Linking microbial activity and soil organic matter transformations in forest soils under elevated CO2

Soil organic matter (SOM) dynamics ultimately govern the ability of soil to provide long‐term C sequestration and the nutrients required for ecosystem productivity. Predicting belowground responses to elevated CO₂ requires an integrated understanding of SOM transformations and the microbial activity that governs them. It remains unclear how the microorganisms upon which these transformations depend will function in an elevated CO₂ world. This study examines SOM transformations and microbial metabolism in soils from the Duke Free Air Carbon Enrichment site in North Carolina, USA. We assessed microbial respiration and net nitrogen (N) mineralization in soils with and without elevated CO₂ exposure during a 100‐day incubation. We also traced the depleted C isotopic signature of the supplemental CO₂ into SOM and the soils' phospholipid fatty acids (PLFA), which serve as biomarkers for living cells. Cumulative net N mineralization in elevated CO₂ soils was 50% that in control soils after a 100‐day incubation. Respiration was not altered with elevated CO₂. C : N ratios of bulk SOM did not change with elevated CO₂, but incubation data suggest that the C : N ratios of mineralized organic matter increased with elevated CO₂. Values of SOM δ¹³C were depleted with elevated CO₂ (?26.7±0.2 vs. ?30.2±0.3‰), reflecting the depleted signature of the supplemental CO₂. We compared δ¹³C of individual PLFA with the δ¹³C of SOM to discern incorporation of the depleted C isotopic signature into soil microbial groups in elevated CO₂ plots. PLFA i15:0, a15:0, and 10Met18:0 reflected significant incorporation of recently produced photosynthate, suggesting that the bacterial groups defined by these biomarkers are active metabolizers in elevated CO₂ soils. At least one of these groups (actinomycetes, 10Met18:0) specializes in metabolizing less labile substrates. Because control plots did not receive an equivalent ¹³C tracer, we cannot determine from these data whether this group of organisms was stimulated by elevated CO₂ compared with these organisms in control soils. Stimulation of this group, if it occurred in the elevated CO₂ plot, would be consistent with a decline in the availability of mineralizable organic matter with elevated CO₂, which incubation data suggest may be the case in these soils.     

Interactions between Carbon and Nitrogen Mineralization and Soil Organic Matter Chemistry in Arctic Tundra Soils

We used long-term laboratory incubations and chemical fractionation to characterize the mineralization dynamics of organic soils from tussock, shrub, and wet meadow tundra communities, to determine the relationship between soil organic matter (SOM) decomposition and chemistry, and to quantify the relative proportions of carbon (C) and nitrogen (N) in tundra SOM that are biologically available for decomposition. In all soils but shrub, we found little decline in respiration rates over 1 year, although soils respired approximately a tenth to a third of total soil C. The lack of decline in respiration rates despite large C losses indicates that the quantity of organic matter available was not controlling respiration and thus suggests that something else was limiting microbial activity. To determine the nature of the respired C, we analyzed soil chemistry before and after the incubation using a peat fractionation scheme. Despite the large losses of soil C, SOM chemistry was relatively unchanged after the incubation. The decomposition dynamics we observed suggest that tundra SOM, which is largely plant detritus, fits within existing concepts of the litter decay continuum. The lack of changes in organic matter chemistry indicates that this material had already decomposed to the point where the breakdown of labile constituents was tied to lignin decomposition. N mineralization was correlated with C mineralization in our study, but shrub soil mineralized more and tussock soil less N than would have been predicted by this correlation. Our results suggest that a large proportion of tundra SOM is potentially mineralizable, despite the fact that decomposition was dependent on lignin breakdown, and that the historical accumulation of organic matter in tundra soils is the result of field conditions unfavorable to decomposition and not the result of fundamental chemical limitations to decomposition. Our study also suggests that the anticipated increases in shrub dominance may substantially alter the dynamics of SOM decomposition in the tundra. Received 31 January 2002; accepted 16 July 2002.     

Soil carbon stabilization in jack pine stands along the Boreal Forest Transect Case Study

Boreal forests, containing >20% of the total organic carbon (OC) present at the surface of the Earth, are expected to be highly vulnerable to global warming. The objective of this study was to compare soil OC stocks and chemistry in jack pine stands located along a latitudinal climatic transect in central Canada. Total OC stocks (0–1 m) increased with decreasing mean annual temperature (MAT). We used a combination of physical fractionation of soil OC pools, ¹³C isotopic determination and cross‐polarization, magic‐angle spinning ¹³C nuclear magnetic resonance (NMR) spectroscopy to further characterize OC composition at all sites. Soil OC was dominated by labile pools. As illustrated by the C/N ratios, δ¹³C data and results from the ¹³C NMR analysis, the light fraction showed little alteration within the soil profiles. Instead, this fraction reflected the importance of fresh litter inputs and showed an increase in root contribution with depth. As opposed to the light fraction, the clay‐ and silt‐stabilized OC exhibited an increase in δ¹³C and a decrease in C/N with depth, indicating an increase in its degree of decomposition. These changes with depth were more marked at the southern than the northern sites. Results hence suggest that if the MAT were to increase in the northern boreal forest the overall jack pine soil OC stocks would decrease but the remaining OC would become more decomposed, and likely more stabilized than what is currently present within the soils.     

Grazing and Ecosystem Carbon Storage in the North American Great Plains

Isotopic signatures of ¹³C were used to quantify the relative contributions of C₃ and C₄ plants to whole-ecosystem C storage (soil+plant) in grazed and ungrazed sites at three distinct locations (short-, mid- and tallgrass communities) along an east–west environmental gradient in the North American Great Plains. Functional group composition of plant communities, the source and magnitude of carbon inputs, and total ecosystem carbon storage displayed inconsistent responses to long-term livestock grazing along this gradient. C₄ plants [primarily Bouteloua gracilis (H.B.K.) Lag ex Steud.] dominated the long-term grazed site in the shortgrass community, whereas the ungrazed site was co-dominated by C₃ and C₄ species; functional group composition did not differ between grazed and ungrazed sites in the mid- and tallgrass communities. Above-ground biomass was lower, but the relative proportion of fine root biomass was greater, in grazed compared to ungrazed sites at all three locations. The grazed site of the shortgrass community had 24% more whole-ecosystem carbon storage compared to the ungrazed site (4022 vs. 3236 g C m⁻²). In contrast, grazed sites at the mid- and tallgrass communities had slightly lower (8%) whole-ecosystem carbon storage compared to ungrazed sites (midgrass: 7970 vs. 8683 g C m⁻²; tallgrass: 8273 vs. 8997 g C m⁻²). Differential responses between the shortgrass and the mid- and tallgrass communities with respect to grazing and whole-ecosystem carbon storage are likely a result of: (1) maintenance of larger soil organic carbon (SOC) pools in the mid- and tallgrass communities (7476–8280 g C m⁻²) than the shortgrass community (2517–3307 g C m⁻²) that could potentially buffer ecosystem carbon fluxes, (2) lower root carbon/soil carbon ratios in the mid- and tallgrass communities (0.06–0.10) compared to the shortgrass community (0.20–0.27) suggesting that variation in root organic matter inputs would have relatively smaller effects on the size of the SOC pool, and (3) the absence of grazing-induced variation in the relative proportion of C₃ and C₄ functional groups in the mid- and tallgrass communities. We hypothesize that the magnitude and proportion of fine root mass within the upper soil profile is a principal driver mediating the effect of community composition on the biogeochemistry of these grassland ecosystems.     

Changes in Soil Carbon and Nitrogen after Contrasting Land-use Transitions in Northeastern Costa Rica

Northeastern Costa Rica is a mosaic of primary and secondary forests, tree plantations, pastures, and cash crops. Many studies have quantified the effects of one type of land-use transition (for example, deforestation or reforestation) on soil properties such as organic carbon (C) storage, but few have compared different land-use transitions simultaneously. We can best understand the effects of land-use change on regional and global ecosystem processes by considering all of the land-use transitions that occur in a landscape. In this study, I examined the changes in total soil C and nitrogen (N) pools (to 0.3 m) that have accompanied different land-use transitions in a 140,000-ha region in northeastern Costa Rica. I paired sites that had similar topography and soils but differed in recent land-use history. The following land-use transitions were represented: 12 conversions of primary forests to banana plantations, 15 conversions of pastures to cash crops, and four conversions of pastures to Vochysia guatemalensis tree plantations. The conversion of forests to bananas decreased soil C concentrations and inventories (Mg C ha^–1) in the surface soil by 37% and 16.5%, respectively. The conversion of pastures to cash crops reduced soil C concentrations and inventories to the same extent that forest-to-banana cropping did. Furthermore, young Vochysia plantations do not appear to increase soil C storage, at least over the 1st decade. When data from all land-use transitions were pooled, the difference in root biomass and leaf litter pools between land-use pairs explained 50% of the differences in soil C concentrations and 36% of the differences in soil C inventories. Thus, reduced productivity or C inputs to the soil is one mechanism that could explain the losses in soil C pools with land-use change. In this landscape, losses of soil C due to cultivation are rapid, whereas re- accumulation rates are slow. Total soil N pools (0–10 cm) were also reduced after the conversion of forests to banana plantations or the conversion of pastures to crops, despite fertilization of the cropped soils. This suggests that the added N fertilizer is not retained but instead is exported via produce, N gas emissions, and hydrologic processes.     

Carbon sequestration under Miscanthus: a study of 13C distribution in soil aggregates

The growing of bioenergy crops has been widely suggested as a key strategy in mitigating anthropogenic CO₂ emissions. However, the full mitigation potential of these crops cannot be assessed without taking into account their effect on soil carbon (C) dynamics. Therefore, we analyzed the C dynamics through four soil depths under a 14‐year‐old Miscanthus plantation, established on former arable land. An adjacent arable field was used as a reference site. Combining soil organic matter (SOM) fractionation with ¹³C natural abundance analyses, we were able to trace the fate of Miscanthus‐derived C in various physically protected soil fractions. Integrated through the whole soil profile, the total amount of soil organic carbon (SOC) was higher under Miscanthus than under arable crop, this difference was largely due to the input of new C. The C stock of the macroaggregates (M) under Miscanthus was significantly higher than those in the arable land. Additionally, the C content of the micro‐within macroaggregates (mM) were higher in the Miscanthus soil as compared with the arable soil. Analysis of the intramicroaggregates particulate organic matter (POM) suggested that the increase C storage in mM under Miscanthus was caused by a decrease in disturbance of M. Thus, the difference in C content between the two land use systems is largely caused by soil C storage in physically protected SOM fractions. We conclude that when Miscanthus is planted on former arable land, the resulting increase in soil C storage contributes considerably to its CO₂ mitigation potential.     

重金属污染对土壤有机质积累的影响

采用田间采样分析与室内培养试验相结合的方法,研究了不同重金属污染土壤中有机质积累的差异及重金属污染强度对土壤有机质矿化动态变化的影响.结果表明：污染土壤中重金属的大量积累可减弱有机物质的矿化速率,增加土壤有机质的积累.土壤中颗粒态有机质及其占总有机碳的比例随重金属积累的增加而增加;而微生物生物量碳占总碳的比例却随土壤重金属污染水平的提高而下降.污染土壤中颗粒态有机质对重金属有显著的富集,这可能是影响土壤有机物质进一步矿化的原因之一.重金属污染可改变土壤有机质的矿化速率,影响土壤有机质的积累与分配.     

Carbon Quality and Stocks in Organic Horizons in Boreal Forest Soils

We investigated the mechanisms that determine the quality and quantity of organic carbon (C) stocks in boreal forest soils by analyzing both qualitative and quantitative changes in the organic fractions in the soil organic matter (OM) in a vertical gradient in the decomposition continuum of the organic horizon [litter layer (L), fermentation layer (F), and humus layer (H)] in forest soils using a sequential fractionation method at two forest types along a climatic gradient in Finland. We predicted that the concentrations of water-soluble (WSE) and non-polar (NPE) extractives should decrease and those of the acid-soluble (AS) fraction and acid-insoluble residue (AIR) should increase from the L to the F, and from the F to the H layers, but the C/N ratio of soil OM should stay constant after reaching the critical quotient. We also predicted that the AIR concentrations should be higher in the south than north boreal, and in sub-xeric than mesic forests. Consistent with our hypothesis, the concentrations of WSE and NPE fractions decreased and concentrations of AIR increased in the vertical soil gradient. The highest concentrations of the AS fraction were found in the F layer. The C/N ratio was lowest in the F layer, and the highest in the H layer, indicating that soil OM is depleted in N in relation to C along the vertical soil gradient. Concentrations of WSE and NPE were lower, and concentrations of AIR were higher in the south than in north boreal forests, which is in agreement with our hypothesis that higher soil temperatures may enhance accumulation of slowly decomposable OM in the soil. The concentrations of AIR were higher in the sub-xeric than mesic forests. Contrary to our expectations, however, the differences in the chemical quality in soil OM between the site types were amplified from the L to the H layer. The size of the C storage was significantly larger in south than north boreal sites, and larger in the mesic than in the sub-xeric sites.     

Effects of Dominant Plant Species on Soils during Succession in Nutrient-poor Ecosystems

During the initial phases of succession on nutrient-poor, mineral substrates dead plant material accumulates rapidly in the soil. This accumulation of soil organic matter can result in a more than 10-fold increase in nitrogen mineralization within a few decades. These changes in soil features have important consequences for plant growth and the competition between plant species. During succession in heathlands an increase in nutrient mineralization leads to species with low maximum growth rates and low biomass loss rates being replaced by species with high potential growth rates and high biomass losses. The plant properties responsible for reduced biomass loss rates appear to result in the litter produced being poorly decomposable, whereas the litter from plants with high potential growth rates decomposes more easily. Model simulations suggest that such combinations of plant features greatly influence the increase in mineralization and the change in plant species composition during ecosystem development. Studies in the field and garden plot experiments confirmed this hypothesis.     

Carbon and oxygen isotope composition of organic compounds in the phloem sap provides a short-term measure for stomatal conductance of European beech (Fagus sylvatica L.)

At eight different dates during the 2000 growing season, δ¹³C and δ¹⁸O were determined in the phloem of adult beech trees growing in natural beech stands in south‐west Germany differing in stand density and local climate. In addition, stand transpiration, precipitation, photosynthetic active radiation, relative air humidity, water pressure deficit of the air, air and soil temperature, soil water potential, and sugar concentration of the phloem sap were determined directly and evaporation and canopy stomatal conductance were modelled. All parameters were related to δ¹³C. The study aimed to identify the time integral within which the δ¹³C of organic compounds transported in the phloem is an indicative measure of these environmental influences. δ¹³C of soluble carbon transported in the phloem was well correlated with mean stomatal conductance in a two‐day integral prior to phloem sampling but did not depend on either light intensity or soil water availability. A strong positive relationship between δ¹³C and δ¹⁸O pointed to observed variation in δ¹³C of phloem sap being a result of variation in stomatal conductance. Bulk leaf δ¹³C was a poor indicator of changes in environmental conditions during the growing season. From these results we conclude that the analysis of δ¹³C in soluble carbon transported in the phloem is a reliable indicator of short‐term changes in C_i/C_a. In contrast, the δ¹³C of structural carbon in beech foliage represents an integration of a range of factors that mask short‐term influences responsible for C_i/C_a.     

Re-evaluation of proposed C4 photosynthetic characteristics in the genus Larix

It has been suggested previously that Japanese larch ( Larix kaempferi ) exhibits characteristics of C₄ photosynthesis. To further evaluate this suggestion, stable carbon isotope ratios were determined for leaf and bark tissue of Larix gmelini, L. kaempferi, L. laricina, L. Iyallii, L. occidentalis , and L. sibirica. All δ¹³C values were more negative than –22‰. Short-term labeling with ¹⁴CO₂ showed that phosphoglyceric acid and other phosphorylated compounds were the first products of photosynthesis in L. sibirica. Both of these results strongly suggest that the initial fixation of atmospheric CO₂ in these six Larix species is accomplished solely via the C₃ photosynthetic pathway.     

Carbon dioxide fixation in orchid aerial roots

Acidity fluctuation, CO₂ gas exchange, δ¹³C value, PEP carboxylase and RuBP carboxylase activities in aerial roots of selected thick-leaved orchid hybrids ( Arachnis and Aranthera ) were studied. Both aerial roots and leaves showed acidity fluctuation over a 24 h period. Dark acidification in aerial roots was enhanced at low temperature (15°C). Aerial roots had δ¹³C values close to those of leaves which have been previously demonstrated to possess crassulacean acid metabolism. Variation in δ¹³C values along the length of the roots was observed; the root tip having a less negative δ¹³C value (—13.34%‰) than the older portions of the roots (—14.55%‰). There was no net CO₂ fixation by aerial root, although ¹⁴³²CO₂ fixation was observed in light and in darkness. The pattern of fluctuation in activities of PEP carboxylase and RuBP carboxylase in aerial roots was similar to that obtained for the leaves. In both aerial roots and leaves, PEP carboxylase activity was several times higher than that of RuBP carboxylase.     

From forest to cropland and pasture systems: a critical review of soil organic carbon stocks changes in Amazonia

The impact of deforestation on soil organic carbon (SOC) stocks is important in the context of climate change and agricultural soil use. Trends of SOC stock changes after agroecosystem establishment vary according to the spatial scale considered, and factors explaining these trends may differ sometimes according to meta‐analyses. We have reviewed the knowledge about changes in SOC stocks in Amazonia after the establishment of pasture or cropland, sought relationships between observed changes and soil, climatic variables and management practices, and synthesized the δ¹³C measured in pastures. Our dataset consisted of 21 studies mostly synchronic, across 52 sites (Brazil, Colombia, French Guiana, Suriname), totalling 70 forest–agroecosystem comparisons. We found that pastures (n = 52, mean age = 17.6 years) had slightly higher SOC stocks than forest (+6.8 ± 3.1 %), whereas croplands (n = 18, mean age = 8.7 years) had lower SOC stocks than forest (?8.5 ± 2.9 %). Annual precipitation and SOC stocks under forest had no effect on the SOC changes in the agroecosystems. For croplands, we found a lower SOC loss than other meta‐analyses, but the short time period after deforestation here could have reduced this loss. There was no clear effect of tillage on the SOC response. Management of pastures, whether they were degraded/nominal/improved, had no significant effect on SOC response. δ¹³C measurements on 16 pasture chronosequences showed that decay of forest‐derived SOC was variable, whereas pasture‐derived SOC was less so and was characterized by an accumulation plateau of 20 Mg SOC ha^?1 after 20 years. The large uncertainties in SOC response observed could be derived from the chronosequence approach, sensitive to natural soil variability and to human management practices. This study emphasizes the need for diachronic and long‐term studies, associated with better knowledge of agroecosystem management.     

Natural 15N abundance of epiphytes depends on the position within the forest canopy: source signals and isotope fractionation

The natural ¹⁵N abundance (δ¹⁵N) of epiphytes and its N sources were studied in the canopy of a lowland rainforest in Costa Rica. Vascular and non‐vascular epiphytes and canopy soils were collected from four canopy zones and analysed for N contents and δ¹⁵N signals. In addition, the N concentrations and δ¹⁵N signatures of bulk precipitation, throughfall and stemflow were measured during the wet and the dry season. The δ¹⁵N values of epiphyte leaves decreased significantly from the lower zones (means of ?3·9 and ?4·3‰) to the upper zones (means of ?5·4 and ?6·1‰) of the canopy. In contrast, δ¹⁵N signatures of canopy soils (average ?0·3‰) differed little between the zones. Bulk deposition was enriched in ¹⁵N (+4·3‰) compared to all other potential N sources and was higher than throughfall and stemflow (+0·5 to ?1·3‰). δ¹⁵N values of atmospheric deposition were inversely related to those of the epiphyte leaves, whereas N isotopic composition of canopy soils did not vary significantly. Consequently, it is concluded that the variations in foliar N isotope composition of epiphytes were not simply caused by utilization of isotopically different N sources, but by different ¹⁵N discrimination during N acquisition.     

土地利用变化对草原生态系统土壤碳贮量的影响

本文概述了长期非持续性土地利用（草原开垦和过度放牧）对草原生态系统土壤碳贮量影响的有关研究成果和进展,指出了今后这方面研究的重点内容和对策。     

蒙古栎叶片及其土壤碳、氮同位素自然丰度对大气CO2浓度升高的响应

大气CO₂浓度升高对土壤氮素转化过程产生重要影响,研究其变化有助于更好地预测陆地生态系统的固碳潜力.氮同位素自然丰度作为生态系统氮素循环过程的综合指标能够有效地指示CO₂浓度升高对土壤氮素转化过程的影响.本研究采用开顶箱CO₂ 熏蒸法研究连续10年的大气CO₂ 浓度升高对我国东北地区蒙古栎及其土壤和微生物生物量碳、氮同位素自然丰度的影响.结果表明: 大气CO₂浓度升高改变了土壤氮循环过程,增加了土壤微生物和植物叶片δ¹⁵N;促进了富¹³C土壤有机碳分解,中和了贫¹³C植物光合碳输入的效果,导致土壤可溶性有机碳和微生物碳δ¹³C在CO₂升高条件下没有发生显著变化.这些结果表明,CO₂浓度升高很可能促进了土壤有机质矿化过程,并加剧了系统氮限制的状态.     

Effects of white clover (Trifolium repens L.) on plant and soil nitrogen and soil organic matter in mixtures with perennial ryegrass (Lolium perenne L.)

To increase our insight into the above- and belowground N flows in grass and grass-clover swards relations between crop and soil parameters were studied in a cutting trial with perennial ryegrass (Lolium perenne) monocultures and ryegrass–white clover (Trifolium repens) mixtures. The effects of clover cultivar on herbage yield, the amount of clover-derived nitrogen, apparent N transfer to companion grass, dynamics of N and organic matter in the soil were estimated.The grass monocultures had very low DM yields (<2.1 t ha^-1) and a low N concentration in the harvested herbage. During 1992–1995 the annual herbage DM yield in the mixtures ranged from 7.0 to 14.3 t ha^-1, the white clover DM yield from 2.4 to 11.2 t ha^-1 and the mean annual clover content in the herbage DM harvested from 34 to 78%. Mixtures with the large-leaved clover cv. Alice yielded significantly more herbage and clover DM and had a higher clover content than mixtures with small/medium-leaved cvs. Gwenda and Retor. Grass cultivar did not consistently affect yield, botanical composition or soil characteristics.The apparent N₂ fixation was very high, ranging from 150 to 545 kg N ha^-1 in the different mixtures. For each tonne of clover DM in the harvested herbage 49 to 63 kg N was harvested, while the apparent N transfer from clover to grass varied between 55 and 113 kg N ha^-1 year^-1.The net N mineralization rate was lower under monocultures than under mixtures. The C mineralization and the amounts of C and N in active soil organic matter fractions were similar for monocultures and mixtures, but the C:N ratio of the active soil organic matter fractions were higher under grass than under mixtures. This explains the lower N mineralization under grass.     

土壤水分和氮添加对华北平原高产农田有机碳矿化的影响

通过105 d的恒温(25℃)控湿室内培养方法,探讨了华北平原高产粮田土壤有机碳矿化特征以及水分和有机、无机氮输入对其影响。试验设4个肥料添加水平和4个水分梯度,分别为对照(S0)、仅添加无机氮(尿素)(S1)、无机氮和有机氮(鸡粪)配施(S2)以及仅添加有机氮(S3)和25%(田间持水量;M0)、50%(M1)、75%(M2)和100%(M3)共16个处理,每处理3次重复。结果表明,各处理有机碳矿化速率均在培养后1 d达第1高峰,之后直线下降,培养7 d时下降幅度达57.2%—75.0%,培养20—30 d时出现第2高峰。有机碳累积矿化量有208.8—1161 mg/kg,主要集中在前30 d,可占整个培养期的59.1%—69.9%,105 d的净矿化率为0.07%—2.01%。根据双指数方程模拟结果,研究了土壤潜在矿化碳库(C1+C2),其中活性碳库(C1)和惰性碳库(C2)分别为53.0—135.1 mg/kg和156.9—1069 mg/kg,潜在矿化率为1.75%—9.66%。土壤含水量显著影响有机碳矿化,且与潜在矿化碳库呈二次函数关系(P0.05)。田间持水量25%—100%范围内,随着土壤含水量的升高,有机碳矿化速率呈增加趋势,但增幅降低,其中M2(田间持水量75%)的有机碳净矿化率最高。有机碳矿化量与土壤微生物碳和矿质氮含量呈线性正相关(P0.05),保持氮水平(200 kg N/hm2)相同,有机氮(鸡粪)和无机氮(尿素)均显著促进土壤有机碳矿化,但两者间差异不显著(P0.05),且有机氮和无机氮对有机碳矿化的影响均与土壤含水量有显著交互作用(P0.05)。