亚热带山区红壤地碳平衡研究进展

碳平衡研究日益成为全球变化与地球科学研究领域的热点问题.亚热带红壤区是我国发展粮食作物和各种热带、亚热带经济作物与林木的重要基地,因该区特殊的生态地理位置,在我国碳平衡研究中占有重要地位.本文论述了亚热带山区红壤地碳平衡研究的重要性,对碳平衡研究中植被、凋落物、土壤碳库和土壤呼吸的研究现状和主要结论等进行阐述,总结了碳平衡的综合研究方法,并对亚热带山区红壤地碳平衡研究中存在的问题和今后的发展方向进行探讨.     

Soil respiration and carbon balance in a subtropical native forest and two managed plantations

From 1999 to 2003, a range of carbon fluxes was measured and integrated to establish a carbon balance for a natural evergreen forest of Castanopsis kawakamii (NF) and adjacent monoculture evergreen plantations of C. kawakamii (CK) and Chinese fir (Cunninghamia lanceolata, CF) in Sanming Nature Reserve, Fujian, China. Biomass carbon increment of aboveground parts and coarse roots were measured by the allometric method. Above- and belowground litter C inputs were assessed by litter traps and sequential cores, respectively. Soil respiration (SR) was determined by the alkaline absorbance method, and the contribution from roots, above- and belowground litters was separated by the DIRT plots. Annual SR averaged 13.742 t C ha⁻¹ a⁻¹ in the NF, 9.439 t C ha⁻¹ a⁻¹ in the CK, and 4.543 t C ha⁻¹ a⁻¹ in the CF. For all forests, SR generally peaked in later spring or early summer (May or June). The contribution of root respiration ranged from 47.8% in the NF to 40.3% in the CF. On average, soil heterotrophic respiration (HR) was evenly distributed between below- (47.3∼54.5%) and aboveground litter (45.5%–52.7%). Annual C inputs (t C ha⁻¹ a⁻¹) from litterfall and root turnover averaged 4.452 and 4.295, 4.548 and 2.313, and 2.220 and 1.265, respectively, in the NF, CK, and CF. As compared to HR, annual net primary production (NPP) of 11.228, 13.264, and 6.491 t C ha⁻¹ a⁻¹ in the NF, CK, and CF brought a positive net ecosystem production (NEP) of 4.144, 7.514, and 3.677 t C ha⁻¹ a⁻¹, respectively. It suggests that native forest in subtropical China currently acts as an important carbon sink just as the timber plantation does, and converting native forest to tree plantations locally during last decades might have caused a high landscape carbon loss to the atmosphere.     

Low stocks of coarse woody debris in a southwest Amazonian forest

The stocks and dynamics of coarse woody debris (CWD) are significant components of the carbon cycle within tropical forests. However, to date, there have been no reports of CWD stocks and fluxes from the approximately 1.3 million km² of lowland western Amazonian forests. Here, we present estimates of CWD stocks and annual CWD inputs from forests in southern Peru. Total stocks were low compared to other tropical forest sites, whether estimated by line-intercept sampling (24.4 ± 5.3 Mg ha⁻¹) or by complete inventories within 11 permanent plots (17.7 ± 2.4 Mg ha⁻¹). However, annual inputs, estimated from long-term data on tree mortality rates in the same plots, were similar to other studies (3.8 ± 0.2 or 2.9 ± 0.2 Mg ha⁻¹ year⁻¹, depending on the equation used to estimate biomass). Assuming the CWD pool is at steady state, the turnover time of coarse woody debris is low (4.7 ± 2.6 or 6.1 ± 2.6 years). These results indicate that these sites have not experienced a recent, large-scale disturbance event and emphasise the distinctive, rapid nature of carbon cycling in these western Amazonian forests.     

Effects of elevated CO2 on growth and carbon/nutrient balance in the deciduous woody shrub Lindera benzoin (L.) Blume (Lauraceae)

We examined the effects of elevated CO₂ on growth and carbon/nutrient balance in a natural population of the deciduous temperate zone shrub Lindera benzoin. Our data concern whole plant, leaf, and stem growth for the first two seasons of a long-term field experiment in which CO₂ levels were manipulated in situ. In addition to growth parameters, we evaluated changes in leaf and stem chemistry, including total nitrogen, nonstructural carbohydrates, and total phenolics. Over the course of this study, L. benzoin appeared to respond to elevated CO₂ primarily by physiological and biochemical changes, with only a slight enhancement in aboveground growth (ramet height). Positive effects on aboveground growth were primarily evident in young (nonreproductive) ramets. Our results suggest that nitrogen limitation may have constrained plants to allocate carbohydrates produced in response to elevated CO₂ primarily to storage and belowground growth, and perhaps to increased secondary chemical production, rather than to increased stem and leaf growth. We discuss our results in terms of changes in carbon/nutrient balance induced by elevated CO₂, and provide predictions for future changes in this system based upon constraints imposed by intrinsic and extrinsic factors and their potential effects on the reallocation of stored reserves.     

Quantitative analysis of carbon balance in the reproductive organs and leaves of <Emphasis Type="Italic">Cinnamomum camphora</Emphasis> (L.) Presl

We investigated seasonal changes in dry mass and CO₂ exchange rate in fruit and leaves of the evergreen tree Cinnamomum camphora with the aim of quantitatively determining the translocation balance between the two organs. The fruit dry mass growth peaked in both August and October: the first increase was due to fruit pulp development and the second to seed development. Fruit respiration also increased with the rapid increase in fruit dry mass. Therefore, the carbohydrates required for fruit development showed two peaks during the reproductive period. Fruit photosynthesis was relatively high in early August, when fruit potentially re-fixed 75% of respired CO₂, indicating that fruit photosynthesis contributed 15–35% of the carbon requirement for fruit respiration. Current-year leaves completed their growth in June when fruit growth began. Current-year leaves translocated carbohydrates at a rate of approximately 10–25 mg dry weight (dw) leaf⁻¹ day⁻¹ into other organs throughout the entire fruit growth period. This rate of translocation from current-year leaves was much higher than the amount of carbohydrate required for reproduction (ca. 3 mg dw fruit⁻¹ day⁻¹). Given the carbon balance between fruit and current-year leaves, carbohydrates for reproduction were produced within the current-year fruit-bearing shoots. C. camphora would be adaptive for steadily supplying enough amount of carbohydrate to the fruits, as there was little competition for carbohydrates between the two organs. As assimilates by leaves are used for processes such as reproduction and the formation of new shoots, photosynthesis by reproductive organs is considered to be important to compensate for reproductive cost.     

A carbon balance model of the sorghum-Striga hermonthica host-parasite association

Abstract. Growth and gas exchange measurements are used to formulate a carbon balance model to describe the sorghum- Striga hermonthica host-Parasite association. S. hermonthica reduces the growth and radically alters the architecture of infected sorghum plants. Grain and stem weight are reduced, whilst leaf and root biomass are maintained. Losses in host productivity result from two processes: export of carbon to the parasite and Parasite-induced reductions in host photosynthesis. The latter occurs before the emergence of the Parasite above ground and accounts for 80% of the Predicted loss in production over the lifecycle of the association. S. hermonthica is dependent on carbon exported from the host, since the plant has low rates of photosynthesis coupled with high rates of respiration. Host-derived carbon accounts for approximately one-third of the total parasite carbon requirement.     

Simulated annual carbon fluxes of grassland ecosystems in extremely arid conditions

In order to understand how changes in climate and land cover affect carbon cycles and structure and function of regional grassland ecosystems, we developed a grassland landscape productivity model, proposed an approach that combined both process-based modeling and spatial analysis with field measurements, and provided an example of semiarid region in Inner Mongolia, China, in extremely arid conditions. The modeled monthly mean and total net primary productivity, and monthly and annual mean respiration rates for Leymus chinensis steppes during the growing seasons in 2002 were mostly within the normal varying ranges of measured values, or similar to the field measurements, conducted in the similarly arid conditions. And the modeled total net ecosystem productivity (NEP) for L. chinensis steppes and Stipa grandis steppes were both between the lower and the higher measurements or within modeled multi-annual data by the other model. The modeled total NEP was 1.91 g C/m²/year over the entire study region. It indicated that if human disturbances were not considered, carbon budget over the entire study region during the growing seasons was nearly in balance or weak carbon sink even under extremely arid conditions. However, the modeled NEP spatially greatly varied not only over the entire study region (−48.28–52.09 g C/m²/year), but also among different land cover types. The modeled results also showed that there were obvious seasonal variations in carbon fluxes, mainly caused by leaf area index; and annual precipitation was the major limiting factor for the obvious spatial patterns of carbon fluxes from the east to the west. The modeled results also revealed the influence of extreme drought on carbon fluxes. The study provides an effective approach to derive useful information about carbon fluxes for different land cover types, and thus can instruct regional land-use planning and resource management based on carbon storage conditions.     

Pollination and breeding systems of woody plant species in tropical dry evergreen forests,southern India

Tropical dry evergreen forests (TDEF) are a unique forest type found along the east coast of India. They mostly occur as small, isolated fragments of varying sizes (0.5 to ≈10 ha) and are considered as endangered forests types in peninsular India. Although plant diversity is well documented in these forests, there is a paucity of ecological studies vital for conservation and for planning restoration activities. We studied reproductive biology of 13 woody species: four trees, six shrubs, and three lianas in fragments of TDEF in southern India. The phenology of reproduction, floral biology, anthesis and sexual system of each species were recorded. The pollination mode was assessed through observations of the visitation frequency of pollinators and from the floral characters. The breeding system was determined by hand-augmented self- and cross-pollination experiments. The plants flowered during the dry season from January to July. Plants of nine species had both flowers and fruits at the same time. Twelve species were hermaphrodites and one was polygamo-dioecious. Flowers of 11 species opened at dawn and two at dusk. Four species were self-incompatible and six were self-compatible. Natural fruit set ranged from 10% to 56%, self-incompatible species having low fruit set. Cross-pollen augmentation increased fruit set, suggesting presence of outcrossing in all species. The majority of plants species (85%) had a generalized pollination system, receiving visits from diverse insects, such as social bees, solitary bees, wasps, moths and flies. However, only few of them were functionally important for the species. Two species namely: Capparis brevispina and C. zeylanica had butterflies and birds, respectively, as their main pollinators. Our data reveal that there is a predominance of outcrossing in plant species and a generalized pollination system in these forests. We suggest that restoration of TDEFs is crucial as habitats, not only for wild plants but also for pollinating insects.     

Influence of management and precipitation on carbon fluxes in great plains grasslands

Suitable management and sufficient precipitation on grasslands can provide carbon sinks. The net carbon accumulation of a site from the atmosphere, modeled as the Net Ecosystem Productivity (NEP), is a useful means to gauge carbon balance. Previous research has developed methods to integrate flux tower data with satellite biophysical datasets to estimate NEP across large regions. A related method uses the Ecosystem Performance Anomaly (EPA) as a satellite-derived indicator of disturbance intensity (e.g., livestock stocking rate, fire, and insect damage). To better understand the interactions among management, climate, and carbon dynamics, we evaluated the relationship between EPA and NEP data at the 250 m scale for grasslands in the Central Great Plains, USA (ranging from semi-arid to mesic). We also used weekly estimates of NEP to evaluate the phenology of carbon dynamics, classified by EPA (i.e., by level of disturbance impact). Results show that the cumulative carbon balance over these grasslands from 2000 to 2008 was a weak net sink of 13.7 g C m⁻² yr⁻¹. Overall, NEP increased with precipitation (R² = 0.39, P < 0.05) from west to east. Disturbance influenced NEP phenology; however, climate and biophysical conditions were usually more important. The NEP response to disturbance varies by ecoregion, and more generally by grassland type, where the shortgrass prairie NEP is most sensitive to disturbance, the mixed-grass prairie displays a moderate response, and tallgrass prairie is the least impacted by disturbance (as measured by EPA). Sustainable management practices in the tallgrass and mixed-grass prairie may potentially induce a period of average net carbon sink until a new equilibrium soil organic carbon is achieved. In the shortgrass prairie, management should be considered sustainable if carbon stocks are simply maintained. The consideration of site carbon balance adds to the already difficult task of managing grasslands appropriately to site conditions. Results clarify the seasonal and interannual dynamics of NEP, specifically the influence of disturbance and moisture availability.     

Translocation and utilization of carbon in wheat (Triticum aestivum)

Wheat ( Triticum aestivum L. cv. SUN 9E) was grown in a growth chamber under conditions of low soil nitrogen. Translocation of carbon to the roots and the subsequent utilization of these carbohydrates was determined. In vegetative plants (22 days old), 21.5 mg C day⁻¹ were translocated to the roots. 29% of this was incorporated into dry matter, 32% was respired (28% via the cytochrome and 4% via a SHAM-sensitive, presumably the alternative nonphosphorylating, pathway) and 39% was translocated back to the shoots, mainly in the form of amino acids. – The rote of root maintenance respiration during the vegetative phase was estimated to be 0.7 mg O₂ h⁻¹ (g dry weight of roots)⁻¹ and the root growth respiration to be 0.41 g O₂ (g dry weight of roots)⁻¹. Total carbohydrate utilization due to root respiration via the alternative, nonphosphorylating pathway during the major part of the growth period was calculated to be only ca 6% of carbohydrate utilization for grain growth. The rate of specific mass transfer (SMT) of sugars in the sieve tubes was estimated from the data on C-translocation and data on the total area occupied by sieve tubes in a cross section of the root system. SMT was calculated to be 0.8 mg sucrose s⁻¹ cm⁻², which is very similar to the published value on SMT for other organs, except roots.     

Responses of dry weight growth under SO2 stress in an SO2-tolerant plant,Polygonum cuspidatum

The effects of exposure to 0.5–0.7 ppm SO₂ for about one month on the dry weight growth and net photosynthesis ofPolygonum cuspidatum were investigate. Furthermore, the carbon and nitrogen concentrations in each plant organ were measured. The results obtained showed no significant decrease in the total dry weight of SO₂-treated plants in comparison with controls. On the other hand, the leaf area (LA) and/or leaf dry weight of SO₂-treated plants were increased, and the root dry weight (RW) was decreased, in comparison with controls. The leaf carbon assimilation rate (CAR) in SO₂-treated plants was slightly decreased in spite of a clear decrease in net photosynthesis, and the value of (SW+RW)/LA (SW stem dry weight) was decreased in comparison with controls, thus minimizing the reduction in CAR. Furthermore, the ratio of total leaf carbon absorption (leaf area x CAR) to total root nitrogen absorption (root dry weight x nitrogen assimilation rate) in SO₂-treated plants was similar to that in controls. From these results, it can be concluded that an increase in leaf area and/or leaf dry weight and a decrease in root dry weight inP. cuspidatum under SO₂ stress may be induced in order to compensate for the decrease in CAR and to maintain the ratio of total leaf carbon absorption to total root nitrogen absorption in the early stage of vegetative growth.     

基于水文平衡的湿地退化驱动因子定量研究

作为地球上最为重要的生态系统类型之一,湿地与森林、海洋并称为地球三大生态系统。但是,近年来湿地生态系统退化速度远大于其他类型生态系统,开展湿地退化的定量评估分析研究对于湿地生态系统的保护和恢复具有重要意义。选择北京城市湿地为研究对象,利用卫星遥感数据分别提取得到1991年和2007年的湿地面积,基于湿地水量平衡理论和湿地水文方程方法,定量评估分析了导致湿地退化的原因和不同驱动因子的贡献率。结果表明:(1)与1991年相比,2007年北京湿地减少约6275.31 hm2,约占1991年北京湿地总面积的24.46%。显著退化区域主要发生在野鸭湖湿地和密云水库湿地,分别减少了约1377.69 hm2和4654.50 hm2。(2)引起湿地退化的自然驱动因子中,以降水减少、入境地表水减少和蒸发量增加为主,驱动湿地退化的贡献率分别为39.22%、14.05%和11.85%。引起湿地退化的人为驱动因子中,以城市扩展为主,驱动湿地退化的贡献率为3.42%,而技术进步所采取的节水措施等有利于湿地保护,贡献率为25.55%。     

Retranslocation of carbon reserves from the woody storage tissues into the fruit as a response to defoliation stress during the ripening period in Vitis vinifera L.

A technique for reliable labeling of the carbon reserves of the trunk and roots without labeling the current year's growth of grapevines was developed in order to study retranslocation of carbon from the perennial storage tissues into the fruit in response to defoliation stress during the ripening period. A special training system with two shoots was used: the lower one (feeding shoot) was cut back and defoliated to one single leaf (¹⁴CO₂-feeding leaf) while the other (main shoot) was topped to 12 leaves. The potted plants were placed in a water bath at 30 °C to increase root temperature and therefore their sink activity. Additionally, a cold barrier (2–4 °C) was installed at the base of the main shoot to inhibit acropetal ¹⁴C translocation. Using this method, we were able to direct labeled assimilates to trunk and roots in preference to the current year's growth. On vines with root and shoot at ambient temperature, 44% of the ¹⁴C activity was found in the main shoot 16 h after feeding whereas only 2% was found in the temperature-treated vines. At the onset of fruit ripening, and at three-week intervals thereafter until harvest, potted grapevines were fed with ¹⁴CO₂ using the temperature treatment described above. Sixteen hours after feeding, half of the vines of each group were defoliated by removing all except the two uppermost main leaves. Three weeks after each treatment, vines were destructively harvested and the dry weight and ¹⁴C incorporation determined for all plant parts. Under non-stressing conditions, there was no retranslocation of carbon reserves to support fruit maturation. Vines responded to defoliation stress by altering the natural translocation pattern and directing carbon stored in the lower parts to the fruit. In the three weeks following veraison (the inception of ripening in the grape berry), 12% of the labeled carbon reserves was translocated to the fruit of defoliated plants compared to 1.6% found in the clusters of control vines. Retranslocation from trunk and roots was highest during the middle of the ripening period, when 32% of the labeled carbon was found in the fruit compared to 0.7% in control plants. Defoliation during this period also caused major changes in dry-matter partitioning: the fruit represented 31% of total plant biomass compared to 21% measured in the control vines. Root growth was reduced by defoliation at veraison and during the ripening period. Defoliation three weeks before harvest did not affect dry matter or ¹⁴C partitioning.     

Seasonal patterns of canopy development and carbon gain in nineteen warm desert shrub species

Summary Canopy development and photosynthetic rate were measured at monthly intervals over a period of one year in 19 shrub and subshrub species of the Mojave and upper Sonoran Deserts. Thirteen of these species realized a substantial fraction of their total net carbon assimilation via twig photosynthesis. The twig contribution to whole plant yearly carbon gain reached a maximum of 83% in species such as Thamnosma montana, Salizaria mexicana, and Baccharis brachyphylla. This large contribution by twigs was due to both low levels of leaf production and the greater longevity of twig tissues. In some other species, however, leaf and twig organs had similar lifespans. During the year of this study (which had an unusually warm, mild winter), no species showed a pattern of winter deciduousness. The reduction in total photosynthetic area between maximal spring canopy development and mid August summer dormancy ranged from 32 to 94%. Some herbaceous perennial species died back to the ground, but none of the woody shrubs were totally without green canopy area at any time of the year. No species studied were capable of high rates of photosynthesis at low plant water potentials in July and August, but, in those species which maintained a substantial canopy area through the drought period, previously stressed tissues showed substantial recovery after fall rains. Photosynthetic rate was significantly correlated with both plant water potential and tissue nitrogen content over the entire year, but only weakly so. This is due in part to the winter months when plant water potentials and tissue nitrogen contents were high, but photosynthetic rates were often low.     

Stoichiometric analysis of dissolved organic carbon flux into storage and growth in aerobic granules culture

This study attempted to address a fundamental question of whether metabolic behaviors of aerobic granules are different from their counterparts, such as activated sludge and biofilms. A series of respirometric experiments were carried out using mature aerobic granules with mean sizes of 0.75–3.4 mm. Results suggested that metabolism of aerobic granules comprised three consecutive phases: (i) conversion of external dissolved organic carbon to a poly-β-hydroxybutyrate-like substance; (ii) growth of aerobic granules on the stored poly-β-hydroxybutyrate-like substance derived from phase I, and (iii) subsequent endogenous metabolism of aerobic granules. The stoichiometric analysis revealed that the conversion yields of external dissolved organic carbon to the poly-β-hydroxybutyrate-like substance, the growth yields of biomass on storage, and the overall growth yields of biomass on external dissolved organic carbon were not significantly correlated to the sizes of aerobic granules, i.e., the metabolism of aerobic granules would be size independent. The conversion coefficients and growth rates of aerobic granules were found to be comparable with those reported in the activated sludge and biofilms cultures, indicating that there would not be significant difference in the metabolisms of aerobic granules over activated sludge and biofilms. This information will be useful for modeling and designing aerobic granular sludge processes.     

Stem respiration and carbon dioxide efflux of young <Emphasis Type="Italic">Populus deltoides</Emphasis> trees in relation to temperature and xylem carbon dioxide concentration

Oxidative respiration is strongly temperature driven. However, in woody stems, efflux of CO₂ to the atmosphere (E _A), commonly used to estimate the rate of respiration (R _S), and stem temperature (T _st) have often been poorly correlated, which we hypothesized was due to transport of respired CO₂ in xylem sap, especially under high rates of sap flow (f _s). To test this, we measured E _A, T _st, f _s and xylem sap CO₂ concentrations ([CO₂*]) in 3-year-old Populus deltoides trees under different weather conditions (sunny and rainy days) in autumn. We also calculated R _S by mass balance as the sum of both outward and internal CO₂ fluxes and hypothesized that R _S would correlate better with T _st than E _A. We found that E _A sometimes correlated well with T _st, but not on sunny mornings and afternoons or on rainy days. When the temperature effect on E _A was accounted for, a clear positive relationship between E _A and xylem [CO₂*] was found. [CO₂*] varied diurnally and increased substantially at night and during periods of rain. Changes in [CO₂*] were related to changes in f _s but not T _st. We conclude that changes in both respiration and internal CO₂ transport altered E _A. The dominant component flux of R _S was E _A. However, on a 24-h basis, the internal transport flux represented 9–18% and 3–7% of R _S on sunny and rainy days, respectively, indicating that the contribution of stem respiration to forest C balance may be larger than previously estimated based on E _A measurements. Unexpectedly, the relationship between R _S and T _st was sometimes weak in two of the three trees. We conclude that in addition to temperature, other factors such as water deficits or substrate availability exert control on the rate of stem respiration so that simple temperature functions are not sufficient to predict stem respiration.