Changes in leaf Δ13C of herbarium plant species during the last 3 centuries of CO2 increase

Δ¹³C were determined for herbarium specimens of 12 C₃ plants (trees, shrubs and herbs) collected during the last 240 years in Catalonia, an area with a Mediterranean climate. Values were 19.91 (S.E. = 0.32, n= 21) for 1750–1760, 19–86 (S.E. = 0.21, n= 49) for 1850–1890 and 19.95 (S.E. = 0.29, n= 25) for 1925–1950, and decreased significantly to 18.87 (S.E. = 0.31, n= 29) for 1982–1988. More irregular temporal changes were found in Δ¹³C of two C₄ species, but they also suggest a decrease in discrimination in recent decades. These results suggest that either carbon assimilation rates have increased or stomatal conductance has decreased, and therefore, that there has been an increase in water use efficiency over the last few decades.     

Summer monsoon intensity controls C4/C3 plant abundance during the last 35 ka in the Chinese Loess Plateau: Carbon isotope evidence from bulk organic matter and individual leaf waxes

The natural abundance of C₃ and C₄ plants is affected by multiple environmental factors including temperature, moisture balance and atmospheric pCO₂. The relative importance of these factors is a subject of considerable debate, and may vary in different natural ecosystems. Previous studies generally focus on single loess sequences in the Chinese Loess Plateau, and conflicting conclusions on C₄/C₃ have been reached when studying carbon isotope ratios in carbonate and organic matter. In this paper we report a comprehensive carbon isotopic characterization of total organic carbon (TOC) and individual higher plant leaf waxes from five loess sequences spanning the last 35 ka from the Chinese Loess Plateau (CLP). The five coring sites encompass large gradients of annual mean temperature (9.2–13.9 °C) and precipitation (402–673 mm), allowing us to assess the controlling mechanisms on C₄/C₃ plant ratios. Glacial–interglacial sequences provide carbon isotope data for comparison with other climatic and environmental proxies such as lithology and magnetic susceptibility. Our results demonstrate that increased C₄/C₃ ratios are positively correlated with higher temperature and increased summer rainfall which characterize stronger summer monsoon in all five sites. We conclude C₄ abundance increases from the last glacial to the Holocene in response to greater monsoon activity and that the C₄ expression is suppressed in the cold and drier intervals.     

Models of carbon metabolism in C3-C4 intermediate plants as applied to the evolution of C4 photosynthesis

Abstract Models developed to explain the biphasic response of CO₂ compensation concentration to O₂ concentration and the C₃-like carbon isotope discrimination in C₃-C₄ intermediate species are used to characterize quantitatively the steps necessary in the evolution of C₄ photosynthesis. The evolutionary stages are indicated by model outputs, CO₂ compensation concentration and δ₁₃C value. The transition from intermediate plants to C₄ plants requires the complete formation of C₄ cycle capacity, expressed by the models as transition from C₄ cycle limitation by phosphoenolpyruvate (PEP) regeneration rate to limitation by PEP carboxylase activity. Other steps refer to CO₂ leakage from bundle sheath cells, to further augmentations of C₄ cycle components, to the repression of ribulose-1,5-bisphos-phate carboxylase in the mesophyll cells, and to a decrease in the CO₂ affinity of the enzyme. Possibilities of extending the suggested approach to other physiological characteristics, and the adaptive significance of the steps envisaged, are discussed.     

Root responses along a subambient to elevated CO2 gradient in a C3–C4 grassland

Atmospheric CO₂ (C_a) concentration has increased significantly during the last 20 000 years, and is projected to double this century. Despite the importance of belowground processes in the global carbon cycle, community‐level and single species root responses to rising C_a are not well understood. We measured net community root biomass over 3 years using ingrowth cores in a natural C₃–C₄ grassland exposed to a gradient of C_a from preglacial to future levels (230–550 μmol mol^?1). Root windows and minirhizotron tubes were installed below naturally occurring stands of the C₄ perennial grass Bothriochloa ischaemum and its roots were measured for respiration, carbohydrate concentration, specific root length (SRL), production, and lifespan over 2 years. Community root biomass increased significantly (P<0.05) with C_a over initial conditions, with linear or curvilinear responses depending on sample date. In contrast, B. ischaemum produced significantly more roots at subambient than elevated C_a in minirhizotrons. The lifespan of roots with five or more neighboring roots in minirhizotron windows decreased significantly at high C_a, suggesting that after dense root growth depletes soil resource patches, plants with carbon surpluses readily shed these roots. Root respiration in B. ischaemum showed a curvilinear response to C_a under moist conditions in June 2000, with the lowest rates at C_a<300 μmol mol^?1 and peak activity at 450 μmol mol^?1 in a quadratic model. B. ischaemum roots at subambient C_a had higher SRLs and slightly higher carbohydrate concentrations than those at higher C_a, which may be related to drier soils at low C_a. Our data emphasize that belowground responses of plant communities to C_a can be quite different from those of the individual species, and suggest that complex interactions between and among roots and their immediate soil environment influence the responses of root physiology and lifespan to changing C_a.     

Comparative response of δ13C, δ18O and δ15N in durum wheat exposed to salinity at the vegetative and reproductive stages

This study compared the performance of the stable isotope composition of carbon (δ¹³C), oxygen (δ¹⁸O) and nitrogen (δ¹⁵N) by tracking plant response and genotypic variability of durum wheat to different salinity conditions. To that end, δ¹³C, δ¹⁸O and δ¹⁵N were analysed in dry matter (dm) and the water‐soluble fraction (wsf) of leaves from plants exposed to salinity, either soon after plant emergence or at anthesis. The δ¹³C and δ¹⁸O of the wsf recorded the recent growing conditions, including changes in evaporative conditions. Regardless of the plant part (dm or wsf), δ¹³C and δ¹⁸O increased and δ¹⁵N decreased in response to stress. When the stress conditions were established just after emergence, δ¹⁵N and δ¹³C correlated positively with genotypic differences in biomass, whereas δ¹⁸O correlated negatively in the most severe treatment. When the stress conditions were imposed at anthesis, relationships between the three isotope signatures and biomass were only significant and positive within the most severe treatments. The results show that nitrogen metabolism, together with stomatal limitation, is involved in the genotypic response to salinity, with the relative importance of each factor depending on the severity and duration of the stress as well as the phenological stage that the stress occurs.     

Role of land cover changes for atmospheric CO2 increase and climate change during the last 150 years

We assess the role of changing natural (volcanic, aerosol, insolation) and anthropogenic (CO₂ emissions, land cover) forcings on the global climate system over the last 150 years using an earth system model of intermediate complexity, CLIMBER‐2. We apply several datasets of historical land‐use reconstructions: the cropland dataset by Ramankutty & Foley (1999) (R&F), the HYDE land cover dataset of Klein Goldewijk (2001) , and the land‐use emissions data from Houghton & Hackler (2002) . Comparison between the simulated and observed temporal evolution of atmospheric CO₂ and δ¹³CO₂ are used to evaluate these datasets. To check model uncertainty, CLIMBER‐2 was coupled to the more complex Lund–Potsdam–Jena (LPJ) dynamic global vegetation model. In simulation with R&F dataset, biogeophysical mechanisms due to land cover changes tend to decrease global air temperature by 0.26°C, while biogeochemical mechanisms act to warm the climate by 0.18°C. The net effect on climate is negligible on a global scale, but pronounced over the land in the temperate and high northern latitudes where a cooling due to an increase in land surface albedo offsets the warming due to land‐use CO₂ emissions. Land cover changes led to estimated increases in atmospheric CO₂ of between 22 and 43 ppmv. Over the entire period 1800–2000, simulated δ¹³CO₂ with HYDE compares most favourably with ice core during 1850–1950 and Cape Grim data, indicating preference of earlier land clearance in HYDE over R&F. In relative terms, land cover forcing corresponds to 25–49% of the observed growth in atmospheric CO₂. This contribution declined from 36–60% during 1850–1960 to 4–35% during 1960–2000. CLIMBER‐2‐LPJ simulates the land cover contribution to atmospheric CO₂ growth to decrease from 68% during 1900–1960 to 12% in the 1980s. Overall, our simulations show a decline in the relative role of land cover changes for atmospheric CO₂ increase during the last 150 years.     

Carbon: terrestrial C4 plants

The carbon isotope composition of terrestrial C₄ plants depends on the primary carboxylation of phosphoenolpyruvate (PEP) and on the diffusion of CO₂ to the carboxylation sites, but is also influenced by the final carboxylation of ribulose-1,5-bisphosphate (RuBP). Several models have been used for reproducing this complex situation. In the present review, a particular model is applied as a means to interpret the effects of environmental and genetically determined factors on carbon isotope discrimination during C₄ photosynthesis. As a new feature, the model considers four types of limitation of the overall CO₂ assimilation rate. Both carboxylation reactions are assumed to be limited by either maximum enzyme activity or maximum substrate regeneration rate. The model is applied to experimental data on the effects of CO₂, irradiance and water stress on short-term discrimination by leaves of several C₄ species measured simultaneously with CO₂ gas exchange characteristics. In particular, different patterns of the influence of low irradiances on carbon isotope discrimination are interpreted as due to variations in that irradiance at which a transition from limitation by PEP regeneration rate and RuBP carboxylase activity to limitation by the regeneration rates of both substrates occurs. After discussing literature data on the effects of environmental conditions on carbon isotope discrimination by C₄ plants seasonal and developmental changes in carbon isotope composition, studies on the systematic and geographic distribution of C₄ plants, evolutionary and genetical aspects, and some ecological implications are reviewed.     

基于稳定碳同位素技术研究大气CO2浓度升高对植物-土壤系统碳循环的影响

大气CO₂浓度升高影响植物光合作用过程和生物量积累,改变植物地上和地下生物量的动态分配.土壤有机质的形成和周转依赖于植物组分的输入,因此,CO₂浓度升高所造成的植物生理和代谢的变化对土壤碳库收支平衡具有重要影响.采用稳定碳同位素(¹³C)技术研究土壤-植物系统的碳循环可阐明大气CO₂浓度升高条件下光合碳在植物各器官的分配特征和时间动态,明确光合碳在土壤中的积累、分解与迁移转化过程以及对土壤有机碳库周转的影响.本文综述了基于¹³C自然丰度法或¹³C示踪技术研究大气CO₂浓度升高对土壤-植物系统碳循环的影响,主要包括:1)对植物光合作用的同位素分馏的影响;2)对植物光合碳(新碳)分配动态的影响;3)对土壤有机碳新老碳库动态以及微生物转化过程的影响.明确上述过程及其调控机制可为预测CO₂浓度升高对陆地生态系统碳循环及源汇效应的长期影响奠定基础.     

Changes in the abundance of C3/C4 species of Inner Mongolia grassland: evidence from isotopic composition of soil and vegetation

Global warming, increasing CO₂ concentration, and environmental disturbances affect grassland communities throughout the world. Here, we report on variations in the C3/C4 pattern of Inner Mongolian grassland derived from soil and vegetation. Soil samples from 149 sites covering an area of approximately 250 000 km² within Inner Mongolia, People's Republic of China were analyzed for the isotopic composition (δ¹³C) of soil organic carbon (SOC). The contrast in δ¹³C between C3 and C4 plants allowed for calculation of the C3/C4 ratio from δ¹³C of SOC with a two‐member mixing model, which accounted for influences of aridity and altitude on δ¹³C of the C3 end‐member and for changes in δ¹³C of atmospheric CO₂. Maps were created geostatistically, and showed a substantially lower C4 abundance in soil than in recent vegetation (?10%). The difference between soil and vegetation varied regionally and was most pronounced within an E–W belt along 44°N and in a mountainous area, suggesting a spread of C4 plants toward northern latitudes (about 1°) and higher altitudes. The areas of high C4 abundance for present vegetation and SOC were well delineated by the isotherms of crossover temperature based on the climatic conditions of the respective time periods. Our study indicates that change in the patterns of C3/C4 composition in the Inner Mongolia grassland was mainly triggered by increasing temperature, which overrode the antagonistic effect of rising CO₂ concentrations.     

Mapping of the α4 subunit gene (GABRA4) to human chromosome 4 defines an α2—α4—β1—γ1 gene cluster: further evidence that modern GABAA receptor gene clusters are derived from an ancestral cluster

We demonstrated previously that an α₁—β₂—γ₂ gene cluster of the γ-aminobutyric acid (GABA_A) receptor is located on human chromosome 5q34–q35 and that an ancestral α—β—γ gene cluster probably spawned clusters on chromosomes 4, 5, and 15. Here, we report that the α₄ gene (GABRA4) maps to human chromosome 4p14–q12, defining a cluster comprising the α₂, α₄, β₁, and γ₁ genes. The existence of an α₂—α₄—β₁—γ₁ cluster on chromosome 4 and an α₁—α₆—β₂—γ₂ cluster on chromosome 5 provides further evidence that the number of ancestral GABA_A receptor subunit genes has been expanded by duplication within an ancestral gene cluster. Moreover, if duplication of the α gene occurred before duplication of the ancestral gene cluster, then a heretofore undiscovered subtype of α subunit should be located on human chromosome 15q11–q13 within an α₅—α_x—β₃—γ₃ gene cluster at the locus for Angelman and Prader—Willi syndromes.     

Dietary variation within and between populations of northeast Atlantic killer whales,Orcinus orca,inferred from δ13C and δ15N analyses

Epidermal skin samples from eastern North Atlantic killer whales, Orcinus orca, were analyzed for carbon and nitrogen stable isotope ratios. From those, comparisons within a data set of 17 samples collected from Tysfjord, Norway, in November suggested that diet is relatively specialized during this time period at this location. There were significant differences between a small set of samples from Iceland and those collected from Norway, which had all been assigned to the same population by a previous population genetics study. The results would be consistent with matrilines feeding on either the Norwegian or Icelandic stocks of Atlantic herring (Clupea harengus). There was no significant difference within Icelandic samples between those assigned to the population known to feed upon herring and those assigned to a population hypothesized to follow Atlantic mackerel (Scomber scombrus). The greatest differences were between the epidermal samples analyzed in this study and tooth and bone collagen samples from the North Sea that were analyzed previously, which also showed significantly more variation in isotopic ratios than found for skin samples. These differences could reflect differences in turnover rate, differences in diet‐tissue fractionation and discrimination due to the amino acid composition of the different tissues, and/or greater competition promoting dietary variation between groups in the North Sea.     

Glacial–interglacial deep-water changes in the NW Pacific inferred from single foraminiferal δO and δC

Oxygen and carbon isotope values of single benthic foraminiferal tests in a core from the Shatsky Rise, NW Pacific Ocean, show greater intra-horizon variance during the Holocene than during the Last Glacial Maximum (LGM). This greater variance is caused by the introduction of glacial specimens some 20 cm upward from their original deposition layer due to bioturbation. In contrast, foraminiferal populations belonging to glacial layers do not include Holocene specimens. The difference in direction of bioturbation greatly modifies climate information in horizons formed during and after deglacial events. After omitting glacial specimens from Holocene sediments, the glacial–interglacial difference in δ¹⁸O suggests that Pacific deep-water temperature changed by 2.4–3.8°C at the most. The δ¹³C values suggest that nutrient concentration was higher during the LGM than the Holocene. The glacial deep North Pacific Ocean apparently was influenced by cold deep waters of southern origin.     

Stomatal responses of C3, C3-C4 and C4Flaveria species to light and intercellular CO2 concentration: implications for the evolution of stomatal behaviour

Stomatal function mediates physiological trade‐offs associated with maintaining a favourable H₂O balance in leaf tissues while acquiring CO₂ as a photosynthetic substrate. The C₃ and C₄ species appear to have different patterns of stomatal response to changing light conditions, and variation in this behaviour may have played a role in the functional diversification of the different photosynthetic pathways. In the current study, we used gain analysis theory to characterize the stomatal conductance response to light intensity in nine different C₃, C₄ and C₃‐C₄ intermediate species Flaveria species. The response of stomatal conductance (g_s) to a change in light intensity represents both a direct (related to a change in incident light intensity, I) and indirect (related to a change in intercellular CO₂ concentration, C_i) response. The slope of the line relating the change in g_s to C_i was steeper in C₄ species, compared with C₃ species, with C₃‐C₄ species having an intermediate response. This response reflects the greater relative contribution of the indirect versus direct component of the g_s versus I response in the C₄ species. The C₃‐C₄ species, Flaveria floridana, exhibited a C₄‐like response whereas the C₃‐C₄ species, Flaveria sonorensis and Flaveria chloraefolia, exhibited C₃‐like responses, similar to their hypothesized position along the evolutionary trajectory of the development of C₄ photosynthesis. There was a positive correlation between the relative contribution of the indirect component of the g_s versus I response and water use efficiency when evaluated across all species. Assuming that the C₃‐C₄ intermediate species reflect an evolutionary progression from fully expressed C₃ ancestors, the results of the current study demonstrate an increase in the contribution of the indirect component of the g_s versus I response as taxa evolve toward the C₄ extreme. The greater relative contribution of the indirect component of the stomatal response occurs through both increases in the indirect stomatal components and through decreases in the direct. Increases in the magnitude of the indirect component may be related to the maintenance of higher water use efficiencies in the intermediate evolutionary stages, before the appearance of fully integrated C₄ photosynthesis.     

Climate and CO2 controls on global vegetation distribution at the last glacial maximum: analysis based on palaeovegetation data, biome modelling and palaeoclimate simulations

The global vegetation response to climate and atmospheric CO₂ changes between the last glacial maximum and recent times is examined using an equilibrium vegetation model (BIOME4), driven by output from 17 climate simulations from the Palaeoclimate Modelling Intercomparison Project. Features common to all of the simulations include expansion of treeless vegetation in high northern latitudes; southward displacement and fragmentation of boreal and temperate forests; and expansion of drought‐tolerant biomes in the tropics. These features are broadly consistent with pollen‐based reconstructions of vegetation distribution at the last glacial maximum. Glacial vegetation in high latitudes reflects cold and dry conditions due to the low CO₂ concentration and the presence of large continental ice sheets. The extent of drought‐tolerant vegetation in tropical and subtropical latitudes reflects a generally drier low‐latitude climate. Comparisons of the observations with BIOME4 simulations, with and without consideration of the direct physiological effect of CO₂ concentration on C₃ photosynthesis, suggest an important additional role of low CO₂ concentration in restricting the extent of forests, especially in the tropics. Global forest cover was overestimated by all models when climate change alone was used to drive BIOME4, and estimated more accurately when physiological effects of CO₂ concentration were included. This result suggests that both CO₂ effects and climate effects were important in determining glacial‐interglacial changes in vegetation. More realistic simulations of glacial vegetation and climate will need to take into account the feedback effects of these structural and physiological changes on the climate.     

Above- and below-ground responses of C3–C4 species mixtures to elevated CO2 and soil water availability

We evaluated the influences of CO₂[Control, ～ 370 µ mol mol ^{? 1}; 200 µ mol mol ^{? 1} above ambient applied by free‐air CO₂ enrichment (FACE)] and soil water (Wet, Dry) on above‐ and below‐ground responses of C₃ (cotton, Gossypium hirsutum) and C₄ (sorghum, Sorghum bicolor) plants in monocultures and two density mixtures. In monocultures, CO₂ enrichment increased height, leaf area, above‐ground biomass and reproductive output of cotton, but not sorghum, and was independent of soil water treatment. In mixtures, cotton, but not sorghum, above‐ground biomass and height were generally reduced compared to monocultures, across both CO₂ and soil water treatments. Density did not affect individual plant responses of either cotton or sorghum across the other treatments. Total (cotton + sorghum) leaf area and above‐ground biomass in low‐density mixtures were similar between CO₂ treatments, but increased by 17–21% with FACE in high‐density mixtures, due to a 121% enhancement of cotton leaf area and a 276% increase in biomass under the FACE treatment. Total root biomass in the upper 1.2 m of the soil was not influenced by CO₂ or by soil water in monoculture or mixtures; however, under dry conditions we observed significantly more roots at lower soil depths ( > 45 cm). Sorghum roots comprised 81–85% of the total roots in the low‐density mixture and 58–73% in the high‐density mixture. CO₂‐enrichment partly offset negative effects of interspecific competition on cotton in both low‐ and high‐density mixtures by increasing above‐ground biomass, with a greater relative increase in the high‐density mixture. As a consequence, CO₂‐enrichment increased total above‐ground yield of the mixture at high density. Individual plant responses to CO₂ enrichment in global change models that evaluate mixed plant communities should be adjusted to incorporate feedbacks for interspecific competition. Future field studies in natural ecosystems should address the role that a CO₂‐mediated increase in C₃ growth may have on subsequent vegetation change.