Hydrogen isotopic compositions of n-alkanes from terrestrial plants correlate with their ecological life forms

Stable hydrogen isotopic compositions (δD) of compound-specific biomarkers, such as n-alkanes from plant leaf waxes, can be used as a proxy for paleoclimatic change. However, the relationship between hydrogen isotopes of plant leaf wax and plant ecological life forms is not well understood. Here, we report the δD of n-alkanes from 34 modern terrestrial plants, including twenty-one C₃ plants and thirteen C₄ plants from northwestern China, determined using gas chromatography/thermal conversion/isotope ratio mass spectrometry. Our data show that the stable hydrogen isotopes are poorly correlated with the plant photosynthetic pathway (C₃ vs. C₄) and that they do not give clear regional precipitation signals. Together with a comparative analysis of published δD values from plant leaf waxes in other regions, we believe that the stable hydrogen isotope of plant leaf waxes is more closely related to ecological life forms of these terrestrial plants (i.e. tree, shrub, and grass). In general, the grasses have more negative δD values than the co-occurring trees and shrubs. Our findings suggest that the δD values of sedimentary leaf waxes from higher plants may record changes of a plant ecosystem under the influence of environmental alteration and imply that reconstruction of the paleoclimate using δD values from plant n-alkanes should be based upon specific plant taxa, and comparison should be made among plants with similar ecological life forms.     

Observations of Hydrogen and Oxygen Isotopes in Leaf Water Confirm the Craig-Gordon Model under Wide-Ranging Environmental Conditions

The Craig-Gordon evaporative enrichment model of the hydrogen (δD) and oxygen (δ¹⁸O) isotopes of water was tested in a controlled-environment gas exchange cuvette over a wide range (400‰ δD and 40‰ δ¹⁸O) of leaf waters. (Throughout this paper we use the term “leaf water” to describe the site of evaporation, which should not be confused with “bulk leaf water” a term used exclusively for uncorrected measurements obtained from whole leaf water extractions.) Regardless of how the isotopic composition of leaf water was achieved (i.e. by changes in source water, atmospheric vapor δD or δ¹⁸O, vapor pressure gradients, or combinations of all three), a modified version of the Craig-Gordon model was shown to be sound in its ability to predict the δD and δ¹⁸O values of water at the site of evaporation. The isotopic composition of atmospheric vapor was shown to have profound effects on the δD and δ¹⁸O of leaf water and its influence was dependent on vapor pressure gradients. These results have implications for conditions in which the isotopic composition of atmospheric vapor is not in equilibrium with source water, such as experimental systems that grow plants under isotopically enriched water regimes. The assumptions of steady state were also tested and found not to be a major limitation for the utilization of the leaf water model under relatively stable environmental conditions. After a major perturbation in the δD and δ¹⁸O of atmospheric vapor, the leaf reached steady state in approximately 2 h, depending on vapor pressure gradients. Following a step change in source water, the leaf achieved steady state in 24 h, with the vast majority of changes occurring in the first 3 h. Therefore, the Craig-Gordon model is a useful tool for understanding the environmental factors that influence the hydrogen and oxygen isotopic composition of leaf water as well as the organic matter derived from leaf water.     

Multiple controls for the variability of hydrogen isotopic compositions in higher plant n-alkanes from modern ecosystems

We performed a global scale analysis of available leaf wax n -alkane δ D data compiled from our new results, as well as from the literature and expressed as average values of D/H ratios from three common lipids of n -alkanes with odd carbon numbers ( n -C₂₇, n -C₂₉, and n -C₃₁) from living higher plants. Our results clearly indicate multiple controls of hydrogen isotope composition and its variability in plants leaf wax. (1) At the global scale, precipitation δ D values play a dominating factor that exercises the first order of control for hydrogen isotopic compositions in plant leaf wax. The hydrogen isotopic composition of plant leaf wax tracks the decreasing trend of precipitation δ D with increasing latitude. (2) Because of different water acquisition systems, plant life form influences the hydrogen isotopic composition of leaf wax n -alkanes with woody plants and grasses having different responses to the change of global precipitation δ D. (3) Physiological difference, due to different photosynthesis pathways or different water usage strategies, can leave an imprint on δ D patterns of plant leaf waxes, causing δ D variations among plants using the same source water. While these results better explain the variability of hydrogen isotope composition in leaf wax, they also have important implications for the interpretation of n -alkane δ D data from fossils and ancient sediments.     

Inverse gradients in leaf wax δD and δ13C values along grass blades of Miscanthus sinensis: Implications for leaf wax reproduction and plant physiology

Compound specific hydrogen and carbon isotopic ratios of higher plant leaf waxes have been extensively used in paleoclimate and paleoenvironmental reconstructions. However, studies so far have focused on the comparison of leaf wax isotopic differences in bulk leaf samples between different plant species. We sampled three different varieties of tall grasses (Miscanthus sinensis) in six segments from base to tip and determined hydrogen and carbon isotopic ratios of leaf waxes, as well as hydrogen and oxygen isotopic ratios of leaf water samples. We found an increasing, base-to-tip hydrogen isotopic gradient along the grass blades that can probably be attributed to active leaf wax regeneration over the growth season. Carbon isotopic ratios, on the other hand, show opposite trends to hydrogen isotopic ratios along the grass blades, which may reflect different photosynthetic efficiencies at different blade locales.     

Effects of Mild Water Stress and Diurnal Changes in Temperature and Humidity on the Stable Oxygen and Hydrogen Isotopic Composition of Leaf Water in Cornus stolonifera L

In this paper we make comparisons between the observed stable isotopic composition of leaf water and the predictions of the Craig-Gordon model of isotopic enrichment when plants (Cornus stolonifera L.) were exposed to natural, diurnal changes in temperature and humidity in a glasshouse. In addition, we determined the effects of mild water stress on the isotopic composition of leaf water. The model predicted different patterns of diurnal change for the oxygen and hydrogen isotopic composition of leaf water. The observed leaf water isotopic composition followed qualitatively similar patterns of diurnal change to those predicted by the model. At midday, however, the model always predicted a higher degree of heavy isotope enrichment than was actually observed in leaves. There was no effect of mild water stress on the hydrogen isotopic composition of leaf water. For the oxygen isotopic composition of leaf water, there was either no significant difference between control and water-stressed plants or the stressed plants had lower δ¹⁸O values, despite the enriched stem water isotopic composition observed for the stressed plants.     

Carbon sources for the Palaeozoic giant fungus Prototaxites inferred from modern analogues

A wide range of carbon isotope values in the Devonian fossil Prototaxites has been interpreted to support heterotrophy and the classification of Prototaxites as a giant fungus. This inference remains controversial because of the huge size of Prototaxites relative to co-occurring terrestrial vegetation and the lack of existing fungal analogues that display equally broad isotopic ranges. Here, we show wide isotopic variability in the modern saprotrophic fungus Arrhenia obscurata collected adjacent to shallow meltwater pools of a sparsely vegetated glacial succession in the Washington Cascades, USA. Soils collected specifically around the edges of these pools were up to 5‰ higher in δ¹³C than adjacent soils consistent with C₃ origin. Microbial sources of primary production appear to cause these high δ¹³C values, and the environment may be analogous to that of the Early Devonian landscapes, where Prototaxites individuals with extreme isotopic variance were found. Carbon isotopes are also compared in Prototaxites, Devonian terrestrial vascular plants, and Devonian algal-derived lake sediments. Prototaxites isotopic values show little correspondence with those of contemporaneous tracheophytes, providing further evidence that non-vascular land plants or aquatic microbes were important contributors to its carbon sources. Thus, a saprotrophic fungal identity is supported for Prototaxites, which may have relied on deposits of algal-derived organic matter in floodplain environments that were less dominated by vascular plants than a straight reading of the macrofossil record might suggest.     

Isotopic Fractionation of Hydrogen in Plants

The naturally-occurring stable isotopes deuterium and hydrogen are fractionated by a number of physical and biological processes. Deuterium has a tendency to precipitate out first from a moist air mass. Thus ground water will become isotopically lighter with an increase in latitude, altitude, or distance inland. Water taken up by the plant from the soil undergoes little change until evapotranspiration results in leaf water becoming isotopically heavier. Thus hydrogen isotopes in plants can reveal something of geography (groundwater) and climate. Hydrogen isotopes undergo little fractionation by passage through the food chain, although plant parasites tend to be enriched in D as compared to their hosts, possibly due to higher rates of transpiration in the parasitic plants. The splitting of water in photosynthesis results in the lighter isotope being incorporated into organic matter. An even larger isotopic fractionation results during lipid synthesis and other processes involving the pyruvate dehydrogenase complex. Differences in metabolic pathway between species can be detected by D/H ratios. Hydrogen isotopic differences can be detected between CAM, C₄, and C₃ species. Within C₄ plants, the NADP-ME plants are isotopically distinguishable from NAD-ME and PEP-CK plants.     

Spatial patterns of leaf δ13C and δ15N of aquatic macrophytes in the arid zone of northwestern China

Analysis of stable isotope composition is an important tool in research on plant physiological ecology. However, large‐scale patterns of leaf‐stable isotopes for aquatic macrophytes have received considerably less attention. In this study, we examined the spatial pattern of stable isotopes of carbon (δ¹³C) and nitrogen (δ¹⁵N) of macrophytes leaves collected across the arid zone of northwestern China (approximately 2.4 × 10⁶ km²) and attempted to illustrate its relationship with environmental factors (i.e., temperature, precipitation, potential evapotranspiration, sediment total carbon and nitrogen). Our results showed that the mean values of the leaf δ¹³C and δ¹⁵N in the macrophytes sampled from the arid zone were −24.49‰ and 6.82‰, respectively, which were far less depleted than those measured of terrestrial plants. The order of averaged leaf δ¹³C from different life forms was as follows: submerged > floating‐leaved > emergent. Additionally, our studies indicated that the values of foliar δ¹³C values of all the aquatic macrophytes were only negatively associated with precipitation, but the foliar δ¹⁵N values were mainly associated with temperature, precipitation, and potential evapotranspiration. Therefore, we speculated that water‐relation factors are the leaf δ¹³C determinant of macrophytes in the arid zone of northwestern China, and the main factors affecting leaf δ¹⁵N values are the complex combination of water and energy factors.     

Biogeochemistry of the stable isotopes of hydrogen and carbon in salt marsh biota

Deuterium to hydrogen ratios of 14 plant species from a salt marsh and lagoon were 55‰ depleted in deuterium relative to the environmental water. Carbon tetrachloride-extractable material from these plants was another 92‰ depleted in deuterium. This gave a fractionation factor from water to CCl₄ extract of 1.147. This over-all fractionation was remarkably constant for all species analyzed. Plants also discriminate against ¹³C, particularly in the lipid fraction. Data suggest that different mechanisms for carbon fixation result in different fractionations of the carbon isotopes. Herbivore tissues reflected the isotopic ratios of plants ingested. Apparently different metabolic processes are responsible for the different degrees of fractionation observed for hydrogen and carbon isotopes.     

Developmental Changes in Composition and Morphology of Cuticular Waxes on Leaves and Spikes of Glossy and Glaucous Wheat (Triticum aestivum L.)

The glossy varieties (A14 and Jing 2001) and glaucous varieties (Fanmai 5 and Shanken 99) of wheat (Triticum aestivum L.) were selected for evaluation of developmental changes in the composition and morphology of cuticular waxes on leaves and spikes. The results provide us with two different wax development patterns between leaf and spike. The general accumulation trend of the total wax load on leaf and spike surfaces is first to increase and then decrease during the development growth period, but these changes were caused by different compound classes between leaf and spike. Developmental changes of leaf waxes were mainly the result of variations in composition of alcohols and alkanes. In addition, diketones were the third important contributor to the leaf wax changes in the glaucous varieties. Alkanes and diketones were the two major compound classes that caused the developmental changes of spike waxes. For leaf waxes, β- and OH-β-diketones were first detected in flag leaves from 200-day-old plants, and the amounts of β- and OH-β-diketones were significantly higher in glaucous varieties compared with glossy varieties. In spike waxes, β-diketone existed in all varieties, but OH-β-diketone was detectable only in the glaucous varieties. Unexpectedly, the glaucous variety Fanmai 5 yielded large amounts of OH-β-diketone. There was a significant shift in the chain length distribution of alkanes between early stage leaf and flag leaf. Unlike C₂₈ alcohol being the dominant chain length in leaf waxes, the dominant alcohol chain length of spikes was C₂₄ or C₂₆ depending on varieties. Epicuticular wax crystals on wheat leaf and glume were comprised of platelets and tubules, and the crystal morphology changed constantly throughout plant growth, especially the abaxial leaf crystals. Moreover, our results suggested that platelets and tubules on glume surfaces could be formed rapidly within a few days.     

Temperature effect on leaf water deuterium enrichment and isotopic fractionation during leaf lipid biosynthesis: results from controlled growth of C3 and C4 land plants

The hydrogen isotopic ratios ((2)H/(1)H) of land plant leaf water and the carbon-bound hydrogen of leaf wax lipids are valuable indicators for climatic, physiological, metabolic and geochemical studies. Temperature will exert a profound effect on the stable isotopic composition of leaf water and leaf lipids as it directly influences the isotopic equilibrium (IE) during leaf water evaporation and cellular water dissociation. It is also expected to affect the kinetics of enzymes involved in lipid biosynthesis, and therefore the balance of hydrogen inputs along different biochemical routes. We conducted a controlled growth experiment to examine the effect of temperature on the stable hydrogen isotopic composition of leaf water and the biological and biochemical isotopic fractionations during lipid biosynthesis. We find that leaf water (2)H enrichment at 20°C is lower than that at 30°C. This is contrary to the expectation that at lower temperatures leaf water should be more enriched in (2)H due to a larger equilibrium isotope effect associated with evapotranspiration from the leaf if all other variables are held constant. A hypothesis is presented to explain the apparent discrepancy whereby lower temperature-induced down-regulation of available aquaporin water channels and/or partial closure of transmembrane water channel forces water flow to "detour" to a more convoluted apoplastic pathway, effectively increasing the length over which diffusion acts against advection as described by the Péclet effect (Farquhar and Lloyd, 1993) and decreasing the average leaf water enrichment. The impact of temperature on leaf water enrichment is not reflected in the biological isotopic fractionation or the biochemical isotopic fractionation during lipid biosynthesis. Neither the biological nor biochemical fractionations at 20°C are significantly different from that at 30°C, implying that temperature has a negligible effect on the isotopic fractionation during lipid biosynthesis.     

Compensation point and isotopic characteristics of c(3)/c(4) intermediates and hybrids in panicum

Leaf CO₂ compensation points and stable hydrogen, oxygen and carbon isotope ratios were determined for Panicum species including C₃/C₄ intermediate photosynthesis plants, hybrids between C₃/C₄ intermediates and C₃ plants, C₃ and C₄ plants in the Panicum genus as well as several other C₃ and C₄ plants. C₃ plants had the highest compensation points, followed by hybrids, C₃/C₄ intermediates, and C₄ plants. δ¹³C values of cellulose nitrate and saponifiable lipids from C₄ plants were about 10‰ higher than those observed for cellulose nitrate and saponifiable lipids of C₃/C₄ intermediates, hybrids, and C₃ plants. Oxygen isotope ratios of cellulose as well as those of leaf water were similar for all plants. There was substantial variability in the δD values of cellulose nitrate among the plants studied. In contrast, such variability was not observed in δD values of water distilled from the leaves, nor in the δD values of the saponifiable lipids. Variability in δD values of cellulose nitrate from C₃/C₄ intermediates, hybrids, C₃, and C₄ plants is due to fractionations occurring during biochemical reactions specific to leaf carbohydrate metabolism.     

Phospholipase D δ knock-out mutants are tolerant to severe drought stress

Phospholipase D (PLD) is involved in different plant processes, ranging from responses to abiotic and biotic stress to plant development. Phospholipase Dδ (PLDδ) is activated in dehydration and salt stress, producing the lipid second messenger phosphatidic acid. In this work we show that pldδ Arabidopsis mutants were more tolerant to severe drought than wild-type plants. PLDδ has been shown to be required for ABA regulation of stomatal closure of isolated epidermal peels. However, there was no significant difference in stomatal conductance at the whole plant level between wild-type and pldδ mutants. Since PLD hydrolyses structural phospholipids, then we looked at membrane integrity. Ion leakage measurements showed that during dehydration of leaf discs pldδ mutant has less membrane degradation compared to the wild-type. We further analyzed the mutants and showed that pldδ have higher mRNA levels of RAB18 and RD29A compared to wild-type plants under normal growth conditions. Transient expression of AtPLDδ in Nicotiana benthamiana plants induced a wilting phenotype. These findings suggest that, in wt plants PLDδ disrupt membranes in severe drought stress and, in the absence of the protein (PLDδ knock-out) might drought-prime the plants, making them more tolerant to severe drought stress. The results are discussed in relation to PLDδ role in guard cell signaling and drought tolerance.     

Water stress enhances expression of an alpha-amylase gene in barley leaves

The amylases of the second leaves of barley seedlings (Hordeum vulgare L. cv Betzes) were resolved into eight isozymes by isoelectric focusing, seven of which were β-amylase and the other, α-amylase. The α-amylase had the same isoelectric point as one of the gibberellin-induced α-amylase isozymes in the aleurone layer. This and other enzyme characteristics indicated that the leaf isozyme corresponded to the type A aleurone α-amylase (low pI group). Crossing experiments indicated that leaf and type A aleurone isozymes resulted from expression of the same genes.

In unwatered seedlings, leaf α-amylase increased as leaf water potential decreased and ABA increased. Water stress had no effect on β-amylase. α-Amylase occurred uniformly along the length of the leaf but β-amylase was concentrated in the basal half of the leaf. Cell fractionation studies indicated that none of the leaf α-amylase occurred inside chloroplasts.

Leaf radiolabeling experiments followed by extraction of α-amylase by affinity chromatography and immunoprecipitation showed that increase of α-amylase activity involved synthesis of the enzyme. However, water stress caused no major change in total protein synthesis. Hybridization of a radiolabeled α-amylase-related cDNA clone to size fractionated RNA showed that water-stressed leaves contained much more α-amylase mRNA than unstressed plants. The results of these and other studies indicate that regulation of gene expression may be a component in water-stress induced metabolic changes.

     

Effects of trophic level and metamorphosis on discrimination of hydrogen isotopes in a plant-herbivore system

The use of stable isotopes in ecological studies requires that we know the magnitude of discrimination factors between consumer and element sources. The causes of variation in discrimination factors for carbon and nitrogen have been relatively well studied. In contrast, the discrimination factors for hydrogen have rarely been measured. We grew cabbage looper caterpillars (Trichoplusia ni) on cabbage (Brassica oleracea) plants irrigated with four treatments of deuterium-enriched water (δD = −131, −88, −48, and −2‰, respectively), allowing some of them to reach adulthood as moths. Tissue δD values of plants, caterpillars, and moths were linearly correlated with the isotopic composition of irrigation water. However, the slope of these relationships was less than 1, and hence, discrimination factors depended on the δD value of irrigation water. We hypothesize that this dependence is an artifact of growing plants in an environment with a common atmospheric δD value. Both caterpillars and moths were significantly enriched in deuterium relative to plants by ∼45‰ and 23‰ respectively, but the moths had lower tissue to plant discrimination factors than did the caterpillars. If the trophic enrichment documented here is universal, δD values must be accounted for in geographic assignment studies. The isotopic value of carbon was transferred more or less faithfully across trophic levels, but δ¹⁵N values increased from plants to insects and we observed significant non-trophic ¹⁵N enrichment in the metamorphosis from larvae to adult.     

Carbon Isotope Composition of Nighttime Leaf-Respired CO2 in the Agricultural-Pastoral Zone of the Songnen Plain,Northeast China

Variations in the carbon isotope signature of leaf dark-respired CO₂ (δ¹³C_R) within a single night is a widely observed phenomenon. However, it is unclear whether there are plant functional type differences with regard to the amplitude of the nighttime variation in δ¹³C_R. These differences, if present, would be important for interpreting the short-term variations in the stable carbon signature of ecosystem respiration and the partitioning of carbon fluxes. To assess the plant functional type differences relating to the magnitude of the nighttime variation in δ¹³C_R and the respiratory apparent fractionation, we measured the δ¹³C_R, the leaf gas exchange, and the δ¹³C of the respiratory substrates of 22 species present in the agricultural-pastoral zone of the Songnen Plain, northeast China. The species studied were grouped into C₃ and C₄ plants, trees, grasses, and herbs. A significant nocturnal shift in δ¹³C_R was detected in 20 of the studied species, with the magnitude of the shift ranging from 1‰ to 5.8‰. The magnitude of the nighttime variation in δ¹³C_R was strongly correlated with the daytime cumulative carbon assimilation, which suggests that variation in δ¹³C_R were influenced, to some extent, by changes in the contribution of malate decarboxylation to total respiratory CO₂ flux. There were no differences in the magnitude of the nighttime variation in δ¹³C_R between the C₃ and C₄ plants, as well as among the woody plants, herbs and graminoids. Leaf respired CO₂ was enriched in ¹³C compared to biomass, soluble carbohydrates and lipids; however the magnitude of enrichment differed between 8 pm and 4 am, which were mainly caused by the changes in δ¹³C_R. We also detected the plant functional type differences in respiratory apparent fractionation relative to biomass at 4 am, which suggests that caution should be exercised when using the δ¹³C of bulk leaf material as a proxy for the δ¹³C of leaf-respired CO₂.     

黑河中游春玉米叶片水δD和δ18O的富集过程和影响因素

植物水的稳定同位素分馏过程是水在土壤-植物-大气连续体中循环的重要环节。以往研究由于叶片水¹⁸O同位素比值(δ¹⁸O _l,b)和氘(D)同位素比值(δD_l,b)(合称δ_l,b)实测数量少只能作为模型验证数据, 导致δ_l,b富集机制研究多集中于模型研究, 缺乏基于野外试验条件的δ_l,b富集的控制机制研究。叶片水δD_l,b和δ¹⁸O _l,b的富集程度(ΔD_l,b和Δ¹⁸O _l,b, 合称Δ_l,b)通常表示为δ_l,b与茎秆水D同位素比值(δD_x)和¹⁸O同位素比值(δ¹⁸O_x) (合称δ_x)之差, 即Δ_l,b = δ_l,b - δ_x。该研究以黑河中游沙漠绿洲春玉米(Zea mays)生态系统为研究对象, 重点采集和分析了季节和日尺度δ_l,b和δ_x数据, 配套开展了大气水汽δ¹⁸O和δD (合称δ_v)等辅助变量的原位连续观测, 探讨了季节和日尺度上的δ_l,b富集特征及其影响因素。结果表明: 叶片水δ_l,b和Δ_l,b的季节变化趋势不明显, 而受蒸腾作用影响表现出白天富集夜间贫化的单峰日变化特征。对于D来说, 无论季节尺度上还是日尺度上, 大气水汽δ_v和相对湿度是δD_l,b和ΔD_l,b的主要环境控制因素; 而对于¹⁸O来说, 无论季节尺度上还是日尺度上, 相对湿度是δ¹⁸O _l,b和Δ¹⁸O _l,b的主要环境控制因素。由于D和¹⁸O在热力学平衡分馏上有约8倍差异, 直接分析叶片水ΔD_l,b和Δ¹⁸O_l,b与影响因素的差异性, 有助于理解叶片水δD和δ¹⁸O富集过程以及对模型发展有一定的指导意义。     

基于稳定同位素技术的塔里木河下游不同林龄胡杨的水分利用来源

水分是制约很多陆地生态系统植物生长和繁殖的重要因素, 在干旱地区尤为明显。利用稳定同位素技术探究塔里木河下游不同林龄胡杨(Populus euphratica)的水分来源情况, 了解生态输水背景下荒漠河岸林的水分利用循环与利用策略, 可为生态输水提供科学依据, 同时也可对同类地区的生态恢复提供借鉴。本研究通过测定塔里木河下游胡杨茎干水和各潜在水源(土壤水、地下水)的稳定氢氧同位素值(δD、δ¹⁸O), 应用多源线性混合模型(IsoSource)分析了各潜在水源对不同林龄胡杨的贡献比例, 并结合3种林龄胡杨不同土壤深度含水量的变化, 分析了胡杨的主要吸水层位。结果表明: (1)不同林龄胡杨样地的不同深度区间上的土壤水δ¹⁸O值存在显著差异(P < 0.05): 胡杨幼龄木、成熟木、过熟木木质部δ¹⁸O分别为-7.83 ± 0.07‰、-8.53 ± 0.11‰、-9.36 ± 0.21‰; 而δD值不存在显著差异(P > 0.05)。可据此来推断胡杨的主要吸水层位。(2)总体上, 三种林龄胡杨土壤水δ¹⁸O值随土壤深度增加而减小, 并趋于接近地下水的δ¹⁸O值。其中, 0-60 cm土壤水受蒸发影响比较大, 其同位素组成经历了强烈的蒸发分馏过程, 土壤含水量极少, 土壤水δ¹⁸O值偏正。(3)不同林龄胡杨所利用的水分来源不同: 胡杨幼龄木对于地表80 cm以下的土壤水以及地下水均有一定程度的利用, 对80-140 cm、140-220 cm和220-340 cm的土壤水平均利用比率依次为16.2%、21.4%和24.6%, 对地下水平均利用比率为24.5%; 成熟木主要利用220-340 cm的土壤水及地下水, 平均利用比率分别为36.9%和42.3%; 过熟木主要利用140-340 cm的土壤水及地下水, 平均利用比率分别为32.8%和49.3%。     

Pronghorn (Antilocapra americana) enamel phosphate δ18O values reflect climate seasonality: Implications for paleoclimate reconstruction

Stable oxygen isotope (δ¹⁸O) compositions from vertebrate tooth enamel are widely used as biogeochemical proxies for paleoclimate. However, the utility of enamel oxygen isotope values for environmental reconstruction varies among species. Herein, we evaluate the use of stable oxygen isotope compositions from pronghorn (Antilocapra americana Gray, 1866) enamel for reconstructing paleoclimate seasonality, an elusive but important parameter for understanding past ecosystems. We serially sampled the lower third molars of recent adult pronghorn from Wyoming for δ¹⁸O in phosphate (δ¹⁸O_PO4) and compared patterns to interpolated and measured yearly variation in environmental waters as well as from sagebrush leaves, lakes, and rivers (δ¹⁸O_w). As expected, the oxygen isotope compositions of phosphate from pronghorn enamel are enriched in ¹⁸O relative to environmental waters. For a more direct comparison, we converted δ¹⁸O_w values into expected δ¹⁸O_PO4* values (δ¹⁸O_W‐_PO4*). Pronghorn δ¹⁸O_PO4 values from tooth enamel record nearly the full amplitude of seasonal variation from Wyoming δ¹⁸O_W‐PO4* values. Furthermore, pronghorn enamel δ¹⁸O_PO4 values are more similar to modeled δ¹⁸O_W‐PO4* values from plant leaf waters than meteoric waters, suggesting that they obtain much of their water from evaporated plant waters. Collectively, our findings establish that seasonality in source water is reliably reflected in pronghorn enamel, providing the basis for exploring changes in the amplitude of seasonality of ancient climates. As a preliminary test, we sampled historical pronghorn specimens (1720 ± 100 AD), which show a mean decrease (a shift to lower values) of 1–2‰ in δ¹⁸O_PO4 compared to the modern specimens. They also exhibit an increase in the δ¹⁸O amplitude, representing an increase in seasonality. We suggest that the cooler mean annual and summer temperatures typical of the 18th century, as well as enhanced periods of drought, drove differences among the modern and historical pronghorn, further establishing pronghorn enamel as excellent sources of paleoclimate proxy data.     

Physiological and isotopic aspects of photosynthesis in peperomia

Physiological and isotopic aspects of several Peperomia species were investigated. All but one species had C₃-like stomatal behavior, in that stomata were open during the day and closed during the night. In these species, most atmospheric CO₂ uptake occurred during the day. Concurrent with this stomatal behavior, there were Crassulacean acid metabolism-like acid fluctuations in most species. Carbon and hydrogen isotope ratios of cellulose nitrate from Peperomia reflect their physiological behavior. The δ¹³C values of cellulose nitrate from Peperomia species were similar to values observed in C₃ plants and consistent with the daytime uptake of exogeneous CO₂ via the C₃ photosynthetic pathway. The δD values of cellulose nitrate from Peperomia species approach those of Crassulacean acid metabolism plants. These elevated δD values are caused by fractionations occurring during biochemical reactions and not as a consequence of water relations.