scholarly journals Evidence for distinct isotopic composition of sap and tissue water in tree stems: consequences for plant water source identification

Author(s):  
Adrià Barbeta ◽  
Régis Burlett ◽  
Paula Martín-Gómez ◽  
Bastien Fréjaville ◽  
Nicolas Devert ◽  
...  

AbstractFor decades, theory has upheld that plants do not fractionate water isotopes as they move across the soil-root interface or along plant stems. This theory is now being challenged by several recent studies reporting that the water held in woody stems has an isotopic composition that cannot be attributed to any potential water source. Isotopic offsets between stem and source water still need to be explained, as they prevent identifying unambiguously tree water’s origin from water isotope measurements. Here we show that isotopic offsets between stem and source water can be explained by micrometer-scale water isotope heterogeneity within woody stems and soil micropores. Using a novel technique to extract sap water in xylem conduits separately from the water held in other xylem tissues, we show that these non-conductive xylem tissues are more depleted in deuterium than sap water. We also report that, in cut stems and well-watered potted plants, the isotopic composition of sap water reflects well that of irrigation water, demonstrating that no isotopic fractionation occurs during root water uptake or the sap water extraction process. Previous studies showed that isotopic heterogeneity also exists in soils at the pore scale where water adsorbed onto soil particles is more depleted than capillary/mobile soil water. Data collected at a beech (Fagus sylvatica) forest indicate that sap water matches best the capillary/mobile soil water from deep soil horizons, indicating that micrometer-scale water isotope heterogeneity in soils and stems must be accounted for to unambiguously identify where trees obtain their water within catchments.Significance StatementForests are prime regulators of the water cycle over land. They return, via transpiration, a large fraction of precipitation back to the atmosphere, influence surface runoff, groundwater recharge or stream flow, and enhance the recycling of atmospheric moisture inland from the ocean. The isotopic composition of water in woody stems can provide unique information on the role forests play in the water cycle only if it can be unambiguously related to the isotopic composition of source water. Here, we report a previously overlooked isotopic fractionation of stem water whereby non-conductive tissues are more depleted in deuterium than sap water, and propose a new technique to extract sap water separately from bulk stem water to unambiguously identify plant water sources.

2021 ◽  
Author(s):  
Javier de la Casa ◽  
Adrià Barbeta ◽  
Asun Rodriguez-Uña ◽  
Lisa Wingate ◽  
Jérôme Ogeé ◽  
...  

<p> </p><p>Long-standing ecological theory establishes that the isotopic composition of the plant water reflects that of the root-accessed sources, at least in non-saline or non-xeric environments. However, a growing number of studies challenge this assumption by reporting plant-source offsets in water isotopic composition, for a wide range of ecosystems. We conducted a global meta-analysis to systematically quantify the magnitude of this plant-source offset in water isotopic composition and its potential explanatory factors. We compiled 108 studies reporting dual water isotopic composition (δ<sup>2</sup>H and δ<sup>18</sup>O) of plant and source water. From these studies, we extracted the δ<sup>2</sup>H and δ<sup>18</sup>O of both plant and source waters for 223 plant species from artic to tropical biomes. For each species and sampling campaign, within each study, we calculated the mean line conditioned excess (LC-excess), with the slope and intercept of the local meteoric water line, and the mean soil water line conditioned excess (SWL-excess), from the slope and intercept of the soil water evaporation line. For each study site and sampling campaign, we obtained land surface temperature and volumetric soil water from the ERA5 database. For each study species, we recorded the functional type, leaf habit and for those available wood density. We found, on average, a significantly negative SWL-excess: plant water was systematically more depleted in δ<sup>2</sup>H than soil water. In > 90% of the cases with significantly negative SWL-excess, we also found negative LC-excess values, meaning that access to sources alternative to soil water was unlikely to explain negative SWL-excess values. </p><p>Calculated SWL-excess was affected by temperature and humidity: there were larger mismatches between plant and source water in isotopic composition in colder and wetter sites. Angiosperms, broadleaved and deciduous species exhibited more negative SWL-excess values than gymnosperms, narrow-leaved and evergreen species. Our results suggest that when using the dual isotopic approach, potential biases in the adscription of plant water sources are more likely in broadleaved forests in humid, and cold regions. Potential underlying mechanism for these isotopic mismatches will be discussed.</p><p> </p>


2019 ◽  
Vol 23 (4) ◽  
pp. 2129-2146 ◽  
Author(s):  
Adrià Barbeta ◽  
Sam P. Jones ◽  
Laura Clavé ◽  
Lisa Wingate ◽  
Teresa E. Gimeno ◽  
...  

Abstract. We investigated plant water sources of an emblematic refugial population of Fagus sylvatica (L.) in the Ciron river gorges in south-western France using stable water isotopes. It is generally assumed that no isotopic fractionation occurs during root water uptake, so that the isotopic composition of xylem water effectively reflects that of source water. However, this assumption has been called into question by recent studies that found that, at least at some dates during the growing season, plant water did not reflect any mixture of the potential water sources. In this context, highly resolved datasets covering a range of environmental conditions could shed light on possible plant–soil fractionation processes responsible for this phenomenon. In this study, the hydrogen (δ2H) and oxygen (δ18O) isotope compositions of all potential tree water sources and xylem water were measured fortnightly over an entire growing season. Using a Bayesian isotope mixing model (MixSIAR), we then quantified the relative contribution of water sources for F. sylvatica and Quercus robur (L.) trees. Based on δ18O data alone, both species used a mix of top and deep soil water over the season, with Q. robur using deeper soil water than F. sylvatica. The contribution of stream water appeared to be marginal despite the proximity of the trees to the stream, as already reported for other riparian forests. Xylem water δ18O could always be interpreted as a mixture of deep and shallow soil waters, but the δ2H of xylem water was often more depleted than the considered water sources. We argue that an isotopic fractionation in the unsaturated zone and/or within the plant tissues could underlie this unexpected relatively depleted δ2H of xylem water, as already observed in halophytic and xerophytic species. By means of a sensitivity analysis, we found that the estimation of plant water sources using mixing models was strongly affected by this δ2H depletion. A better understanding of what causes this isotopic separation between xylem and source water is urgently needed.


2019 ◽  
Author(s):  
Adrià Barbeta ◽  
Sam P. Jones ◽  
Laura Clavé ◽  
Lisa Wingate ◽  
Teresa E. Gimeno ◽  
...  

Abstract. We investigated plant water sources of an emblematic refugial population of Fagus sylvatica (L.) in the Ciron river gorges in South-Western France using stable water isotopes. It is generally assumed that no isotopic fractionation occurs during root water uptake, so that the isotopic composition of xylem water effectively reflects that of source water. However, this assumption has been called into question by recent studies that found that, at least at some dates during the growing season, plant water did not reflect any mixture of the potential water sources. In this context, highly resolved datasets covering a range of environmental conditions could shed light on possible plant-soil fractionation processes responsible for this phenomenon. In this study, the hydrogen (δ2H) and oxygen (δ18O) isotope compositions of all potential tree water sources and xylem water were measured fortnightly over an entire growing season. Using a Bayesian isotope mixing model (MixSIAR), we then quantified the relative contribution of water sources for F. sylvatica and Quercus robur (L.) trees. Based on δ18O data alone, both species used a mix of top and deep soil water over the season, with Q. robur using soil water relatively deeper than F. sylvatica. The contribution of stream water appeared to be marginal despite the proximity of the trees to the stream, as already reported for other riparian forests. Xylem water δ18O could always be interpreted as a mixture of deep and shallow soil waters, but the δ2H of xylem water was often more depleted than the considered water sources. We argue that an isotopic fractionation in the unsaturated zone and/or within the plant tissues could underlie this unexpected relatively depleted δ2H of xylem water, as already observed in halophytic and xerophytic species. By means of a sensitivity analysis, we found that the estimation of plant water sources using mixing models was largely affected by this δ2H depletion. A better understanding of what causes this isotopic separation between xylem and source water is urgently needed.


2021 ◽  
Author(s):  
Anam Amin ◽  
Giulia Zuecco ◽  
Chiara Marchina ◽  
Michael Engel ◽  
Daniele Penna ◽  
...  

<p>Plant transpiration is a main component of the global water cycle and plays a key role in regulating ecohydrological process. Stable isotopes of oxygen and hydrogen are often used for the identification and quantification of plant water sources in ecohydrology. However, the isotopic tracing technique assumes that the isotopic signal in the water taken up by the plants remains unaltered during uptake at the soil-roots interface and transport to the distal twigs, i.e., isotopic fractionation does not occur during the water uptake and along the transport pathway. Nevertheless, recent studies showed that isotopic fractionation can occur under different environmental conditions. In this study, we performed a simple experiment with two olive (<em>Olea europaea</em>) trees utilizing labelled water to test isotopic fractionation of plant water during uptake and transport within the plants under controlled conditions. In addition, we performed the cryogenic vacuum distillation in two different laboratories to examine any possible effects of the extraction system on the isotopic composition of plant water extracts.</p><p>We set up the olive trees in pots inside a glasshouse and measured sap flow rates with Granier thermal dissipation probe, and shallow soil moisture by using a portable soil moisture probe. Air temperature, global solar radiation, and relative humidity were measured by a weather station installed inside the glasshouse nearby the olive trees. We irrigated the two plants with water of known isotopic composition and sampled the twigs, wood cores, roots, and soils at different depths (0-5, 5-15, and 15-25 cm). We extracted plant and soil waters by means of cryogenic vacuum distillation performed in two different laboratories.</p><p>Our results showed that the plant water samples reflected the isotopic signature of labelled water and mobile soil water, suggesting no isotopic fractionation during water transport. No significant differences were detected for twigs and wood cores extracted from distinct sections of the tree. However, only significant differences were obtained between plant tissue water (twigs, cores) and cryogenically-extracted deep soil water (i.e., >15 cm depths). Furthermore, we found no significant effects of the two cryogenic extraction systems on the isotopic composition of water extracts. Our results indicate that isotopic fractionation might not occur during root water uptake and transport processes in olive trees, at least under the specified experimental conditions, validating the conventional isotope-tracing approach. Further work both in the field and under controlled conditions, and on different plant species, is needed to check for this consistency, as well as testing other plant water extraction methods.</p><p> </p><p>Keywords: olive tree; stable isotope analysis; plant water; cryogenic vacuum distillation; fractionation; labelled water.</p>


2018 ◽  
Author(s):  
Adrià Barbeta ◽  
Sam P. Jones ◽  
Laura Clavé ◽  
Lisa Wingate ◽  
Teresa E. Gimeno ◽  
...  

Abstract. We investigated plant-water sources of an emblematic refugial population of Fagus sylvatica (L.) in the Ciron river gorges in South-Western France using stable isotopes. The stable isotopes of water are a powerful tracer of water fluxes in the soil-plant-atmosphere continuum. It is generally assumed that no isotopic fractionation occurs during root water uptake, and that xylem water isotopes effectively reflect source water isotopes. However, recent studies showed that under certain conditions the isotopes in plant water do not reflect any of the potential sources considered. Highly resolved datasets covering a range of environmental conditions could shed light on possible plant-soil fractionations processes. In this study, the hydrogen (δ2H) and oxygen (δ18O) isotope compositions of all potential tree water sources and xylem water were measured bi-weekly over an entire growing season. Using Bayesian isotope mixing models (MixSIAR), we then quantified the contribution of the considered sources to xylem water of F. sylvatica and Quercus robur (L.) trees. Based on δ18O data alone, both species used a mix of top and deep soil water over the season, with Q. robur using soil water relatively deeper than F. sylvatica. The contribution of stream water appeared to be marginal despite the proximity of the trees to the stream, as already reported for other riparian forests. Xylem water δ18O could always be interpreted as a mixture of deep and shallow soil waters, but the δ2H of xylem water was often more depleted than any other possible water source. We argue that an isotopic fractionation in the unsaturated zone and/or within the plant tissues could underlie this unexpected δ2H depletion of xylem water, as already observed in halophytic and xerophytic species. By means of a sensitivity analysis, we found that the estimation of plant-water sources using isotope mixing models was largely affected by this isotopic δ2H depletion. A better understanding of what causes this isotopic separation between xylem and source water is urgently needed.


2021 ◽  
Author(s):  
Javier de la Casa ◽  
Adrià Barbeta ◽  
Asun Rodríguez-Uña ◽  
Lisa Wingate ◽  
Jérôme Ogée ◽  
...  

Abstract. Isotope-based approaches to study plant water sources rely on the assumption that root water uptake and within-plant water transport are non-fractionating processes. However, a growing number of studies have reported offsets between plant and source water stable isotope composition, for a wide range of ecosystems. These isotopic offsets can result in the erroneous attribution of source water used by plants and potential overestimations of groundwater uptake by the vegetation. We conducted a global meta-analysis to quantify the magnitude of these plant-source water isotopic offsets and explore whether their variability could be explained by either biotic or abiotic factors. Our database compiled 112 studies, spanning arctic to tropical biomes that reported the dual water isotope composition (δ2H and δ18O) of plant (stem) and source water, including soil water. We calculated 2H offsets in two ways: a line conditioned excess (LC-excess) that describes the 2H deviation from the local meteoric water line, and a soil water line conditioned excess (SW-excess), that describes the deviation from the soil water line, for each sampling campaign within each study. We tested for the effects of climate (air temperature and soil water content), soil class and plant traits (growth form, leaf habit, wood density and parenchyma fraction and mycorrhizal habit) on LC-excess and SW-excess. Globally, stem water was more depleted in 2H than soil water (SW-excess < 0) by 3.02 ± 0.65 ‰. In 95 % of the cases where SW-excess was negative, LC-excess was negative, indicating that the uptake of water from mobile pools was unlikely to explain the observed soil-plant water isotopic offsets. SW-excess was more negative in cold and wet sites, whereas it was more positive in warm sites. Soil class and plant traits did not have any significant effect on SW-excess. The climatic effects on SW-excess suggest that methodological artefacts are unlikely to be the sole cause of observed isotopic offsets. Instead, our results support the idea that these offsets are caused by isotopic heterogeneity within plant stems whose relative importance will depend on soil and plant water status and evaporative demand. Our results would imply that plant-source water isotopic offsets may lead to inaccuracies when using the isotopic composition of bulk stem water as a proxy to infer plant water sources.


2016 ◽  
Vol 13 (10) ◽  
pp. 3175-3186 ◽  
Author(s):  
Guo Chen ◽  
Karl Auerswald ◽  
Hans Schnyder

Abstract. Hydrophilic surfaces influence the structure of water close to them and may thus affect the isotope composition of water. Such an effect should be relevant and detectable for materials with large surface areas and low water contents. The relationship between the volumetric solid : water ratio and the isotopic fractionation between adsorbed water and unconfined water was investigated for the materials silage, hay, organic soil (litter), filter paper, cotton, casein and flour. Each of these materials was equilibrated via the gas phase with unconfined water of known isotopic composition to quantify the isotopic difference between adsorbed water and unconfined water. Across all materials, isotopic fractionation was significant (p<0.05) and negative (on average −0.91 ± 0.22 ‰ for 18∕16O and −20.6 ± 2.4 ‰ for 2∕1H at an average solid : water ratio of 0.9). The observed isotopic fractionation was not caused by solutes, volatiles or old water because the fractionation did not disappear for washed or oven-dried silage, the isotopic fractionation was also found in filter paper and cotton, and the fractionation was independent of the isotopic composition of the unconfined water. Isotopic fractionation became linearly more negative with increasing volumetric solid : water ratio and even exceeded −4 ‰ for 18∕16O and −44 ‰ for 2∕1H. This fractionation behaviour could be modelled by assuming two water layers: a thin layer that is in direct contact and influenced by the surface of the solid and a second layer of varying thickness depending on the total moisture content that is in equilibrium with the surrounding vapour. When we applied the model to soil water under grassland, the soil water extracted from 7 and 20 cm depth was significantly closer to local meteoric water than without correction for the surface effect. This study has major implications for the interpretation of the isotopic composition of water extracted from organic matter, especially when the volumetric solid : water ratio is larger than 0.5 or for processes occurring at the solid–water interface.


2019 ◽  
Author(s):  
Magali F. Nehemy ◽  
Paolo Benettin ◽  
Mitra Asadollahi ◽  
Dyan Pratt ◽  
Andrea Rinaldo ◽  
...  

Abstract. The stable isotopes of oxygen and hydrogen (δ2H and δ18O) have been widely used to investigate plant water source partitioning. These tracers have shed new light on patterns of plant water use in time and space. However, this black box approach has limited our source water interpretations and mechanistic understanding. Here, we combine measurements of stable isotope composition in xylem and soil water pools with measurements of plant hydraulics, fine root distribution and soil matric potential to investigate mechanism(s) driving tree water source partitioning. We used a 2 m3 lysimeter planted with a small willow tree (Salix viminalis) to conduct a high spatial-temporal resolution experiment. We found that tree water source partitioning was driven mainly by tree water status and not by patterns of fine root distribution. Source water partitioning was regulated by plant hydraulic response to changing atmospheric demand and soil matric potential. The depth distribution of soil matric potential appeared to be the largest control on the patterns of soil water partitioning during periods of tree water deficit. Contrary to the common steady state assumption in ecohydrological source water investigations, our results show that tree water use is a dynamic process, driven by tree water deficit. Overall, our findings suggest new research foci for future plant water isotopic investigations, highlighting the importance of hydrometric measurements from the plant perspective.


2020 ◽  
Author(s):  
John Marshall ◽  
Matthias Cuntz ◽  
Matthias Beyer ◽  
Maren Dubbert ◽  
Kathrin Kuehnhammer

&lt;p&gt;Forest water use has been difficult to quantify. One promising approach is to measure the isotopic composition of plant water, e.g.&lt;br&gt;the transpired water vapor or xylem water, which often differs from that of other water vapor sources. Traditionally such&lt;br&gt;measurements have relied on the extraction of wood samples, which provide limited time resolution at great expense, and risk&lt;br&gt;possible artefacts. Utilizing a borehole drilled through a trees&amp;#8217; stem, we propose a new method based on the notion that water&lt;br&gt;vapor in a slow-moving airstream approaches equilibration with the much greater mass of liquid water in the xylem. We present&lt;br&gt;two empirical data sets showing that the method can work in practice. We then present theoretical models estimating the&lt;br&gt;equilibration times and exploring the limits at which the approach will fail. Given long enough boreholes and slow enough flows,&lt;br&gt;the method provides a simple, cheap, and accurate means of continuously estimating the isotopic composition of the source water&lt;br&gt;for transpiration.&lt;/p&gt;


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