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Soil Science Society of America Journal 64:184-189 (2000)
© 2000 Soil Science Society of America

DIVISION S-3-SOIL BIOLOGY & BIOCHEMISTRY

Soil Organic Matter Pools and Their Associations with Carbon Mineralization Kinetics

R. Alvareza and C.R. Alvareza

a Dep. de Suelos, Facultad de Agronomía, Univ. de Buenos Aires, Av. San Martín 4453 (1417), Buenos Aires, Argentina

ralvarez{at}mail.agro.uba.ar


    ABSTRACT
 TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 Materials and methods
 Results
 Discussion
 REFERENCES
 
The labile component of soil organic matter (SOM) plays an important role in short-term nutrient turnover. Our objectives were (i) to establish the relationships between carbon in soil density fractions with carbon mineralization and the microbial biomass under contrasting conditions, (ii) to compare the goodness of fit of different mathematical models to describe carbon mineralization, and (iii) to evaluate the relationships of the SOM pools and the mineralization parameters estimated by the best kinetic model. Twenty-eight soil samples were collected from a fine, illitic, thermic Typic Argiudoll localized in Argentina. These samples differed in the soil management (pasture and agriculture), tillage systems (chisel tillage, plough tillage, and no-tillage), crop rotation, or depths. Microbial biomass was highly correlated with total carbon and carbon in the SOM light density fraction (density < 1.59 g mL-1) but less strongly correlated to medium (density 1.59–2.0 g mL-1) and heavy (density > 2.0 g mL-1) soil fractions. Carbon in the soil light fraction was strongly related to the carbon mineralized at 10 and 160 d of incubation. The exponential and hyperbolic models showed a good description of the mineralization data (r2 > 0.982). The application of models which considered two organic matter pools could not describe the mineralization of some samples. The hyperbolic model estimated higher potentially carbon mineralizable pools (C0) and semidecomposition time periods than the exponential one. The C0 estimated by the exponential model were similar to the carbon content in the soil light fraction. This soil organic component seemed to be the driving variable of microbial activity and a good predictor of soil potential carbon mineralization.

Abbreviations: SOM, soil organic matter


    INTRODUCTION
 TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 Materials and methods
 Results
 Discussion
 REFERENCES
 
THE LABILE COMPONENT of soil organic matter plays an important role in the short-term nutrient turnover and is responsible for the temporary soil structural stability (Tisdall and Oades, 1982). One possible way to characterize this fraction is through densimetry. SOM can be divided into (i) light fraction, which consists of mineral-free organic matter composed of partly decomposed plant and animal residues, which turn over rapidly and have a specific density considerably lower than that of soil minerals; and (ii) heavy fraction, composed of more processed decomposition products, which turn over more slowly and have a high specific density because of their intimate association to soil minerals (Christensen, 1992; Barrios et al., 1996). Dalal and Mayer (1986) found for Australian clay soils that the rate of loss of carbon from the light fraction (density < 2 g mL-1) was 2 to 11 times greater than that of the heavy fraction. In contrast, Janzen et al. (1992) observed in Canadian Mollisolls that carbon in the light fraction (density < 1.7 g mL-1) was higher in treatments which include perennial forage in the rotation and lower in those with summer fallow. In Pampean Mollisolls of Argentina, the implementation of no tillage produced an accumulation of carbon in soil light fraction (density < 1.6 g mL-1) at the soil surface (0–5 cm layer) in relation to plowed plots (Alvarez et al., 1995, 1998a).

Carbon and nutrient turnover are mediated by the soil microbial biomass, which responds to residues or tillage management (Dalal et al., 1991). Microbial biomass is usually related to the carbon in soil light fraction and to the in vitro carbon mineralization (Bremer et al., 1994; Alvarez et al., 1995, 1998a). Because soil management generally affects these variables more than total organic carbon, many authors have suggested that they could be early indicators of future trends in total SOM (Bremer et al., 1994).

The mathematical description of in vitro carbon and nitrogen mineralization is another interesting approach to characterizing SOM. The exponential model was widely used to describe the carbon and nitrogen mineralization process (Stanford and Smith, 1972; Riffaldi et al., 1996). From this model the potentially mineralizable carbon pool of soils (C0) may be estimated. C0 is assumed to be a readily mineralizable carbon component which mineralized at a constant rate (k) proportional to the size of the pool. Another single-component model is the hyperbolic model, which considers that the time to mineralize 50% of C0 pool (t1/2) increases as the incubation time gets longer, according to the rise of carbon chemical protection. Alternatives to these one-component models are those which consider two organic matter pools with different stability to microbial attack (Riffaldi et al., 1996). Several authors (e.g., Bonde and Rosswall, 1987) proposed the use of the double-exponential model to improve the agreement with experimental mineralization data. This model assumes that the organic matter pool can be divided into two components, a labile pool (CL) decomposing exponentially with a constant rate (kL), and a resistant pool (CR) also decomposing exponentially at a much lower constant rate (kR). A simplification of this model is the exponential and linear version, which consider a labile pool decomposing with an exponential kinetics and a resistant pool decomposing linearly, according to the relative shortness of the incubation periods compared with the turnover of the resistant pool (Bonde and Rosswall, 1987). Other authors have found that the exponential plus a constant model could be useful to describe an initial mineralization flush present in some soil samples (Bonde and Lindberg, 1988). This model contains a parameter (CL) defined as an easy decomposable organic matter which produced an initial mineralization flush during the first stage of the incubation (Riffaldi et al., 1996), and a resistant pool (CR) decomposing exponentially. This initial mineralization flush was attributed to the drying and rewetting of samples or other type of sample handling (Beauchamp et al., 1986).

The relationships between SOM pools isolated by physical or chemical techniques and the potentially mineralizable organic pool obtained through the mathematical modeling of carbon mineralization have not been widely investigated under different soil managements. Our objectives were (i) to establish the relationships between carbon in soil density fractions with carbon mineralization and the microbial biomass in a Mollisoll under contrasting cropping conditions, (ii) to compare the goodness of fit of different mathematical models to describe the kinetics of carbon mineralization, and (iii) to evaluate the relationships of the SOM pools and the mineralization parameters estimated by the best kinetic model.


    Materials and methods
 TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 Materials and methods
 Results
 Discussion
 REFERENCES
 
Soil and Experimental Sites
The samples were obtained from the INTA Experimental Station located at Pergamino (Argentina, 33°56'S; 60°34'W). The climate is humid and temperate, with an annual rainfall of 1000 mm and a mean temperature of 16.5°C. The soil is a Pergamino series (fine, illitic, thermic, Typic Argiudoll). The main characteristics of the top 20 cm were 27% of clay, 57% of silt, and pH (soil:water 1:2.5) 5.8, with no statistically significant differences of these soil parameters between depths throughout this layer. Twenty-eight soil samples were collected from different field experiments. These samples differed on the basis of the soil management, tillage systems, crop rotation, or depths (Table 1) . The values of each sample are the mean of four plots. The total C content of the 28 samples ranged from 13.8 to 36.9 g C kg-1 soil.


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Table 1 Organic carbon content (g kg-1 dry soil) of samples from a fine, illitic, thermic Typic Argiudoll under contrasting soil managements

 
Analytical and Statistical Methods
Fresh soil samples were homogenized by hand. Soil microbial biomass (biomassC) was determined by the fumigation-incubation method (Jenkinson and Powlson, 1976) as described by Alvarez et al. (1995). A k factor of 0.45 was used to convert CO2 C production to biomass C (Oades and Jenkinson, 1979).

Soil density fractionation was performed with both carbon tetrachloride (density = 1.59 g mL-1) and bromoform–ethanol mixture (density = 2.00 g mL-1). Air-dry soil samples were sieved (<500 µg) and plant residues were forced to pass the sieve. Five grams of soil was weighed into a 50-mL beaker, and after adding 30 mL of the separation liquid, was vigorously agitated 1 min by hand and then centrifuged at for 5 min. The supernatant was filtered through fiberglass under suction. Carbon in whole soil and in the two light density fractions was determined by wet digestion (Amato, 1983).

The SOM light fraction (light fraction C) was defined as the carbon contained in the supernatant of soil:carbon tetrachloride mixture . Soil organic matter heavy fraction (heavy fraction C) was estimated as the difference between total C and the carbon quantified in the supernatant of soil:bromoform:ethanol mixture . Carbon content of the SOM medium fraction (medium fraction C) was calculated as the difference between carbon in fractions with density < 2 g mL-1 and the light fraction C.

The metabolic ratio was calculated as the ratio between the carbon respired in 10 d of incubation (respired C) from non fumigated controls and the biomass C. In vitro aerobic carbon mineralization was measured during 160 d (mineralized C), at 30°C and 50% of soil water holding capacity. The equivalent of 100 g of dry soil were incubated in a 400-mL flask and the CO2 C production was periodically (10, 20, 40, 70, 100, 130, and 160 d of incubation) determined by alkali absorption (Alvarez et al., 1995). The cumulative carbon production was fitted to different mathematical models (Table 2) . The models were fitted to carbon mineralization data by the non-linear regression with the Statgraphics software package (Manugistics, Inc., Rockville, MD). The relationships between the different variables were evaluated by regression analysis and tested by their F values.


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Table 2 Models used to described carbon mineralization kinetics

 

    Results
 TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 Materials and methods
 Results
 Discussion
 REFERENCES
 
Biomass C was positively and highly correlated with total C and light fraction C, but had low relationships with medium and heavy fraction C (Table 3) . Microbial activity, evaluated as respired C, was positively related to total C and carbon present in the different soil density fractions, but the light fraction C showed the highest correlation with respired C (Table 3). Poor relationships were observed between the different SOM density fractions.


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Table 3 Coefficients of determination (r2) between the SOM pools. All coefficients are significant at P < 0.001

 
The mineralized C was associated with biomass C and carbon present in the soil density fractions. The mineralized C presented the highest correlation with the light fraction C and lowest with the medium fraction C (Fig. 1) . The amount of carbon mineralized in 160 d of incubation was around five-fold the biomass C and 50% higher than the light fraction C. In contrast, medium and heavy fraction C were 1.8 and 14 times higher than mineralized C. When the biomass C and the SOM density fractions were expressed as proportion of total C, a positive and high relationship was found between mineralized C/total C ratio and the biomass C/ total C ratio and the light fraction C/total C ratio (Fig. 2) . However, the light fraction C/total C showed a higher correlation with the proportion of total C mineralized than did the biomass C/total C. Otherwise, the total C mineralized was negatively related with the ratio heavy fraction C/total C. No statistically significant relationship was found between the medium fraction C/total C and the proportion of total C mineralized.



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Fig. 1 Relationships between the carbon mineralized in 160 d (mineralized C) with carbon in the different organic matter pools. Depths: • 0–5 cm, {circ} 5–10 cm, § 10–15 cm, {square} 15–20 cm

 


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Fig. 2 Relationships between the fraction of the total carbon mineralized in 160 d and the fraction of the total carbon in the different soil organic matter fractions. Depths: • 0–5 cm, {circ} 5–10 cm, § 10–15 cm, {square} 15–20 cm

 
Both the exponential and hyperbolic models provided satisfactory fits of the cumulative CO2 C production data in all samples (Table 4) . The double-exponential model gave higher coefficients of correlation than the one-component models, but it could only be fitted to the mineralization data of 20 soil samples. The exponential plus linear model showed a good fit to the carbon mineralization data, but estimated negative resistant pool mineralization constants (C) for eight samples. The exponential plus a constant model also described satisfactory the in vitro mineralization, but calculated negative values for the carbon labile pool (C0) in 10 samples.


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Table 4 Fit of different models to carbon mineralization data for the 28 soil samples

 
The potentially mineralizable carbon pool estimated by the exponential model (exponential C0) and the hyperbolic model (hyperbolic C0) were highly correlated between each other (Fig. 3) . Additionally, their semidecomposition times () were positive and highly associated too. The hyperbolic model gave C0 values that were about 55% greater and values 2 to 2.8 times greater than those estimated by the exponential model. When these functions were adjusted to mineralization data sets corresponding to shorter incubation time periods (i.e., 130 or 100 d), both models estimated lower potentially mineralizable carbon pools (C0), but still gave high r2 values (not shown).



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Fig. 3 Correlations between potentially mineralizable carbon pool (C0) and semidecomposition time (t1/2) estimated by the exponential and hyperbolic kinetic models. t1/2 for the exponential model was estimated as: 0.693/exponential k. Depths: • 0–5 cm, {circ} 5–10 cm, § 10–15 cm, {square} 15–20 cm

 
The exponential model was selected because of its simplicity and the goodness of fit for all samples. The exponential C0 was closely and linearly related to the mineralized C, with a slope of 1.07 (Fig. 4) . Thus, similar to the relationships previously noted (Fig. 1 and 2) for mineralized C, the exponential C0 was highly correlated with biomass C ( , P < 0.001; not shown) and with light fraction C (Fig. 4). Otherwise, the exponential C0 showed low correlation with the heavy fraction C and with the medium fraction . The mineralization constant of the exponential model (exponential k) presented a positive but weak correlation with the ratio light fraction C/total , and a negative association with the ratio heavy fraction C/total .



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Fig. 4 Correlation between C0 of the exponential with the carbon mineralized in 160 d of incubation (mineralized C) and the light fraction C. Depths: • 0–5 cm, {circ} 5–10 cm, § 10–15 cm, {square} 15–20 cm

 

    Discussion
 TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 Materials and methods
 Results
 Discussion
 REFERENCES
 
Soil Organic Matter Fractions, Microbial Activity, and Carbon Mineralization
The biomass C, mineralized C, and light fraction C have been proposed by many authors to be used as indicators to evaluate the effect of different soil management practices because changes in these fractions may precede future changes in soil organic matter (Janzen et al., 1992; Bremer et al., 1994). In our case, the biomass C varied from 67 to 1270 µg C g-1 soil (a 19-fold difference), light fraction C from 160 to 6630 µg C g-1 soil (a 41-fold difference), and respired C from 30 to 1060 µg C g-1 soil (a 35-fold difference), showing more sensitivity to the different soil managements or depths than the total C which presented only a 2.7-fold difference between the samples. The biomass C and the microbial activity (respired C) were highly correlated with light fraction C; so these parameters were probably regulated by this SOM soil density fraction. Janzen et al. (1992) found a similar relationship between the light fraction C (density < 1.7 g mL-1) and the carbon mineralized in 14 d of incubation for cold Canadian soils under different rotations, but in that study the biomass C was not related with the carbon mineralized during the incubation.

The metabolic ratio (respired C/biomass C) showed a positive and potential relationship with the availability of light fraction C per unit of biomass C [respired C/biomass C = 0.09 x (light fraction C /biomass C)0.53, ; P < 0.001]. Conversely, the relationships of the metabolic ratio with the medium or heavy fractions C per unit of biomass C were not statistically significant. Only a small proportion of the biomass C is in an active state (McGill et al., 1986; van der Werf and Verstraete, 1987). Probably as the availability of labile carbon source increases per unit of biomass C, the proportion of this biomass in an active state increases. Additionally, the presence of more labile substrate could induce changes in soil microbial biomass composition or its physiological state resulting in a higher production of CO2 C per unit of biomass C (Jans-Hammermeister, 1996). In a previous study from this Pampean soil, a strong relationship was found between the metabolic ratio of the active soil microbial biomass and the availability of light fraction C per unit of the active microbial biomass (Alvarez et al., 1998a).

The light fraction consists principally of plant residues and appreciable amounts of microbial and microfaunal debris, which have a rapid turnover (Spycher et al., 1983). According to these characteristics the light fraction C was closely correlated to the carbon mineralization in 160 d. But the amount of carbon mineralized was higher than the biomass C and the light fraction C. The amount of SOM present in the light fraction is usually affected by land use (e.g., years under cultivation, rotations, tillage systems) (Dalal and Mayer, 1986; Janzen et al., 1992; Alvarez et al., 1995). The higher amounts of light fraction C corresponded to the samples from the upper 5 cm of soil profile (Fig. 2), under pasture or conservation tillage treatments (not shown). When the SOM density fractions were expressed as a proportion of the total C, the mineralized C/total C ratio was highly correlated with the light fraction C/total C, independent of soil management or depth. Otherwise, an increase in the amount of total carbon in the heavy fraction C produced a decrease in the mineralized C/total C ratio. Organic compounds associated with clay particles are chemically recalcitrant and are more physically protected than the light fraction C (Cambardella and Elliot, 1993). In contrast to the association observed between the mineralized C and light fraction C, in this soil, the percentage of total nitrogen mineralized in 84 d was principally related to the percentage of total nitrogen present in the medium fraction (Alvarez et al., 1998b). These results could be a consequence of the higher the C:N ratio of the light SOM, which may cause nitrogen immobilization during the incubation.

Kinetics Parameters and Their Relationships with the Soil Organic Matter Fractions
The exponential and hyperbolic models fit the in vitro mineralization of all studied samples. As observed by other authors, the hyperbolic model estimated higher C0 and t1/2 than the hyperbolic one. Otherwise the two-component models could not be adjusted in some samples (Table 4). The duration of the incubation could affect the goodness of fitting of the two component models. As the incubation time increased these models seemed to better describe the pattern of mineralization because the contribution of carbon mineralized from the resistant pool increases. Dou et al. (1996) found that the mean square error of fitting the exponential plus linear model to nitrogen mineralization decreased in some of the studied treatments as the incubation time decreases from 30 to 15 wk. In the latter case, this model gave negative constant of mineralization, and the introduction to the program of the constraint that this pool should be >=0; gave potentially mineralizable nitrogen pools and mineralization constants similar to those estimated by the simple model. In some of our samples, where negative C0 values were obtained, an initial delay phase was present possibly resulting from microbial regrouping or acclimation (Ellert and Bettany, 1988).

We applied the kinetic models to accumulated data of CO2 C production during 160 d, using integrated equations. Many authors suggested that using accumulated data also accumulate errors while dampening the noise and giving a false sense of security (Ellert and Bettany, 1988; Hess and Smith, 1995). Hess and Smith (1995) fitted different models to their mineralization data, expressed in differential or integral form. The differential form showed a random pattern; meanwhile, the integral form had distinctly non-random residuals, showing the superiority of the differential approach. In our study, we also analyzed the mineralization data with the differential form of the exponential and doubled-exponential models (Colores et al., 1996). We obtained the same performance as analyzing the cumulative CO2 C production. The exponential model adjusted to all samples and the doubled exponential could not be fit to 10 samples. The differential form of the simple exponential gave lower coefficient of correlations (r2 from 0.310–0.986) than those obtained by the integral form. Exponential C0 estimated by differential equations were highly and linearly correlated with those estimated by the integrated models, but the regression slope was 1.1 (P < 0.05). This discrepancy between the potentially mineralizable carbon pools estimated by the two forms of the same model could be a consequence of large intervals (1.5–4 wk) between CO2 C determinations, in relation to the duration of the incubation (23 wk). In experiments where the differential form was applied, the interval between determination was very short (about 1 h), compared with the incubation duration (50 h) (Colores et al., 1996). The exponential model (using the differential or integrated forms) was capable of describing the carbon mineralization patterns in a wide range of soil management practices and depths.

The carbon in soil light fraction was very strongly correlated with the microbial biomass and its activity. This soil carbon pool was also closely related with the mineralized C in long-term incubations. The exponential model described the mineralization pattern of all samples, from a wide range of soil managements and different depth. The C0 estimated by this model were similar to the carbon content in the soil light fraction. This soil organic component seemed to be the driving variable of microbial activity and a good predictor of soil potential carbon mineralization.


    NOTES
 TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 Materials and methods
 Results
 Discussion
 REFERENCES
 
This research was supported by the UBACYT AG-089 program of the University of Buenos Aires.

Received for publication December 22, 1997.


    REFERENCES
 TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 Materials and methods
 Results
 Discussion
 REFERENCES
 




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