DIVISION S-6SOIL & WATER MANAGEMENT & CONSERVATION

Soil Structure and Organic Matter: I. Distribution of Aggregate-Size Classes and Aggregate-Associated Carbon
J. Six,* K. Paustian, E. T. Elliott, and C. Combrink ABSTRACT
Cultivation reduces soil C content and changes the distribution and stability of soil aggregates. We investigated the effect of cultivation intensity on aggregate distribution and aggregate C in three soils dominated by 2:1 clay mineralogy and one soil characterized by a mixed (2:1 and 1:1) mineralogy. Each site had native vegetation (NV), no-tillage (NT), and conventional tillage (CT) treatments. Slaked (i.e., air-dried and fast-rewetted) and capillary rewetted soils were separated into four aggregate-size classes (53, 53250, 2502000, and 2000 m) by wet sieving. In rewetted soils, the proportion of macroaggregates accounted for 85% of the dry soil weight and was similar across management treatments. In contrast, aggregate distribution from slaked soils increasingly shifted toward more microaggregates and fewer macroaggregates with increasing cultivation intensity. In soils dominated by 2:1 clay mineralogy, the C content of macroaggregates was 1.65 times greater compared to microaggregates. These observations support an aggregate hierarchy in which microaggregates are bound together into macroaggregates by organic binding agents in 2:1 clay-dominated soils. In the soil with mixed mineralogy, aggregate C did not increase with increasing aggregate size. At all sites, rewetted macro- and microaggregate C and slaked microaggregate C differed in the order NV NT CT. In contrast, slaked macroaggregate C concentration was similar across management treatments, except in the soil with mixed clay mineralogy. We conclude that increasing cultivation intensity leads to a loss of C-rich macroaggregates and an increase of C-depleted microaggregates in soils that express aggregate hierarchy.

isdall and Oades (1982) presented a conceptual model for aggregate hierarchy that described how primary mineral particles are bound together with bacterial, fungal, and plant debris into microaggregates. These microaggregates, in turn, are bound together into macroaggregates by transient binding agents (i.e., microbial- and plant-derived polysaccharides) and temporary binding agents (roots and fungal hyphae). Three consequences of this aggregate hierarchy are: (i) a gradual breakdown of macroaggregates into microaggregates before they dissociate into primary particles, as increasing dispersive energy is applied to soil (Oades and Waters, 1991), (ii) an increase in C concentration with increasing aggregate-size class because large aggregate-size classes are composed of small aggregate-size classes plus organic binding agents (Elliott, 1986), and

Natural Resource Ecology Lab., Colorado State Univ., Fort Collins, CO 80523. Received 21 Dec. 1998. *Corresponding author (johan@ nrel.colostate.edu). Published in Soil Sci. Soc. Am. J. 64:681689 (2000).

(iii) younger and more labile organic matter is contained in macroaggregates than in microaggregates (Elliott, 1986; Puget et al., 1995; Jastrow et al., 1996). Oades and Waters (1991) tested the aggregate hierarchy theory in different soils by applying a range of treatments to disaggregate soils. They concluded that aggregate hierarchy existed in two Alfisols and a Mollisol because organic materials were the major binding agents for aggregate formation and stabilization in these soils. In contrast, they found that an Oxisol did not express any hierarchical aggregate structure, probably because oxides, rather than organic materials, were the dominant stabilizing agents. Elliott (1986) observed more organic matter associated with macroaggregates than with microaggregates in a temperate grassland soil. He also found that organic matter associated with macroaggregates was more labile than organic matter associated with microaggregates. Jastrow et al. (1996) reported greater amounts of recently incorporated organic matter as aggregates became larger, supporting the idea that microaggregates are bound together by young organic matter into larger macroaggregates. Aggregate hierarchy theory has been used by many authors to explain the correlation between a reduction in aggregation and loss of soil organic matter (SOM) with cultivation (Elliott, 1986; Cambardella and Elliott, 1993; Beare et al., 1994). A breakdown of macroaggregates results in a release of labile SOM (Elliott, 1986) and its increased availability for microbial decomposition. The increased microbial activity depletes SOM, which eventually leads to lower microbial biomass and activity and consequently a lower production of microbial-derived binding agents and a loss of aggregation (Jastrow, 1996; Six et al., 1998). Reduced aggregation (and subsequent lower levels of SOM) in conventional tillage (CT) vs. no-tillage (NT) (Paustian et al., 1999) is a result of several indirect effects on aggregation. Tillage brings subsurface soil to the surface where it is then exposed to wetdry and freezethaw cycles and subjected to raindrop impact (Beare et al., 1994; Paustian et al., 1997), thereby increasing the susceptibility of aggregates to disruption (Willis, 1955; Hadas, 1990; Edwards, 1991). Plowing changes the soil conditions (e.g., temperature, moisture, and aeration) and increases the decomposition rates
Abbreviations: CT, conventional tillage; IPOM, intra-aggregate particulate organic matter; LF, light fraction; NV, native vegetation; NT, no-tillage; POM, particulate organic matter; SOM, soil organic matter.

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Table 1. General characteristics of the agricultural experiment field sites.

Sidney, NE Soil classification Soil series Texture Mineralogy MAT (C) MAP (mm) Crop rotation Prior vegetation Year of establishment Pachic Haplustoll Duroc loam illite, chlorite 8.5 440 winter wheatfallow shortgrass prairie 1969 Wooster, OH Typic Fragiudalf Wooster silt loam chlorite, illite 9.1 905 continuous corn grass meadow 1962 KBS, MI Typic Hapludalf Kalamazoo and Oshtemo sandy loam chlorite, illite 9.2 920 cornsoybeanwinter wheat grassland 1986 Lexington, KY Typic Paleudalf Maury silty clay loam vermiculite, kaolinite, illite 13.1 1127 continuous corn (84 kg N) bluegrass pasture 1971

Dominant clay minerals in order of dominance. Data adopted from Six et al. (1999b). MAT mean annual air temperature. MAP mean annual precipitation.

of litter (Rovira and Greacen, 1957; Cambardella and Elliott, 1993). In NT, residues accumulate at the surface where the litter decomposition rate is slowed due to drier conditions and reduced contact between soil microorganisms and litter (Salinas-Garcia et al., 1997). Finally, the proportion of the microbial biomass composed of total fungi (Frey et al., 1999) and mycorrhizal fungi (OHalloran et al., 1986) is generally higher in NT compared to CT and it has been observed that fungi (especially mycorrhizal) contribute to macroaggregate formation and stabilization (Tisdall and Oades, 1982). The objectives of this study were to (i) test the validity of the aggregate hierarchy theory over a range of soils and (ii) study the affect of increased cultivation intensity on aggregation and aggregate-associated C. MATERIALS AND METHODS Sampling
Soils from four long-term agricultural field experiments (Table 1) were sampled at two depths (05 cm and 520 cm) in November 1995. The four sites are located in Sidney, NE (4114N, 10300W), Wooster, OH (4048N, 8200W), W. K. Kellogg Biological Station (KBS), MI (4224N, 8524W), and Lexington, KY (3807N, 8429W). All four sites have NV, NT, and CT treatments with either three or four replicates. At Sidney, all management treatments were established directly from the native shortgrass prairie. At KBS, NT and CT plots were under long-term cultivation before establishment, whereas the adjacent NV plot was in grass vegetation on a former woodlot that had never been cultivated. At Wooster, the NV plot was in nearby forest, but the NT and CT plots were established in a 6-yr-old grass meadow that was cultivated for many years prior to establishment. At Lexington, the experimental plots were initially under long-term cultivation, but for 50 yr prior to treatment establishment were in bluegrass pasture. The Lexington soil is characterized by a mixed-clay mineralogy dominated by kaolinite (1:1 layer type) and vermiculite (2:1 layer type). In addition, a 2 to 16 times higher concentration of amorphous and poorly crystalline Fe- and Al-oxides was reported for the Lexington soil compared to the Sidney, Wooster, and KBS soils (Six et al., 1999b). The Sidney, KBS, and Wooster soils have only 2:1 minerals, predominantly illite and chlorite (Six et al., 1999b).

capillary-rewetted before immersion in water (rewetted treatment). For capillary rewetting, dried soil was placed on filter paper that was slowly moistened until a water content of 1.05 times field capacity was reached. The volumetric water content of the soil at field capacity was determined for all individual samples. A higher amount of disruptive energy occurs upon slaking because rapid wetting of dry soil leads to an entrapment of air and a buildup of air pressure within the aggregates (Kemper et al., 1985). Aggregates of lower stability disrupt because they cannot withstand this pressure. In contrast, soil rewetted to 1.05 times field capacity is at maximum stability (Hofman and De Leenheer, 1975). The method used for aggregate-size separation was adapted from Elliott (1986). Briefly a 100-g subsample (air-dried or capillary-rewetted) was submerged for 5 min on a 2000-m sieve. Aggregates were separated by moving the sieve (by hand) up and down 3 cm with 50 repetitions during 2 min. The 2000-m aggregates were collected and sieving was repeated for the 2000-m fraction with the next smallersized sieve. This procedure was repeated for every sieve size (250 and 53 m). All aggregate fractions were oven-dried (50C) and weighed. Sand content (53 m) of the aggregates was determined on a subsample of aggregates that were dispersed with sodium hexametaphosphate (5 g L1).

Free Light Fraction and Mineral-Associated Fraction Analysis

The free light fraction (POM outside of the aggregates, or free LF) and mineral-associated organic matter fraction were isolated from the three largest aggregate-size classes according to Six et al. (1998). Briefly, free LF was isolated by density flotation in 1.85 g cm3 sodium polytungstate. The free LF probably includes both LF outside of aggregates and some LF released from the aggregates upon submersion in sodium polytungstate. However, the dispersion of aggregates in sodium polytungstate is minimal and therefore the released fraction was only a small proportion of the free LF. Sodium polytungstate was recycled according to Six et al. (1999c) to avoid cross- contamination of C and N between samples. After isolation of free LF, aggregates were dispersed in 5 g L1 sodium hexametaphosphate by shaking for 18 h on a reciprocal shaker. Intra-aggregate (within aggregate) particulate organic matter (IPOM) was isolated by sieving. Aggregate-associated C and mineral-associated C were calculated by difference

aggregate-associated C total aggregate C free LF C mineral-associated C aggregate-associated C IPOM C [2] [1]

Aggregate Separation
Field-moist soil was gently broken to pass an 8-mm sieve and air-dried. Aggregate separations and soil stability assessments were done by wet sieving. Two pretreatments were applied before wet sieving: (i) air-dried soil was rapidly immersed in water (slaked treatment) and (ii) air-dried soil was

where total aggregate C is total C measured in aggregates prior

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to free LF flotation. Aggregate-associated C and mineralassociated C were only determined for the slaked microaggregates (53250 m) and small macroaggregates (2502000 m). The yield of large macroaggregates (2000 m) after slaking was often too small to be analyzed. For the 53-m fraction, the LF yield was too small for C analysis and by definition (Cambardella and Elliott, 1992) there is no IPOM in particles 53 m.

Carbon, Nitrogen, and Isotope Analyses

Isotope and organic C and N analyses were performed according to Six et al. (1998). The natural abundance 13C methodology for SOM studies was only done at the Sidney site. The other locations did not have a single shift in the dominance of plant species with different metabolic pathways (C3 vs. C4) or archived soil samples from the beginning of the experiment. At Sidney, the delta 13C values () were used to calculate the proportion of wheat-derived C ( f ) in each organic matter fraction:

1 0 w 0

[3]

where t 13C at time t, w 13C of wheat straw (crop), 0 13C of original grassland-derived SOM, and f fraction of wheat-derived C in the soil. The proportions of wheatderived C vs. grassland-derived C (1 f ) provide a measure of the relative age of the organic matter in the different size fractions. The proportions of crop-derived C are only presented for the 0- to 5-cm layer because changes in the 13C signature with depth confound interpretations at the 5- to 20-cm depth. In addition, differences in C concentrations were mainly observed in the 0- to 5-cm layer.

Statistical Analyses
The experimental field design was a randomized complete block design for all sites. However, the NV were not replicated within the experiments at Wooster, KBS, and Lexington and therefore not included in the statistical analysis. Analysis of variance (ANOVA-GLM, SAS Institute, 1990) was performed with multiple comparisons within a depth. Tillage treatment was the main factor in the model, with aggregate size and replicate as secondary factors. Separation of means was tested using Tukeys honestly significant difference at a level of P 0.05.

2000 m), which on average accounted for 85% of the dry soil weight (Fig. 1). A nonsignificant higher proportion of large macroaggregates was found in NT compared to NV and CT, especially at Sidney and KBS. Otherwise, there were no major differences among treatments in the rewetted aggregate distribution. In contrast, the slaked aggregate-size distribution differed between management treatments at all sites (Fig. 1). Proportions of macroaggregates decreased in the order NV NT CT, except that NV and NT were similar at Wooster. The lack of differences in aggregation between NV and NT at Wooster (Fig. 1) was consistent with their similar values for total C (Table 1) and total POM (Six et al., 1999a). At all sites, there was a reduction in large macroaggregates and an increase in microaggregates with slaking compared to rewetting. The proportion of silt and clay particles (53 m) increased from 0.05 in rewetted samples to 0.15 in slaked samples and this increase was greatest in the Lexington soil (Fig. 1). At Sidney and Wooster, tilled soils showed a substantial decrease in small and large macroaggregates concomitant with an increase in microaggregates and a small increase of silt and clay particles, compared with NT (Fig. 1). The small difference in slaked aggregate distribution between NT and CT at KBS may be due to the short duration (9 yr) of the experiment at KBS. At Lexington, CT had fewer large macroaggregates (1.4% vs. 18.0%), more microaggregates (37.3% vs. 16.8%), and more silt and clay particles (15.5% vs. 9.3%) than in NT, but there was no difference in the proportion of small macroaggregates between tillage treatments.

Aggregate Carbon Concentrations

It is frequently observed that the major differences in organic matter content between NT and CT soils are in the upper few centimeters of soil (Doran, 1987; Dick et al., 1997). Similarly, we found few differences in aggregate C among treatments at the 5- to 20-cm depth (data not shown). The only exception was KBS, where the NV treatment had substantially higher aggregate C concentrations at 5 to 20 cm compared to NT and CT (data not shown). This trend may be due to the longterm cultivation of NT and CT plots before establishment of the experiment. While not reported, trends across aggregate-size classes within treatments were the same in the subsurface layer and surface layer. Therefore, further comparisons are made only for the 0- to 5-cm layer. In general, sand-free C concentrations of all rewetted aggregate-size classes differed in the order NV NT CT (Fig. 2). At KBS, NT and CT did not differ significantly in this respect, which may again be a result of the young age of the experiment. The apparent large differences between rewetted (and slaked) aggregate C concentrations in NV versus NT and CT at KBS is probably also a result of the long-term cultivation of the NT and CT plots prior to establishment of the experiment. In contrast to the other sites, rewetted aggregate C concentrations at Wooster (the only site with forest vegetation) were not different between NV and NT,

RESULTS Whole Soil Characteristics

Total organic C and N (020 cm) generally decreased (but not always significantly) in the order NV NT CT (Table 2). At all sites, significant differences in total organic C and N between treatments were observed in the 0- to 5-cm depth, except at KBS where NT and CT were not significantly different. Organic C and N levels were on average 38% lower in CT compared to NT in the 0- to 5-cm depth (Table 2). In contrast, there were no significant differences observed in total C and N in the 5- to 20-cm depth at any site. Bulk density was not significantly affected by tillage at any of the sites (Table 2).

Aggregate-Size Distribution
At all sites, rewetted aggregate-size distributions were dominated by macroaggregates (2502000 m and

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Table 2. Organic carbon, nitrogen, and bulk density in four agricultural experiment sites with native vegetation (NV), no-tillage (NT), and conventional tillage (CT) treatments.
Organic C Site Sidney, NE Depth cm 05 520 020 05 520 020 05 520 020 05 520 020 NV 1437a 2653a 4090a 1418 2591 4008 1540 1404 2944 2324 2711 5036 NT g C m 2 1129b 2299a 3428ab 1598a 2209a 3806a 791a 1631a 2422a 1295a 2448a 3742a CT 699c 2208a 2907b 843b 2537a 3380b# 627a 1582a 2209a 662b 2463a 3125b NV 131a 265a 396a 106 194 300 125 116 241 211 277 789 Organic N NT g N m2 108b 258a 366a 150a 196a 346a 68a 150a 218a 136a 305a 441a CT 82c 230a 312a 76b 219a 294b 46a 157a 204a 72b 275a 348b NV 0.82a 1.12a* 1.02 1.19 1.02 1.38 1.02 1.37 Bulk density NT g cm3 1.05b 1.22b* 1.19a 1.30a 1.62a 1.69a 1.07a 1.26a CT 1.18b 1.28b 1.37a 1.30a 1.53a 1.60a 1.05a 1.26a

Wooster, OH

KBS, MI

Lexington, KY

For all sites, total organic C (020 cm) is adopted from Six et al. (1999a) and all data for Sidney is adopted from Six et al. (1998). Tillage treatments within a site and depth followed by a different lowercase letter are significantly different (P 0.05) according to Tukeys HSD mean separation test. Values within a site and treatment followed by * at the 520 cm depth are significantly different compared with corresponding values at 05 cm. No statistics because only one replicate for this treatment. # Significance at the 10% level.

except for the microaggregates. This suggests that forest vegetation is not as beneficial as grassland vegetation for the accumulation of aggregate C. At Sidney, Wooster, and KBS (sites with 2:1 claydominated soils), slaked aggregate C content increased with increasing aggregate size within a management treatment (except NV at KBS), although the C content of large macroaggregates was similar to that of small macroaggregates (Fig. 2). The C content of small macroaggregates was on average 1.65 times greater compared to microaggregates. This trend of greater C content in small macroaggregates compared to microaggregates is also apparent in the aggregate-associated C and mineral-associated C of NT and CT (Table 3). Aggregateassociated C and mineral-associated C concentrations were more similar across tillage treatments for slaked macroaggregates compared to slaked microaggregates, in these 2:1 clay-dominated soils (Table 3). The C concentrations of slaked microaggregates differed in the order NV NT CT. At Sidney, both grassland- and especially crop-derived aggregate C increased with increasing aggregate size, except for the largest size which had equal or lower concentrations than the next-smaller size (Table 4). In addition, the percentage of crop-derived C increased about 32% and 38% with increasing aggregate size in NT and CT, respectively (Table 4). This observation is in agreement with Puget et al. (1995) and Jastrow et al. (1996) who also observed increasing proportions of young C with increasing aggregate size. In contrast to the 2:1 clay-dominated soils, aggregate C (Fig. 2), aggregate-associated C, and mineral-associated C (Table 3) concentrations for slaked aggregates did not differ between macro- and microaggregates within a management treatment in the Lexington soil (mixed-clay mineralogy). Feller et al. (1996) also observed similar C concentrations in different aggregatesize classes in a low-activity (1:1 clay) soil. Elliott et al. (1991) also found no significant differences in aggregate

C concentrations among aggregate-size classes in a Ultisol from the Amazon Basin of Peru.

DISCUSSION
In soils where SOM is the major binding agent an aggregate hierarchy has been observed (Tisdall and Oades, 1982; Oades and Waters, 1991). SOM is expected to be the primary binding agent in 2:1 clay-dominated soils because polyvalent-organic matter complexes form bridges between the negatively charged clay platelets. In contrast, SOM is not the only binding agent in oxideand 1:1 clay-dominated soils. Electrostatic attractions occur between and among oxides and kaolinite platelets due to simultaneous existence of positive and negative charges at field pH (Schofield and Samson, 1954; ElSwaify, 1980). Thus in those soils aggregate formation is partly induced by electrostatic interactions and aggregate hierarchy should be less pronounced (Oades and Waters, 1991). The increased aggregate- and mineral-associated C content of small macroaggregates vs. microaggregates (within treatment) at Sidney, Wooster, and KBS (Table 3) indicates that both IPOM C and mineral-associated C are incorporated during formation of macroaggregates. This also suggests that IPOM C is a major C source for microbial activity and thereby induces the binding of clay- and silt-sized particles and microaggregates into macroaggregates (Jastrow, 1996; Six et al., 1998, 1999a) in these 2:1 clay-dominated soils. In addition, the similarity of aggregate-associated C concentrations of slaked macroaggregates across management treatments indicates the stability of slaked macroaggregates is correlated to their C content (Elliott, 1986; Cambardella and Elliott, 1993). The stability of microaggregates, in contrast, does not seem to be correlated to C content, because there is a difference in slaked microaggregate C content among treatments at all sites (Table 3). Perhaps the physical characteristics of microaggregates such as

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Fig. 1. Slaked and rewetted aggregate-size distributions in the surface layer (05 cm) of four long-term agricultural experiment sites (SID Sidney, NE; WO Wooster, OH; KBS Kellogg Biological Station, MI; LX Lexington, KY) with three management treatments (NV native vegetation; NT no-tillage; CT conventional tillage). Bars are standard deviations.

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Fig. 2. Slaked and rewetted aggregate C concentrations in the surface layer (05 cm) of four long-term agricultural experiment sites (SID Sidney, NE; WO Wooster, OH; KBS Kellogg Biological Station, MI; LX Lexington, KY) with three management treatments (NV native vegetation; NT no-tillage; CT conventional tillage). All C concentrations are corrected for sand content. Bars are standard deviations.

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Table 3. Aggregate-associated C and mineral-associated C concentrations for slaked microaggregates and small macroaggregates in four agricultural experiment sites with native vegetation (NV), no-tillage (NT), and conventional tillage (CT) treatments (05 cm depth).
Aggregate-associated C Site Aggregate-size class m Sidney, NE Wooster, OH KBS, MI Lexington, KY 53250 2502000 53250 2502000 53250 2502000 53250 2502000 37.37a 38.05a 30.98 33.27 64.52 47.59 41.84 39.70 24.95b 33.83a 22.16c 32.73b 15.52b 22.61a 25.76a 26.34a NV NT CT NV Mineral-associated C NT CT

g C kg1 sand-free aggregates 19.27b* 31.99a* 11.07b* 25.63a 11.61a* 15.08a* 12.09a* 14.56a* 27.66a 26.57a 25.74 27.58 37.25 38.12 27.21 30.70 21.55b 26.15a 17.32b 21.93a 13.36b 18.38a 19.77a 21.12a 17.71b* 25.96a 9.36a* 19.45b 10.44a 12.11a* 11.34a* 12.44a*

Values within a site and treatment followed by a different lowercase letter are significantly different (P 0.05) according to Tukeys HSD mean separation test. * Indicates that there are significant differences (P 0.05) between treatments within a site and an aggregate-size class according to Tukeys HSD mean separation test. There are no statistics for this treatment because there were no replicates available.

lower porosity and higher bulk density (Oades and Waters, 1991) are the main factors that confer resistance to slaking rather than their C content. The comparison of rewetted and slaked aggregate distribution and the aggregate-associated C concentrations provides information on the degree of aggregate hierarchy exhibited by the different soils. The aggregate hierarchy theory seems applicable to the 2:1 clay-dominated soils at Sidney, Wooster, and KBS because of (i) small increases in silt and clay particles, but large increases in microaggregates, upon disruption of the macroaggregate (Fig. 1) (Elliott, 1986; Oades and Waters, 1991), (ii) small differences in silt and clay proportion between NT and CT (Fig. 1) (Elliott, 1986), and (iii) increased aggregate-associated C concentrations with increasing aggregate sizes in slaked soils (Table 3) (Elliott, 1986). Additional support for the aggregate hierarchy at Sidney is provided by the 13C natural abundance data (Table 4). The increase in proportions of young C with aggregate size indicates that microaggregates are bound together into larger macroaggregates by cropderived C (Puget et al., 1995; Jastrow et al., 1996). Elliott (1986) used the aggregate hierarchy theory as a basis to explain reduced SOM level in a stubble mulch agroecosystem compared to native sod. We apply this theory to explain the decreasing SOM content in the order NV NT CT at Sidney, Wooster, and KBS (soils that express aggregate hierarchy). However, NV at KBS had much higher slaked aggregate C contents

than NT and CT, probably a result of the difference in history of the NV vs. the NT and CT plots; therefore the NV treatment is ignored in the discussion below. Increasing cultivation intensity led to a loss of C-rich macroaggregates and an increase of C-depleted microaggregates. There were no consistent significant differences in the proportions of rewetted macroaggregates (250-m fractions) among management treatments (Fig. 1), but the rewetted large and small macroaggregate C concentrations differed in the order NV NT CT (Fig. 2). In contrast, the proportions of slaked macroaggregates differed in the order NV NT CT (Fig. 1), but the slaked macroaggregate C concentrations were similar across management treatments (Fig. 2 and Table 3). These observations suggest that the C lost with increasing cultivation intensity is responsible for the higher proportions of stable macroaggregates (i.e., slaked macroaggregates) in the order NV NT CT; it is the C of binding agents that binds individual microaggregates into stable macroaggregates (Tisdall and Oades, 1982; Elliott, 1986). In our study, increasing cultivation intensity increased the proportion of slaked microaggregates, which were depleted in C compared to macroaggregates and increasingly depleted in C with increasing cultivation intensity (Fig. 2). Therefore we conclude that increasing cultivation intensity leads to a loss of C-rich macroaggregates and an increase of C-depleted microaggregates in soils that express aggregate hierarchy. This shift results in a loss of total

Table 4. Grassland-derived aggregate C concentrations and crop-derived aggregate C proportions and concentrations in Sidney, NE, as determined by 13C natural abundance analysis (05 cm; slaked).
Treatment No-tillage Aggregate-size class m 53 53250 2502000 2000 53 53250 2502000 2000 Grassland-derived C Crop-derived C Percentage crop-derived C % 12c 19b 32a 44a 0.4c 8b 39a NA g C kg1 sand-free aggregates 16.57c 2.20b 21.01b 5.00b 25.97a 12.49a 19.05bc 14.85a 20.34b 0.30b 18.72a 1.69b 26.44a 16.72a NA NA

Conventional tillage

Values within a treatment followed by a different lowercase letter are significantly different (P 0.05) according to Tukeys HSD mean separation test. Grassland- and crop-derived C data for the 53250 and 2502000 m aggregate-size class are adopted from Six et al. (1999a). NA not analyzed.

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organic C. The C lost was that which binds microaggregates into macroaggregates. These observations made at Sidney, Wooster, and KBS extend Elliotts results from stubble mulch agroecosystems to NT and CT agroecosystems characterized by 2:1 clay-dominated soils. The aggregate hierarchy theory seems to be less applicable to the Lexington soil (mixed mineralogy) because within management treatments (i) similar total aggregate C, aggregate-associated C, and mineral-associated C concentrations were observed across aggregate-size classes (Table 3), (ii) the proportion of silt- and claysized particles increased the most from the rewetted to slaked aggregate distribution at Lexington (Fig. 1), and (iii) large macroaggregates broke up into silt and clay particles and microaggregates with increasing cultivation intensity, whereas the proportion of small macroaggregates was about the same. Beare et al. (1994) also observed only small differences in aggregate distribution between NT and CT in a kaolinitic soil in Georgia. The largest difference between NT and CT was in the proportions of large macroaggregates, which primarily fell apart into siltand clay-sized particles. The proportions of small macroaggregates were similar across tillage regimes (Beare et al., 1994). As previously mentioned, Feller et al. (1996) and Elliott et al. (1991) did not observe increased C concentrations with increasing aggregate size in 1:1 claydominated soils. The less-pronounced aggregate hierarchy in the Lexington soil is probably a result of the presence of Fe- and Al-oxides and kaolinite (1:1 clay) which contribute to soil stability through electrostatic interactions (Oades and Waters, 1991). We conclude that the Lexington soil does not express as much aggregate hierarchy as the 2:1 clay-dominated soils because of the presence of oxides and 1:1 clays. In capillary-wetted aggregates from a kaolinitic soil, Beare et al. (1994) observed a higher C content in microaggregates than in macroaggregates. We observed the same trend of increasing C content from large macroaggregates to microaggregates in rewetted soils at Lexington. However, at the other sites no significant differences in C content between macroaggregates and microaggregates in rewetted soils within a management treatment were observed. Other authors also found no differences in misted or rewetted aggregate C contents within a treatment (Elliott, 1986; Cambardella and Elliott, 1993). The higher C content observed in nonslaked microaggregates may therefore be specific for soils with 1:1 clay minerals.

aggregates and a gain of C-depleted microaggregates, resulting in an overall loss of SOM C.

ACKNOWLEDGMENTS Thanks are extended to Scott Pavey and Matt Nemeth for their many hours of sieving, weighing, and C and N analysis. Dan Reusss help during the laboratory work is greatly appreciated. We acknowledge the assistance of Drew Lyon (Univ. of Nebraska, Panhandle Research and Extension Center, Scottsbluff) at the Sidney site, Edmund Perfect and Robert Blevins (Univ. of Kentucky, Lexington) at the Lexington site, H.P. Collins and G.P. Robertson (Univ. of Michigan, W.K. Kellogg Biological Station, Hickory Corners) at the Kellogg site, and W.A. Dick (Ohio State Univ., Wooster) at the Wooster site. Comments on the manuscript by three anonymous reviewers and the associate editor are acknowledged. This research was supported by grant DEB-9419854 from the National Science Foundation. REFERENCES
Beare, M.H., P.F. Hendrix, and D.C. Coleman. 1994. Water-stable aggregates and organic matter fractions in conventional- and notillage soils. Soil Sci. Soc. Am. J. 58:777786. Cambardella, C.A., and E.T. Elliott. 1992. Particulate organic matter across a grassland cultivation sequence. Soil Sci. Soc. Am. J. 56: 777783. Cambardella, C.A., and E.T. Elliott. 1993. Carbon and nitrogen distribution in aggregates from cultivated and native grassland soils. Soil Sci. Soc. Am. J. 57:10711076. Dick, W.A., W.M. Edwards, and E.L. McCoy. 1997. Continuous application of no-tillage to Ohio soils: Changes in crop yields and organic matter-related soil properties. p. 171182. In E.A. Paul et al. (ed.) Soil organic matter in temperate agroecosystems: Long-term experiments in North America. CRC Press, Boca Raton, FL. Doran, J.W. 1987. Microbial biomass and mineralizable nitrogen distributions in no-tillage and plowed soils. Biol. Fert. Soils 5:6875. Edwards, L.M. 1991. The effect of alternate freezing and thawing on aggregate stability and aggregate size distribution of some Prince Edward Island soils. J. Soil Sci. 42:193204. Elliott, E.T. 1986. Aggregate structure and carbon, nitrogen, and phosphorus in native and cultivated soils. Soil Sci. Soc. Am. J. 50: 627633. Elliott, E.T., C.A. Palm, D.E. Reuss, and C.A. Monz. 1991. Organic matter contained in soil aggregates from a tropical chronosequence: Correction for sand and light fraction. Agric. Ecosyst. Environ. 34: 443451. El-Swaify, S.A. 1980. Physical and mechanical properties of oxisols. p. 303324. In B.K.G. Theng (ed.) Soils with variable charge. Offset Publ., Palmerston North, New Zealand. Feller, C., A. Albrecht, and D. Tessier. 1996. Aggregation and organic matter storage in kaolinitic and smectitic tropical soils. p. 309359. In M.R. Carter and B.A. Stewart (ed.) Structure and organic matter storage in agricultural soils. CRC Press, Boca Raton, FL. Frey, S.D., E.T. Elliott, and K. Paustian. 1999. Bacterial and fungal abundance and biomass in conventional and no-tillage agroecosystems along two climatic gradients. Soil Biol. Biochem. 31:573585. Hadas, A. 1990. Directional strength in aggregates as affected by aggregate volume and by a wet/dry cycle. J. Soil Sci. 41:8593. Hofman, G., and L. De Leenheer. 1975. Influence of soil prewetting on aggregate instability. Pedologie 25:190198. Jastrow, J.D. 1996. Soil aggregate formation and the accrual of particulate and mineral associated organic matter. Soil Biol. Biochem. 28: 656676. Jastrow, J.D., T.W. Boutton, and R.M. Miller. 1996. Carbon dynamics of aggregate-associated organic matter estimated by carbon-13 natural abundance. Soil Sci. Soc. Am. J. 60:801807. Kemper, W.D., R. Rosenau, and S. Nelson. 1985. Gas displacement and aggregate stability of soils. Soil Sci. Soc. Am. J. 49:2528. Oades, J.M., and A.G. Waters. 1991. Aggregate hierarchy in soils. Aust. J. Soil Res. 29:815828.

CONCLUSIONS
Aggregate hierarchy was observed in soils from Sidney, KBS, and Wooster, all dominated by 2:1 clay mineralogies. The Lexington soil expressed less aggregate hierarchy, which may be due to the presence of oxides and low-activity clays (kaolinite). There was a clear relationship between loss of soil structure and loss of SOM in the soils that expressed aggregate hierarchy. Increasing cultivation intensity induced a loss of C-rich macro-

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Soil Organic Matter Content as a Function of Different Land Use History

M. M. Pulleman,* J. Bouma, E. A. van Essen, and E. W. Meijles ABSTRACT
A regional survey of management and crop type and soil organic matter (SOM) content was conducted in one soil series in the Netherlands (loamy, mixed, mesic, Fluventic Eutrudept). The objective was to determine the effects of land use history on SOM contents in a prime agricultural soil, using available soil survey information and statistical analyses. Soil organic matter content is a relatively stable, integrating soil characteristic that reflects long-term land use and is an important indicator of soil quality. The SOM contents and information about past land use were obtained from 45 fields. Land use history was expressed in terms of (i) tillage; (ii) crop type; and (iii) use of chemical fertilizers, (iv) manure, and (v) biocides, for six successive periods (6331, 3115, 157, 73, 31, and 10 yr before sampling). Only four land use types occurred: conventional-arable, conventionalgrass, organic-arable, and organic-grass. The SOM contents ranged between 17 and 88 g kg1. Regression models of the actual SOM content as a function of crop type and management in the different periods showed that SOM contents were increased under long-term grass or, to a lesser extent, by organic farming, when compared with conventional-arable use. The regression model depends on the nature of land use history in any particular region and on the length of the selected periods, but it provides an easy method to predict SOM content as a function of management in a given soil series. The method can be an alternative to simulation modeling in situations where detailed data records from long-term field experiments are not available.

raditionally, activities of soil survey were focused on the production of soil maps and soil classification systems. Those activities are nearly completed and soil survey expertise is increasingly being used to study sustainable management systems (Bouma, 1988). Because maintenance and improvement of soil quality

Laboratory of Soil Science and Geology, Dep. of Environ. Sci., Wageningen Agricultural Univ., P.O. Box 37, 6700 AA Wageningen, the Netherlands. Received 22 Mar. 1999. *Corresponding author (mirjam. pulleman@oio.beng.wau.nl). Published in Soil Sci. Soc. Am. J. 64:689693 (2000).

is critical to sustaining agricultural productivity (Reeves, 1997), the soil quality concept recently has received much attention. Karlen et al. (1997) defined soil quality as the capacity of a specific kind of soil to function, within natural or managed ecosystem boundaries, to sustain plant and animal productivity, maintain or enhance water and air quality, and support human health and habitation. In order to express soil quality, different indicators have been used. They can be static, using soil parameters such as bulk density, porosity, and SOM content (Lal and Kimble, 1997), or dynamic, using simulation modeling (Bouma and Droogers, 1998). A given soil series cannot be considered to have a static set of characteristics. Land use influences soil properties and therefore soil functioning. In this respect, Droogers and Bouma (1997) distinguished between soil genoform and soil phenoform. The former has been defined as the genetically defined soil series, and the latter indicates differences in a certain genoform as a result of a different land use history. They found that different types of agricultural land use have resulted in different soil conditions within one soil series in marine clay deposits in the southwestern part of the Netherlands (a loamy, mixed, mesic, Fluventic Eutrudept; Soil Survey Staff, 1998). Based on significant differences in total SOM contents, bulk densities, and porosities within one genoform, three different phenoforms were distinguished, resulting from: (i) organic management, (ii) conventional management, and (iii) permanent grassland. Simulation modeling was used to translate differences in static soil properties into a dynamic quality indicator, which was expressed as the ratio between actual production and potential production (Bouma and Droogers, 1998). Criteria to characterize phenoforms and soil quality are still poorly defined. Soil structure parameters and
Abbreviations: SOM, soil organic matter.