Soil organic matter, microbial biomass and enzyme activities in a tropical agroforestry system

 The effects of growing trees in combination with field crops on soil organic matter, microbial biomass C, basal respiration and dehydrogenase and alkaline phosphatase activities were studied in soils under a 12-year-old Dalbergia sissoo (a N₂-fixing tree) plantation intercropped with a wheat (Triticum aestivum) – cowpea (Vigna sinensis) cropping sequence. The inputs of organic matter through D. sissoo leaf litter increased and crop roots decreased with the increase in tree density. Higher organic C and total N, microbial biomass C, basal soil respiration and activities of dehydrogenase and alkaline phosphatase were observed in treatments with tree-crop combination than in the treatment without trees. Soil organic matter, microbial biomass C and soil enzyme activities increased with the decrease in the spacing of the D. sissoo plantation. The results indicate that adoption of the agroforestry practices led to an improved organic matter status of the soil, which is also reflected in the increased nutrient pool and microbial activities necessary for long-term productivity of the soil. However, tree spacing should be properly maintained to minimize the effects of shading on the intercrops. Received: 21 February 1997     

Effect of cattle slurry in grassland on microbial biomass and on activities of various enzymes

We examined the long-term effects of cattle slurry, applied at high rates, on microbial biomass, respiration, the microbial quotient (qCO₂) and various soil enzyme activities. In March, June, July, and October 1991, slurry-amended grassland soils (0–10 cm) contained significantly higher levels of microbial biomass, N mineralization and enzyme activities involved in N, P, and C cycling. With microbial biomass as the relative value, the results revealed that the slurry treatment influenced enzyme production by the microbial biomass. High levels of urease activity were the result not only of a larger microbial biomass, but also of higher levels of enzmye production by this microbial biomass. The ratio of alkaline phosphatase and xylanase to microbial biomass was nearly constant in the different treatments. The metabolic quotient (qCO₂) declined with increased levels of slurry application. Therefore it appears that microorganisms in slurry-amended soils require less C and energy if there is no competition for nutrients. The results of this study suggest that urease activity, nitrification, and respiration (metabolic quotient) can be used as indicators of environmental stress, produced by heavy applications of cattle slurry.     

Effects of pasture improvement and intensive cultivation on microbial biomass,enzyme activities,and composition and size of earthworm populations

We investigated the quantity and distribution of organic C, microbial biomass C, protease, arylsulphatase and arylphosphatase activity, and earthworm numbers and biomass in the soil from a 37-year-old grazed pasture supplied with superphosphate at rates of 0, 188, and 376 kg ha^-1 annually. The results were compared with a non-irrigated wilderness site which had not been used for agriculture and an arable site that had been intensively cultivated for 11 consecutive years. In the 0- to 5-cm layer, organic C followed the trend arableAporrectodea caliginosa (77–89% of total numbers) although Lumbricus rubellus made an increasing contribution to the population with increasing superphosphate rates. In the unirrigated wilderness site the population consisted of 56% A. caliginosa and 44% L. rubellus. While Octolasion cyaneum and A. rosea made up a small proportion of the population in the improved pasture sites, they were not present in the wilderness or arable sites. A. caliginosa was the only species present in the arable site. The mean fresh weight of individuals followed the order arable相似文献   

Organic matter, microbial biomass and enzyme activity of soils under different crop rotations in the tropics

Soil organic matter level, soil microbial biomass C, ninhydrin-N, C mineralization, and dehydrogenase and alkaline phosphatase activity were studied in soils under different crop rotations for 6 years. Inclusion of a green manure crop of Sesbania aculeata in the rotation improved soil organic matter status and led to an increase in soil microbial biomass, soil enzyme activity and soil respiratory activity. Microbial biomass C increased from 192 mg kg^–1 soil in a pearl millet-wheat-fallow rotation to 256 mg kg^–1 soil in a pearl millet-wheat-green manure rotation. Inclusion of an oilseed crop such as sunflower or mustard led to a decrease in soil microbial biomass, C mineralization and soil enzyme activity. There was a good correlation between microbial biomass C, ninhydrin-N and dehydrogenase activity. The alkaline phosphatase activity of the soil under different crop rotations was little affected. The results indicate the green manuring improved the organic matter status of the soil and soil microbial activity vital for the nutrient turnover and long-term productivity of the soil. Received: 7 January 1996     

Response of soil microbial biomass and enzyme activities to the transient elevation of carbon dioxide in a semi-arid grassland

Although elevation of CO₂ has been reported to impact soil microbial functions, little information is available on the spatial and temporal variation of this effect. The objective of this study was to determine the microbial response in a northern Colorado shortgrass steppe to a 5-year elevation of atmospheric CO₂ as well as the reversibility of the microbial response during a period of several months after shutting off the CO₂ amendment. The experiment was comprised of nine experimental plots: three chambered plots maintained at ambient CO₂ levels of 360 μmol mol⁻¹ (ambient treatment), three chambered plots maintained at 720 μmol mol⁻¹ CO₂ (elevated treatment) and three unchambered plots of equal ground area used as controls to monitor the chamber effect.Elevated CO₂ induced mainly an increase of enzyme activities (protease, xylanase, invertase, alkaline phosphatase, arylsulfatase) in the upper 5 cm of the soil and did not change microbial biomass in the soil profile. Since rhizodeposition and newly formed roots enlarged the pool of easily available substrates mainly in the upper soil layers, enzyme regulation (production and activity) rather than shifts in microbial abundance was the driving factor for higher enzyme activities in the upper soil. Repeated soil sampling during the third to fifth year of the experiment revealed an enhancement of enzyme activities which varied in the range of 20-80%. Discriminant analysis including all microbiological properties revealed that the enzyme pattern in 1999 and 2000 was dominated by the CO₂ and chamber effect, while in 2001 the influence of elevated CO₂ increased and the chamber effect decreased.Although microbial biomass did not show any response to elevated CO₂ during the main experiment, a significant increase of soil microbial N was detected as a post-treatment effect probably due to lower nutrient (nitrogen) competition between microorganisms and plants in this N-limited ecosystem. Whereas most enzyme activities showed a significant post-CO₂ effect in spring 2002 (following the conclusion of CO₂ enrichment the previous autumn, 2001), selective depletion of substrates is speculated to be the cause for non-significant treatment effects of most enzyme activities later in summer and autumn, 2002. Therefore, additional belowground carbon input mainly entered the fast cycling carbon pool and contributed little to long-term carbon storage in the semi-arid grassland.     

Effects of long-term waste water irrigation on soil organic matter, soil microbial biomass and its activities in central Mexico

 The effect of long-term waste water irrigation (up to 80 years) on soil organic matter, soil microbial biomass and its activities was studied in two agricultural soils (Vertisols and Leptosols) irrigated for 25, 65 and 80 years respectively at Irrigation District 03 in the Valley of Mezquital near Mexico City. In the Vertisols, where larger amounts of water have been applied than in the Leptosols, total organic C (TOC) contents increased 2.5-fold after 80 years of irrigation. In the Leptosols, however, the degradability of the organic matter tended to increase with irrigation time. It appears that soil organic matter accumulation was not due to pollutants nor did microbial biomass:TOC ratios and qCO₂ values indicate a pollutant effect. Increases in soil microbial biomass C and activities were presumably due to the larger application of organic matter. However, changes in soil microbial communities occurred, as denitrification capacities increased greatly and adenylate energy charge (AEC) ratios were reduced after long-term irrigation. These changes were supposed to be due to the addition of surfactants, especially alkylbenzene sulfonates (effect on denitrification capacity) and the addition of sodium and salts (effect on AEC) through waste water irrigation. Heavy metals contained in the sewage do not appear to be affecting soil processes yet, due to their low availability. Detrimental effects on soil microbial communities can be expected, however, from further increases in pollutant concentrations due to prolonged application of untreated waste water or an increase in mobility due to higher mineralization rates. Received: 28 April 1999     

Microbial biomass,activity, and organic matter accumulation in soils contaminated with heavy metals

Chemical characteristics and some parameters related to biological components were determined in 16 soils from a fairly homogeneous area in the north of Italy, contaminated with different levels of heavy metals. Correlation analysis of the parameters studied showed close positive relationships among the metals and with the organic C content in the soils studied. Negative relationships were observed among the heavy metals, soil respiration, and the ratio between evolved CO₂–C and microbial biomass C per unit time (specific respiratory activity). This was ascribed to an adverse heavy metal effect on the soil microflora, which appeared to increase the accumulation of organic matter as the heavy metal content increased, probably because the biomass was less effective in mineralising soil organic matter under these conditions.     

Characteristics of microbial biomass and soil organic matter in newly reclaimed wetland rice soils

Microbial biomass C, N, total organic C, N and mineralizable N were measured in newly reclaimed wetland sandy loam rice soil with a very low nutrient status. Microbial biomass C increased 5.4–10.4 times due to application of barnyard manure, but decreased drastically to 24–27% during rice cultivation. Organic C and N contents also decreased during cultivation, but to a lesser extent to 59–76%. At the tillering stage of the rice plant, microbial biomass N was highly correlated with mineralizable N (r=0.986).     

The combined effects of earthworms and arbuscular mycorrhizal fungi on microbial biomass and enzyme activities in a calcareous soil spiked with cadmium

Earthworms and arbuscular mycorrhizal fungi (AMF) are known to independently affect soil microbial and biochemical properties, in particular soil microbial biomass (SMB) and enzymes. However, less information is available about their interactive effects, particularly in soils contaminated with heavy metals such as cadmium (Cd). The amount of soil microbial biomass C (MBC), the rate of soil respiration (SRR) and the activities of urease and alkaline phosphatase (ALP) were measured in a calcareous soil artificially spiked with Cd (10 and 20 mg Cd kg⁻¹), inoculated with earthworm (Lumbricus rubellus L.), and AMF (Glomus intraradices and Glomus mosseae species) under maize (Zea mays L.) crop for 60 days. Results showed that the quantity of MBC, SRR and enzyme activities decreased with increasing Cd levels as a result of the elevated exchangeable Cd concentration. Earthworm addition increased soil exchangeable Cd levels, while AMF and their interaction with earthworms had no influence on this fraction of Cd. Earthworm activity resulted in no change in soil MBC, while inoculation with both AMF species significantly enhanced soil MBC contents. However, the presence of earthworms lowered soil MBC when inoculated with G. mosseae fungi, showing an interaction between the two organisms. Soil enzyme activities and SRR values tended to increase considerably with the inoculation of both earthworms and AMF. Nevertheless, earthworm activity did not affect ALP activity when inoculated with G. mosseae fungi, while the presence of earthworm enhanced urease activity only with G. intraradices species. The increases in enzyme activities and SRR were better ascribed to changes in soil organic carbon (OC), MBC and dissolved organic carbon (DOC) contents. In summary, results demonstrated that the influence of earthworms alone on Cd availability is more important than that of AMF in Cd-polluted soils; and that the interaction effects between these organisms on soil microorganism are much more important than on Cd availability. Thus, the presence of both earthworms and AMF could alleviate Cd effects on soil microbial life.     

Soil respiration is not limited by reductions in microbial biomass during long-term soil incubations

Declining rates of soil respiration are reliably observed during long-term laboratory incubations. However, the cause of this decline is uncertain. We explored different controls on soil respiration to elucidate the drivers of respiration rate declines during long-term soil incubations. Following a long-term (707 day) incubation (30 °C) of soils from two sites (a cultivated and a forested plot at Kellogg Biological Station, Hickory Corners, MI, USA), soils were significantly depleted of both soil carbon and microbial biomass. To test the ability of these carbon- and biomass-depleted (“incubation-depleted”) soils to respire labile organic matter, we exposed soils to a second, 42 day incubation (30 °C) with and without an addition of plant residues. We controlled for soil carbon and microbial biomass depletion by incubating field fresh (“fresh”) soils with and without an amendment of wheat and corn residues. Although respiration was consistently higher in the fresh versus incubation-depleted soil (2 and 1.2 times higher in the fresh cultivated and fresh forested soil, respectively), the ability to respire substrate did not differ between the fresh and incubation-depleted soils. Further, at the completion of the 42 day incubation, levels of microbial biomass in the incubation-depleted soils remained unchanged, while levels of microbial biomass in the field-fresh soil declined to levels similar to that of the incubation-depleted soils. Extra-cellular enzyme pools in the incubation-depleted soils were sometimes slightly reduced and did not respond to addition of labile substrate and did not limit soil respiration. Our results support the idea that available soil organic matter, rather than a lack microbial biomass and extracellular enzymes, limits soil respiration over the course of long-term incubations. That decomposition of both wheat and corn straw residues did not change after major changes in the soil biomass during extended incubation supports the omission of biomass values from biogeochemical models.     

Microbial biomass turnover and enzyme activities following the application of farmyard manure to field soils with and without previous long-term applications

Summary We studied the build-up and turnover of microbial biomass following the addition of farmyard manure to an unmanured soil and to soils from a long-term experiment in which different levels of farmyard manure had been applied for the last 23 years. The application of farmyard manure at 15–90 t ha^-1 to previously unmanured soil increased the microbial biomass during the first 3 months of incubation but a gradual decline occurred with further incubation for up to 12 months. Microbial biomass C was positively correlated with soil organic C and ranged from 1.8% to 2.2% of organic C after 12 months of farmyard manure applications. Biomass turnover increased with the application of farmyard manure, ranging from 0.81 to 0.87 year^-1 with various levels of manure. Amendment of soils from the long-term manure experiment with various levels of farmyard manure led to a build-up and decline in biomass C as seen in the unmanured soils, but biomass C was higher in all treatments compared to the corresponding unmanured soil treatments. Biomass turnover was greater compared to the unmanured soil treatments and it decreased with increasing levels of farmyard manure. The average soil respiratory activity increased with increasing levels of farmyard manure, but respiratory activity per unit of biomass C decreased with increasing levels of manure. Enzyme activities were greater in long-term manured soils compared to unmanured soils amended with various levels of manure. There was a significant correlation between biomass C and enzyme activities.     

Impact of agricultural practices on the size and activity of the microbial biomass in a long-term field experiment

The Dehérain long-term field experiment was initiated in 1875 to study the impact of fertilization on a wheat-sugarbeet rotation. In 1987, the rotation was stopped to be replaced by continuous maize. Crop residues were soil-incorporated and the mineral fertilization was doubled in some plots. The impact of those changes on the microbial biomass and activity are presented. In spring 1987, the soil was still in a steady-state condition corresponding to the rotation. The microbial biomass was correlated with total organic C and decreased in the order farmyard manure>mineral NPK>unfertilized control. Microbial specific respiratory activity was higher in the unfertilized treatments. The soil biomass was closely related to soil N plant uptake. In 1989, after 2 years of maize and crop residue incorporation, the steady-state condition corresponding to the previous agricultural practices disappeared. So did the relationship between the biomass and total organic C, and the soil N plant uptake. Biomass specific respiratory activity increased because of low efficiency in the use of maize residues by microbes under N stress.     

Examination of microbial biomass in beech forest moder profiles

Microbial biomass C, ATP, and substrate-induced respiration were measured in the organic layers and the mineral A horizon of three beech forest soils with moder humus differing in Ca and Mg supply. Analyses of variance showed that horizon-specific differences explained most of the variance in the three microbial parameters. All three were significantly interrelated, with Spearman rank correlation coefficients of between 0.86 and 0.93. However, differences in the decline of these parameters with depth led to horizon-specific differences in their ratios. Thus, the ratios were not markedly interrelated. The mean ATP: microbial C ratio was 5.2 mol ATP g^-1 C in the L 2 layer, 19.5 in the F layers, and 9.6 in the H and A horizons. The ratio of substrate-induced respiration to microbial C varied between 39.3 and 82.2 O₂h^-1 g^-1 C in the F1 layers and between 5.3 and 32.1 l in the other layers. It is concluded that the use of different parameters can help to analyze both horizonand site-specific differences in microbial performance.     

添加生物质炭对红壤水稻土有机碳矿化和微生物生物量的影响

通过室内培育试验,研究了添加生物质炭对江西红壤水稻土有机碳矿化和微生物生物量碳、氮含量的影响。结果表明:红壤有机碳矿化速率在培育第2天达最大值后迅速降低,培养7天后下降缓慢并趋于平稳;添加生物质炭降低了土壤有机碳的矿化速率和累积矿化量,培养结束时,不加生物质炭的对照处理中有机碳的累积矿化量分别比添加0.5%和1.0%生物质炭的处理高10.0%和10.8%。此外,生物质炭的加入显著提高了土壤微生物生物量,添加0.5%生物质炭处理的土壤微生物生物量碳、氮含量分别比对照高111.5%～250.6%和11.6%～97.6%,添加1.0%生物质炭处理的土壤微生物生物量碳、氮含量分别比对照高58.9%～243.6%和55.9%～110.4%。相同处理中,干旱的水分条件下(40%田间持水量)微生物生物量要高于湿润的水分条件(70%田间持水量)。同时,添加0.5%和1.0%的生物质炭使土壤代谢熵分别降低2.4%和26.8%,微生物商减少了43.7%和31.7%。     

Biochemical properties in managed grassland soils in a temperate humid zone: modifications of soil quality as a consequence of intensive grassland use

Although soil biochemical properties are considered to be good indicators of changes in soil quality, few studies have been made of the changes in biochemical properties brought about by anthropogenic disturbance of grassland ecosystems. In the present study, several biochemical properties were analysed in 31 grassland soils subjected to a high level of management, and the values obtained were compared with known values corresponding to native grasslands from the same region (Galicia, NW Spain). The 31 managed grasslands were divided into two groups (re-sown grasslands and improved grasslands) according to their management and past land use. The biochemical properties studied were: labile carbon, microbial biomass carbon, microbial respiration, metabolic quotient, net nitrogen mineralisation and the activities of dehydrogenase, catalase, phosphodiesterase, phosphomonoesterase, casein hydrolysing proteases, benzoyl arginamide (BAA)-hydrolysing proteases, urease, cellulase, ß-glucosidase, invertase and arylsulphatase. Managed grasslands exhibited lower values of soil biochemical properties than native grasslands. Three biochemical equilibrium equations were used to compare soil quality in managed and native grasslands. One of the equations did not show any significant difference between the groups of grassland soils considered. In contrast, two of the equations showed similar soil quality for improved and native grasslands, while re-sown grasslands exhibited a loss of soil quality when compared to native grassland soils.     

Barley yield and soil microbial and enzyme activities as affected by contamination of two soils with lead, zinc or copper

The effects of increasing rates of Pb, Zn and Cu on extractable heavy metal levels, barley yields, basal respiration and the activities of catalase, urease, invertase and acid phosphatase were investigated in two soils in a 2-year greenhouse experiment. In the first year, barley yields were decreased by increasing additions of Pb, Zn and Cu. In the second year, increased yields were recorded at lower rates of addition of all three metals in both the chestnut and chernozem soils. Yield depressions were most marked for added Cu and least marked for Pb but, in contrast, accumulation of heavy metals in grain, in excess of recommended limits, was most pronounced for Pb and did not occur for Cu. Increasing rates of all three metals caused a decrease in basal respiration; the degree of inhibition was generally greater in the second than in the first year. After 1 year of incubation, increasing rates of addition of metals reduced all tested enzyme activities. However, after 2 years, the pattern of response was more complex, with increases in enzyme activities being noted at lower rates of addition of all three metals. In general, invertase and urease activities were more markedly inhibited by heavy metal contamination than those of catalase and phosphatase. Ammonium acetate-extractable heavy metal concentrations in soils were less after 2 years than 1 years reaction time due to their transformation into less labile forms. Significant negative correlations between grain yield, basal respiration and enzyme activities were observed in both years.     

Soil microbial biomass and organic matter composition in soils under cultivation

To determine whether there is a relationship between the composition of soil organic matter and the activity of the soil microbial biomass, the composition of the organic matter in 12 typical arable soils in Northwest Germany was investigated by wet chemical analysis and CPMAS cross polarization magic angle spinning ¹³C-NMR spectroscopy. The data were correlated with the microbial biomass as estimated by substrate-induced respiration. A strong correlation between the microbial biomass and alkylic C compounds was observed (r=-0.960^***). Recalcitrant substances were enriched in this fraction, which were classified as humic acids according to the wet chemical procedure. The microbial decomposition of these humic acids is probably retarded, due to their chemical structure and/or physical bonding, when the soil microbial biomass activity is limited.     

Tillage effects on microbial biomass and nutrient distribution in soils under rain-fed corn production in central-western Mexico

Quantifying how tillage systems affect soil microbial biomass and nutrient cycling by manipulating crop residue placement is important for understanding how production systems can be managed to sustain long-term soil productivity. Our objective was to characterize soil microbial biomass, potential N mineralization and nutrient distribution in soils (Vertisols, Andisols, and Alfisols) under rain-fed corn (Zea mays L.) production from four mid-term (6 years) tillage experiments located in central-western, Mexico. Treatments were three tillage systems: conventional tillage (CT), minimum tillage (MT) and no tillage (NT). Soil was collected at four locations (Casas Blancas, Morelia, Apatzingán and Tepatitlán) before corn planting, at depths of 0–50, 50–100 and 100–150 mm. Conservation tillage treatments (MT and NT) significantly increased crop residue accumulation on the soil surface. Soil organic C, microbial biomass C and N, potential N mineralization, total N, and extractable P were highest in the surface layer of NT and decreased with depth. Soil organic C, microbial biomass C and N, total N and extractable P of plowed soil were generally more evenly distributed throughout the 0–150 mm depth. Potential N mineralization was closely associated with organic C and microbial biomass. Higher levels of soil organic C, microbial biomass C and N, potential N mineralization, total N, and extractable P were directly related to surface accumulation of crop residues promoted by conservation tillage management. Quality and productivity of soils could be maintained or improved with the use of conservation tillage.     

Microbial biomass and activity in urban soils contaminated with Zn and Pb

Summary The effects of heavy metals on microbial biomass and activity were investigated in 30 urban soils, contaminated mainly with Zn and Pb to different extents, in terms of the physicochemical and biological characteristics of the soils. Evaluated by simple and multiple regression analyses, the microbial biomass was not affected significantly by easily soluble Zn + Pb (extractable with 0.1 NHCI). The biomass was accounted for as a function of cation exchange capacity (CEC), total organic C and the numbers of fungal colonies present (R ² = 0.692). Carbon dioxide evolution from soils, which reflected microbial activity, was studied on soils incubated with microbial-promoting substrates (glucose and ammonium sulfate) or without. Carbon dioxide evolution was negatively related to Zn+Pb, and this inhibitory effect of the metals was greater in the soils incubated with substrates. Carbon dioxide evolution in soils with substrates was closely related to Zn+Pb, bacterial numbers and the numbers of fungal colonies (R ² = 0.718). Carbon dioxide evolution in soils without substrates was accounted for as a function of Zn + Pb, biomass and the C/N ratio (R ² = 0.511). Using these relationships, the effects of heavy metals on soil microorganisms are discussed in terms of metabolically activated and dormant populations.     

The proportional mineralisation of microbial biomass and organic matter caused by air-drying and rewetting of a grassland soil

During the first few days after rewetting of an air-dried soil (AD-RW), microbial activity increases compared to that in the original moist soil, causing increased mineralisation (a flush) of soil organic carbon (C) and other nutrients. The AD-RW flush is believed to be derived from the enhanced mineralisation of both non-biomass soil organic matter (due to its physical release and enhanced availability) and microbial biomass killed during drying and rewetting. Our aim was to determine the effects of AD-RW on the mineralisation of soil organic matter and microbial biomass during and after repeated AD-RW cycles and to quantify their proportions in the CO₂-C flushes that resulted. To do this, a UK grassland soil was amended with ¹⁴C-labelled glucose to label the biomass and then given five AD-RW cycles, each followed by 7 d incubation at 25 °C and 50% water holding capacity. Each AD-RW cycle increased the amount of CO₂-C evolved (varying from 83 to 240 μg g⁻¹ soil), compared to the control with, overall, less CO₂-C being evolved as the number of AD-RW cycles increased. In the first cycle, the amount of biomass C decreased by 44% and microbial ATP by 70% while concentrations of extractable C nearly doubled. However, all rapidly recovered and within 1.3 d after rewetting, biomass C was 87% and ATP was 78% of the initial concentrations measured prior to air-drying. Similarly, by 2 d, extractable organic C had decreased to a similar concentration to the original. After the five AD-RW cycles, the amounts of total and ¹⁴C-labelled biomass C remaining in the soil accounted for 60 and 40% of those in the similarly incubated control soil, respectively. Soil biomass ATP concentrations following the first AD-RW cycle remained remarkably constant (ranging from about 10 to 14 μmol ATP g⁻¹ biomass C) and very similar to the concentration in the fresh soil prior to air-drying. We developed a simple mathematical procedure to estimate the proportion of CO₂-C derived from biomass C and non-biomass C during AD-RW. From it, we estimate that, over the five AD-RW cycles, about 60% of the CO₂-C evolved came from mineralisation of non-biomass organic C and the remainder from the biomass C itself.