Ecoenzymatic stoichiometry of microbial nutrient acquisition in tropical soils

The relative activities of soil enzymes involved in mineralizing organic carbon (C), nitrogen (N), and phosphorus (P) reveal stoichiometric and energetic constraints on microbial biomass growth. Although tropical forests and grasslands are a major component of the global C cycle, the effects of soil nutrient availability on microbial activity and C dynamics in these ecosystems are poorly understood. To explore potential microbial nutrient limitation in relation to enzyme allocation in low latitude ecosystems, we performed a meta-analysis of acid/alkaline phosphatase (AP), β-1,4-glucosidase (BG), and β-1,4-N-acetyl-glucosaminidase (NAG) activities in tropical soils. We found that BG:AP and NAG:AP ratios in tropical soils are significantly lower than those of temperate ecosystems overall. The lowest BG:AP and NAG:AP ratios were associated with old or acid soils, consistent with greater biological phosphorus demand relative to P availability. Additionally, correlations between enzyme activities and mean annual temperature and precipitation suggest some climatic regulation of microbial enzyme allocation in tropical soils. We used the results of our analysis in conjunction with previously published data on soil and biomass C:N:P stoichiometry to parameterize a biogeochemical equilibrium model that relates microbial growth efficiency to extracellular enzyme activity. The model predicts low microbial growth efficiencies in P-limited soils, indicating that P availability may influence C cycling in the highly weathered soils that underlie many tropical ecosystems. Therefore, we suggest that P availability be included in models that simulate microbial enzyme allocation, biomass growth, and C mineralization.     

Above-ground biomass and structure of 260 African tropical forests

We report above-ground biomass (AGB), basal area, stem density and wood mass density estimates from 260 sample plots (mean size: 1.2 ha) in intact closed-canopy tropical forests across 12 African countries. Mean AGB is 395.7 Mg dry mass ha⁻¹ (95% CI: 14.3), substantially higher than Amazonian values, with the Congo Basin and contiguous forest region attaining AGB values (429 Mg ha⁻¹) similar to those of Bornean forests, and significantly greater than East or West African forests. AGB therefore appears generally higher in palaeo- compared with neotropical forests. However, mean stem density is low (426 ± 11 stems ha⁻¹ greater than or equal to 100 mm diameter) compared with both Amazonian and Bornean forests (cf. approx. 600) and is the signature structural feature of African tropical forests. While spatial autocorrelation complicates analyses, AGB shows a positive relationship with rainfall in the driest nine months of the year, and an opposite association with the wettest three months of the year; a negative relationship with temperature; positive relationship with clay-rich soils; and negative relationships with C : N ratio (suggesting a positive soil phosphorus–AGB relationship), and soil fertility computed as the sum of base cations. The results indicate that AGB is mediated by both climate and soils, and suggest that the AGB of African closed-canopy tropical forests may be particularly sensitive to future precipitation and temperature changes.     

不同林龄胡杨活立木枯枝生物量和化学计量特征

基于不同林龄胡杨天然林样地活立木枯枝(DBST)生物量调查,探讨不同发育阶段胡杨活立木枯枝化学计量和碳储量的时空变化规律。研究结果表明:(1)不同林龄胡杨林活立木枯枝生物量随着林龄增加呈先增加后减小趋势,其中成熟林活立木枯枝生物量最大(10.93 t/hm~2),占地上生物量的17.79%。对于单株胡杨而言,随林龄增加活立木枯枝生物量增大,年平均活立木枯枝生物量为0.22 kg/株。同时胡杨枝径级越大越易形成枯枝,生物量也随枯枝径级的增加而增加。(2)胡杨活立木枯枝C、N、P、K平均含量分别为491.01、4.13、2.75、1.83 g/kg,活立木枯枝C、N、P元素含量高于活体枝,而活体枝K元素含量高于活立木枯枝。胡杨群落活立木枯枝平均C∶N、C∶P、N∶P比分别为:128.58、232.79、1.95,其活立木枯枝化学计量比均高于活体枝。林龄对化学元素含量和化学计量比影响均差异不显著(P0.05),说明了胡杨活立木枯枝的元素含量随龄级变化不大,具有稳定的特点。(3)活立木枯枝碳密度随林龄的增加先增加后降低,其中成熟林碳密度最大;胡杨群落平均碳密度为2.63 t/hm~2,占地上活立木碳密度的20.74%。研究认为胡杨群落枯枝生物量和碳密度随林龄增加的变化规律明显,胡杨活立木枯枝在荒漠森林生态系统中是不可忽略的C库。     

Sediment nutrient accumulation and nutrient availability in two tidal freshwater marshes along the Mattaponi River, Virginia, USA

Sediment deposition is the main mechanism of nutrient delivery to tidal freshwater marshes (TFMs). We quantified sediment nutrient accumulation in TFMs upstream and downstream of a proposed water withdrawal project on the Mattaponi River, Virginia. Our goal was to assess nutrient availability by comparing relative rates of carbon (C), nitrogen (N), and phosphorus (P) accumulated in sediments with the C, N, and P stoichiometries of surface soils and above ground plant tissues. Surface soil nutrient contents (0.60–0.92% N and 0.09–0.13% P) were low but within reported ranges for TFMs in the eastern US. In both marshes, soil nutrient pools and C, N, and P stoichiometries were closely associated with sedimentation patterns. Differences between marshes were more striking than spatial variations within marshes: both C, N, and P accumulation during summer, and annual P accumulation rates (0.16 and 0.04 g P m^–2 year^–1, respectively) in sediments were significantly higher at the downstream than at the upstream marsh. Nitrogen:P ratios <14 in above ground biomass, surface soils, and sediments suggest that N limits primary production in these marshes, but experimental additions of N and/or P did not significantly increase above ground productivity in either marsh. Lower soil N:P ratios are consistent with higher rates of sediment P accumulation at the downstream site, perhaps due to its greater proximity to the estuarine turbidity maximum.     

C:N:P stoichiometry in soil: is there a “Redfield ratio” for the microbial biomass?

Well-constrained carbon:nitrogen:phosphorus (C:N:P) ratios in planktonic biomass, and their importance in advancing our understanding of biological processes and nutrient cycling in marine ecosystems, has motivated ecologists to search for similar patterns in terrestrial ecosystems. Recent analyses indicate the existence of “Redfield-like” ratios in plants, and such data may provide insight into the nature of nutrient limitation in terrestrial ecosystems. We searched for analogous patterns in the soil and the soil microbial biomass by conducting a review of the literature. Although soil is characterized by high biological diversity, structural complexity and spatial heterogeneity, we found remarkably consistent C:N:P ratios in both total soil pools and the soil microbial biomass. Our analysis indicates that, similar to marine phytoplankton, element concentrations of individual phylogenetic groups within the soil microbial community may vary, but on average, atomic C:N:P ratios in both the soil (186:13:1) and the soil microbial biomass (60:7:1) are well-constrained at the global scale. We did see significant variation in soil and microbial element ratios between vegetation types (i.e., forest versus grassland), but in most cases, the similarities in soil and microbial element ratios among sites and across large scales were more apparent than the differences. Consistent microbial biomass element ratios, combined with data linking specific patterns of microbial element stoichiometry with direct evidence of microbial nutrient limitation, suggest that measuring the proportions of C, N and P in the microbial biomass may represent another useful tool for assessing nutrient limitation of ecosystem processes in terrestrial ecosystems.     

C and N storage in living trees within Finland since 1950s

Living biomass contains 45 to 60% carbon and 0.05 to 3% nitrogen, in dry weight. Like throughout Europe, the amount of living biomass in Finnish forests has increased on average over the last decades, largely because of changes in forest management. The storage of organic C and N in biomass has also increased.Changes in biomass vary between regions. Data were analysed on changes in the last 30–40 years in C and N storage in living trees in Finland, subdivided into 20 regions. Tree biomass increased in 17 regions, and decreased in 3 regions. The storage rate varied between -170 and +480 kg C ha^-1 a^-1, and between –0.5 and +1.2 kg N ha^-1 a^-1.Nitrogen accumulation in trees was less than 15% of atmospheric N deposition in all regions. Although the eventual increase of the nitrogen concentration in tree tissues was omitted, it is not possible that living biomass has been the major sink for atmospheric N deposition to forests. A hypothesis is presented that the main sink is litter layer and organic soil. Carbon can also be accumulating in soils essentially faster than hitherto estimated in analyses of carbon budgets of European forests.Died on September 2, 1994.     

The carbon budget of the Spanish forests

A model for the calculation of anthropogenic CO₂ emissions from perturbed forests (extraordinary emissions) is described. Timber production as well as wildfire statistics are used, and relevant physicochemical parameters are derived from the literature, to calculate the annual amounts of perturbed biomass and extraordinary emissions to the atmosphere from the Spanish forests — including soils — in the years 1960–1990; these emissions increased from 5.3 10⁶ t (metric tons) of carbon in 1970 to 10.6 10⁶ t C in 1990. A sensitivity analysis of the results has been performed to identify the most critical parameters. Contributions of observed timber growth and natural vegetal detritus and soil organic matter to the net forest carbon flux have been estimated to calculate the carbon budget in the Spanish forests between 1966 and 1974, which represents a net gain of carbon, ranging from 9.2 to 18 millions t. Finally, the methods used to calculate biospheric carbon balance and their results are compared and discussed.     

Changes in carbon, nutrients and stoichiometric relations under different soil depths, plant tissues and ages in black locust plantations

To investigate influences of forest plantations on soil nutrient properties, biomass accumulation, major nutrient elements (NPK) and their stoichiometric couplings in different tissues and aged plants, and correlations between major nutrient contents in soils and in foliage of plants, 5-, 10-, 15- and 20-year-old plantations of black locust (Robinia pseudoacacia L.) and farmland were selected. Black locust plantations increased soil organic carbon (SOC) and N stocks by 23–327 and 23–119 %, respectively, in the 0–10 cm top soil layer compared to those in farmland. Soil C:N, C:P, C:K, N:P, N:K and P:K ratios were 10.1, 22.9, 0.7, 2.2, 0.7 and 0.03, respectively. These ratios were higher in the 0–10 cm soil layer than those in the 10–20 cm soil layer and increased under older plantations. Higher C contents in stem, N contents in leaf, the largest C pools in stem and N pools in root in 20-year-old plantation were observed. Correspondingly, the highest C:N, C:P and C:K and the lowest N:P and N:K ratios in stem, decreased C:N and C:P ratios in older trees were found. No strong correlations were observed between element contents in soils and in leaves of black locust trees. These results suggest that black locust plantations can increase soil nutrient concentrations, SOC and N stocks resulting in changes in element stoichiometric relations. CNPK contents and their stoichiometries vary with tissues and tree ages of black locust. No strong coupling relations exist between major nutrient element contents in the top soil and in foliage of black locust.     

Variation in Carbon Storage and Its Distribution by Stand Age and Forest Type in Boreal and Temperate Forests in Northeastern China

The northeastern forest region of China is an important component of total temperate and boreal forests in the northern hemisphere. But how carbon (C) pool size and distribution varies among tree, understory, forest floor and soil components, and across stand ages remains unclear. To address this knowledge gap, we selected three major temperate and two major boreal forest types in northeastern (NE) China. Within both forest zones, we focused on four stand age classes (young, mid-aged, mature and over-mature). Results showed that total C storage was greater in temperate than in boreal forests, and greater in older than in younger stands. Tree biomass C was the main C component, and its contribution to the total forest C storage increased with increasing stand age. It ranged from 27.7% in young to 62.8% in over-mature stands in boreal forests and from 26.5% in young to 72.8% in over-mature stands in temperate forests. Results from both forest zones thus confirm the large biomass C storage capacity of old-growth forests. Tree biomass C was influenced by forest zone, stand age, and forest type. Soil C contribution to total forest C storage ranged from 62.5% in young to 30.1% in over-mature stands in boreal and from 70.1% in young to 26.0% in over-mature in temperate forests. Thus soil C storage is a major C pool in forests of NE China. On the other hand, understory and forest floor C jointly contained less than 13% and <5%, in boreal and temperate forests respectively, and thus play a minor role in total forest C storage in NE China.     

Rooting depth,water availability,and vegetation cover along an aridity gradient in Patagonia

Above-and belowground biomass distribution, isotopic composition of soil and xylem water, and carbon isotope ratios were studied along an aridity gradient in Patagonia (44–45°S). Sites, ranging from those with Nothofagus forest with high annual rainfall (770 mm) to Nothofagus scrub (520 mm), Festuca (290 mm) and Stipa (160 mm) grasslands and into desert vegetation (125 mm), were chosen to test whether rooting depth compensates for low rainfall. Along this gradient, both mean above-and belowground biomass and leaf area index decreased, but average carbon isotope ratios of sun leaves remained constant (at-27), indicating no major differences in the ratio of assimilation to stomatal conductance at the time of leaf growth. The depth of the soil horizon that contained 90% of the root biomass was similar for forests and grasslands (about 0.80–0.50 m), but was shallower in the desert (0.30 m). In all habitats, roots reached water-saturated soils or ground water at 2–3 m depth. The depth profile of oxygen and hydrogen isotope ratios of soil water corresponded inversely to volumetric soil water contents and showed distinct patterns throughout the soil profile due to evaporation, water uptake and rainfall events of the past year. The isotope ratios of soil water indicated that high soil moisture at 2–3 m soil depth had originated from rainy periods earlier in the season or even from past rainy seasons. Hydrogen and oxygen isotope ratios of xylem water revealed that all plants used water from recent rain events in the topsoil and not from water-saturated soils at greater depth. However, this study cannot explain the vegetation zonation along the transect on the basis of water supply to the existing plant cover. Although water was accessible to roots in deeper soil layers in all habitats, as demonstrated by high soil moisture, earlier rain events were not fully utilized by the current plant cover during summer drought. The role of seedling establishment in determining species composition and vegetation type, and the indirect effect of seedling establishment on the use of water by fully developed plant cover, are discussed in relation to climate change and vegetation modelling.     

C:N:P stoichiometry in Australian soils with respect to vegetation and environmental factors

Aims

We estimate organic carbon (C): total nitrogen (N): total phosphorus (P) ratios in soils under Australia’s major native vegetation groups.

Methods

We use digital datasets for climate, soils, and vegetation created for the National Land and Water Resources Audit in 2001. Analysis-of-variance is used to investigate differences in nutrient ratios between ecosystems. Linear discriminant analysis and logistic regression are used to investigate the relative importance of climatic variables and soil nutrients in vegetation patterns.

Results

We find that the N:P and C:P ratios have a greater range of values than the C:N ratio, although major vegetation groups tend to show similar trends across all three ratios. Some apparently homeostatic groupings emerge: those with very low, low, medium, or high N:P and C:P. Tussock grasslands have very low soil N, N:P, and C:P, probably due to frequent burning. Eucalypt woodlands have low soil N:P and C:P ratios, although their total P level varies. Rainforests and Melaleuca forests have medium soil N:P and C:P ratios, although their total P level is different. Heathlands, tall open eucalypt forests, and shrublands occur on soils with low levels of total P, and high N:P and C:P ratios that reflect foliar nutrient ratios and recalcitrant litter.

Conclusions

Certain plant communities have typical soil nutrient stoichiometries but there is no single Redfield-like ratio. Vegetation patterns largely reflect soil moisture but for several plant communities, eucalypt communities in particular, soil N and P (or N:P) also play a significant role. Soil N:P and the presence of Proteaceae appear indicative of nutrient constraints in ecosystems.     

Not all forests are expanding over central Brazilian savannas

Recently we reported on the expansion of riparian forests into savannas in central Brazil. To enlarge the scope of the earlier study we investigated whether upland deciduous and xeromorphic forests behaved similarly. We investigated past vegetation changes that occurred in forest/savanna transitions using carbon isotope ratios (δ¹³C) measured in the soil organic matter as a tracer. We analyzed the ¹⁴C activity where δ¹³C showed major shifts in vegetation. The role of soil chemical and physical attributes in defining vegetation distribution is discussed. Structural changes in vegetation were found to be associated with shifts in the isotope composition (δ¹³C) of soil organic matter. This was attributed to intrinsic differences in the biomass of trees and grasses and allowed for the determination of past shifts in vegetation by evaluating δ¹³C at different depths. The deciduous forest decreased in area approximately 980 years ago. Tree cover increased in the xeromorphic forest, but the border stayed stable through time. The deciduous forest and adjacent savanna have eutrophic soils while the xeromorphic forest and adjacent savanna have dystrophic soils. However, greater organic carbon, nitrogen and phosphorus concentrations are observed in the forests. We provide concrete evidence of deciduous forest retreat unlike the stability observed in the xeromorphic forest/savanna boundary. These results contrast with the expansion of riparian forests recently reported in the same region.     

Root biomass allocation in the world's upland forests

Because the world's forests play a major role in regulating nutrient and carbon cycles, there is much interest in estimating their biomass. Estimates of aboveground biomass based on well-established methods are relatively abundant; estimates of root biomass based on standard methods are much less common. The goal of this work was to determine if a reliable method to estimate root biomass density for forests could be developed based on existing data from the literature. The forestry literature containing root biomass measurements was reviewed and summarized and relationships between both root biomass density (Mg ha⁻¹) and root:shoot ratios (R/S) as dependent variables and various edaphic and climatic independent variables, singly and in combination, were statistically tested. None of the tested independent variables of aboveground biomass density, latitude, temperature, precipitation, temperature:precipitation ratios, tree type, soil texture, and age had important explanatory value for R/S. However, linear regression analysis showed that aboveground biomass density, age, and latitudinal category were the most important predictors of root biomass density, and together explained 84% of the variation. A comparison of root biomass density estimates based on our equations with those based on use of generalized R/S ratios for forests in the United States indicated that our method tended to produce estimates that were about 20% higher. Received: 3 July 1996 / Accepted: 23 January 1997     

Incorporation of Plant Residues into Soil Organic Matter Fractions With Grassland Management Practices in the North American Midwest

Disturbed grassland soils are often cited as having the potential to store large amounts of carbon (C). Fertilization of grasslands can promote soil C storage, but little is known about the generation of recalcitrant pools of soil organic matter (SOM) with management treatments, which is critical for long-term soil C storage. We used a combination of soil incubations, size fractionation and acid hydrolysis of SOM, [C], [N], and stable isotopic analyses, and biomass quality indices to examine how fertilization and haying can impact SOM dynamics in Kansan grassland soils. Fertilized soils possessed 113% of the C possessed by soils subjected to other treatments, an increase predominantly harbored in the largest size fraction (212–2,000 μm). This fraction is frequently associated with more labile material. Haying and fertilization/haying, treatments that more accurately mimic true management techniques, did not induce any increase in soil C. The difference in ¹⁵N-enrichment between size fractions was consistent with a decoupling of SOM processing between pools with fertilization, congruent with gains of SOM in the largest size fraction promoted by fertilization not moving readily into smaller fractions that frequently harbor more recalcitrant material. Litterfall and root biomass C inputs increased 104% with fertilization over control plots, and this material possessed lower C:N ratios. Models of incubation mineralization kinetics indicate that fertilized soils have larger pools of labile organic C. Model estimates of turnover rates of the labile and recalcitrant C pools did not differ between treatments (65.5 ± 7.2 and 2.9 ± 0.3 μg C d⁻¹, respectively). Although fertilization may promote greater organic inputs into these soils, much of that material is transformed into relatively labile forms of soil C; these data highlight the challenges of managing grasslands for long-term soil C sequestration.     

甘肃省森林碳储量现状与固碳速率

针对森林碳平衡再评估的重要性和区域尺度森林生态系统碳库量化分配的不确定性, 该研究依据全国森林资源连续清查结果中甘肃省各森林类型分布的面积与蓄积比重以及林龄和起源等要素, 在甘肃省布设212个样地, 经野外调查与采样、室内分析, 并对典型样地信息按照面积权重进行尺度扩展, 估算了甘肃省森林生态系统碳储量及其分布特征。结果表明: 甘肃省森林生态系统总碳储量为612.43 Tg C, 其中植被生物量碳为179.04 Tg C, 土壤碳为433.39 Tg C。天然林是甘肃省碳储量的主要贡献者, 其值为501.42 Tg C, 是人工林的4.52倍。天然林和人工林的植被碳密度均表现为随林龄的增加而增加的趋势, 同一龄组天然林植被碳密度高于人工林。天然林土壤碳密度从幼龄林到过熟林逐渐增加, 但人工林土壤碳密度最大值主要为近熟林。全省森林植被碳密度均值为72.43 Mg C·hm^-2, 天然林和人工林分别为90.52和33.79 Mg C·hm^-2。基于森林清查资料和标准样地实测数据, 估算出全省天然林和人工林在1996年的植被碳储量为132.47和12.81 Tg C, 2011年分别为152.41和26.63 Tg C, 平均固碳速率分别为1.33和0.92 Tg C·a^-1。甘肃省幼、中龄林面积比重较大, 占全省的62.28%, 根据碳密度随林龄的动态变化特征, 预测这些低龄林将发挥巨大的碳汇潜力。     

Succession May Maintain High Leaf Area: Sapwood Ratios and Productivity in OldSubalpine Forests

Old forests are generally believed to exhibit low net primary productivity (NPP) and therefore to be insignificant carbon sinks. This relationship between age and NPP is based, in part, on the hypothesis that the biomass of respiratory tissues such as sapwood increases with age to a point where all photosynthate is required just to maintain existing tissue. However, this theoretical connection between respiration:assimilation ratios and forest productivity is based on age-dependent trends in the sapwood:leaf ratios of individual trees and even-aged stands; it does not take into account such processes in natural forests as disproportional increases in shade-tolerant species over time and multiple-age cohorts. Ignoring succession and structural complexity may lead to large underestimates of the productivity of old forests and inaccurate estimates of the ages at which forest productivity declines. To address this problem, we compared biomass allocation and productivity between whitebark pine, a shade-intolerant, early-successional tree species, and subalpine fir, a shade-tolerant, late-successional species, by harvesting 14 whitebark pines and nine subalpine firs that varied widely in dbh and calculating regression models for dbh vs annual productivity and biomass allocation to leaves, sapwood, and heartwood. Late-successional subalpine fir allocated almost twice as much biomass to leaves as early-successional whitebark pine. Subalpine firs also had a much lower allocation to sapwood and higher growth rates across all tree sizes. We then modeled biomass allocation and productivity for 12 natural stands in western Montana that were dominated by subalpine fir and whitebark pine varying in age from 67 to 458 years by applying the regressions to all trees in each stand. Whole-stand sapwood:leaf ratios and stand productivity increased asymptotically with age. Sapwood:leaf ratios and productivity of whitebark pine in these stands increased for approximately 200–300 years and then decreased slowly over the next 200 years. In contrast, sapwood:leaf ratios of all sizes of subalpine fir were lower than those of pine and productivity was higher. As stands shifted in dominance from pine to fir with age, subalpine fir appeared to maintain gradually increasing rates of whole-forest productivity until stands were approximately 400 years old. These results suggest that forests such as these may continue to sequester carbon for centuries. If shade-tolerant species that predominate late in succession maintain high assimilation-to-respiration ratios in other forests, we may be underestimating production in old forests, and current models may underestimate the importance of mature forests as carbon sinks for atmospheric CO₂ in the global carbon cycle. Received 16 February 1999; accepted 24 November 1999.     

Nutrient Limitation on Ecosystem Productivity and Processes of Mature and Old-Growth Subtropical Forests in China

Nitrogen (N) is considered the dominant limiting nutrient in temperate regions, while phosphorus (P) limitation frequently occurs in tropical regions, but in subtropical regions nutrient limitation is poorly understood. In this study, we investigated N and P contents and N:P ratios of foliage, forest floors, fine roots and mineral soils, and their relationships with community biomass, litterfall C, N and P productions, forest floor turnover rate, and microbial processes in eight mature and old-growth subtropical forests (stand age >80 yr) at Dinghushan Biosphere Reserve, China. Average N:P ratios (mass based) in foliage, litter (L) layer and mixture of fermentation and humus (F/H) layer, and fine roots were 28.3, 42.3, 32.0 and 32.7, respectively. These values are higher than the critical N:P ratios for P limitation proposed (16–20 for foliage, ca. 25 for forest floors). The markedly high N:P ratios were mainly attributed to the high N concentrations of these plant materials. Community biomass, litterfall C, N and P productions, forest floor turnover rate and microbial properties were more strongly related to measures of P than N and frequently negatively related to the N:P ratios, suggesting a significant role of P availability in determining ecosystem production and productivity and nutrient cycling at all the study sites except for one prescribed disturbed site where N availability may also be important. We propose that N enrichment is probably a significant driver of the potential P limitation in the study area. Low P parent material may also contribute to the potential P limitation. In general, our results provided strong evidence supporting a significant role for P availability, rather than N availability, in determining ecosystem primary productivity and ecosystem processes in subtropical forests of China.     

Differential Nutrient Limitation of Soil Microbial Biomass and Metabolic Quotients (qCO2): Is There a Biological Stoichiometry of Soil Microbes?

Background

Variation in microbial metabolism poses one of the greatest current uncertainties in models of global carbon cycling, and is particularly poorly understood in soils. Biological Stoichiometry theory describes biochemical mechanisms linking metabolic rates with variation in the elemental composition of cells and organisms, and has been widely observed in animals, plants, and plankton. However, this theory has not been widely tested in microbes, which are considered to have fixed ratios of major elements in soils.

Methodology/Principal Findings

To determine whether Biological Stoichiometry underlies patterns of soil microbial metabolism, we compiled published data on microbial biomass carbon (C), nitrogen (N), and phosphorus (P) pools in soils spanning the global range of climate, vegetation, and land use types. We compared element ratios in microbial biomass pools to the metabolic quotient qCO₂ (respiration per unit biomass), where soil C mineralization was simultaneously measured in controlled incubations. Although microbial C, N, and P stoichiometry appeared to follow somewhat constrained allometric relationships at the global scale, we found significant variation in the C∶N∶P ratios of soil microbes across land use and habitat types, and size-dependent scaling of microbial C∶N and C∶P (but not N∶P) ratios. Microbial stoichiometry and metabolic quotients were also weakly correlated as suggested by Biological Stoichiometry theory. Importantly, we found that while soil microbial biomass appeared constrained by soil N availability, microbial metabolic rates (qCO₂) were most strongly associated with inorganic P availability.

Conclusions/Significance

Our findings appear consistent with the model of cellular metabolism described by Biological Stoichiometry theory, where biomass is limited by N needed to build proteins, but rates of protein synthesis are limited by the high P demands of ribosomes. Incorporation of these physiological processes may improve models of carbon cycling and understanding of the effects of nutrient availability on soil C turnover across terrestrial and wetland habitats.     

Forest vegetation as related to climate and soil conditions at varying altitudes on a humid subtropical mountain, Mount Emei, Sichuan, China

Altitudinal variations in temperature and soils were analysed on a humid subtropical mountain, Mt Emei (3,099 m a.s.l., 29°34.5N, 103°21.5E), in Sichuan, China, to see how the vegetation varies with the environmental factors. As a principal finding, the coldest mean monthly temperature –1°C, rather than the warmth index of 85°C·months, emerged as the primary factor that delimited the evergreen broadleaved forest. With regard to soils, properties such as organic C, total N, available P, exchangeable K tended to increase with altitude. The highest values in organic C (26.6%), total N (1.34%) and available P (45.39 ppm) were recorded in surface soils of the mixed forest (2,210 m a.s.l.) including all three tree life forms, i.e. evergreen/deciduous broadleaved and coniferous trees. The high pH and contents of exchangeable Ca and Mg in the surface soils derived from the parent material, limestone and dolomite, between 900 and 1,200 m, where several Tertiary tree species existed. The C/N ratios of surface soils in the coniferous forests (2,500–3,099 m) were higher than those of the evergreen broadleaved forests (600–1,500 m) and the mixed forests (1,500–2,500 m).     

Patterns of Biomass and Carbon Distribution across a Chronosequence of Chinese Pine (Pinus tabulaeformis) Forests

Patterns of biomass and carbon (C) storage distribution across Chinese pine (Pinus tabulaeformis) natural secondary forests are poorly documented. The objectives of this study were to examine the biomass and C pools of the major ecosystem components in a replicated age sequence of P. tabulaeformis secondary forest stands in Northern China. Within each stand, biomass of above- and belowground tree, understory (shrub and herb), and forest floor were determined from plot-level investigation and destructive sampling. Allometric equations using the diameter at breast height (DBH) were developed to quantify plant biomass. C stocks in the tree and understory biomass, forest floor, and mineral soil (0–100 cm) were estimated by analyzing the C concentration of each component. The results showed that the tree biomass of P. tabulaeformis stands was ranged from 123.8 Mg·ha^–1 for the young stand to 344.8 Mg·ha^–1 for the mature stand. The understory biomass ranged from 1.8 Mg·ha^–1 in the middle-aged stand to 3.5 Mg·ha^–1 in the young stand. Forest floor biomass increased steady with stand age, ranging from 14.9 to 23.0 Mg·ha^–1. The highest mean C concentration across the chronosequence was found in tree branch while the lowest mean C concentration was found in forest floor. The observed C stock of the aboveground tree, shrub, forest floor, and mineral soil increased with increasing stand age, whereas the herb C stock showed a decreasing trend with a sigmoid pattern. The C stock of forest ecosystem in young, middle-aged, immature, and mature stands were 178.1, 236.3, 297.7, and 359.8 Mg C ha^–1, respectively, greater than those under similar aged P. tabulaeformis forests in China. These results are likely to be integrated into further forest management plans and generalized in other contexts to evaluate C stocks at the regional scale.