▪ Abstract The use of stable isotope techniques in plant ecological research has grown steadily during the past two decades. This trend will continue as investigators realize that stable isotopes can serve as valuable nonradioactive tracers and nondestructive integrators of how plants today and in the past have interacted with and responded to their abiotic and biotic environments. At the center of nearly all plant ecological research which has made use of stable isotope methods are the notions of interactions and the resources that mediate or influence them. Our review, therefore, highlights recent advances in plant ecology that have embraced these notions, particularly at different spatial and temporal scales. Specifically, we review how isotope measurements associated with the critical plant resources carbon, water, and nitrogen have helped deepen our understanding of plant-resource acquisition, plant interactions with other organisms, and the role of plants in ecosystem studies. Where possible we also...

Stable Isotopes in Plant Ecology

During drought periods, sugar maple (Acer saccharum) demonstrates “hydraulic lift”; nocturnal uptake of water by roots from deep soil layers that is released from shallow roots into upper soil layers. Using standard water relations methods and stable hydrogen isotope analysis of both source-water and plant-water, I investigated (1) the magnitude and radial extent of hydraulic lift by mature, relatively open-grown trees, of A. saccharum, (2) the proportion of hydraulically-lifted water (HLW) used by shallow-rooted neighbors growing at different distances from target trees, and (3) the influence that this water source had on stomatal conductance to water vapor (g), water balance and growth of these neighbors. Soil water potentials (ψs) at −20 and −35 cm showed a distinct diel fluctuation. Soil pits dug beneath three mature trees revealed a distinct hard-pan (e.g. fragipan) layer at a depth of approximately 50 cm. Examination of root distributions obtained from soil cores and soil pits revealed that some larger diameter roots (1.9–3.7 cm) did penetrate the fragipan and were established in the ground water table. The presence of the fragipan indicated that the “rewetting” of the upper soil layer during the night could not be explained by capillary rise from the shallow water table; it was the trees that were taking up ground water and then redepositing it at night into the upper 35 cm of soil, above the fragipan. The greatest fluctuations in ψs occurred within 2.5 m of trees and only extended out to approximately 5 m. Application of a two-end-member linear mixing model which used stable hydrogen isotopic data obtained from environmental water sources and xylem-sap demonstrated that all neighbors used some fraction (3–60%) of HLW supplied by sugar maple trees. Plants that used a high proportion of HLW (e.g. rhizomatous or stoloniferous perennials) maintained significantly higher leaf water potentials and g, and showed greater aboveground growth when compared with (i) neighbors that used little or no HLW or (ii) conspecifics found growing at distances greater than about 3 m away from maple trees. Three important conclusions can be drawn from the results of this investigation that have not been demonstrated before: (1) hydraulic lift need not only occur in arid or semi-arid environments where chronic water deficits prevail, but can be important in relatively mesic environments when subjected to periodic soil water deficits, (2) that plants neighboring trees which conduct hydraulic lift can use a significant proportion of this water source, and (3) that the HLW source can effectively ameliorate the influence of drought on the performance and growth of neighboring vegetation. The results are also discussed in terms of their influence on plant nutrient relations (including plant-mycorrhizal associations), the nature of plant-plant interactions and the water balance of individuals, communities and floristic regions.

Hydraulic lift and water use by plants: implications for water balance, performance and plant-plant interactions.

Groundwater depletion threatens many riparian ecosystems in arid and semi- arid regions of the world. The aquifer that sustains Arizona's San Pedro River riparian ecosystem, for example, is threatened by regional groundwater declines and localized pump- ing from the alluvial aquifer. This paper demonstrates the important role of shallow ground- water in structuring the San Pedro River plant community, portions of which function as reference areas that indicate site potential for a globally rare forest type (Sonoran riparian Populus-Salix forests). Several ecological indicators varied with depth to groundwater, including a weighted average wetland indicator score calculated for herbaceous and woody plant species, cover of plants within wetland indicator groups, and frequency of indicator plant species. These relationships can be used in a space-for-time substitution to predict consequences of groundwater decline. For example, the wetland indicator score changed sharply as depth to groundwater ranged from 0 to 4 m, and abundance of obligate wetland herbs (the group most sensitive to groundwater changes) declined sharply at groundwater depths below -0.25 m. Such sequential "desertification" of the riparian flora (i.e., loss or reduction in cover of species based on their probability of occurrence in wetlands) is one predicted response to groundwater decline. Other predicted impacts of groundwater decline include reduced establishment of Populus fremontii-Salix gooddingii forests, and reduced cover of herbaceous species associated with the fine-textured soils and shady conditions of floodplain terraces stabilized by these early seral tree species. High floodplain terraces (depth to groundwater of 5-8 m) had wetland indicator scores below those of upland sites and were vegetated by species (e.g., Prosopis velutina and Sporobolus wrightii) with low sensitivity to groundwater changes.

Effects of Groundwater Decline on Riparian Vegetation of Semiarid Regions: The San Pedro, Arizona

A natural abundance hydrogen stable isotope technique was used to study seasonal changes in source water utilization and water movement in the xylem of dimorphic root systems and stem bases of several woody shrubs or trees in mediterranean-type ecosystems of south Western Australia. Samples collected from the native treeBanksia prionotes over 18 months indicated that shallow lateral roots and deeply penetrating tap (sinker) roots obtained water of different origins over the course of a winter-wet/summer-dry annual cycle. During the wet season lateral roots acquired water mostly by uptake of recent precipitation (rain water) contained within the upper soil layers, and tap roots derived water from the underlying water table. The shoot obtained a mixture of these two water sources. As the dry season approached dependence on recent rain water decreased while that on ground water increased. In high summer, shallow lateral roots remained well-hydrated and shoots well supplied with ground water taken up by the tap root. This enabled plants to continue transpiration and carbon assimilation and thus complete their seasonal extension growth during the long (4–6 month) dry season. Parallel studies of other native species and two plantation-grown species ofEucalyptus all demonstrated behavior similar to that ofB. prionotes. ForB. prionotes, there was a strong negative correlation between the percentage of water in the stem base of a plant which was derived from the tap root (ground water) and the amount of precipitation which fell at the site. These data suggested that during the dry season plants derive the majority of the water they use from deeper sources while in the wet season most of the water they use is derived from shallower sources supplied by lateral roots in the upper soil layers. The data collected in this study supported the notion that the dimorphic rooting habit can be advantageous for large woody species of floristically-rich, open, woodlands and heathlands where the acquisition of seasonally limited water is at a premium.

Seasonal water uptake and movement in root systems of Australian phraeatophytic plants of dimorphic root morphology: a stable isotope investigation

Soil water is usually assumed to be equally available for all purposes, supplying plant transpiration as well as groundwater and streamflow; however, a study of hydrogen and oxygen isotopes from 47 globally distributed sites shows that in fact the water used by plants tends to be isotopically distinct from the water that feeds streamflow. Soil water is usually assumed to be available for all purposes in equal measure, supplying plant transpiration as well as groundwater and streamflow. Building on prior but limited studies, Jaivime Evaristo et al. have assembled a dataset of hydrogen and oxygen isotopes — drawn from widely distributed sites — and show that ecohydrological separation is the rule. Water used by plants tends to be isotopically distinct from that used for streamflow, suggesting that hydrological separation of precipitation inputs creates distinct pools of water resources. This finding implies that that existing land surface model parameterizations of plant physiological processes and streamflow can be made more realistic through the incorporation of ecohydrological separation. Current land surface models assume that groundwater, streamflow and plant transpiration are all sourced and mediated by the same well mixed water reservoir—the soil. However, recent work in Oregon1 and Mexico2 has shown evidence of ecohydrological separation, whereby different subsurface compartmentalized pools of water supply either plant transpiration fluxes or the combined fluxes of groundwater and streamflow. These findings have not yet been widely tested. Here we use hydrogen and oxygen isotopic data (2H/1H (δ2H) and 18O/16O (δ18O)) from 47 globally distributed sites to show that ecohydrological separation is widespread across different biomes. Precipitation, stream water and groundwater from each site plot approximately along the δ2H/δ18O slope of local precipitation inputs. But soil and plant xylem waters extracted from the 47 sites all plot below the local stream water and groundwater on the meteoric water line, suggesting that plants use soil water that does not itself contribute to groundwater recharge or streamflow. Our results further show that, at 80% of the sites, the precipitation that supplies groundwater recharge and streamflow is different from the water that supplies parts of soil water recharge and plant transpiration. The ubiquity of subsurface water compartmentalization found here, and the segregation of storm types relative to hydrological and ecological fluxes, may be used to improve numerical simulations of runoff generation, stream water transit time and evaporation–transpiration partitioning. Future land surface model parameterizations should be closely examined for how vegetation, groundwater recharge and streamflow are assumed to be coupled.

https://www.fs.fed.us/pnw/pubs/journals/pnw_2016_evaristo001.pdf

Global separation of plant transpiration from groundwater and streamflow

Water sources of Eucalyptus camaldulensis Dehn. trees were investigated on a semiarid floodplain in south-eastern Australia. The trees investigated ranged in distance from 0.5 to 40 m from a stream, with electrical conductivity 0.8 dSm−1, and grew over groundwater with electrical conductivity ranging from 30 to 50 dSm−1. The sources of water being used by the trees were investigated using the naturally occurring stable isotopes of water and measurements of soil water potential. Xylem water potential and leaf conductance were also examined to identify the trees' response to using these sources of water. Trees at distances greater than about 15 m from the stream used no stream water. The trees used groundwater in summer and a combination of groundwater and rain-derived surface-soil water (0.05–0.15 m depth) in winter. In doing so they suffered water stress at electrical conductivities higher than approximately 40 dSm−1 (equivalent to approximately −1.4 MPa). Trees adjacent to the stream used stream water directly in summer, but may have used stream water from the soil profile in winter, after the stream had risen and recharged the soil water. E. camaldulensis appeared to be partially opportunistic in the sources of water they used.

Sources of water used by riparian Eucalyptus camaldulensis overlying highly saline groundwater.

Over the last ten years, there has been a large increase in the number of vegetation studies that have incorporated measurements of the stable isotopic composition of water. There are many methods for measuring the amount of water being used by plants, but until recently it has been difficult to determine from where plants obtained their water. This has been particularly difficult where there is more than one available water source (e.g. where groundwater is shallow or streams are nearby).

The Use of Stable Isotopes of Water for Determining Sources of Water for Plant Transpiration

The isotopic composition of water extracted from various parts of alfalfa (Medicago saliva) plants was compared with that transpired by the plants to validate water extraction procedures, and to define the degree of alteration of the isotopic composition of water that occurred in different plant parts. The mean δ2H and δ18O values of water extracted from the upper tap root and lower crown regions were < 1‰ and 0.3‰ different from that of water transpired, respectively, with an analytical precision similar to that found in previous studies on soils and trees. Water from other plant parts closer to the leaves was enriched in 2H (up to 12.6 in upper stems) and 18O (4.2‰) with respect to that transpired. The enrichment was associated with a greater variability in δ2H values between replicate plants. δ2H‐δ18O data indicated that the isotopic enrichment was due to evaporation during transpiration. The study showed that azeotropic distillation is an accurate means of extracting water from alfalfa, altho...

Sampling water from alfalfa (Medicago sativa) for analysis of stable isotopes of water

Water sources used by creek-side river red gums in the semi-arid and saline Chowilla region of South Australia were determined from stable isotope measurements. Trees in two landscape positions, on creek banks and approximately 40m from creeks, were studied. The trees distant from the creek were only sampled when flooded. The {2}H and {18}O concentrations of water sampled from the trees were significantly different from those of the creek water, indicating that generally some water came from the saline soils or water table. This was found even in trees overlying high saline groundwater. Trees distant from the creek also showed little evidence of surface water uptake. The results indicated that the water relations of these trees are complex, and the trees may be less affected by changes in creek flow and/or salinity than previously anticipated.

Lisa J. Mensforth

Papers

Sources of water used by riparian Eucalyptus camaldulensis overlying highly saline groundwater.

The Use of Stable Isotopes of Water for Determining Sources of Water for Plant Transpiration

Sampling water from alfalfa (Medicago sativa) for analysis of stable isotopes of water

Uptake of Groundwater by Creek-side River Red Gums