Respiration in a future, higher-CO2 world

Abstract. Apart from its impact on global warming, the annually increasing atmospheric [CO₂] is of interest to plant scientists primarily because of its direct influence on photosynthesis and photorespiration in C₃ species. But in addition, 'dark' respiration, another major component of the carbon budget of higher plants, may be affected by a change in [CO₂] independent of an increase in temperature. Literature pertaining to an impact of [CO₂] on respiration rate is reviewed. With an increase in [CO₂], respiration rate is increased in some cases, but decreased in others. The effects of [CO₂] on respiration rate may be direct or indirect. Mechanisms responsible for various observations are proposed. These proposed mechanisms relate to changes in: (1) levels of nonstructural carbohydrates, (2) growth rate and structural phytomass accumulation, (3) composition of phytomass, (4) direct chemical interactions between CO₂ and respiratory enzymes, (5) direct chemical interactions between CO₂ and other cellular components, (6) dark CO₂ fixation rate, and (7) ethylene biosynthesis rate. Because a range-of (possibly interactive) effects exist, and present knowledge is limited, the impact of future [CO₂] on respiration rate cannot be predicted. Theoretical considerations and types of experiments that can lead to an increase in the understanding of this issue are outlined.     

Effects of climate change on labile and structural carbon in Douglas-fir needles as estimated by δ13C and Carea measurements

Models of photosynthesis, respiration, and export predict that foliar labile carbon (C) should increase with elevated CO₂ but decrease with elevated temperature. Sugars, starch, and protein can be compared between treatments, but these compounds make up only a fraction of the total labile pool. Moreover, it is difficult to assess the turnover of labile carbon between years for evergreen foliage. Here, we combined changes in foliar C_area (C concentration on an areal basis) as needles aged with changes in foliar isotopic composition (δ¹³C) caused by inputs of ¹³C‐depleted CO₂ to estimate labile and structural C in needles of different ages in a four‐year, closed‐chamber mesocosm experiment in which Douglas‐fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings were exposed to elevated temperature (ambient + 3.5 °C) and CO₂ (ambient + 179 ppm). Declines in δ ¹³C of needle cohorts as they aged indicated incorporation of newly fixed labile or structural carbon. The δ ¹³C calculations showed that new C was 41 ± 2% and 28 ± 3% of total needle carbon in second‐ and third‐year needles, respectively, with higher proportions of new C in elevated than ambient CO₂ chambers (e.g. 42 ± 2% vs. 37 ± 6%, respectively, for second‐year needles). Relative to ambient CO₂, elevated CO₂ increased labile C in both first‐ and second‐year needles. Relative to ambient temperature, elevated temperature diminished labile C in second‐year needles but not in first‐year needles, perhaps because of differences in sink strength between the two needle age classes. We hypothesize that plant‐soil feedbacks on nitrogen supply contributed to higher photosynthetic rates under elevated temperatures that partly compensated for higher turnover rates of labile C. Strong positive correlations between labile C and sugar concentrations suggested that labile C was primarily determined by carbohydrates. Labile C was negatively correlated with concentrations of cellulose and protein. Elevated temperature increased foliar %C, possibly due to a shift of labile constituents from low %C carbohydrates to relatively high %C protein. Decreased sugar concentrations and increased nitrogen concentrations with elevated temperature were consistent with this explanation. Because foliar constituents that vary in isotopic signature also vary in concentrations with leaf age or environmental conditions, inferences of c_i/c_a values from δ ¹³C of bulk leaf tissue should be done cautiously. Tracing of ¹³C through foliar carbon pools may provide new insight into foliar C constituents and turnover.     

Maximum impacts of future reforestation or deforestation on atmospheric CO2

There is scope for land‐use changes to increase or decrease CO₂ concentrations in the atmosphere over the next century. Here we make simple but robust calculations of the maximum impact of such changes. Historical land‐use changes (mostly deforestation) and fossil fuel emissions have caused an increase in atmospheric concentration of CO₂ of 90 ppm between the pre‐industrial era and year 2000. The projected range of CO₂ concentrations in 2100, under a range of emissions scenarios developed for the IPCC, is 170–600 ppm above 2000 levels. This range is mostly due to different assumptions regarding fossil fuel emissions. If all of the carbon so far released by land‐use changes could be restored to the terrestrial biosphere, atmospheric CO₂ concentration at the end of the century would be about 40–70 ppm less than it would be if no such intervention had occurred. Conversely, complete global deforestation over the same time frame would increase atmospheric concentrations by about 130–290 ppm. These are extreme assumptions; the maximum feasible reforestation and afforestation activities over the next 50 years would result in a reduction in CO₂ concentration of about 15–30 ppm by the end of the century. Thus the time course of fossil fuel emissions will be the major factor in determining atmospheric CO₂ concentrations for the foreseeable future.     

The input and fate of new C in two forest soils under elevated CO2

The aim of this study was to estimate (i) the influence of different soil types on the net input of new C into soils under CO₂ enrichment and (ii) the stability and fate of these new C inputs in soils. We exposed young beech–spruce model ecosystems on an acidic loam and calcareous sand for 4 years to elevated CO₂. The added CO₂ was depleted in ¹³C, allowing to trace new C inputs in the plant–soil system. We measured CO₂‐derived new C in soil C pools fractionated into particle sizes and monitored respiration as well as leaching of this new C during incubation for 1 year. Soil type played a crucial role in the partitioning of C. The net input of new C into soils under elevated CO₂ was about 75% greater in the acidic loam than in the calcareous sand, despite a 100% and a 45% greater above‐ and below‐ground biomass on the calcareous sand. This was most likely caused by a higher turnover of C in the calcareous sand as indicated by 30% higher losses of new C from the calcareous sand than from the acidic loam during incubation. Therefore, soil properties determining stabilization of soil C were apparently more important for the accumulation of C in soils than tree productivity. Soil fractionation revealed that about 60% of the CO₂‐derived new soil C was incorporated into sand fractions. Low natural ¹³C abundance and wide C/N ratios show that sand fractions comprise little decomposed organic matter. Consistently, incubation indicated that new soil C was preferentially respired as CO₂. During the first month, evolved CO₂ consisted to 40–55% of new C, whereas the fraction of new C in bulk soil C was 15–23% only. Leaching of DOC accounted for 8–23% of the total losses of new soil C. The overall effects of CO₂ enrichment on soil C were small in both soils, although tree growth increased significantly on the calcareous sand. Our results suggest that the potential of soils for C sequestration is limited, because only a small fraction of new C inputs into soils will become long‐term soil C.     

Soil carbon inventories and δ13C along a moisture gradient in Botswana

We present a study of soil organic carbon (SOC) inventories and δ¹³C values for 625 soil cores collected from well‐drained, coarse‐textured soils in eight areas along a 1000 km moisture gradient from Southern Botswana, north into southern Zambia. The spatial distribution of trees and grass in the desert, savannah and woodland ecosystems along the transect control large systematic local variations in both SOC inventories and δ¹³C values. A stratified sampling approach was used to smooth this variability and obtain robust weighted‐mean estimates for both parameters. Weighted SOC inventories in the 0–5 cm interval of the soils range from 7 mg cm^?2 in the driest area (mean annual precipitation, MAP=225 mm) to 41±12 mg cm^?2 in the wettest area (MAP=910 mm). For the 0–30 cm interval, the inventories are 37.8 mg cm^?2 for the driest region and 157±33 mg cm^?2 for the wettest region. SOC inventories at intermediate sites increase as MAP increases to approximately 400–500 mm, but remain approximately constant thereafter. This plateau may be the result of feedbacks between MAP, fuel load and fire frequency. Weighted δ ¹³C values decrease linearly in both the 0–5 and 0–30 cm depth intervals as MAP increases. A value of –17.5±1.0‰ characterizes the driest areas, while a value of ?25±0.7‰ characterizes the wettest area. The decrease in δ ¹³C value with increasing MAP reflects an increasing dominance of C₃ vegetation as MAP increases. SOC in the deeper soil (5–30 cm depth) is, on average, 0.4±0.3‰ enriched in ¹³C relative to SOC in the 0–5 cm interval.     

The effects of reduced and elevated CO2 and O2 on the seaweed Lomentaria articulata

We grew a non-bicarbonate using red seaweed, Lomentaria articulata (Huds.) Lyngb., in media aerated with four O₂ concentrations between 10 and 200% of current ambient [O₂] and four CO₂ concentrations between 67 and 500% of current ambient [CO₂], in a factorial design, to determine the effects of gas composition on growth and physiology. The relative growth rate of L. articulata increased with increasing [CO₂] up to 200% of current ambient [CO₂] but was unaffected by [O₂]. The relative growth enhancement, on a carbon basis, was 52% with a doubling of [CO₂] but fell to 23% under 5× ambient [CO₂]. Plants collected in winter responded more extremely to [CO₂] than did plants collected in the summer, although the overall pattern was the same. Discrimination between stable carbon isotopes (Δ¹³C) increased with increasing [CO₂] as would be expected for diffusive CO₂ acquisition. Tissue C and N were inversely related to [CO₂]. Growth in terms of biomass appeared to be limited by conversion of photosynthate to new biomass rather than simply by diffusion of CO₂, suggesting that non-bicarbonate-using macroalgae, such as L. articulata, may not be directly analogous to C3 higher plants in terms of their responses to changing gas composition.     

葡萄园生态系统碳源/汇及碳减排策略研究进展

葡萄园生态系统是农业生态系统的重要组成部分, 集中连片栽培的葡萄园具有重要的生态价值。开展葡萄园生态系统碳源/汇的研究, 是完整探讨葡萄园生态系统碳循环必不可少的内容。随着葡萄生态学研究的进一步深入, 如何直观地揭示葡萄园生态系统碳循环规律和碳汇功能已经成为葡萄生态学领域关注的热点问题。研究发现, 葡萄园生态系统固定大量碳, 将碳封存在葡萄果实等一年生器官、主干等多年生器官以及土壤碳库中。葡萄园生态系统碳输入量大于碳输出量, 是碳汇; 土壤是葡萄园生态系统最大的碳库, 占总碳储量的70%, 尤其是土藤界面; 覆盖和免耕作为葡萄园的碳减排策略, 可以减少碳排放, 提高葡萄园土壤肥力。基于此, 为了阐明葡萄园生态系统的碳汇价值, 该文围绕葡萄生态学最新研究进展, 系统回顾了葡萄园生态系统中碳循环规律、碳汇研究进展及碳减排策略, 为葡萄生态学的研究提供理论基础, 并对本领域未来的研究方向和应用前景进行展望。     

Interannual variability in net CO2 exchange of a native tallgrass prairie

Year‐round eddy covariance flux measurements were made in a native tallgrass prairie in north‐central Oklahoma, USA during 1997–2000 to quantify carbon exchange and its interannual variability. This prairie is dominated by warm season C₄ grasses. The soil is a relatively shallow silty clay loam underlined with a heavy clay layer and a limestone bedrock. During the study period, the prairie was burned in the spring of each year, and was not grazed. In 1997 there was adequate soil moisture through the growing season, but 1998 had two extended periods of substantially low soil moisture (with concurrent high air temperatures and vapor pressure deficits), one early and one later in the growing season. There was also moisture stress in 1999, but it was less severe and occurred later in the season. The annual net ecosystem CO₂ exchange, NEE (before including carbon loss during the burn) was 274, 46 and 124 g C m ^{? 2} yr ^{? 1} in 1997, 1998, and 1999, respectively (flux toward the surface is positive), and the associated variation seemed to mirror the severity of moisture stress. We also examined integrated values of NEE during different periods (e.g. day/night; growing season/senescence). Annually integrated carbon dioxide uptake during the daytime showed the greatest variability from year to year, and was primarily linked to the severity of moisture stress. Carbon loss during nighttime was a significant part of the annual daytime NEE, and was fairly stable from year to year. When carbon loss during the burn (estimated from pre‐ and post‐burn biomass samples) was incorporated in the annual NEE, the prairie was found to be approximately carbon neutral (i.e. net carbon uptake/release was near zero) in years with no moisture stress (1997) or with some stress late in the season (1999). During a year with severe moisture stress early in the season (1998), the prairie was a net source of carbon. It appears that moisture stress (severity as well as timing of occurrence) was a dominating factor regulating the annual carbon exchange of the prairie.     

Vertical partitioning of CO2 production within a temperate forest soil

The major driving factors of soil CO₂ production – substrate supply, temperature, and water content – vary vertically within the soil profile, with the greatest temporal variations of these factors usually near the soil surface. Several studies have demonstrated that wetting and drying of the organic horizon contributes to temporal variation in summertime soil CO₂ efflux in forests, but this contribution is difficult to quantify. The objectives of this study were to partition CO₂ production vertically in a mixed hardwood stand of the Harvard Forest, Massachusetts, USA, and then to use that partitioning to evaluate how the relative contributions of CO₂ production by genetic soil horizon vary seasonally and interannually. We measured surface CO₂ efflux and vertical soil profiles of CO₂ concentration, temperature, water content, and soil physical characteristics. These data were applied to a model of effective diffusivity to estimate CO₂ flux at the top of each genetic soil horizon and the production within each horizon. A sensitivity analysis revealed sources of uncertainty when applying a diffusivity model to a rocky soil with large spatial heterogeneity, especially estimates of bulk density and volumetric water content and matching measurements of profiles and surface fluxes. We conservatively estimate that the O horizon contributed 40–48% of the total annual soil CO₂ efflux. Although the temperature sensitivity of CO₂ production varied across soil horizons, the partitioning of CO₂ production by horizon did not improve the overall prediction of surface CO₂ effluxes based on temperature functions. However, vertical partitioning revealed that water content covaried with CO₂ production only in the O horizon. Large interannual variations in estimates of O horizon CO₂ production indicate that this layer could be an important transient interannual source or sink of ecosystem C.     

四川长宁毛竹林碳储量与碳汇能力估测

利用生物量法研究了四川长宁毛竹林(Phyllostachys edulis)碳密度、碳储量及其空间分配格局,并对毛竹林碳汇能力进行了估算。结果表明:(1)毛竹立竹各器官的平均含碳率波动范围为462.37—480.68 g/kg,不同龄级毛竹各器官含碳率差异不显著。土壤有机碳含量为15.77 g/kg,不同土层差异极显著;(2)毛竹立竹碳储量为40.92 t/hm2,其中竹竿碳储量所占比例为51.49%,竹杆、竹枝、竹叶地上部分碳储量为26.76 t/hm2,占立竹碳储量的65.39%,地上碳储量为地下碳储量的1.89倍;(3)毛竹林总碳储量为156.57 t/hm2,其中土壤是其最大的碳库,为113.54 t/hm2,占总碳储量的72.52%,立竹碳储量所占比例为26.14%,林下植被碳库最小,为0.52 t/hm2,只占总碳储量的0.33%,可忽略不计;(4)毛竹林年生产量为20.28 t/hm2,年固碳量为9.43 t/hm2,相当于每年固定CO2量34.57 t/hm2,固碳能力较强。     

CO2 and intracellular pH

Abstract The experimental determination of cytoplasmic and vacuolar pH values is discussed. Despite variation in these values evidence indicates that intracellular pH values are normally regulated within narrow limits. The regulatory mechanisms proposed involve the metabolic consumption of OH^& and the active efflux of H ⁺. The evidence for intracellular pH modification in response to CO₂ hydration and the production of HCO^?₃ and H⁺ is examined. Theoretical calculations and experimental data indicate that CO₂ concentrations as high as 5% will lower intracellular pH. Conversely, variation in CO₂ levels around atmospheric concentrations is unlikely to perturb intracellular pH. High CO₂ levels are found in bulky tissues, and flooded root systems. Evidence is presented that the slow diffusion of dissolved CO₂ compared to gaseous CO₂ results in its accumulation. It is proposed that the accumulation of respiratory CO₂ may reduce intracellular pH values when plant tissues, cells or protoplasts are maintained in a liquid culture medium. Finally, the possible role of dark CO₂ fixation and organic acid synthesis in the regulation of intracellular pH is examined.     

Hierarchical saturation of soil carbon pools near a natural CO2 spring

Soil has been identified as a possible carbon (C) sink to mitigate increasing atmospheric CO₂ concentration. However, several recent studies have suggested that the potential of soil to sequester C is limited and that soil may become saturated with C under increasing CO₂ levels. To test this concept of soil C saturation, we studied a gley and organic soil at a grassland site near a natural CO₂ spring. Total and aggregate‐associated soil organic C (SOC) concentration showed a significant increase with atmospheric CO₂ concentration. An asymptotic function showed a better fit of SOC and aggregation with CO₂ level than a linear model. There was a shift in allocation of total C from smaller size fractions to the largest aggregate fraction with increasing CO₂ concentration. Litter inputs appeared to be positively related to CO₂ concentration. Based on modeled function parameters and the observed shift in the allocation of the soil C from small to large aggregate‐size classes, we postulate that there is a hierarchy in C saturation across different SOC pools. We conclude that the asymptotic response of SOC concentration at higher CO₂ levels indicates saturation of soil C pools, likely because of a limit to physical protection of SOC.     

The respiratory source of CO2

Abstract Approximately half of the carbon plants fix in photosynthesis is lost in dark respiration. The major pathways for dark respiration and their control are briefly discussed in the context of a growing plant. It is suggested that whole-plant respiration may be largely ADP-limited and that fine control of the respiratory network serves to select the respiratory substrate and to partition carbon between the numerous possible fates within the network. The striking stoichiometry between whole-plant growth and respiration is reviewed, and the relationships between substrate-limited growth and ADP-limited respiration are discussed.     

陕北农田作物生产碳源/汇及碳足迹空间特征

农作物生产过程既是碳源,也是碳汇。研究作物生产过程中碳吸收、碳排放特征对区域农业碳减排具有重要意义。以陕北区域为例,采用高分辨率遥感数据,结合GEE遥感云平台和随机森林算法,获取了作物种植分布信息,并建立碳吸收排放测算模型,分析了陕北地区2021年农田作物的碳源/汇效应、碳足迹及其空间分布格局。结果表明:①陕北种植的粮食作物主要为玉米、稻谷、薯类、豆类,经济作物主要为蔬菜、苹果、枣树,这七类作物集中分布在延安南部河谷区域和榆林西北部区域。②除枣类外,陕北地区其余作物的碳吸收量均高于碳排放量,以碳汇功能为主,其中,玉米和苹果分别对该地区碳吸收、碳排放的贡献率最高,碳吸收、排放量分别达到了189.74×10⁴ t和11.41×10⁴ t,苹果、薯类和枣类碳足迹较高,分别达到了9.92×10⁴ hm²、8.77×10⁴ hm²和21.65×10⁴ hm²,其余作物碳足迹处于0.26-1.49×10⁴ hm²之间。③从空间上看,研究区单位面积农田碳吸收量呈现西北高、南部低的分布格局,而碳排放量、碳足迹分布正好相反,南部高、西北低。④研究区可通过培育高产品种、优化施肥量、控制农膜农药用量、调整作物种植结构等措施,提高作物固碳效应,促进农业生产碳减排。     

Possible effects of rising C02 on climate

Abstract Two factors will determine the rate at which CO₂ levels in the atmosphere increase in the future: the rate of input to the atmosphere, primarily from fossil fuel burning, and the way in which this CO₂ is partitioned between atmosphere, ocean and biosphere. A brief review is given of the current state of knowledge of these aspects of the CO₂ issue prior to a discussion of the changes in climate that might be expected from increased levels of CO₂, whenever these might occur. The basis of climate modelling upon which our expectations rest is explained, indicating the nature of the uncertainty that currently exists in the model results. While some of the gross features of the likely climatic change seem reasonably well established qualitatively, considerable model development will be needed before reliable information on the likely regional effects is forthcoming. Observations have yet to confirm the occurrence of temperature change attributable to CO₂ increases. Nevertheless, the possibility exists of a change in climate during the coming century that may be substantial relative to past experience. Although direct measures to control CO₂ emissions would certainly be premature, long-term planning of infrastructures, closely tuned to present climatic conditions, should ensure their robustness in the face of the uncertain climatic changes that may lie ahead.     

Canopy-scale δ13C of photosynthetic and respiratory CO2 fluxes: observations in forest biomes across the United States

The δ¹³C values of atmospheric carbon dioxide (CO₂) can be used to partition global patterns of CO₂ source/sink relationships among terrestrial and oceanic ecosystems using the inversion technique. This approach is very sensitive to estimates of photosynthetic ¹³C discrimination by terrestrial vegetation (Δ_A), and depends on δ¹³C values of respired CO₂ fluxes (δ¹³C_R). Here we show that by combining two independent data streams – the stable isotope ratios of atmospheric CO₂ and eddy‐covariance CO₂ flux measurements – canopy scale estimates of Δ_A can be successfully derived in terrestrial ecosystems. We also present the first weekly dataset of seasonal variations in δ¹³C_R from dominant forest ecosystems in the United States between 2001 and 2003. Our observations indicate considerable summer‐time variation in the weekly value of δ¹³C_R within coniferous forests (4.0‰ and 5.4‰ at Wind River Canopy Crane Research Facility and Howland Forest, respectively, between May and September). The monthly mean values of δ¹³C_R showed a smaller range (2–3‰), which appeared to significantly correlate with soil water availability. Values of δ¹³C_R were less variable during the growing season at the deciduous forest (Harvard Forest). We suggest that the negative correlation between δ¹³C_R and soil moisture content observed in the two coniferous forests should represent a general ecosystem response to the changes in the distribution of water resources because of climate change. Shifts in δ¹³C_R and Δ_A could be of sufficient magnitude globally to impact partitioning calculations of CO₂ sinks between oceanic and terrestrial compartments.     

The utilization of pre-anthesis reserves in grain filling of wheat. Assessment by steady-state 13CO2/12CO2 labelling

δ, C isotope composition relative to Pee Dee Belemnite
WSC, water-soluble carbohydrates
N, nitrogen
C, carbon
cv, cultivar
ME, efficiency of mobilized pre-anthesis C utilization in grain filling (g C g^–1C)

Significant mobilization of protein and carbohydrates in vegetative plant parts of wheat regularly occurs during grain filling. While this suggests a contribution of reserves to grain filling, the actual efficiency of mobilized assimilate conversion into grain mass (ME) is unknown. In the present study the contribution of pre-anthesis C (C fixed prior to anthesis) to grain filling in main stem ears of two spring wheat (Triticum aestivum L.) cultivars was determined by ¹³C/¹²C steady-state labelling. Mobilization of pre-anthesis C in vegetative plant parts between anthesis and maturity, and the contributions of water-soluble carbohydrates (WSC) and protein to pre-anthesis C mobilization were also assessed. Experiments were performed with two levels of N fertilizer supply in each of 2 years. Pre-anthesis reserves contributed 11–29% to the total mass of C in grains at maturity. Pre-anthesis C accumulation in grains was dependent on both the mass of pre-anthesis C mobilized in above-ground vegetative plant parts (r² = 0·87) and ME (defined as g pre-anthesis C deposited in grains per g pre-anthesis C mobilized in above-ground vegetative plant parts; r² = 0·40). ME varied between 0·48 and 0·75. The effects of years, N fertilizer treatments and cultivars on ME were all related to differences in the fractional contribution of WSC to pre-anthesis C mobilization. Multiple regression analysis indicated that C from mobilized pre-anthesis WSC may be used more efficiently in grain filling than C present in proteins at anthesis and mobilized during grain filling. Possible causes for variability of ME are discussed.