We hypothesized that P. edulis was dependent on summer precipitation, utilizing summer precipitation even during extremely dry summers, whereas J. osteosperma was not dependent, using summer precipitation only when amounts were above some minimum threshold. Using sap flux and stable isotopic methods to assess seasonal water sources and water use efficiency, we examined the response of these two species to seasonal variations in moisture at a site located near the northern limits of the North American monsoon. Both sap flux and isotopic results indicated that P. edulis was responsive to summer rain, while J. osteosperma was not. Following summer rain events, sap flux density increased in P. edulis for several days, but not in J. osteosperma. Isotopic evidence indicated that P. edulis took up summer-derived moisture to a greater extent than J. osteosperma. Values of the natural abundance stable isotope ratio of carbon of leaf soluble carbohydrates increased over the summer for P. edulis, indicative of assimilation at higher water use efficiency, but were invariant for J. osteosperma. Our results supported the hypothesis that P. edulis and J. osteosperma are differentially sensitive to summer precipitation and are discussed in the light of potential changes in the seasonality of precipitation associated with climate change.     

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

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

Monosoonal precipitation responses of shrubs in a cold desert community on the Colorado Plateau

South-eastern Utah forms a northern border for the region currently influenced by the Arizona monosoonal system, which feeds moisture and summer precipitation into western North America. One major consequence predicted by global climate change scenarios is an intensification of monosoonal (summer) precipitation in the aridland areas of the western United States. We examined the capacity of dominant perennial shrubs in a Colorado Plateau cold desert ecosystem of southern Utah, United States, to use summer moisture inputs. We simulated increases of 25 and 50 mm summer rain events on Atriplex canescens, Artemisia filifolia, Chrysothamnus nauseosus, Coleogyne ramosissima, and Vanclevea stylosa, in July and September with an isotopically enriched water (enriched in deuterium but not ¹⁸O). The uptake of this artificial water source was estimated by analyzing hydrogen and oxygen isotope ratios of stem water. The predawn and midday xylem water potentials and foliar carbon isotope discrimination were measured to estimate changes in water status and water-use efficiency. At. canescens and Ch. nauseosus showed little if any uptake of summer rains in either July or September. The predawn and midday xylem water potentials for control and treatment plants of these two species were not significantly different from each other. For A. filifolia and V. stylosa, up to 50% of xylem water was from the simulated summer rain, but the predawn and midday xylem water potentials were not significantly affected by the additional summer moisture input. In contrast, C. ramosissima showed significant uptake of the simulated summer rain (>50% of xylem water was from the artificial summer rain) and an increase in both predawn and midday water potentials. The percent uptake of simulated summer rain was greater when those rains were applied in September than in July, implying that high soil temperature in midsummer may in some way inhibit water uptake. Foliar carbon isotope discrimination increased significantly in the three shrubs taking up simulated summer rain, but pre-treatment differences in the absolute discrimination values were maintained among species. The ecological implications of our results are discussed in terms of the dynamics of this desert community in response to changes in the frequency and dependability of summer rains that might be associated with a northward shift in the Arizona monsoon boundary.     

Limited uptake of summer precipitation by bigtooth maple (Acer grandidentatum Nutt) and Gambel's oak (Quereus gambelii Nutt)

Winter and spring precipitation that saturates to deep soil layers precedes summer droughts in the Intermountain West. Occasional summer convection storms relieve summer drought, but are infrequent and unreliable from year to year, leading to the hypothesis that dominant tree species might not invest limited carbon reserves to surface roots to take up summer precipitation in these regions. We compared the hydrogen (D) and oxygen (¹⁸O) isotope ratios of winter, spring and summer precipitation to that of xylem sap water in Acer grandidentatum and Quercus gambelii, two dominant trees of this region. By this method we could identify water sources utilized throughout the growing season. Xylem D and ¹⁸O values changed significantly when each species leafed-out; this change was not associated with changes in either soil or plant water status (as measured by predawn and midday water potentials). This shift is apparently related increased transpirational flux, which may flush out residual stem water from the previous growing season. D values of xylem sap of both species matched winter precipitation input values throughout most of the summer, indicating a reliance on deep-soil moisture sources throughout the growing season. Mature Q. gambelii did not take up summer precipitation, whereas A. grandidentatum responded slightly to the largest summer rain event. Small trees of both species, particularly A. grandidentatum, showed a limited uptake of summer rains.     

Photosynthetic gas exchange and the stable isotope composition of leaf water: comparison of a xylem-tapping mistletoe and its host

Photosynthetic gas exchange and the stable isotopic composition of foliage water were measured for a xylem tapping mistletoe, Phoradendron juniperinum, and its host tree, Juniperus osteosperma, growing in southern Utah. The observed isotopic composition of water extracted from foliage was compared to predictions of the Craig-Gordon model of isotopic enrichment at evaporative sites within leaves. Assimilation rates of juniper were higher and stomatal conductance was lower than the values observed for the mistletoe. This resulted in lower intercellular/ ambient CO₂ values in the juniper tree relative to its mistletoe parasite. For mistletoe, the observed foliage water hydrogen and oxygen isotopic enrichment was less than that predicted by the model. In juniper, foliage water hydrogen isotopic enrichment was also lower than that predicted by the evaporative enrichment model. In contrast, the oxygen isotopic enrichment in juniper foliage water was slightly greater than that predicted for the evaporative sites within leaves. Hydrogen isotopic enrichment in mistletoe foliage shows systematic variation with stem segment, being highest near the tips of the youngest stems and decreasing toward the base of the mistletoe, where isotopic composition is close to that of stem water in the host tree. In a correlated pattern, mid-day stomatal conductance declined abruptly in mistletoe foliage of increasing age.     

Extensive summer water pulses do not necessarily lead to canopy growth of Great Basin and northern Mojave Desert shrubs

Plant species and functionally related species groups from arid and semi-arid habitats vary in their capacity to take up summer precipitation, acquire nitrogen quickly after summer precipitation, and subsequently respond with ecophysiological changes (e.g. water and nitrogen relations, gas exchange). For species that respond ecophysiologically, the use of summer precipitation is generally assumed to affect long-term plant growth and thus alter competitive interactions that structure plant communities and determine potential responses to climate change. We assessed ecophysiological and growth responses to large short-term irrigation pulses over one to three growing seasons for several widespread Great Basin and northern Mojave Desert shrub species: Chrysothamnus nauseosus, Sarcobatus vermiculatus, Atriplex confertifolia, and A. parryi. We compared control and watered plants in nine case studies that encompassed adults of all four species, juveniles for three of the species, and two sites for two of the species. In every comparison, plants used summer water pulses to improve plant water status or increase rates of functioning as indicated by other ecophysiological characters. Species and life history stage responses of ecophysiological parameters (leaf N, ¹⁵N, ¹³C, gas exchange, sap flow) were consistent with several previous short-term studies. However, use of summer water pulses did not affect canopy growth in eight out of nine comparisons, despite the range of species, growth stages, and site conditions. Summer water pulses affected canopy growth only for C. nauseosus adults. The general lack of growth effects for these species might be due to close proximity of groundwater at these sites, co-limitation by nutrients, or inability to respond due to phenological canalization. An understanding of the connections between short-term ecophysiological responses and growth, for different habitats and species, is critical for determining the significance of summer precipitation for desert community dynamics.     

STEM AND LEAF GAS EXCHANGE OF TWO ARID LAND SHRUBS

Gas exchange studies were conducted on two shrub species found in cool shrub-steppe communities of the American West, big sagebrush (Artemisia tridentata subsp. tridentata Nutt.) and broom snakeweed (Gutierrezia sarothrae [Pursh] Britt. and Rusby), with a goal of evaluating characteristics and relative contributions of green stem and leaf material to total shoot CO₂ exchange at different temperatures. Variations in tissue temperature exerted a pronounced effect on CO₂ exchange—net photosynthesis and dark respiration—of green stems and leaves of both species. Definite temperature optima of net photosynthesis were noted, and dark respiration rates consistently increased with increases in temperature. Green stems of both species exhibited sizable dark respiration rates, although stem rates at all temperatures were lower than corresponding leaf rates. Artemisia tridentata did not exhibit sizeable green stem net photosynthesis even under conditions of optimal temperature and water availability, and leaf net photosynthesis rates were much lower than those of G. sarothrae. However, A. tridentata in general possessed a greater leaf biomass than G. sarothrae. Green stems of G. sarothrae exhibited considerable rates of net photosynthesis under both optimal and sub-optimal temperature and water availability conditions. A higher optimum temperature of net photosynthesis was noted for stems than for leaves of G. sarothrae. The adaptive significance of these interspecific differences in CO₂ exchange characteristics is discussed.     

Productivity and species richness in an arid ecosystem: a long-term perspective

To examine the different effects of rain pulse size on uptake of summer rains by three dominant desert plants in field conditions of desertified grasslands on the Ordos Plateau of northwestern China, we studied relationships between precipitation event size and rainwater uptake using stable isotopes of hydrogen in plant and soil water. Four natural precipitation events that represented precipitation sizes of 5.3, 8.3, 13.3, and 65.3 mm in the summer were chosen for the experiment. The perennial grass Stipa bungeana, the shrub Artemisia ordosia, and the herb Cynanchum komarovii – the dominant species in the communities – were compared for their use of summer rains with different pulse sizes based on the changes in the hydrogen isotope ratios (δD) of their stem water 7 days following each natural rain event. We found that S. bungeana and C. komarovii took advantage of shallow water sources derived from small (< 10 mm) rain events, A. ordosia took advantage of deeper soil water recharged by large (> 65 mm) rain events, and C. komarovii relied primarily on rain events of intermediate (10–20 mm) size. These different responses to rain pulses among species suggested that more frequent small rain events will promote the dominance of S. bungeana and C. komarovii, medium-sized events will facilitate development of C. komarovii, and large events will advance A. ordosia in this community. The rainwater utilization patterns of the three species would allow the coexistence of S. bungeana and A. ordosia or the coexistence of A. ordosia and C. komorovii in various successional serals following the disturbances. With an increase in variability of summer rain pulse size as predicted by climate change models, we expect that the structure of this community will undergo significant change in the future. Altered precipitation regimes, especially in combination with anthropogenic-related disturbances such as over-grazing, are likely to accelerate rates of degradation in northwestern China.     

Seasonal variability of groundwater level effects on the growth of Carex cinerascens in lake wetlands

Groundwater level is crucial for wetland plant growth and reproduction, but the extent of its effect on plant growth can vary along with changed precipitation and temperature at different seasons. In this context, we investigated the effect of two groundwater levels (10 cm vs. 20 cm depth) on growth and reproductive parameters of Carex cinerascens, a dominant plant species in the Poyang Lake wetland, during three seasons (spring, summer, and autumn) and during two consecutive years (2015 and 2016). Carex cinerascens showed low stem number, height, and individual and population biomass in summer compared to spring and autumn. 10 cm groundwater level was overall more suitable for plant growth resulting in higher stem height and biomass. However, the interactive effect between groundwater level and season clearly demonstrated that the effect of groundwater level on plant growth occurred mainly in autumn. After the withering of the plant population in summer, we observed that C. cinerascens growth recovered in autumn to similar values observed in spring only with 10 cm groundwater level. Consequently, we could deduce that lowering groundwater level in the studied Poyang Lake wetland will negatively impact C. cinerascens regeneration and growth particularly during the second growth cycle occurring in autumn. Additionally, our results showed that, independently of the season and groundwater level, population biomass of C. cinerascens was lower during drier year. Altogether, our findings suggest that water limitation due to both reduction in precipitation and decreased groundwater level during the year can strongly impact plant communities.     

Tree distribution on a steep environmental gradient in an arid savanna

Aim The structure of savannas ranges from scrub to woodland over broad geographical gradients. Here we examine the hypothesis that water availability is a major determinant of these structural differences by ascertaining the relationship between water availability and tree growth across a steep moisture gradient. Location The study site is a sub‐tropical savanna, with a mean annual precipitation of 400 mm, located in the Phugwane river basin, Kruger National Park, Limpopo Province, South Africa. Methods We determined plant moisture stress using xylem pressure potentials, stem growth using dendrometer bands, and estimated the water sources available to plants using stable hydrogen and oxygen isotope ratios. The primary objective was to understand tree growth relative to available water along an environmental gradient. Results We found that Philenoptera violacea trees growing close to the stream have lower water stress (least negative xylem pressure potentials) and higher cumulative growth than those growing away from the stream. The stem growth of P. violacea was characterized by steady incremental growth and could not be related to antecedent rainfall. Colophospermum mopane trees experienced higher water stress, yet trees growing adjacent to the stream achieved the highest cumulative growth rates over the study period. The growth of C. mopane could be clearly linked to antecedent rainfall, and most growth was achieved during short growth pulses that followed rainfall events. δ¹⁸O values become progressively more enriched in the heavier isotope with distance from the stream, suggesting that access to groundwater decreases with distance from the stream side. The isotopic data suggest that P. violacea has access to groundwater, but that C. mopane does not. Main conclusions Our results show that water stress increases with distance from the stream side as a result of reduced access to groundwater. Trees without access to deep water adopt an opportunistic growth strategy.     

Hydrogen isotope fractionation during water uptake by woody xerophytes

Stable isotope measurements are employed extensively in plant–water relations research to investigate physiological and hydrological processes from whole plant to ecosystem scales. Stable isotopes of hydrogen and oxygen are routinely measured to identify plant source water. This application relies on the assumption that no fractionation of oxygen and hydrogen isotopes in water occurs during uptake by roots. However, a large fraction of the water taken up through roots in halophytic and xerophytic plants transverses cell membranes in the endodermis before entering the root xylem. Passage of water through this symplastic pathway has been hypothesized to cause fractionation leading to a decrease in ²H of root xylem water relative to that in the surrounding soil medium. We examined 16 woody halophytic and xerophytic plant species in controlled conditions for evidence of hydrogen isotope fractionation during uptake at the root–soil interface. Isotopic separation (Δ²H = δ²H_{soil water} − δ²H_{xylem water}) ranging from 3‰ to 9‰ was observed in 12 species. A significant positive correlation between salinity tolerance and the magnitude of Δ²H was observed. Water in whole stem segments, sapwood, and roots had significantly lower δ²H values relative to soil water in Prosopis velutina Woot., the species expressing the greatest Δ²H values among the 16 species examined. Pressurized water flow through intact root systems of Artemisia tridentata Nutt. and Atriplex canescens (Pursh) Nutt. caused the δ²H values to decrease as flow rate increased. This relationship was not observed in P. velutina. Destroying the plasma membranes of root cells by excessive heat from boiling did not significantly alter the relationship between δ²H of expressed water and flow rate. In light of these results, care should be taken when using the stable isotope method to examine source-water use in halophytic and xerophytic species.     

Variation in carbon isotope discrimination within and among Sphagnum species in a temperate wetland

Field samples of bryophytes are highly variable in carbon isotope discrimination values (Δ, a measure of ¹³CO₂ uptake relative to ¹²CO₂), but it is unknown what affects Δ under field conditions, or how variation in Δ relates to bryophyte performance. This study employed field and greenhouse common garden studies to evaluate the influence of microsite, seasonal, and genetic variation on Δ in peatmosses. Three species of Sphagnum that occupy hollow (S. recurvum), carpet (S. palustre), and hummock (S. tenerum) habitats were sampled for relative growth rates (RGR), C:N ratio, and Δ throughout a growing season. Values of Δ ranged from 19.0 to 27.1‰. This variation was unrelated to species (P=0.61). However, Δ varied seasonally (P<0.001), with lower discrimination in the spring (mean 22.5‰), followed by summer (23.8‰) and winter (24.7‰). There was also significant microsite variation (P=0.015) which disappeared when plants were grown in a common garden. In both spring and summer, microsite variation in Δ was inversely related to RGR (P<0.001), but unrelated to C:N ratios (P>0.08). These results suggest that environmental, not genetic, variation at microsites affects Δ in non-vascular plants. However, environmental control of Δ is unlike that in vascular plants where water limitation lowers chloroplastic demand and increases resistance to carbon uptake. In non-vascular plants, water limitation lowers chloroplastic demand and decreases resistance to carbon uptake. These processes have additive effects and generate high spatial and seasonal variability in Δ. Received: 29 April 1999 / Accepted: 8 November 1999     

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

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

Hydrogen isotope response to changing salinity and rainfall in Australian mangroves

Hydrogen isotope ratios (²H/¹H, δ²H) of leaf waxes covary with those in precipitation and are therefore a useful paleohydrologic proxy. Mangroves are an exception to this relationship because their δ²H values are also influenced by salinity. The mechanisms underlying this response were investigated by measuring leaf lipid δ²H and leaf and xylem water δ²H and δ¹⁸O values from three mangrove species over 9.5 months in a subtropical Australian estuary. Net ²H/¹H fractionation between surface water and leaf lipids decreased by 0.5–1.0‰ ppt^?1 for n‐alkanes and 0.4–0.8‰ ppt^?1 for isoprenoids. Xylem water was ²H depleted relative to surface water, reflecting ²H discrimination of 4–10‰ during water uptake at all salinities and opportunistic uptake of freshwater at high salinity. However, leaf water ²H enrichment relative to estuary water was insensitive to salinity and identical for all species. Therefore, variations in leaf and xylem water δ²H values cannot explain the salinity‐dependent ²H depletion in leaf lipids, nor the 30‰ range in leaf lipid δ²H values among species. Biochemical changes in direct response to salt stress, such as increased compatible solute production or preferential use of stored carbohydrates, and/or the timing of lipid production and subsequent turnover rates, are more likely causes.     

Influence of groundwater depth on the seasonal sources of water accessed by <Emphasis Type="Italic">Banksia</Emphasis> tree species on a shallow,sandy coastal aquifer

In Mediterranean ecosystems vegetation overlying shallow, transient aquifers is often dominated by woody phreatophytes, trees and shrubs that have been shown to be dependent on groundwater for their water requirements. Natural and anthropogenic alterations of groundwater tables (abstraction) are of clear importance to phreatophytic vegetation as reduction of water tables may sever these plants from their natural water sources. Seasonal water sources were determined for species growing on a coastal dune system that overlies a shallow sandy aquifer in south-western Australia. The plants studied grew over groundwater that ranged in depth from 2.5 to 30 m. The naturally occurring stable isotope of hydrogen (deuterium, '²H) was used to distinguish potential water sources. Isotopic ratios from vascular water of the dominant species of the study area (Banksia ilicifolia R. Br. and Banksia attenuata R. Br. trees) were compared with those of potential sources of precipitation, soil moisture and groundwater. A relatively shallow-rooted perennial shrub, Hibbertia hypericoides Benth., was also included as an isotopic reference. The results suggest that both B. attenuata and B. ilicifolia are phreatophytic as they derived some of their water from groundwater throughout the dry-wet cycle, with the exception of B. attenuata at the site of greatest depth to groundwater (30 m) which did not use groundwater. A high proportion (>50%) of groundwater use was not maintained throughout all seasons. With the onset of the hot Mediterranean summer, progressive drying of the surface soils resulted in increased use of groundwater and deep soil moisture. During the wet winter plants used proportionately more water from the upper layers of the soil profile. The degree to which groundwater was utilised by the study species was dependent on the proximity of groundwater, availability of moisture in shallower horizons of the soil profile, root system distribution and maximum root depth.     

Root hydraulic conductivity of Larrea tridentata and Helianthus annuus under elevated CO2

While investigations into shoot responses to elevated atmospheric CO₂ are extensive, few studies have focused on how an elevated atmospheric CO₂ environment might impact root functions such as water uptake and transport. Knowledge of functional root responses may be particularly important in ecosystems where water is limiting if predictions about global climate change are true. In this study we investigated the effect of elevated CO₂ on the root hydraulic conductivity (L_p) of a C₃ perennial, Larrea tridentata, and a C₃ annual, Helianthus annuus. The plants were grown in a glasshouse under ambient (360 μmol mol^–1) and elevated (700 μmol mol^–1) CO₂. The L_p through intact root systems was measured using a hydrostatic pressure-induced flow system. Leaf gas exchange was also determined for both species and leaf water potential (ψ_leaf) was determined in L. tridentata. The L_p of L. tridentata roots was unchanged by an elevated CO₂ growth environment. Stomatal conductance (g_s) and transpiration (E) decreased and photosynthetic rate (A_net) and Ψ_leaf increased in L. tridentata. There were no changes in biomass, leaf area, stem diameter or root : shoot (R : S) ratio for L. tridentata. In H. annuus, elevated CO₂ induced a nearly two-fold decrease in root L_p. There was no effect of growth under elevated CO₂ on A_net, g_s, E, above- and below-ground dry mass, R : S ratio, leaf area, root length or stem diameter in this species. The results demonstrate that rising atmospheric CO₂ can impact water uptake and transport in roots in a species-specific manner. Possible mechanisms for the observed decrease in root L_p in H. annuus under elevated CO₂ are currently under investigation and may relate to either axial or radial components of root L_p.     

不同降水条件下两种荒漠植物的水分利用策略

自然降水是干旱、半干旱地区荒漠植物重要的水分来源。为了说明自然降水量的变化对干旱、半干旱地区荒漠植物水分利用策略的影响, 研究了两种常见荒漠植物油蒿(Artemisia ordosica)和白刺(Nitraria tangutorum)在3个不同自然降水地区(内蒙古的杭锦旗和磴口县及甘肃的民勤县)的水分来源、水分利用效率及植物的抗逆能力的变化。测定了不同地区的植物茎水、各潜在水源(降水、地下水和土壤水)的δD和δ¹⁸O值, 并利用IsoSource模型分析了这两种植物在不同地区对这些潜在水源的选择性利用情况; 同时测定了叶片的δ¹³C和游离脯氨酸浓度。结果表明: 在年降水量最高的杭锦旗, 这两种植物对浅层土壤水的利用比例最高, 其中油蒿主要利用0-50 cm土层中的水源; 在年降水量相对较低的磴口和民勤, 植物利用的主要水源为深层土壤水和地下水。随着年降水量的增加, 这两种植物的水分利用效率逐渐降低。白刺的脯氨酸浓度大于油蒿, 与水分利用效率无关, 但油蒿的水分利用效率和脯氨酸浓度成正比。研究表明, 荒漠植物能通过改变其水分利用策略和其他生理特性适应自然降水量的变化, 但不同植物种采用的策略可能有所不同。     

Effect of slight vegetation degradation on soil properties in Brachypodium pinnatum grasslands

Water is the most important factor controlling plant growth, primary production, and ecosystem stability in arid and semi-arid grasslands. Here we conducted a 2-year field study to explore the contribution of winter half-year (i.e. October through April) and summer precipitation (May through September) to the growth of coexisting plant species in typical steppe ecosystems of Inner Mongolia, China. Hydrogen stable isotope ratios of soil water and stem water of dominant plant species, soil moisture, and plant water potential were measured at three steppe communities dominated by Stipa grandis, Caragana microphylla, and Leymus chinensis, respectively. The fraction of water from winter half-year precipitation was an important water source, contributing 45% to plant total water uptake in a dry summer after a wet winter period (2005) and 15% in a summer where subsoil moisture had been exploited in the previous year (2006). At species level, Caragana microphylla exhibited a complete access to deep soil water, which is recharged by winter precipitation, while Cleistogenes squarrosa completely depended on summer rains. Leymus chinensis, Agropyron cristatum, and Stipa grandis showed a resource-dependent water use strategy, utilizing deep soil water when it was well available and shifting to rain water when subsoil water had been exploited. Our findings indicate that differentiation of water sources among plants improves use of available soil water and lessens the interspecific competition for water in these semi-arid ecosystems. The niche complementarity in water sources among coexisting species is likely to be the potential mechanism for high diversity communities with both high productivity and high resilience to droughts.     

ROOT PROFILES AND COMPETITION BETWEEN THE INVASIVE,EXOTIC PERENNIAL,CARPOBROTUS EDULIS,AND TWO NATIVE SHRUB SPECIES IN CALIFORNIA COASTAL SCRUB

The mat-forming succulent, Carpobrotus edulis (Aizoaceae), surrounds and grows over many native plant species in California coastal communities. Two shrub species, Haplopappus ericoides and H. venetus var. sedoides, were found to have shallow root systems that occupied the same soil depths as those of C. edulis. In the presence of C. edulis, the normal rooting profiles of the shrubs were displaced downward, although partial overlap with C. edulis remained. Removal of C. edulis from around individuals of both shrub species resulted in higher predawn xylem pressure potentials in shrubs from the removal treatments as compared to controls, suggesting that the surrounding C. edulis was utilizing water that would otherwise have been available to these shrubs. In H. ericoides, predawn xylem pressure potentials of removal treatment shrubs remained higher than those of the controls throughout the remainder of the dry season even though these shrubs showed a marked increase in canopy area after removal of surrounding C. edulis. Removal of C. edulis from around H. venetus also initially led to higher predawn xylem pressure potentials in the removal shrubs as compared to the controls and increased production of new leaves. After this initial period the predawn xylem pressure potentials of the removal shrubs were not significantly different from those of controls, suggesting that the demand for water by the increased leaf areas eventually matched the enhanced water availability resulting from C. edulis removal. Also, the morphology of H. venetus shrubs changed to that more typical of unaffected individuals. Thus, C. edulis significantly affected not only the water relations of these two shrub species but also their shoot sizes and overall morphologies.     

Validating the use of woodrat (Neotoma) middens for documenting natural invasions

Aim Plant macrofossils in fossil woodrat (Neotoma) middens are the primary source of information on late Quaternary biogeographical history of plants in arid and semi‐arid regions of North America. Macrofossil records from middens are playing particularly important roles in documenting spatial and temporal patterns of plant migrations and invasions since the last glacial maximum 21,000 years ago. However, relatively few actualistic studies comparing contents of modern middens with surrounding flora and vegetation have been carried out. The primary aim of this study is to assess the reliability of midden assemblages in detecting the presence of tree, shrub and several other plant species growing on the surrounding landscape. The secondary aims are to determine whether probability of occurrence of species in middens is related to abundance in vegetation, and whether representation of individual species in middens is contingent on presence/absence or abundance of other species. Location Our five study sites were bedrock escarpments or canyons in the central Rocky Mountains (north‐eastern Utah, central Wyoming and south‐central Montana). All sites were in conifer woodland or forest/woodland variously dominated by Juniperus osteosperma, J. scopulorum, Pinus edulis, P. flexilis and Pseudotsuga menziesii. Neotoma cinerea is the only woodrat species in the region. Methods Macrofossil assemblages from 59 modern middens (all showing clear signs of recent or ongoing woodrat activity) were compared with floristic composition of vegetation within 50 m of the middens, and with percent cover of vegetation within 30 m of the middens. Results Coniferous trees and shrubs were well‐represented in middens, occurring consistently even when abundance in the local vegetation was very low. Juniperus osteosperma and J. scopulorum were particularly well‐represented, occurring in middens regardless of local abundance. Other conifers (P. edulis, P. flexilis, P. ponderosa, Pseudotsuga menziesii) were occasionally absent from middens when abundance in vegetation was low (< 20% canopy cover). Occurrence of dicot shrubs and graminoids was less consistent. Main conclusions Middens constructed by N. cinerea are highly reliable sensors of presence/absence of J. osteosperma and J. scopulorum, and hence can be used to infer invasions and past biogeographical distributions of these species. The middens are also reliable in registering populations of other conifers, although presence in middens may be contingent on local abundance. Additional comparative studies are needed to develop a sound empirical basis for using middens of N. cinerea and other species to infer past presence/absence of plant species on the landscape, and to explore the vegetation‐sensing properties of midden assemblages.