Stable isotopes in leaf water of terrestrial plants

Leaf water contains naturally occurring stable isotopes of oxygen and hydrogen in abundances that vary spatially and temporally. When sufficiently understood, these can be harnessed for a wide range of applications. Here, we review the current state of knowledge of stable isotope enrichment of leaf water, and its relevance for isotopic signals incorporated into plant organic matter and atmospheric gases. Models describing evaporative enrichment of leaf water have become increasingly complex over time, reflecting enhanced spatial and temporal resolution. We recommend that practitioners choose a model with a level of complexity suited to their application, and provide guidance. At the same time, there exists some lingering uncertainty about the biophysical processes relevant to patterns of isotopic enrichment in leaf water. An important goal for future research is to link observed variations in isotopic composition to specific anatomical and physiological features of leaves that reflect differences in hydraulic design. New measurement techniques are developing rapidly, enabling determinations of both transpired and leaf water δ¹⁸O and δ²H to be made more easily and at higher temporal resolution than previously possible. We expect these technological advances to spur new developments in our understanding of patterns of stable isotope fractionation in leaf water.     

Simulations of the Ontogeny of Spongiophyton, a Devonian Plant

Mathematical representations of the cellular organization anddimensions of Spongiophyton Krusel, a Middle Devonian thallophytewere projected by means of a computer to simulate patterns ofdevelopment and organization. Extrapolation of the cellularpatterns observed on the surface of the fossil may be comparedwith those derived from living plants of which the ontogenycan be directly observed. Spongiophyton is compared in thisrespect with the growth of Protosalvinia (an Upper Devonianplant of enigmatic affinity), Pellia (a thallose liverwort)and Cutleria (a brown alga). The growth pattern of Spongiophytondeveloped by computer shows a closer similarity to that of thepseudoparenchymatous alga Cutleria than to the truly parenchymatousPellia or the fossil Protosalvinia. Computer simulations ofthe growth process throw light on the affinity of Spongiophytonwhich cannot be derived from direct observation of the fossil.Broader applications of computer simulations of tissue organizationand gross morphology are suggested with regard to the studyof living and fossil plants.     

Stable isotopes and the metabolism of the European cave bear

Isotopic analyses of fossil bones of the extinct European cave bear indicate that this animal was a hibernator with the same unusual metabolic processes as some modern bear species. This finding provides useful biological and archaeological information on an extinct species, and the methods themselves may prove generally useful in studies of the metabolisms of modern bears, other hibernators, and perhaps of starving animals. Received: 6 October 1997 / Accepted: 31 March 1998     

Stable isotopes in the management and diagnosis of inborn errors of metabolism

Stable isotope techniques offer advantages over older methods in safety, sensitivity, specificity, and reduction in number of subjects required for analytic determinations in some types of studies in "inborn errors of metabolism." In addition to their use as internal standards for gas chromatography - mass spectrometry, quantitation of plasma substrates, and their urinary metabolites, stable isotopes have been successfully employed in studies of metabolite identification, enzyme activity, nutrient turnover and requirements, and diagnosis of inborn errors of metabolism.     

Stable isotopes of carbon and nitrogen in soil ecological studies

The development of stable isotope techniques is one of the main methodological advances in ecology of the last decades of the 20th century. Many biogeochemical processes are accompanied by changes in the ratio between stable isotopes of carbon and nitrogen (¹²C/¹³C and ¹⁴N/¹⁵N), which allows different ecosystem components and different ecosystems to be distinguished by their isotopic composition. Analysis of isotopic composition makes it possible to trace matter and energy flows through biological systems and to evaluate the rate of many ecological processes. The main concepts and methods of stable isotope ecology and patterns of stable isotope fractionation during organic matter decomposition are considered with special emphasis on the fractionation of isotopes in food chains and the use of stable isotope studies of trophic relationships between soil animals in the field.     

Fourier transform infrared characterization of the Middle Devonian non‐vascular plant Spongiophyton

Spectroscopy information (functional groups and semiquantitative data) by Fourier transform infrared (FTIR) spectroscopy of Spongiophyton Kräusel emend. Chaloner et al. is reported for the first time, in an attempt to identify spectroscopic patterns that would differentiate species of these taxa. A total of 33 specimens identified as S. lenticularis, S. nanum and S. minutissimum and preserved as compressions were analysed. They come from the same fossiliferous horizon of one outcrop situated in the uppermost Middle Devonian São Domingos Formation of the Paraná Basin in southern Brazil. The results reveal that functional groups in the 3000–2800 and 1800–700 cm^?1 wavenumber regions are present in all specimens. Infrared‐derived ratios (CH₂/CH₃, Al/Ox, Ox1/Ox2, C=O cont., C=C cont., Al2 and Ar/Al) and types of kerogen led to a comparison of the morphological characteristics of the studied material and indicate that compressions of S. lenticularis, S. nanum and S. minutissimum contain different organic compounds. S. nanum and S. minutissimum show higher oxidation and lower aromaticity of its organic matter than S. lenticularis. Additionally, S. lenticularis shows low A‐factor values, suggesting a composition similar to type III kerogen, while S. nanum and S. minutissimum showed intermediate to high A‐factor values, suggesting a composition similar to type II kerogen. In conclusion, the chemical‐analytical techniques and statistical analysis (using PCA technique) reveal a closer taxonomic relationship between S. nanum and S. minutissimum than either has with S. lenticularis, supporting their taxonomic separation, previously proposed in the literature on the basis of morphological features. Lastly, the chemical signatures support the hypothesis that links these enigmatic fossils to a type of lichen or other early inland plant.     

Stable carbon isotopes in tree rings of beech: climatic versus site-related influences

 Stable carbon isotopes in tree rings are a promising tool in palaeoclimate research, provided attempts are made to disentangle climatic from local effects (e.g. soil properties, competition, light). The ¹³C/¹²C variations in cellulose of tree rings of beech (Fagus sylvatica) were determined at several sites in the Swiss Central Plateau covering the last 50 years. We chose sites which differ in moisture conditions and sampled cores from four to six trees per site. The mean ¹³C/¹²C series from the different dry sites (distant by up to 40 km) are closely interrelated suggesting a common external cause. Correlation analysis with climate data proved the total precipitation in the months May, June and July to have the strongest effect on the carbon isotopes (r =  – 0.73). This result is in agreement with the commonly used model which relates the isotope discrimination to the water use efficiency. On the other hand, the isotope series of the wet sites are not as well correlated to the climate. At two of the sites (a dry and a humid) tree ring width suddenly increased. We used this effect as a test-case to study the influence of local growth conditions on the climate-isotope relationship. Received: 17 April 1996 / Accepted: 2 September 1996     

Stable carbon isotopes of lipid biomarkers: analysis of metabolites and metabolic fates of environmental microorganisms

Lipid biomarkers are specific compounds that are characteristic of certain groups or species of microorganisms. The use of natural or labeled carbon isotopes of lipid biomarkers has enabled a better understanding of carbon flow pathways at the molecular level. Recent advances include, but are not limited to, the elucidation of mechanisms of anaerobic methane oxidation mediated by syntrophic sulfate-reducing bacteria and Archaea, linking microbial populations with specific microbial processes or bacterial transport mechanisms in natural or contaminated environments, and elucidation of the biosynthetic pathways of cellular material.     

Stable carbon and oxygen isotopes in human tooth enamel: Identifying breastfeeding and weaning in prehistory

This paper investigates the utility of stable carbon and oxygen isotopes in human dental enamel to reveal patterns of breastfeeding and weaning in prehistory. Enamel preserves a record of childhood diet that can be studied in adult skeletons. Comparing different teeth, we used δ¹³C to document the introduction of solid foods to infant diets and δ¹⁸O to monitor the decline of breastfeeding. We report enamel carbonate δ¹³C and δ¹⁸O of 33 first molars, 35 premolars, and 25 third molars from 35 burials from Kaminaljuyú, an early state in the valley of Guatemala. The skeletons span from Middle Preclassic through Late Postclassic occupations, ca. 700 B.C. to 1500 A.D. Sections of enamel were removed from each tooth spanning from the cusp to the cemento-enamel junction. Stable isotope ratios were measured on CO₂ liberated by reaction of enamel with H₃PO₄ in an automated carbonate system attached to a VG Optima mass spectrometer. Within a skeleton, teeth developing at older ages are more enriched in ¹³C and more depleted in ¹⁸O than teeth developing at younger ages. Premolars average 0.5% higher in δ¹³C than first molars from the same skeleton (P = 0.0001), but third molars are not significantly enriched over premolars. The shift from first molars to premolars may be due to the shift to solid foods from lipid-rich milk. After 2 years, when premolars begin to mineralize, the δ¹³C in childhood diets did not change systematically. First molars and premolars are similar in δ¹⁸O, but third molars average 0.7% lower than first molars (P = 0.0001) and 0.5% lower than premolars (P = 0.0003). First molar and premolar δ¹⁸O is heavier, because breast milk is more enriched in ¹⁸O than is drinking water. Hence, many children continued to nurse during the period of premolar formation. Together, these results indicate that Kaminaljuyú children had begun to eat solid maize foods before the age of 2 years but continued to drink breast milk until much later. Am J Phys Anthropol 106:1–18, 1998. © 1998 Wiley-Liss, Inc.     

Trophic fractionation of carbon and nitrogen stable isotopes in Chironomus riparius reared on food of aquatic and terrestrial origin

1. Trophic fractionation was studied in short‐term laboratory feeding experiments with larvae of the deposit‐feeding midge Chironomus riparius. Larvae were fed food of terrestrial (oats, peat) and aquatic origin (Spirulina, Tetraphyll^®). 2. By analysing both whole larvae and isolated gut contents we were able to distinguish between the isotopic signature of recently ingested food and that of assimilated carbon and nitrogen in body tissue. Additionally we studied the effects of microbial conditioning, i.e. the colonisation and growth on food particles of microbes, on the isotopic signal of food resources. 3. Nitrogen fractionation for the different food types ranged from 0.67‰ to 2.68‰ between consumer and diet and showed that isotopic fractionation can be much lower than the value of 3.4‰ that is commonly assumed. 4. Microbial degradation of food particles resulted in an approximate doubling of the δ¹⁵N in 8 days, from 6.24 ± 0.05‰ to 11.36 ± 0.56‰. Values for δ¹³C increased only marginally, from ?20.66 ± 0.11‰ to ?20.34 ± 0.12‰. These results show that microbial conditioning of food may affect dietary isotope signatures (in particular N) and, unless accounted for, could introduce an error in measures of trophic fractionation. Microbial conditioning could well account for some of the variation in fractionation reported in the literature.     

Predictability of the terrestrial carbon cycle

Terrestrial ecosystems sequester roughly 30% of anthropogenic carbon emission. However this estimate has not been directly deduced from studies of terrestrial ecosystems themselves, but inferred from atmospheric and oceanic data. This raises a question: to what extent is the terrestrial carbon cycle intrinsically predictable? In this paper, we investigated fundamental properties of the terrestrial carbon cycle, examined its intrinsic predictability, and proposed a suite of future research directions to improve empirical understanding and model predictive ability. Specifically, we isolated endogenous internal processes of the terrestrial carbon cycle from exogenous forcing variables. The internal processes share five fundamental properties (i.e., compartmentalization, carbon input through photosynthesis, partitioning among pools, donor pool‐dominant transfers, and the first‐order decay) among all types of ecosystems on the Earth. The five properties together result in an emergent constraint on predictability of various carbon cycle components in response to five classes of exogenous forcing. Future observational and experimental research should be focused on those less predictive components while modeling research needs to improve model predictive ability for those highly predictive components. We argue that an understanding of predictability should provide guidance on future observational, experimental and modeling research.     

Stable carbon isotopes of methane for real‐time process monitoring in anaerobic digesters

Efficient operation and stability of biogas plants requires continuous monitoring of the digester content. Traditional laboratory analysis of digester sludge is often complex and time‐consuming and shows a delayed response to disruptions within the fermentation process. As a new approach, we applied an online measurement technique (laser absorption spectroscopy) for real‐time monitoring of stable carbon isotopes of methane () in a pilot‐scale biogas digester (3500 L) regularly fed with maize silage. Generally, isotopic composition of methane gives information about specific substrate degradation, that is, methanogenic pathways that reflect the actual digester state. First results of a 2‐wk monitoring experiment show that stable carbon isotopes of methane respond promptly and highly dynamic to changes in the process state of the digester. In combination with other monitoring parameters (methane production rate, concentration of volatile fatty acids, and pH) the fluctuations in can be interpreted as a change in methanogenic pathways due to a high organic loading rate. In this context, might be used as a new parameter tool for monitoring and characterization of the process state of the digester.     

Productivity, organism size, and the trophic structure of the major terrestrial biomes

I present a hypothesis suggesting that terrestrial biomes' productivity determines the size of the major organisms and plant physiognomy in them and consequently determines their trophic structure. I suggest a more comprehensive set of patterns over current hypotheses, which consider productivity alone. Based on my hypothesis, I predict that the number of trophic levels decreases with increasing productivity from four in hot deserts to two in productive grasslands and that there are three to four trophic levels in all types of forested biomes. I also suggest that productivity is the limiting factor for the number of trophic levels at the lower end of the terrestrial productivity gradient, herbivore size is the limiting factor at intermediate productivities, and plant physiognomy is the limiting factor in the high productivity range. This contradicts existing hypotheses that predict either an increase (Am Nat 118:240–261, 1981; Am Nat 155:703–723, 2000) or no change (Am Nat 142:379–411, 1993) in the number of trophic levels with the increase in biome productivity.     

Stable oxygen and carbon isotopes of live benthic foraminifera from the Bay of Biscay: Microhabitat impact and seasonal variability

We determined the stable oxygen and carbon isotopic composition of live (Rose Bengal stained) benthic foraminifera (> 150 μm size fraction) of seven taxa sampled along a downslope transect between 140 to 2000 m water depth in the Bay of Biscay. At the five stations, Hoeglundina elegans, Cibicidoides pachydermus, Uvigerina peregrina, Uvigerina mediterranea preferentially occupy shallow infaunal niches, whereas Melonis barleeanus and Uvigerina elongatastriata occupy an intermediate infaunal microhabitat, and Globobulimina spp. live in a deep infaunal niche close to the zero oxygen boundary.When compared with δ¹⁸O values of calcite formed in equilibrium with bottom waters, U. peregrina forms its test in close equilibrium with bottom water δ¹⁸O. All other foraminiferal taxa calcify with a constant offset to calculated equilibrium calcite. There is no systematic relationship between the foraminiferal microhabitat depth and the Δδ¹⁸O between foraminiferal and equilibrium calcite. We calculated correcting factors for the various taxa, which are needed for constructing multispecies-based oxygen isotope records in paleoceanographic studies of the study area.The δ¹³C values of foraminiferal taxa investigated in this study do neither record bottom water δ¹³C_DIC in a 1 : 1 relationship nor with a constant offset, but appear to be mainly controlled by microhabitat effects. The increase of δ¹³C values of shallow infaunal taxa with increasing water depth reflects the decrease of the exported flux of organic carbon along the bathymetric transect and early diagenetic processes in the surface sediment. This is particularly the case for the shallow infaunal U. peregrina. The δ¹³C values of deep infaunal Globobulimina spp. are much less dependent on the exported organic matter flux. We suggest that the Δδ¹³C between U. peregrina and Globobulimina spp. can shed light on the various pathways of past degradation of organic detritus in the benthic environments.At a station in 550 m water depth, where periodic eutrophication of sediment surface niches was demonstrated previously, we performed a two-year seasonal survey of the isotopic composition of foraminiferal faunas. No marked seasonal changes of the stable carbon isotopic composition of shallow, intermediate and deep infaunal foraminiferal taxa were observed. Thus, the δ¹³C values of foraminiferal individuals belonging to the > 150 μm fraction may result from rather long-term calcification processes lasting for several weeks or months, which limit the impact of ephemeral ¹²C enrichment of shallow infaunal niches on the isotope chemistry of adult individuals during eutrophic periods. Only highly opportunistic taxa reproducing or calcifying during phytoplankton bloom periods and the subsequent deposits of phytoplankton remains in the benthic environment may exhibit a particularly low δ¹³C, indicative of such short productive periods.