Oxygen Isotopic Composition Of Cellulose Research Articles

Effects of increasing atmospheric CO2 on leaf water δ18O values are small and are attenuated in grasses and amplified in dicotyledonous herbs and legumes when transferred to cellulose δ18O values.

The oxygen isotope composition of cellulose (δ18O values) has been suggested to contain information on stomatal conductance (gs) responses to rising pCO2. The extent by which pCO2 affects leaf water and cellulose δ18O values (δ18OLW and δ18OC) and the isotope processes that determine pCO2 effects on δ18OLW and δ18OC are, however, unknown. We tested the effects of pCO2 on gs, δ18OLW and δ18OC in a glasshouse experiment, where six plant species were grown under pCO2 ranging from 200 to 500 ppm. Increasing pCO2 caused a decline in gs and an increase in δ18OLW, as expected. Importantly, the effects of pCO2 on gs and δ18OLW were small and pCO2 effects on δ18OLW were not directly transferred to δ18OC but were attenuated in grasses and amplified in dicotyledonous herbs and legumes. This is likely because of functional group-specific pCO2 effects on the model parameter pxpex. Our study highlights important uncertainties when using δ18OC as a proxy for gs. Specifically, pCO2-triggered gs effects on δ18OLW and δ18OC are possibly too small to be detected in natural settings and a pCO2 effect on pxpex may render the commonly assumed negative linkage between δ18OC and gs to be incorrect, potentially confounding δ18OC based gs reconstructions.

Cellulose δ18O as a hydrological proxy: A case study for willows growing in Indigirka river lowland, Northeastern Siberia

During the last few decades, stable oxygen isotopic composition of cellulose (δ18Ocell) has been employed as a potential tool for understanding hydrological cycles on modern and past environments. In the present study, we measured the δ18O values of stem water (δ18Osw) and the δ18Ocell values of willows, which are growing on transects along the river with multiple water sources and different levels of humidity in Northeastern Siberia, further to evaluate the contribution of (1) the δ18Osw values, (2) the ratio of ambient/intercellular water (i.e., ea/ei), and (3) the δ18O values of water vapor (δ18Ov), to the δ18Ocell values. The results well demonstrate that the cross-plot of δ18Ocell/δ18Osw can be used to evaluate the contributions of δ18Osw values and ea/ei to the variation of δ18Ocell values. Also, the contribution of the δ18Ov values on the δ18Ocell values are significantly increased, where the water vapor near the plants is isotopically equilibrated to large water pools (e.g., rivers, lakes, ponds, etc.) rather than the source water (e.g., precipitation) of plants. Based on these findings, we predict that full consideration of the above (1)–(3) permits us to enhance accuracy in our understanding of plant δ18Ocell values with respect to hydrological cycles.

An oxygen isotope chronometer for cellulose deposition: the successive leaves formed by tillers of a C4 perennial grass.

Multiannual time series of (palaeo)hydrological information can be reconstructed from the oxygen isotope composition of cellulose (δ18 OCel ) in biological archives, for example, tree rings, but our ability to temporally resolve information at subannual scale is limited. We capitalized on the short and predictable leaf appearance interval (2.4d) of a perennial C4 grass (Cleistogenes squarrosa), to assess its potential for providing highly time-resolved δ18 OCel records of vapour pressure deficit (VPD). Plants grown at low (0.63kPa) or high (1.58kPa) VPD were swapped between VPD environments and exposed to the new environment for 7d with simultaneous 13 CO2 labelling. Then, leaves were sampled by age/position along individual tillers. Five leaves at different developmental stages were growing simultaneously. The period of most-active leaf elongation, from 10 to 90% of final length, lasted 6.6d, and ~80% of all carbon and oxygen incorporation in whole-leaf cellulose occurred within 7d. Cellulose deposition stopped at (or shortly after) full leaf expansion. The direction of change, low-to-high or high-to-low VPD, had no differential effect on new oxygen and carbon incorporation in cellulose. Successive leaves produced by tillers of C.squarrosa provide a δ18 OCel record useful for reconstructions of short-term hydrological dynamics.

Nitrogen fertilization and δ18 O of CO2 have no effect on 18 O-enrichment of leaf water and cellulose in Cleistogenes squarrosa (C4 ) - is VPD the sole control?

The oxygen isotope composition of cellulose (δ18 OCel ) archives hydrological and physiological information. Here, we assess previously unexplored direct and interactive effects of the δ18 O of CO2 (δ18 OCO2 ), nitrogen (N) fertilizer supply and vapour pressure deficit (VPD) on δ18 OCel , 18 O-enrichment of leaf water (Δ18 OLW ) and cellulose (Δ18 OCel ) relative to source water, and pex px , the proportion of oxygen in cellulose that exchanged with unenriched water at the site of cellulose synthesis, in a C4 grass (Cleistogenes squarrosa). δ18 OCO2 and N supply, and their interactions with VPD, had no effect on δ18 OCel , Δ18 OLW , Δ18 OCel and pex px . Δ18 OCel and Δ18 OLW increased with VPD, while pex px decreased. That VPD-effect on pex px was supported by sensitivity tests to variation of Δ18 OLW and the equilibrium fractionation factor between carbonyl oxygen and water. N supply altered growth and morphological features, but not 18 O relations; conversely, VPD had no effect on growth or morphology, but controlled 18 O relations. The work implies that reconstructions of VPD from Δ18 OCel would overestimate amplitudes of VPD variation, at least in this species, if the VPD-effect on pex px is ignored. Progress in understanding the relationship between Δ18 OLW and Δ18 OCel will require separate investigations of pex and px and of their responses to environmental conditions.

Oxygen isotope ratios (18O/16O) of hemicellulose-derived sugar biomarkers in plants, soils and sediments as paleoclimate proxy I: Insight from a climate chamber experiment

The oxygen isotopic composition of cellulose is a valuable proxy in paleoclimate research. However, its application to sedimentary archives is challenging due to extraction and purification of cellulose. Here we present compound-specific δ18O results of hemicellulose-derived sugar biomarkers determined using gas chromatography–pyrolysis–isotope ratio mass spectrometry, which is a method that overcomes the above-mentioned analytical challenges. The biomarkers were extracted from stem material of different plants (Eucalyptus globulus, Vicia faba and Brassica oleracea) grown in climate chamber experiments under different climatic conditions.The δ18O values of arabinose and xylose range from 31.4‰ to 45.9‰ and from 28.7‰ to 40.8‰, respectively, and correlate highly significantly with each other (R=0.91, p<0.001). Furthermore, δ18Ohemicellulose (mean of arabinose and xylose) correlate highly significantly with δ18Oleaf water (R=0.66, p<0.001) and significantly with modeled δ18Ocellulose (R=0.42, p<0.038), as well as with relative air humidity (R=−0.79, p<0.001) and temperature (R=−0.66, p<0.001). These findings confirm that the hemicellulose-derived sugar biomarkers, like cellulose, reflect the oxygen isotopic composition of plant source water altered by climatically controlled evapotranspirative 18O enrichment of leaf water. While relative air humidity controls most rigorously the evapotranspirative 18O enrichment, the direct temperature effect is less important. However, temperature can indirectly exert influence via plant physiological reactions, namely by influencing the transpiration rate which affects δ18Oleaf water due to the Péclet effect. In a companion paper (Tuthorn et al., this issue) we demonstrate the applicability of the hemicellulose-derived sugar biomarker δ18O method to soils and provide evidence from a climate transect study confirming that relative air humidity exerts the dominant control on evapotranspirative 18O enrichment of leaf water.Finally, we present a conceptual model for the interpretation of δ18Ohemicellulose records and propose that a combined δ18Ohemicellulose and δ2Hn-alkane biomarker approach is promising for disentangling δ18Oprecipitation variability from evapotranspirative 18O enrichment variability in future paleoclimate studies.

Differing source water inputs, moderated by evaporative enrichment, determine the contrasting δ18OCELLULOSE signals in maritime Antarctic moss peat banks

AbstractOxygen isotope palaeoclimate records, preserved in moss tissue cellulose, are complicated by environmental influences on the relationships between source water inputs and evaporative conditions. We carried out stable isotope analyses of precipitation collected from the maritime Antarctic and cellulose extracted from co‐located Chorisodontium aciphyllum dominated moss peat bank deposits accumulated since 1870 A.D. Analyses of stable oxygen and hydrogen isotope composition of summer precipitation on Signy Island (60.7°S, 45.6°W) established a local meteoric water line (LMWL) similar to both the global MWL and other LMWLs, and almost identical to the HadAM3 isotope‐enabled global circulation model output. The oxygen isotopic composition of cellulose (δ18OC) revealed little temporal variation between four moss peat banks on Signy Island since 1870. However, δ18OC followed two patterns with Sites A and D consistently 3‰ enriched relative to δ18OC values from Sites B and C. The growing moss surfaces at Sites A and D are likely to have been hydrated by isotopically heavier summer precipitation, whilst at Sites B and C, the moss banks are regularly saturated by the isotopically depleted snow melt streams. Laboratory experiments revealed that evaporative enrichment of C. aciphyllum moss leaf water by 5‰ occurred rapidly following saturation (ecologically equivalent to post‐rainfall or snow melt periods). In addition to the recognized source water‐cellulose fractionation extent of 27 ± 3‰, such a shift would account for the 32‰ difference measured between δ18O of Signy Island precipitation and cellulose.

Secular temperature trends for the southern Rocky Mountains over the last five centuries

Pre‐instrumental surface temperature variability in the Southwestern United States has traditionally been reconstructed using variations in the annual ring widths of high altitude trees that live near a growth‐limiting isotherm. A number of studies have suggested that the response of some trees to temperature variations is non‐stationary, warranting the development of alternative approaches towards reconstructing past regional temperature variability. Here we present a five‐century temperature reconstruction for a high‐altitude site in the Rocky Mountains derived from the oxygen isotopic composition of cellulose (δ18Oc) from Bristlecone Pine trees. The record is independent of the co‐located growth‐based reconstruction while providing the same temporal resolution and absolute age constraints. The empirical correlation betweenδ18Ocand instrumental temperatures is used to produce a temperature transfer function. A forward‐model for cellulose isotope variations, driven by meteorological data and output from an isotope‐enabled General Circulation Model, is used to evaluate the processes that propagate the temperature signal to the proxy. The cellulose record documents persistent multidecadal variations inδ18Octhat are attributable to temperature shifts on the order of 1°C but no sustained monotonic rise in temperature or a step‐like increase since the late 19th century. The isotope‐based temperature history is consistent with both regional wood density‐based temperature estimates and some sparse early instrumental records.

Improved Method for the Determination of the Oxygen Isotopic Composition of Cellulose

A modification of a chemical preparation procedure for oxygen isotope analysis of cellulose has been developed which yields greater precision and requires less time for sample preparation. In the place of isoquinoline, zinc is used to remove HCl from the reaction products of the sample and HgCl 2 . The two methods give systematically different results because isoquinoline biases measurements by contaminating the sample with additional CO 2 . For the zinc method, precision is ±0.2% (1σ standard deviation) compared with ±0.9% for the isoquinoline method

Oxygen and Hydrogen Isotope Fractionation during Cellulose Metabolism in Lemna gibba L.

Lemna gibba L. B3 was grown under heterotrophic, photoheterotrophic, and autotrophic conditions in water having a variety of hydrogen and oxygen isotopic compositions. The slopes of the linear regression lines between the isotopic composition of water and leaf cellulose indicated that under the three growth conditions about 40, 70, and 100% of oxygens and carbon-bound hydrogens of cellulose exchanged with those of water prior to cellulose formation. Using the equations of the linear relationships, we estimated the overall fractionation factors between water and the exchanged oxygen and carbon bound-hydrogen of cellulose. At least two very different isotope effects must determine the hydrogen isotopic composition of Lemna cellulose. One reflects the photosynthetic reduction of NADP, while the second reflects exchange reactions that occur subsequent to NADP reduction. Oxygen isotopic composition of cellulose apparently is determined by a single type of exchange reaction with water. Under different growth conditions, variations in metabolic fluxes affect the hydrogen isotopic composition of cellulose by influencing the extent to which the two isotope effects mentioned above are recorded. The oxygen isotopic composition of cellulose is not affected by such changes in growth conditions.

Oxygen-18 Content of Atmospheric Oxygen Does Not Affect the Oxygen Isotope Relationship between Environmental Water and Cellulose in a Submerged Aquatic Plant, Egeria densa Planch

We determined that the oxygen isotopic composition of cellulose synthesized by a submerged plant, Egeria densa Planch., is related to the isotopic composition of environmental water by a linear function, delta(18)O cellulose = 0.48 delta(18)O water + 24.1% per thousand. The observation of a slope of less than 1 indicates that a portion of cellulose oxygen is derived from an isotopically constant source other than water. We tested whether this source might be molecular oxygen by growing plants in the presence of high concentrations of (18)O in the form of O(2) bubbled into the bottom of an aquarium. Cellulose synthesized during this experiment did not have significantly different oxygen isotope ratios than that synthesized by control plants exposed to O(2) of normal (18)O abundance. We propose that oxygen in organic matter recycled from senescent portions of the plant is incorporated into cellulose. Our findings indicate that paleoclimatic models linking the oxygen isotope composition of environmental water to cellulose from fossil plants will have to be modified to account for contributions of oxygen from this or other sources besides water.

Climatic information from 18O/16O analysis of cellulose, lignin and whole wood from tree rings

MEASUREMENTS of stable isotope ratios on tree rings have proved a promising way of determinating past climate1–5. Recent discussions have centred on which component of wood provides the most reliable record6,7. We have measured the oxygen isotopic composition of cellulose, whole wood and lignin from tree rings in a white spruce (Picea glauca) which grew in the Edmonton area from 1882 to 1969. Using meteorological records we have evaluated the responses of the 18O/16O ratios of the three components of the tree rings to seasonal temperatures. Results show a high correlation between cellulose 18O/16O ratios and mean annual temperature, a poorer, though still significant correlation between whole wood 18O/16O ratios and mean annual temperature and no significant correlation between lignin 18O/16O ratios and mean annual temperature.

Climatic information from 18O/16O ratios of cellulose in tree rings

THE high precision of dendrochronology makes possible the construction of short (∼ 9,000 yr), detailed records of past climates. This paper explores the possibility of using the oxygen isotopic composition of cellulose from tree rings as a ‘thermometer’ to measure past temperatures. Using meteorological data from Edmonton we have shown that temperatures can be measured with a precision of ∼ ±0.15°C when averaged over a 5-yr period.