Stable Isotopes and Carbon Cycle Processes in Forests and Grasslands

Abstract: Scaling and partitioning are frequently two difficult challenges facing ecology today. With regard to ecosystem carbon balance studies, ecologists and atmospheric scientists are often interested in asking how fluxes of carbon dioxide scale across the landscape, region and continent. Yet at the same time, physiological ecologists and ecosystem ecologists are interested in dissecting the net ecosystem CO₂ exchange between the biosphere and the atmosphere to achieve a better understanding of the balance between photosynthesis and respiration within a forest. In both of these multiple-scale ecological questions, stable isotope analyses of carbon dioxide can play a central role in influencing our understanding of the extent to which terrestrial ecosystems are carbon sinks. In this synthesis, we review the theory and present field evidence to address isotopic scaling of CO₂ fluxes. We first show that the ¹³C isotopic signal which ecosystems impart to the atmosphere does not remain constant over time at either temporal or spatial scales. The relative balances of different biological activities and plant responses to stress result in dynamic changes in the ¹³C isotopic exchange between the biosphere and atmosphere, with both seasonal and stand-age factors playing major roles influencing the ¹³C biosphere-atmosphere exchange. We then examine how stable isotopes are used to partition net ecosystem exchange fluxes in order to calculate shifts in the balance of photosynthesis and respiration. Lastly, we explore how fundamental differences in the ¹⁸O isotopic gas exchange of forest and grassland ecosystems can be used to further partition terrestrial fluxes.     

Physical and chemical basis of carbon isotope fractionation in plants

Naturally-occurring variations in the abundances of the stable isotopes of carbon and other elements can be used to understand the dynamics of natural processes in chemistry, biochemistry, biology, medicine, ecology and other fields. The use of carbon-13 isotopic abundances as an indicator of photosynthetic function in plants has become common. The purpose of this article is to describe the physical and chemical processes that contribute to the abundances of carbon-13 in plant materials, and to provide a framework for understanding how those processes control the isotopic contents of natural materials.     

Studies on the reactivity of malondialdehyde. I. Effects of acid concentration and solvent composition on deuterium exchange reactions

The rates of deuterium exchange reactions of malondialdehyde (MDA) and deuterated malondialdehyde (MDAd) have been studied as a function of acidity and the content of dimethyl sulfoxide (DMSO) in binary mixtures with D₂O . MDA incorporates deuterium from D₂O solutions in a first-order reaction with a rate constant (k_obs) that depends on the acid concentration. From this dependence, a catalytic constant, k_cat, can be derived (k_cat^MDA = 2.25 × 10⁵M^?s^?1). Similar kinetic behavior was found for MDAd in H₂O solutions, and in this case, k_cat^MDA = 1.56 × 10⁵M^?1s^?1. Results from reactions of MDA and MDAd in identical H₂OD₂O mixtures show that primary and secondary isotope effects are small (k_H/k_D = 1.13) and that solvent isotope effects cause most of the differences found between reactions in D₂O and H₂O. Reactions in binary DMSOd₆D₂O mixtures show a six-fold rate increase as the proportion of DMSOd₆ increases from 50% to 90%. These results also illustrate the relatively high reactivity of MDA at pH values well above its pK_a and the importance of medium composition on its reaction rate.     

利用反硝化细菌法测试水体硝酸盐氮氧同位素

反硝化细菌方法作为测试硝酸盐氮氧同位素组成的最新方法,具有可测试低浓度水样、对样品无需特殊处理、不会交叉污染和需样量少等诸多优点而得到迅速发展。本研究率先在国内实验室利用反硝化细菌法成功测试了硝酸盐氮氧同位素组成,将经过5～10d培养的反硝化细菌Pseudomonas aureofaciens离心,然后将菌液浓缩5倍,再向顶空进样瓶注入3mL菌液,密封后利用高纯氮气吹扫3h以上,注入50nmolNO3-水样经过夜培养灭活后,使用TraceGasPre-concentrator-Isoprime测试N2O同位素组成,结果表明,重现性和测试精度与国际上类似研究接近。该方法的建立对于国内开展河流及湖泊(水库)、降水等氮的生物地球化学循环将起到促进作用。     

碳与锶同位素在六盘水地下水研究中的应用

分季节采集贵州省六盘水市地下水,并分析其水化学组分、溶解无机碳及其碳同位素和锶同位素组成,研究其地下水的水质情况和污染特征。结果表明:研究区水化学组成主要是以HCO3-Ca为主,属于典型的喀斯特区域地下水组成特征;地下水方解石饱和指数SIcalcite接近稳态,具有弱侵蚀性,与pCO2之间呈负相关关系,说明有机污染物降解CO2对于水岩反应起到了相当程度的影响;枯水季节溶解无机碳含量高于丰水季节,而δ13C季节变化并不明显,同位素与水化学的分析表明,区内城镇居民区和农田区地下水受人为活动污染明显;研究区87Sr/86Sr变化不大,而污水的87Sr/86Sr在0.7080。通过水化学和同位素分析表明,人为输入对于地下水的影响与地质背景、污染源特征及水文条件等有关。     

Influence of water flow on skeletal isotopic composition in the coral Pocillopora damicornis

Fragments of branching Pocillopora damicornis coral colonies were grown in experimental flumes under two water flow regimes. Colony size and buoyant weight increased most rapidly in the fast-flow regime. Branch tips from the upper and outer parts of the colonies showed the lowest and most consistent skeletal oxygen isotope ratios. Flow regime had little influence on the lowest oxygen isotope ratios, which were at least 3.5‰ lighter than the calculated oxygen isotopic equilibrium. These “kinetic” isotope effects are comparable to those observed in Porites corals. Relatively more branch tips showed extreme ¹⁸O depletions under low-flow conditions, and among small coral colonies. Isotopic variability was greater among branch tips from the lower and inner parts of the colonies and at high flow. Skeletal oxygen and carbon isotope ratios generally showed positive correlations. Despite the particularly large offsets from isotopic equilibrium, the isotopically lightest branches showed the greatest isotopic consistency and therefore would make the best isotopic thermometers. Isotopic variability within the colony may provide an indication of flow regime.     

Regional differences in the carbon isotopic compositions of teak from two monsoonal regimes of India

Inter and intra-annual carbon isotope compositions (δ¹³C) of several annual growth rings of teak trees from two monsoonal regimes from India were studied and compared with the corresponding oxygen isotopic (δ¹⁸O) variations. In teak from both the regimes, amplitudes of intra-annual δ¹³C were ∼2-3 times lower than that observed in δ¹⁸O. Seasonal cycle with lower δ¹³C values at the middle and higher at ring boundaries was observed for teak from central India, dominated by the southwest monsoon. Positive correlations of intra-annual δ¹³C values with the corresponding δ¹⁸O values of the same rings and with relative humidity (RH) of the concurrent period suggest a dominant role of RH in controlling δ¹³C values of teak from central India. Intra-annual δ¹³C variations of teak from southern India, receiving both the southwest and northeast monsoons, revealed an initial decreasing trend followed by an increasing trend before culminating in depleted ¹³C values at the end of the growing season. No correlation was observed between intra-annual δ¹³C and δ¹⁸O variations of teak trees from southern India. Regional differences in the climatology of δ¹³C of atmospheric CO₂ or the lengths of growing season could be likely reasons for differing intra-annual δ¹³C variations of teak from the two climatic regimes.     

Intraspecific variation in hair delta(13)C and delta(15)N values of ring-tailed lemurs (Lemur catta) with known individual histories, behavior, and feeding ecology

Stable carbon and nitrogen isotope compositions were analyzed from hair samples of 30 sympatric ring-tailed lemurs (Lemur catta) inhabiting the Beza Mahafaly Special Reserve, Madagascar. All lemurs were known individuals involved in a longitudinal study, which allowed us to explore the degree to which group membership, sex, health status, and migration influenced their stable isotope compositions. The differences in delta(13)C and delta(15)N values between groups were small (<1.5 per thousand) but highly significant. In fact, each group was tightly clustered, and discriminant function analysis of the stable isotope data assigned individuals to the group in which they were originally collared with over 90% accuracy. In general, the differences between groups reflected the degree to which they utilized forested versus open habitats. As open habitats at Beza Mahafaly often correspond to areas of anthropogenic disturbance, these data suggest that isotopic data can be useful for addressing questions of lemur conservation. There were few sex differences, but significant differences did occur between individuals of normal and suboptimal health, with those in poor health (especially those in the worst condition) being enriched in (15)N and to a lesser degree (13)C compared with healthy individuals. Moreover, lemurs that had emigrated between 2003 and 2004 had different delta(13)C and delta(15)N compositions than their original groups.     

Ecological and cultural shifts of hunter‐gatherers of the Jomon period paralleled with environmental changes

Objectives

Holocene hunter‐gatherers adapted to climatic and environmental changes over time. Carbon and nitrogen stable isotope analysis of human skeletal remains from the Inariyama shell mound of the Final Jomon period have revealed large dietary variations in the population. This study analyzed radiocarbon dates of these individuals to test temporal changes in diet and its relationship with tooth ablation.

Materials and Methods

Twenty‐nine human skeletal remains from Inariyama were included in this study. Extracted bone collagen samples were purified to graphite. Then, radiocarbon dating of these samples was performed using the accelerator mass spectrometer.

Results

The radiocarbon ages of Inariyama ranged about, 3,230–2,140 cal BP and showed three peaks of occupation. In the early and late phases, terrestrial resource consumption and incisor extraction were observed, while marine resource consumption and canine extraction were observed in the middle phase.

Discussion

These temporal changes of diet and tooth ablation types occurred in parallel with climatic cooling and environmental change and help reveal how Holocene hunter‐gatherers adapted to the changing environments.

     

Spatial variability in Florida Bay particulate organic matter composition: combining flow cytometry with stable isotope analyses

Long-term management plans for restoration of natural flow conditions through the Everglades increase the importance of understanding potential nutrient impacts of increased freshwater delivery on Florida Bay biogeochemistry. Planktonic communities respond quickly to changes in water quality, thus spatial variability in community composition and relationships to nutrient parameters must be understood in order to evaluate future downstream impacts of modifications to Everglades hydrology. Here we present initial results combining flow cytometry analyses of phytoplankton and bacterial populations (0.1–50 μm size fraction) with measurements of δ¹³C and δ¹⁵N composition and dissolved inorganic nutrient concentrations to explore proxies for planktonic species assemblage compositions and nutrient cycling. Particulate organic material in the 0.1–50 μm size fraction was collected from five stations in Northeastern and Western Florida Bay to characterize spatial variability in species assemblage and stable isotopic composition. A dense bloom of the picocyanobacterium, Synechococcus elongatus, was observed at Western Florida Bay sites. Smaller Synechococcus sp. were present at Northeast sites in much lower abundance. Bacteria and detrital particles were also more abundant at Western Florida Bay stations than in the northeast region. The highest abundance of detritus occurred at Trout Creek, which receives freshwater discharge from the Everglades through Taylor Slough. In terms of nutrient availability and stable isotopic values, the S. elongatus population in the Western bay corresponded to low DIN (0.5 μM NH ₄ ⁺ ; 0.2 μM NO ₃ ⁻ ) concentrations and depleted δ¹⁵N signatures ranging from +0.3 to +0.8‰, suggesting that the bloom supported high productivity levels through N₂-fixation. δ¹⁵N values from the Northeast bay were more enriched (+2.0 to +3.0‰), characteristic of N-recycling. δ¹³C values were similar for all marine Florida Bay stations, ranging from −17.6 to −14.4‰, however were more depleted at the mangrove ecotone station (−25.5 to −22.3‰). The difference in the isotopic values reflects differences in carbon sources. These findings imply that variations in resource availability and nutrient sources exert significant control over planktonic community composition, which is reflected by stable isotopic signatures.     

海洋生态系统稳定同位素基线的选取

稳定同位素技术在海洋食物网研究中得到了广泛的应用,但在分析海洋生物食性和营养级时,需要选择某种生物或物质的稳定同位素值作为基线.本文综述了河口和海湾、浅海、大洋及深海4类典型海洋生态系统稳定同位素基线的选取,分析了影响稳定同位素基线选择的因素,以及特定化合物稳定同位素在消除基线时空异质性中的潜在价值,并对目前存在的问题以及今后的研究方向进行了展望,以期为国内学者进一步深入开展海洋生态系统的稳定同位素生态学研究提供有益参考.     

稳定同位素技术在植物水分利用研究中的应用

近20a稳定同位素技术在植物生态学研究中的应用得到了长足发展,使得对植物与水分关系也有了更深一步的了解。介绍稳定同位素性碳、氢、氧同位素在研究植物水分关系中的应用及进展,以期能为国内植物水分利用研究提供参考。由于植物根系从土壤中吸收水分时并不发生同位素分馏,对木质部水分同位素分析有助于对植物利用水分来源,生态系统中植物对水分的竞争和利用策略的研究,更好地了解生态系统结构与功能。稳定碳同位素作为植物水分利用效率的一个间接指标,在不同水分梯度环境中,及植物不同代谢产物与水分关系中有着广泛的应用。同位素在土壤-植被-大气连续体水分中的应用,有助于了解生态系统的水分平衡。随着稳定同位素方法的使用,植物与水分关系的研究将取得更大的进展。     

Experimental Evidence for the Isotope Effect in Photorespiration

Recent data on carbon isotope fractionation in photosynthesis are reviewed. Analysis of the carbon isotope composition in photosynthates and derivative products supports the hypothesis of the isotope effect in photorespiration. This hypothesis envisages the existence in a photosynthesizing cell of two carbon flows differing in isotope composition. One of these flows is enriched in ¹²C and associated with the assimilation pathway of photosynthesis. The other flow enriched in ¹³C circulates in the photorespiratory pathway. The relation between stimulated photorespiration and the carbon isotope composition of biomass supports this view. Our hypothesis of two interrelated isotope effects in photosynthesis leads to the conclusion that photosynthesis and photorespiration are coupled processes subject to periodical oscillations, where Rubisco acts as a main switch regulating these two pathways.     

四川裂腹鱼耳石碳、氧稳定同位素特征

为探讨耳石碳(δ¹³C)、氧(δ¹⁸O)稳定同位素在淡水鱼类群体识别中的作用,本研究以养殖条件下不同年龄组四川裂腹鱼为对象,采用稳定同位素质谱仪进行碳、氧同位素测定,揭示耳石中碳、氧稳定同位素特征,探讨其与环境间的关系. 结果表明:1⁺龄四川裂腹鱼δ¹³C和δ¹⁸O值均与耳石质量无显著相关关系,但在微耳石和星耳石之间存在显著差异;不同年龄四川裂腹鱼微耳石δ¹³C和δ¹⁸O平均值分别为(-9.58±0.06)‰、(-8.33±0.17)‰,其在雌雄个体之间均无显著性差异,但在不同年龄组间存在显著差异. 耳石δ¹⁸O和δ¹³C的关联分析能有效区分四川裂腹鱼不同养殖年龄群体,可作为一种识别淡水鱼类养殖群体的手段.     

KATBERGIA GEN. NOV., A NEW TRACE FOSSIL FROM UPPER PERMIAN AND LOWER TRIASSIC ROCKS OF THE KAROO BASIN: IMPLICATIONS FOR PALAEOENVIRONMENTAL CONDITIONS AT THE P/TR EXTINCTION EVENT

Abstract:  A new ichnogenus and ichnospecies of burrow, Katbergia carltonichnus , are described from Upper Permian and Lower Triassic rocks of the Karoo Basin, South Africa, where they are preserved in pedogenically modified overbank deposits that are interpreted as inceptisols subsequently gleysol overprinted. Sigmoidal burrows consist of a long (≥0.5 m) cylindrical tube, ranging from 1–2 cm in diameter, terminating in a slightly larger living chamber. The burrows are unlined and passively filled, preserving a hierarchy of scratch patterns on the burrow walls. Scratch patterns include longitudinal, transverse, and crescent-shaped markings found around the circumference of the burrow, but which are less densely concentrated on the burrow floor. Calcareous concretions are associated with burrowed siltstone intervals, generally restricted to the lowermost decimetre, with nodules nucleating around burrows. Stable δ¹³C and δ¹⁸O isotope data on calcite cement in the burrow fill, entombing siltstone, and associated calcareous nodules all cluster together when plotted, indicating that nodule formation occurred following burrow horizon abandonment and a rise in regional water table. Isotopic data reflect calcite precipitation under a semi-closed system in saturated conditions. A model for burrow emplacement, abandonment and infill, and subsequent cementation by calcite is presented demonstrating that previous interpretations of Late Permian and Early Triassic palaeosol types associated with the P/Tr extinction event must be re-evaluated.     

Leaf carbon isotope discrimination and nitrogen content for riparian trees along elevational transects

 Leaf carbon isotope discrimination (Δ), seasonal estimates of the leaf-to-air water vapor gradient on a molar basis (ω), and leaf nitrogen contents were examined in three riparian tree species (Populus fremontii, P. angustifolia, and Salix exigua) along elevational transects in northern and southern Utah USA (1500–2670 m and 600–1820 m elevational gradients, respectively). The ω values decreased with elevation for all species along transects. Plants growing at higher elevations exhibited lower Δ values than plants at lower elevations (P. fremontii, 22.9‰ and 19.5‰, respectively; P. angustifolia, 23.2‰ and 19.2‰, respectively; and S.␣exigua, 21.1‰ and 19.1‰, respectively). Leaf nitrogen content increased with elevation for all species, suggesting that photosynthetic capacity at a given intercellular carbon dioxide concentration was greater at higher elevations. Leaf Δ and nitrogen content values were highly correlated, implying that leaves with higher photosynthetic capacities also had lower intercellular carbon dioxide concentrations. No significant interannual differences were detected in carbon isotope discrimination. Received: 25 February 1996 / Accepted: 8 September 1996     

特定化合物同位素分析技术在树木非结构性碳水化合物研究中的应用

特定化合物同位素分析(CSIA)可以实现对复杂基质中特定化合物稳定碳同位素组成(δ¹³C)的精确测定。应用此方法测定树木非结构性碳水化合物(NSC)中特定成分(如糖类、有机酸和糖醇)的δ¹³C,不仅能够追踪新同化的光合产物在树木中的运移及与外界的碳交换,还能够更敏感地指示树木生理状况对环境变化的响应。本文首先系统介绍了CSIA从样品采集、处理到δ¹³C测定的方法,然后综述了树木NSC中各成分之间及各成分在不同器官之间的δ¹³C差异,阐述了树木NSC的δ¹³C时间动态变化特征及内在机制,最后分析了NSC作为主要呼吸底物,其δ¹³C与树木呼吸释放CO₂的δ¹³C(δ¹³C_R)之间的联系,并针对CSIA分析技术在后光合分馏、树木逆境生理和年轮δ¹³C形成机制等研究的应用前景提出了展望。