Geoderma 124 (2005) 3 – 22
www.elsevier.com/locate/geoderma
Soil structure and management: a review
C.J. Bronick, R. Lal *
Carbon Management and Sequestration Center, OARDC/FAES, School of Natural Resources, The Ohio State University,
2021 Coffey Road, Columbus, OH 43210-1085, USA
Available online 27 April 2004
Abstract
Soil structure exerts important influences on the edaphic conditions and the environment. It is often expressed as the degree of
stability of aggregates. Aggregation results from the rearrangement, flocculation and cementation of particles. It is mediated by soil
organic carbon (SOC), biota, ionic bridging, clay and carbonates. The complex interactions of these aggregants can be synergistic
or disruptive to aggregation. Clay-sized particles are commonly associated with aggregation by rearrangement and flocculation,
although swelling clay can disrupt aggregates. Organo-metallic compounds and cations form bridges between particles. The SOC
originates from plants, animals and microorganisms, and their exudates. It enhances aggregation through the bonding of primary
soil particles. The effectiveness of SOC in forming stable aggregates is related to its decomposition rate, which in turn is influenced
by its physical and chemical protection from microbial action. Soil inorganic carbon (SIC) increases aggregation in arid and semiarid environments, and the formation of secondary carbonates is influenced by the presence of SOC and Ca2 + and Mg2 +. Soil biota
release CO2 and form SOC which increase dissolution of primary carbonates while cations increase precipitation of secondary
carbonates. The precipitation of (hydr)oxides, phosphates and carbonates enhances aggregation. Cations such as Si4 +, Fe3 +, Al3 +
and Ca2 + stimulate the precipitation of compounds that act as bonding agents for primary particles. Roots and hyphae can enmesh
particles together while realigning them and releasing organic compounds that hold particles together, a process with a positive
impact on soil C sequestration. Soil structure can be significantly modified through management practices and environmental
changes. Practices that increase productivity and decrease soil disruption enhance aggregation and structural development.
D 2004 Elsevier B.V. All rights reserved.
Keywords: Soil aggregation; Soil structure; Aggregate dynamics; Land use management; Carbon sequestration
1. Introduction
Abbreviations: AMF, arbuscular mycorrhizal fungi; CEC, cation
exchange capacity; CI, inert carbon; CL, labile fraction; Cl, clay;
Cmic, C attributed to microbial biomass; CR, recalcitrant C; DOC,
dissolved organic matter; ESP, exchangeable sodium percentage;
FA, fulvic acid; HA, humic acid; HS, humic substance; LF, light
fraction; MWD, mean weight diameter; OM, organic matter; OOC,
occluded organic C; OOM, occluded organic matter; P, polyvalent
cations; POM, particulate organic matter; SIC, soil inorganic C;
SOC, soil organic C; SOM, soil organic matter; WSA, waterstable
aggregates.
* Corresponding author. Tel.: +1-614-292-9069; fax: +1-614292-7432.
E-mail address:
[email protected] (R. Lal).
0016-7061/$ - see front matter D 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.geoderma.2004.03.005
Soil structure is a key factor in the functioning of
soil, its ability to support plant and animal life, and
moderate environmental quality with particular emphasis on soil carbon (C) sequestration and water quality.
Aggregate stability is used as an indicator of soil
structure (Six et al., 2000). Aggregation results from
the rearrangement of particles, flocculation and cementation (Duiker et al., 2003). Aggregation is mediated by
soil organic carbon (SOC), biota, ionic bridging, clay
and carbonates. The SOC acts as a binding agent and as
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C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
a nucleus in the formation of aggregates. Biota and
their organic products contribute to the development of
soil structure; which in turn exert a significant control
over SOC dynamics. The SOC residence time and
decomposition rate are key factors influencing its
effectiveness in increasing aggregation. Crystalline
and amorphous metal oxides and hydroxides are important aggregants in soils. Metal ions form bridges
between mineral and organo-mineral particles. Clay
also acts as an aggregant, binding particles together and
influencing SOC decomposition and turnover. Longterm stability of aggregates is often related to the
presence of recalcitrant C (CR) compounds and metal
ions. Formation of secondary carbonates in arid and
semi-arid regions is also linked to aggregate dynamics.
The decline in soil structure is increasingly seen as a
form of soil degradation (Chan et al., 2003) and is often
related to land use and soil/crop management practices.
Soil structure influences soil water movement and
retention, erosion, crusting, nutrient recycling, root
penetration and crop yield. Externalities such as runoff,
surface- and ground-water pollution and CO2 emissions are influenced by soil structure. Reduction in
tillage and fertilization can reduce CO2 emissions by
reducing cultivation and production inputs both of
which are dependant on fossil fuels. Species biodiversity is affected by management practices; generally
high-input agricultural practices decrease biodiversity
while the low-input practices enhance biodiversity
(Munyanziza et al., 1997; Lupwayi et al., 2001). With
increases in population and urbanization, it is important
to identify methods to enhance food production while
maintaining environmental quality.
This review collates and synthesizes available
information on soil structure and its dynamics in
relation to management. The emphasis of this review
is on a wide range of environmental and anthropogenic factors, and the dynamic interaction among
them. Soil structure is discussed in relation to the
environment, soil biological components, soil properties, and soil management practices with the intent of
better understanding soil structural dynamics.
2. Soil structure
Soil structure refers to the size, shape and arrangement of solids and voids, continuity of pores and
voids, their capacity to retain and transmit fluids and
organic and inorganic substances, and ability to support vigorous root growth and development (Lal,
1991). Favorable soil structure and high aggregate
stability are important to improving soil fertility,
increasing agronomic productivity, enhancing porosity and decreasing erodibility.
2.1. Basic concepts of aggregation
Aggregates are secondary particles formed through
the combination of mineral particles with organic and
inorganic substances. The complex dynamics of aggregation are the result of the interaction of many
factors including the environment, soil management
factors, plant influences and soil properties such as
mineral composition, texture, SOC concentration,
pedogenic processes, microbial activities, exchangeable ions, nutrient reserves, and moisture availability
(Kay, 1998). Aggregates occur in a variety of manners
and sizes. These are often grouped by size: macroaggregates (>250 Am) and microaggregates ( < 250
Am) with these groups being further divided by size
(Tisdall and Oades, 1982). Different size groups differ
in properties such as binding agents and carbon and
nitrogen (N) distribution.
2.2. Mechanisms of aggregation
There are several mechanisms of aggregation. Aggregates are formed in stages, with different bonding
mechanisms dominating at each stage (Tisdall and
Oades, 1982). Hierarchical theory of aggregation proposes that microaggregates join together to form macroaggregates and the bonds within microaggregates are
stronger than the bonds between microaggregates
(Edwards and Bremner, 1967). Microaggregates
( < 250 Am) are formed from organic molecules (OM)
attached to clay (Cl) and polyvalent cations (P) to form
compound particles (Cl – P – OM), which are joined
with other particles (Cl –P – OM) to form macroaggregates [(Cl – P –OM)x]y (Edwards and Bremner, 1967;
Tisdall, 1996). Alternatively, macroaggregates can
form around particulate organic matter (POM). As
POM is decomposed and microbial exudates are released, the macroaggregate becomes more stable, the
C:N ratio decreases, and microaggregates form inside.
The internally formed microaggregates contain more
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
recalcitrant SOC pool (Plante and McGill, 2002a;
Beare et al., 1994). As the more labile SOC pool is
utilized and microbial activity decreases, the supply of
exudates decreases and the macroaggregate loses stability, eventually disrupts and releases more stable
microaggregates (Jastrow et al., 1998; Six et al., 1999).
Roots and hyphae enmesh and release organic
compounds that act as glue to hold particles together.
Particles can be rearranged during enmeshment,
while wet – dry cycles help to stabilize the aggregates. Bacterial microaggregates form as bacterial
colonies and their exudates form a polysaccharide
capsule around which clay particles are aligned and
pulled in by drying and shrinkage. The clay shell
forms a protective coating for the bacterial colony
inhibiting decomposition of the SOC inside. Concentric theory of aggregation suggests that external
layers are concentrically built upon the external
surface of the aggregate, with younger C in outer
layers of aggregates than in aggregate interiors
(Santos et al., 1997).
The precipitation of (hydr)oxides, phosphates and
carbonates enhances aggregation. Cations such as
Si4 +, Fe3 +, Al3 + and Ca2 + stimulate the precipitation
of compounds that act as bonding agents for primary
particles. Cations also form bridges between clay and
SOM particles resulting in aggregation. Dissolved
organic compounds (DOC) can complex with Fe3+
and Al3+ at low pHs forming mobile, organo-metallic
compounds that can be precipitated elsewhere in the
soil. The complexation decreases microbial access to
SOC and mineralization.
It is possible that aggregates form through a
combination of these processes (Fig. 1). Macroaggregates may initially form through accumulation of
Fig. 1. Some possible scenarios of aggregation. Organic matter
(OM), particulate organic matter (POM), clay (Cl), particle (P).
5
Fig. 2. Factors affecting soil aggregation.
microaggregates or around POM or bacterial cores,
decomposing or breaking down later into microaggregates. Microaggregates may initially form by
the progressive bonding of clay, SOM and cations,
or as turnover products from macroaggregates. Primary particles can accumulate on outer layers of
aggregates.
2.3. Aggregate dynamics and turnover
The on-going interactive effects of soil-forming
processes, soil properties and exogenous factors
such as terrain and climate establish a dynamic
equilibrium in soil structure (Fig. 2). Aggregates
can be disrupted by a variety of mechanisms
depending on the nature of the bonding agents.
The activities of soil organisms influence C retention time and turnover in soil, which in turn affect
C stabilization, aggregation and turnover. Decomposition is effected by the activity of soil organisms,
soil properties and environmental factors such as
temperature, gaseous concentration, nutrient availability and moisture gradients (Christensen, 2001).
The C:N ratio is commonly used as an indicator of
SOC turnover, although for CR and inert C (CI)
fractions, the lignin/N ratio or other more resistant
compounds may be more appropriate (Vanlauwe et
al., 1997). Feedback within the system suggests that
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C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
decomposition of SOM is affected by soil structure,
through porosity, gaseous exchange and soil moisture, as well as by the physical location of C such
as depth and occlusion. Turnover dynamics vary
spatially within the soil and within individual aggregates as well as over time. The SOC dynamics may
be slower towards the end of the growing season,
resulting in slow turnover rates in macroaggregates
(Plante and McGill, 2002b). Inorganic compounds,
low-activity clays and CR bonding agents are generally resistant to breakdown. The mobilization and
precipitation of substances can promote the dissolution and breakdown of aggregates as well as the
formation of new aggregates. Ingestion by soil
fauna can disrupt aggregates, although it generally
increases aggregate stability (Schrader and Zhang,
1997). Aggregates are also susceptible to disruption
by physical disturbances such as clay swelling,
tillage and rainfall impact.
2.4. Pedogenic processes
Soil structural development and aggregation occur within the context of natural pedogenic processes and anthropogenic activities. The complex series
of additions and losses from the soil, action of
organisms and climatic effects determine soil properties. Weathering alters materials, which are translocated within soils through leaching, bioturbation,
eluviation, and illuviation resulting in horizonation.
Dissolved organic matter from eluvial horizons is
immobilized in illuvial, B horizons (De Coninck,
1980; Guggenberger and Kaiser, 2003; Jansen et al.,
2003). Precipitated materials increase binding in the
B horizons.
2.5. Soil structure and plant growth
Soil structure affects plant growth by influencing
root distribution and the ability to take up water and
nutrients (Rampazzo et al., 1998; Pardo et al., 2000).
Soil structure facilitates oxygen and water infiltration
and can improve water storage (Franzluebbers,
2002). Increased water transfer through soil can
reduce fertilizer retention in the soil matrix and
fertilizer use efficiency in plants (Franzluebbers,
2002). Disturbance of soil structure through compaction or tillage can result in the rapid recycling of
nutrients, crusting, reduced water and air availability
to roots.
2.6. Climate and exogenous factors
Climate and landscape position influence soil
structure by factors such as temperature, precipitation,
elevation, slope gradient, and aspect. Soil properties
such as texture, mineralogy, SOC and organisms
moderate the influence of the climate.
2.6.1. Climate
Climate affects soil aggregation through alterations
in temperature and moisture regimes and wet –dry and
freeze –thaw cycles, which can reorient particles (Singer et al., 1992) possibly resulting in improved aggregation and increased isolation of SOC within aggregates. Changes in temperature and moisture levels
affect microbial and biotic activity, which alter decomposition rates. The relationship between temperature
and decomposition is highly variable due to the influence of a variety of other factors. Warmer temperatures
result in higher respiration and biological activity in
soil, while lower temperatures result in higher standing
stock of SOC (Franzluebbers et al., 2001). Frigid and
wet soils tend to have more unavailable SOC than
warm and dry soil (Franzluebbers et al., 2001). In
moist, temperate regions, aggregation is affected by
freeze –thaw cycles (Dalal and Bridge, 1996).
Soils undergo continuous changes in moisture
regime. Under rainfed conditions, wet – dry cycles
are most commonly related to climatic factors. On a
localized level, water uptake by plant roots as influenced by evapotranspiration can lead to drying in the
root zone. Soil moisture and wet –dry cycles have a
variable effect on aggregation. Wet –dry cycles can
disrupt aggregation in swelling clays. As clay particles
swell they separate from other particles, decreasing
aggregate stability (Singer et al., 1992). Wet – dry
cycles have a more positive influence in the initial
stages in soils with non-swelling clays and in macroaggregates (Fig. 3). During wetting, clay particles tend
to disperse and then form bridges and coatings while
drying (Attou and Bruand, 1998). This leads to closer
contact between particles and increased clay bridging
(Singer et al., 1992). Wet –dry cycles also affect the
amount of POM incorporated into aggregates (Denef
et al., 2001) and porosity (Czarnes et al., 2000). Wet –
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
7
Fig. 3. Wet – dry cycles on aggregate stability for 0.5 – 1.0 mm aggregates with 25% kaolinite or smectite clay (redrawn from Singer et al., 1992).
dry cycles are important to aggregation in soils of
arid, semi-arid and subhumid regions (Dalal and
Bridge, 1996).
Arid conditions have variable effects on aggregation. Factors such as carbonates, earthworms and
crusting can increase aggregate stability in arid environments (Boix-Fayos et al., 2001). While crusting
reduces water infiltration, it also reduces detachment
and erosion, which has a positive impact on aggregation (Amezketa, 1999). Some soils of arid regions
have higher levels of aggregation and stable microaggregates than those in humid regions of the Mediterranean (Boix-Fayos et al., 2001). Decrease in soil
moisture and the ensuing reduced vegetation can
result in decreased structural development and aggregation and increased erosion. Decreased aggregate
stability may be due to increased erosion and runoff
resulting in lower SOC, clay content, and cation
exchange capacity (CEC) (Boix-Fayos et al., 1998).
Temperature and moisture regimes can be modified
by management practices such as irrigation, cover
cropping, and mulching. Conventional tillage increases
exposure to air, sun, and wind. Management practices
moderate the impact of wet – dry cycles, no-till soils
experience less intense wet – dry cycles due to protection by surface residue (Denef et al., 2001). Dispersion
or slaking caused by wet –dry cycles can be reduced by
amending soils with humic substances (Piccolo et al.,
1997).
2.6.2. Terrain
Geographic region, elevation, aspect and slope
gradient influence vegetation and erosion. Elevation
influences the rate of weathering in soils, having an
indirect effect on soil structure. North-facing slopes in
the Mediterranean soils have higher aggregation than
south-facing slopes, perhaps due to vegetative differences caused by differences in micro-climate (BoixFayos et al., 1998). The increased aggregate stability
and vegetation feeds back to increased infiltration and
reduced erosion (Boix-Fayos et al., 1998). Sloping
soils are more susceptible to erosion, particularly in
regions of intense rainfall. Erosion tends to preferentially remove low density or light particles including
clay and SOC that are two of the primary bonding
agents in aggregation. Erosion of SOC may also
increase the rate of mineralization (Jacinthe et al.,
2002a).
2.7. Soil properties
2.7.1. Soil types
Aggregation is controlled by different mechanisms
in different soil types (Table 1). The rate and stability
of aggregation generally increases with SOC and clay
surface area and CEC. In soils low in SOC or clay
concentration, aggregation may be dominated by
cations, while the role of cations in aggregation
may be minimal in soils with high SOC or clay
concentration. In Oxisols and Ultisols Al – humus
complexes and noncrystalline Al3 + hydroxides are
predominant aggregants as these compounds are able
to protect SOC from microbial decomposition and
stabilize aggregation. Aridisols display high aggregate stability associated with carbonates (Boix-Fayos
et al., 2001).
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C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
containing polyvalent cations (Ca2 +, Al3 + and Fe3 +)
are resistant to slaking (Tisdall, 1996).
Table 1
Soil types and aggregation factors
Soil orders
Aggregation factors
Authors
Alfisols
SOM
Andisols
Allophane clay,
non-crystalline clay
SIC, carbonates and
weathered clay minerals
Dalal and Bridge, 1996;
Oades and Waters, 1991
Torn et al., 1997
Aridisols
Entisols
Inceptisols
Oxisols
SOM
Amorphous clay
Al3 + and Fe3 + oxides
Ultisols
Non-crystalline
Al hydroxides
Plant roots and
rhizosphere
Hydrophobic SOM
Organo-metallic
complexes
Metallic hydroxides
SOM
Vertisols
Non-crystalline
sesquioxides
Clay-size fractions
Spodosols
Boix-Fayos et al., 1998;
Boettinger and Southard,
1995
Dalal and Bridge, 1996
Dalal and Bridge, 1996
Oades and Waters, 1991;
Dalal and Bridge, 1996
De Coninck, 1980
Dalal and Bridge, 1996;
Zhang and Horn, 2001
Leinweber et al., 1999;
Dalal and Bridge, 1996
Polycationic bridges,
wet – dry cycles
SOM, soil organic matter; SIC, soil inorganic matter.
2.7.2. Texture
Soil texture has a significant influence on aggregation. In coarse-textured soils, the SOC has a greater
influence on structure; while with increasing clay
content the type of clay is more important than the
amount in determining aggregation (Kay, 1998). Clay
concentration physically affects aggregation through
swelling and dispersion. The potential of swellinginduced disaggregation is reduced at low clay levels
(Attou et al., 1998). Increasing clay concentration is
associated with increased SOC stabilization (Sollins et
al., 1996).
2.7.3. Cation exchange capacity (CEC)
The CEC is often related to stable aggregates
(Dimoyiannis et al., 1998). Aggregation is stimulated
by the interaction of polycationic bridging in which
the repulsive forces between negatively charged clay
and/or SOC are reduced (Tisdall, 1996). Aggregates
2.7.4. Soil pH
In addition to effects on plant growth, metal ion
solubility, microbial activity and clay dispersion are
also influenced by soil pH (Haynes and Naidu,
1998). The negative surface charge on clay particles
increases with pH increasing particle repulsion.
Therefore, controlling soil pH is important in soils
with dispersive clays (Chorom et al., 1994), as clay
particles often flocculate at high pH values (Haynes
and Naidu, 1998). Large aggregates form in soils of
high pH and high carbonate concentration (BoixFayos et al., 2001). Lime is commonly added to soil
to increase pH often resulting in increased microbial
activity and crop yields, and contributing to higher
SOM and increased aggregation (Haynes and Naidu,
1998).
2.7.5. Porosity
A wide range of pore sizes exists in well-aggregated soils both between and within aggregates (Dalal
and Bridge, 1996). Large pores (>30 Am) include
biopores, cracks and interaggregate pores. Pore space,
size and amount can influence SOC and its turnover;
conversely, SOC and soil texture can influence porosity (Thomsen et al., 1999). Biopores are important
for improving gaseous and water diffusion influencing
decomposition. Small pores can protect SOC from
decomposition by limiting microbial access and
through control of gas diffusivity and water availability (Dalal and Bridge, 1996; Kay, 1998; Thomsen et
al., 1999, 2003). Clay dispersion, tillage, compaction
and crusting can result in decreased porosity (Dalal
and Bridge, 1996). In swelling soils, porosity is
related to soil moisture content and swell/shrink
characteristics (Kay, 1998). Tillage causes short-term
increases in porosity, but long-term decreases in
aggregation.
2.7.6. Soil water
Soil structure and texture influence soil water flow,
availability and storage (Pachepsky and Rawls, 2003).
Aggregation and interconnected pores increase bypass
flow in soil; this can result in increased infiltration and
reduced runoff, the movement of water deeper into the
soil profile and increased leaching (Franzluebbers,
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
2002; Nissen and Wander, 2003). Reduced matrix
flow can lead to water stress in arid conditions (Nissen
and Wander, 2003).
2.8. Agents of aggregation
2.8.1. Carbon
The source of C, whether it is SOC or SIC,
influences its composition and concentration in soil,
which in turn influences its effectiveness in aggregation through associations with cations and soil
particles. The composition of SOC reflects its decomposition rate and release of cations into soil
solution as well as its ability to complex with cations
in soil solution.
2.8.1.1. Soil inorganic carbon. The SIC exists in
soil as primary and secondary minerals. Primary or
lithogenic carbonates originate from parent rock material. Primary carbonates are the source material for
the formation of secondary carbonates when they are
dissolved and translocated by water with organic acids
and/or CO2 from soil and atmosphere (Fig. 4). Secondary or pedogenic carbonates form when dissolved
CO2 precipitates carbonate and bicarbonate with Ca2 +
and Mg2 + from outside the system. Under conditions
of decreased moisture or increased pH, cations, bicarbonate (HCO3 ), dissolved carbonates and CO2 can
react with available cations to form secondary carbonate coatings on primary soil particles.
The effect of carbonates on structure is moderated
by SOC. Increases in SOC results in increased dissolution and reprecipitation of carbonates in soil. The
Fig. 4. Relationship between carbonates and CO2 in the soil via
weathering and fixation.
9
SOC increases microbial respiration and CO2 and is a
source of Ca2 + and Mg2 +. At low SOC concentration,
macroaggregate stability is enhanced by carbonates
(Boix-Fayos et al., 2001). High carbonate concentration enhances SOC protection, probably because of
decreased SOC mineralization and increased Ca2+
(Clough and Skjemstad, 2000). High carbonate concentration in the silt fraction decreases aggregation,
suggesting that particle size may also influence the role
of carbonates in aggregation (Dimoyiannis et al.,
1998). Carbonates may increase tensile strength of
aggregates but decrease stability of microaggregates
(Boix-Fayos et al., 2001; Schrader and Zhang, 1997).
Organisms influence carbonate-mediated aggregation:
earthworm activity can alter carbonate concentration,
although the effects are inconsistent and may depend
on earthworm species (Schrader and Zhang, 1997).
Irrigation and fertility management increase SOC
thereby accelerating formation of secondary carbonates in arid and semi-arid soils through the addition
of carbonic and organic acids that can react with soil
silicates to sequester C (Schwartzman et al., 1994)
(Fig. 5).
2.8.1.2. Soil organic carbon. The SOC creates
regions of heterogeneity in the soil, leading to ‘‘hot
spots’’ of aggregation. Increased SOC is related to
increased aggregation (Fig. 6). The chemical properties of SOC determine their charge and complexation
capacities and influence decomposition rates which
have direct effects on aggregation (Schulten and
Leinweber, 2000). The aggregate binding effect of
labile SOC is rapid but transient (Kay, 1998) while
slower decomposing SOC has subtler effects on
aggregation, but the effects may be longer lived
(Martens, 2000). The isolation of the readily decomposable labile C (CL) fraction within aggregates
increases stability and durability by reducing its
decomposition (Fig. 6). Increases in microbial biomass are associated with increases in aggregate stability (Haynes and Beare, 1997).
Particulate organic matter. The POM is comprised of large particles of organic matter (250 –2000
Am) that exist as free POM light fraction (LF) or
encrusted with soil particles, which in turn offers
physical protection from decomposition (Plante and
McGill, 2002a). The LF in soil is generally associated
with clay and polyvalent cations to form aggregates
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C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
Fig. 5. SOC and MWD (redrawn from Chenu et al., 2000). SOC, soil organic carbon; MWD, mean weight diameter.
(Jastrow, 1996). The increased SOC and aggregation
in no-till soils may be due to increases in LF-POM
(Six et al., 1999). The POM can act as a nucleus for
macroaggregate formation with material accumulating
around the POM. Macroaggregates have a large
concentration of low density POM (Jastrow, 1996).
Within macroaggregates, the decomposition of CL in
occluded organic matter (OOM) may lead to relative
enrichment of CR. Disruption of OOM within aggregates results in the exposure of CL, making it available
for microbial decomposition (Plante and McGill,
2002a). The POM may be an important agent in
binding microaggregates to form macroaggregates.
As microorganisms decompose the POM, they produce extracellular polysaccharides that act as a binding agent (Jastrow, 1996).
Carbohydrates. The role of carbohydrates in
improving soil structure is variable, relating to the
source and nature of the carbohydrates as well as
aggregate properties and environmental factors. Carbohydrates originating from plants are often coarser
sized and occur in the sand fraction, while carbohydrates produced by microbial activities are finer and
are present in clay and silt fractions (Schulten and
Leinweber, 2000). The microbially derived carbohydrates tend to be resistant to decomposition. Reduced
tillage (conservation till or no-till), manure additions
and cover crops are associated with increased concentrations of carbohydrates and improved structure
in soil (Shepherd et al., 2001; Debosz et al., 2002b).
Polysaccharides. Polysaccharides are readily
mineralizable and act as transient binding agents
Fig. 6. SOC pool flow chart. SOC, soil organic carbon; CI, inert carbon; CL, labile fraction; CR, recalcitrant C.
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
initiating aggregation, but may not have long-term
stability (Kay, 1998; Tisdall and Oades, 1982). Polysaccharides are strongly adsorbed onto mineral surfaces, acting as ‘‘bridges’’ to bind soil particles (Kay,
1998; Martens, 2000). Polysaccharides can also form
a gel-like substance that acts as a glue to bind particles
into aggregates (Haynes and Beare, 1997).
Phenols. Phenolic molecules can complex with
cations to form cationic bridges. Phenols and polyphenols are precursors of humic substances and increase
aggregation. Lower aggregation in soils growing soybeans is attributed to low phenolic contents in the
residues, and high aggregation is often associated with
plants high in phenols (Martens, 2000). Phenolic acids
increase aggregation interactions with microbial products (Martens, 2000).
Lignin. The low rate of lignin decomposition is
considered a key factor in C turnover rates, enhancing
aggregation (Palm and Rowland, 1997; Magill and
Aber, 1998). Mucilage and other compounds from
lignin decomposing fungus contribute to soil aggregation (Caesar-TonThat, 2002).
Lipids. In soils, lipids are often enriched in the
clay fraction (Schulten and Leinweber, 2000). In some
soils lipids can improve aggregate stability against
slaking and dispersion, probably related to their hydrophobic nature (Dinel et al., 1997; Pare et al., 1999).
The effectiveness of lipids in improving aggregate
stability may be related to clay mineralogy, lipids play
a more important role in bonding in illite and mixed
clay soils than in kaolinite- or smectite-dominated
soils (Dinel et al., 1997). Increased total extractable
lipids are present in plow-till as compared to no-till
soils, which generally exhibit higher levels of SOM
biodegradability (Pare et al., 1999).
Humic substances. The recalcitrance of humic
substances (HS) is a result of their chemical resistance
and their association with the soil matrix, which offers
physical protection. Tarchitzky et al. (2000) hypothesized that the distribution of HS, including fulvic acid
(FA), humic acid (HA) and humin, in aggregates may
reflect the initial coating of particles with HS than
additional HS serving to bond particles together. The
HAs can be adsorbed onto clay particles by polyvalent
cations, making them especially effective in overcoming clay dispersion. The HSs can increase aggregate
stability and reduce dispersion due to wet –dry cycles
(Piccolo et al., 1997).
11
Hydrophobicity. The hydrophobicity of SOC,
along with its charge properties and the interaction
with soil particles, affects soil physicochemical properties including structural stability (D’Acqui et al.,
1999). Hydrophobic molecules cause water repellency
in soil aggregates and decrease wettability, which may
result in increased cohesiveness, decreased dispersivity and reduced SOC decomposition rates. Long-term
aggregate stability can be enhanced by the addition of
hydrophobic organic materials such as organic wastes
containing hydrophobic materials and SOC from plant
types such as Cistus rather than Pinus (Ternan et al.,
1996). No-till (Hallett et al., 2001) and burning
(Poulenard et al., 2001) affect soil hydrophobicity.
High temperature burning may result in water repellency (Poulenard et al., 2001) and increased structural
stability.
2.8.2. Clay and clay mineralogy
Clay minerals influence properties that affect aggregation: surface area, CEC, charge density, dispersivity and expandability, and these in turn affect SOC
decomposition rates (Dimoyiannis et al., 1998; Schulten and Leinweber, 2000). The interaction of clay,
SOC and aggregates is affected by soil pH, CEC, ions
(Na+, Ca2 +, Mg2 +), all of which are related to the
amount and type of clay present in the soil (Amezketa,
1999). Clay mineralogical composition is modified
with soil development (Powers and Schlesinger,
2002). Low-activity clays such as kaolinite and halloysites are often present in Alfisols, Ultisols and
Oxisols while high-activity clays with smectites are
present in Vertisols.
Clays occur as crystalline, layered structures or as
non-crystalline minerals with amorphous structure. In
some soils non-crystalline clay is an important factor
for aggregation, such as in volcanic soils where SOC
and aggregation are associated with allophanic clay
(Powers and Schlesinger, 2002). Non-crystalline clay
minerals, such as allophane and imogolite, have high
surface areas, and highly variable and pH-dependant
charge properties that generally increase aggregation
(Powers and Schlesinger, 2002). Non-expanding, crystalline clays, such as kaolinite (1:1), have low CEC and
surface area, which tend to decrease aggregate stability.
Kaolinite has a good flocculation capacity due to
electrostatic charges between platelets and SOC. In
comparison, aggregation is generally high in high-
12
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
activity clays such as smectites and other 2:1 clays,
which are associated with high CEC, large surface
areas and high SOC (Seta and Karathanasis, 1996;
Amezketa, 1999; Schulten and Leinweber, 2000; Six
et al., 2000). The expandability of smectites can disrupt
aggregates during wet –dry cycles. Illitic and smectitic
clays tend to decrease aggregate stability during wet –
dry cycles although there are variable effects possibly
due to the amount of clay and number of shrink – swell
cycles (Piccolo et al., 1997; Singer et al., 1992).
Dispersivity of soil aggregates is affected by the
nature of SOC and the clay type. Cations, particularly
Ca2 + and Na+, electrolytes and pH (Chorom et al.,
1994; Haynes and Naidu, 1998) can influence clay
dispersion. At high concentrations, NH4+ fertilizer can
lead to dispersion of clay (Haynes and Naidu, 1998).
The addition of gypsum to soil decreases clay dispersivity through changes in the electrolyte concentration and composition (Sumner, 1993; Baldock et
al., 1994). The CEC and surface area of clay minerals,
SOC and other soil properties interact to form intraparticle interactions that influence dispersivity (Nelson et al., 1999). Less reactive clays, like kaolinite,
are less dispersive. The SOC and some roots can
increase dispersion of clay particles while others have
the opposite effect (Tisdall, 1996). Negatively charged
SOC can be disruptive to aggregate structure, possibly
by repelling negative charges in the soil minerals
(D’Acqui et al., 1999). Differences in SOC concentrations exist between dispersive and non-dispersive
clays, suggesting that decomposing organic residues
may promote dispersion (Baldock et al., 1994). Less
dispersive clay contains more aliphatic materials in
the topsoils and carbohydrates in the subsoil. These
compounds may act as ‘glue’ to hold particles together (Nelson et al., 1999). Easily dispersed clays contain
higher concentrations of amino acids which may act
as dispersants (Nelson et al., 1999).
Soils dominated by variable charge clay minerals
such as 1:1 clay and oxides have higher aggregation at
lower SOC levels whereas soils with mixed mineralogy clays have higher aggregation at higher SOC
(Denef et al., 2002). Clay type also affects the rate
of decomposition of SOM. Montmorillonite results in
a rapid decomposition of SOC with the production of
more low molecular weight FAs compared to kaolinite, which results in slow decomposition and more
residual C and HAs (D’Acqui et al., 1998). Low pH
levels associated with chlorite may result in low
decomposition rates, accounting for high C associated
with these clays (Schulten and Leinweber, 2000).
2.8.3. Cations
2.8.3.1. Calcium and magnesium. Bivalent Ca2 +
and Mg2 + cations improve soil structure through
cationic bridging with clay particles and SOC. As
previously mentioned, in arid and semi-arid conditions, Ca2 + and Mg2 + carbonates precipitate to form
secondary carbonate coatings and bind primary soil
particles together. Generally, Ca2 + is more effective
than Mg2 + in improving soil structure (Zhang and
Norton, 2002). Among bivalent cations, Ca2 + can
inhibit clay dispersion and the associated disruption
of aggregates by replacing Na+ and Mg2 + in clay and
aggregates, thereby adding to aggregate stability
(Armstrong and Tanton, 1992).
In comparison to Ca2 +, Mg2 + may have a deleterious effect on soil aggregate stability by increasing
clay dispersion (Zhang and Norton, 2002). The extent
of the negative effect of Mg2 + compared to Ca2 + may
depend on the type of clay and electrolyte concentration in the soil (Zhang and Norton, 2002). Further,
Mg2 + may result in high swelling by expanding clays,
resulting in disruption of aggregates.
The use of soil amendments containing Ca2 + and
Mg2 +, such as lime and gypsum, can have profound
effects on aggregation. Increased aggregate stability in
limed soils suggests formation of strong bonding
involving Ca2 + bridges (Chan and Heenan, 1999a).
2.8.3.2. Iron and aluminum. Polyvalent Al3 + and
Fe3 + cations improve soil structure through cationic
bridging and formation of organo-metallic compounds
and gels (Amezketa, 1999). The solubility and mobility of these cations is pH-dependent, with higher
solubility at lower pH. Aggregates containing Al3 +
and Fe3 + and high-CEC clays tend to increase SOC
incorporation. The interaction of Al3 + and Fe3 + with
kaolinite can synergistically promote aggregation with
limited impact on SOC (Six et al., 2000) while oxides
and hydroxides of Al3 + interact synergistically with
SOC and dispersible clay to improve aggregate stability (Molina et al., 2001). Both Al3 + and Fe3 +
control aggregation in acidic soils with low clay and
SOC contents, such as Oxisols (Oades and Waters,
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
1991; Barral et al., 1998). Amorphous Fe3 + and SOM
form fine stable particles in soils with high SOC
content (Barral et al., 1998). Coarse-grained particles
are formed from Fe3 + oxides (Barral et al., 1998),
which increase the tensile strength of aggregates in
Oxisols and improve aggregate stability in others
(Barral et al., 1998).
2.8.3.3. Phosphorus. The effects of P on soil aggregation maybe indirect, as P availability affects shoot
and root growth, and increases plant production and
ground cover. The availability of P also influences
colonization of arbuscular mycorrhizal fungi (AMF)
(Facelli and Facelli, 2002), which affect root morphology and aggregation. Application of P as fertilizer
and phosphoric acid can lead to the formation of Al3 +
or Ca2 + phosphates, which act as aggregate bonding
agents (Haynes and Naidu, 1998).
2.8.3.4. Sodium. The Na+ is a highly dispersive agent
resulting directly in the breakup of aggregates and
indirectly affecting aggregation through decreased
plant productivity. Exchangeable Na+ in the soil solution and at exchange sites contribute to repulsive
charges that disperse clay particles. Increased dispersivity from Na+ can break up aggregates, making SOM
more available for decomposition. Sodic soils occur
mostly in arid and semi-arid regions. Management
practices that replace Na+ from cation exchange sites
with Ca2 + help to ameliorate high Na+ and its adverse
affects. Gypsum is used as a soil amendment to
overcome sodicity by reducing dispersion, pH and
exchangeable sodium percentage, (ESP) and increase
in microbial C (Cmic) (Armstrong and Tanton, 1992;
Batra et al., 1997). In calcareous-saline soils, the use of
salt-tolerant plants improves soil physical properties
due to favorable root growth and increased microbial
respiration. The increase in CO2 partial pressure in the
rhizosphere increases calcium carbonate solubility that
counters the adverse effects of high Na+ (Qadir and
Oster, 2002).
2.9. Biotic influences
2.9.1. Plants
2.9.1.1. Plant species. The combined effects of the
biochemical composition and amount of plant residues
13
returned to soils and chemicals released from plants
affect the rate and stability of aggregation, and the rate
of aggregate turnover. Water-stable aggregates (WSA),
mean aggregate size and mean weight diameter
(MWD) are correlated with biochemical composition
of plant residues: phenols, lignin, proteins, monosaccharide sugars, saccharides, phenols and alkaline extractable HAs in the soil and phenolic acids such as
vanillin-vanillic acid in the residue (Martens and
Frankenberger, 1992; Martens, 2000). Corn (Zea mays)
residues are high in phenols and increase aggregation
compared with other crops, although continuous corn
decreases microaggregates compared to corn grown in
rotation (Raimbault and Vyn, 1991; Martens, 2000).
Soil aggregate stability is high under continuous alfalfa
(Medicago sativa) (Raimbault and Vyn, 1991). The
low aggregation of soil cultivated to soybean is attributed to low concentration of phenols (Martens, 2000)
along with low residue return to the soil.
2.9.1.2. Roots. Plant roots and their rhizosphere have
many effects on soil aggregation. Roots enmesh and
realign soil particles and release exudates, which result
in physical, chemical and biological alterations that
influence aggregation. Aggregation tends to increase
with increasing root length density, microbial associations, glomalin, and percent cover significantly affect
soil aggregate stabilization (Rillig et al., 2002). Aggregate stability is greater in rhizosphere soil than in nonrhizosphere soil (Caravaca et al., 2002) due to rhizodeposition, root mass, root density, size distribution,
root turnover, root length, and hyphal growth (Haynes
and Beare, 1997). The rhizosphere hosts a large population of micro- and macro-organisms that contribute
to SOC and aggregation. Chemically, roots enhance
aggregation by releasing a variety of compounds,
which have a cementing effect on soil particles. Root
mucilage such as polygalacturonic acid may stabilize
aggregates by increasing bond strength and decreasing
wetting rate (Czarnes et al., 2000). Roots increase the
wet – dry cycling of adjacent soil that can increase
aggregate stability in some cases and decrease in
others, possibly related to clay type (Angers and Caron,
1998). Roots can also alter the ionic and osmotic
balance in the rhizosphere through nutrient uptake
and rhizodeposition, which can affect aggregation.
Different root systems affect aggregation differently,
relating to different root properties, exudates and func-
14
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
tions (Chan and Heenan, 1999b). Generally, extensive
fibrous roots produce high levels of macroaggregation
(Harris et al., 1966; Chan and Heenan, 1996). Leguminous plant roots are associated with higher microbial
biomass, increased aggregation and WSA than nonlegumes (Chan and Heenan, 1996; Haynes and Beare,
1997). Aggregate stability in non-legumes is related to
root mass (Haynes and Beare, 1997).
2.9.2. Microorganisms
Roots, fungi and bacteria enhance aggregation by
enmeshing soil particles and providing extracellular
compounds that bind particles together. It is often
difficult to separate the multiple effects of organisms
on aggregation.
2.9.2.1. Microbial activity. The effect of aggregate
size on microbial activity depends on numerous
factors. In some cases, microbial activity may depend
on aggregate size (Mendes et al., 1999) and in others
it may not (Schutter and Dick, 2002). Bacteria are
often associated with clay and polysaccharides in
microaggregates, resulting in lower microbial biomass
in microaggregates than macroaggregates (Tisdall and
Oades, 1982; Lupwayi et al., 2001). Lower bacteria/
fungi ratio in macroaggregates than microaggregates
suggests that bacterial activity may dominate in
microaggregation while fungal activity dominates in
macroaggregate formation (Schutter and Dick, 2002;
Tisdall and Oades, 1982). Increases in macroaggregates are associated with increases in fungal activities
and fresh residues (Denef et al., 2001).
Microbial activity varies with aggregate size, seasons, cropping activities, management, residue quality
and quantity and soil type (Schutter and Dick, 2002;
Mendes et al., 1999). Microbial influence is most
pronounced in sandy soils where soil microorganisms
produce readily available C-source for the rapid
stabilization of aggregates (Kiem and Kandeler,
1997). Fungicide treatments may lower Cmic in soil
(Hu et al., 1995).
2.9.2.2. Fungi, arbuscular mycorrhizal fungi (AMF)
and glomalin. Fungal hyphae improve aggregate
stability (Ternan et al., 1996) by reorientation of clay
particles, binding particles with extracellular polysaccharides, and enmeshing particles. Hyphae also enmesh microaggregates to form macroaggregates,
suggesting that aggregation increases with hyphal
density (Haynes and Beare, 1997).
The AMF is one of the most important biotic
influences on soil aggregation (Jastrow et al., 1998).
Glomalin is a glycoprotein released from AMF. It is
present in soils at high concentrations and is an
important factor in stablizing aggregates, possibly
due to its recalcitrant nature and high concentration
in some soils (Wright and Upadhyaya, 1998; Rillig et
al., 2002). Soils high in expanding clays have more
total glomalin than those containing low-activity
clays. It is possible that the hydrophobic, recalcitrant
nature of the glomalin molecule may protect other
aggregating agents (Wright and Upadhyaya, 1998).
Wet – dry cycles may increase polymerization and
hydrophobicity in glomalin.
2.9.3. Soil fauna
Many immature and mature insects, other arthropods, earthworms, nematodes and larger macro-organisms live in the soil and have an important influence on
soil structure. They ingest and egest soil material,
relocate plant material and form burrows (Amezketa,
1999). Biogenic structures formed by soil organisms
such as termite mounds and casts are often protected
from mineralization. The effects of these activities are
variable. Macro-organisms improve aeration, porosity,
infiltration, aggregate stability, litter mixing, improved
N and C stabilization, C turnover and carbonate reduction and N mineralization, nutrient availability and
metal mobility (Amezketa, 1999; Winsome and
McColl, 1998; Brown et al., 2000). These effects can
degrade soil properties by removing DOC and breaking
down bonds between particles during ingestion. The
dispersion is often compensated for during reformation
of aggregates and egestion of CR compounds. Ingested
soil undergoes many alterations including physical
realignment of clay particles and breaking of bonds
within soil aggregates to alter microbial accessibility of
SOC (Wolters, 2000). Feeding, mixing ejecta with soil,
reworking and biosynthesis of SOC generally result in
an increase in soil CR (Wolters, 2000). Activity of soil
fauna is important in the formation of organo-mineral
complexes and aggregation.
2.9.3.1. Earthworms. Earthworms increase soil aggregation through biological and physicochemical
changes (Brown et al., 2000). Earthworms exert
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
considerable direct and indirect influences on soil
structure and SOC (Brown et al., 2000; Jegou et al.,
2001). The impact of earthworms on aggregation
varies with the species of earthworm, quality of litter
and parent soil (Winsome and McColl, 1998; Schrader
and Zhang, 1997). In turn, species and population of
earthworm are influenced by the quality and quantity
of plant material (Flegel and Schrader, 2000). Some
earthworms digest soil and in the process break bonds,
thereby destabilizing aggregates; however, biochemical processes stabilize aggregates (Schrader and
Zhang, 1997). The reorganization of soil particles
and release of enzymes by earthworms influence
mineralization, water flow and microbial activities
(Jegou et al., 2001). Earthworms increase SOC in
casts and, to a lesser extent, in burrow-wall materials
(Jegou et al., 2001). Earthworm activity increases
water-stable macroaggregates, which are actually
casts (Winsome and McColl, 1998).
2.9.3.2. Termites. The effect of termites on soil
structure and SOC varies with species and soil.
Positive (Al-Houty, 1998) and negative impacts (Fall
et al., 2001) or none at all (Hyodo et al., 2001) have
been reported on SOC. In most cases, ingestion of
SOC and its transformation during digestion results in
more stable SOC forms in termitosphere (Brauman,
2000). Termite-induced microaggregation is related to
soil composition, promoted by concentrations of
gibbsite and Fe-oxide in the tropics, however, not in
kaolinitic soils (Schaefer, 2001).
15
phenolics, which decompose slowly (Hu et al.,
1999). In environments with adequate nutrient
supplies the increases in atmospheric CO2 may result
in increased SOM and aggregation, while in nutrientlimited environments the balance between availability
of nutrients, plant demand and C turnover may limit
the effect.
Changes in dominant species and C allocation are
primary mechanisms of changes in C input and
decomposition (Dukes and Hungate, 2002). Changes
in plant species influence microbial population distribution and microbial compounds involved in soil
structural development (Dukes and Hungate, 2002).
Allocation of photosynthate to root and root exudates
tends to increase SOM and microbes in the rhizosphere (Swift, 2001). Increases in Cmic may occur
under elevated CO2 as an indirect response to changes
in plant growth (Elhottova et al., 1997; Hu et al.,
1999). Increases in AMF, root length and root longevities may occur (Rillig et al., 2001). Increased
levels of CO2 have resulted in increased WSA,
glomalin and AMF hyphal lengths with sorghum
(Rillig et al., 2001). Increased atmospheric CO2 may
result in lower hyphal density possibly due to decreased oxygen diffusion in soils (Schack-Kirchner et
al., 2000). Increased decomposition rates due to
increased temperatures, moisture and microbial activity may have a greater influence on rapid turnover of
SOC pool comprising plant and surface litter, than of
slower fractions such as root, macro and microbial C
resulting in the accumulation of SOM and higher
proportion of microaggregates (Niklaus et al., 2001).
3. Environmental impacts of soil structure
3.2. Enhancing the soil organic carbon pool
3.1. High atmospheric carbon dioxide concentrations
and soil structure
The type of soil, aggregating agents, soil management and environmental conditions interact to determine aggregation in a soil. Soil management practices
that minimize disturbance maximize SOC return to
the soil, optimize productivity and increase the SOC
pool. Appropriate use of soil amendments such as
fertilizer, lime, manure and compost can increase the
SOC pool and aggregation. Sustainable agricultural
practices such as mixed cropping, cover crops and
crop rotations also can increase the SOC pool. Management practices that reduce decomposition rates
and CO2 emissions also help to increase the SOC
storage.
The overall effect of elevated atmospheric CO2 on
soil structure is not very well understood. Enhanced
atmospheric CO2 may result in the increase in photosynthesis and ensuing increase in photosynthate, roots
and microbial communities. The concurrent increases
in nutrient demands may exceed the ability to meet the
demands, resulting in nutrient-limited systems, increased symbiotic communities or altering the chemical composition of plant compounds, resulting in
wider C/N ratio and increased lignin, tannins and
16
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
4. Management for enhancing soil structure
4.1. Soil management
Soil management to increase aggregation must aim
at increasing primary plant production, increasing the
amount of C input into the soil, decreasing disturbances and decreasing the rate of C loss by processes
such as decomposition and erosion. In this regard,
improved management practices include tillage methods, residue management, amendments, soil fertility
management and nutrient cycling.
4.1.1. Tillage
Tillage disrupts soil aggregates, compacts soil and
disturbs plant and animal communities that contribute
to aggregation and lowers SOM, CEC, nutrients,
microbial activity and faunal activities that contribute
to aggregation (Plante and McGill, 2002a). In comparison with plowing no-till management systems
have more stable aggregates and SOC (Filho et al.,
2002). Reduced tillage can result in higher macropores and biochannels that influence water movement
and availability, raising concerns about water quality
(Logan et al., 1991; Warkentin, 2001). In turn, this
impacts leaching and loss of nutrients and chemicals
in drainage and ground water. The intensity and
timing of tillage determine the extent of the effect of
tillage on SOC (Studdert and Echeverria, 2000).
4.1.2. Mulching and residue management
Mulches improve soil structure through a variety of
methods. The addition of mulch to soil surface
decreases erosion, reduces evaporation, protects
against raindrop impact and increases aggregate stability (Layton et al., 1993). Mulches increase the
amount of SOC pool (Duiker and Lal, 1999; Sharma
and Acharya, 2000; Jacinthe et al., 2002b), modify
temperature and moisture regimes and impact soil
fauna. The return of plant residues to soil improves
soil structure (Martens, 2000), which is related to the
amount and quality of the residue.
4.1.3. Manuring
Manuring improves soil structure and MWD,
increases macroaggregation and resistance to slaking
but may decrease stability of soil aggregates against
the dissolution and dispersive actions (Pare et al.,
1999; Whalen and Chang, 2002; Hao and Chang,
2002). The increase in SOC results in increased
biological activity, which in turn results in increased
porosity and the attendant decrease in bulk density
(Kay, 1998). Increased microbial activity due to
increases in soil C from manure applications results
in increases in aggregate stability (Martens and Frankenberger, 1992). Manured soils also have high earthworm population (Hansen and Engelstad, 1999).
Unmanured soils usually contain less SOC and microbial biomass, and are denser than manured soil,
and aggregates are strong when dry, and weak when
wet (Munkholm et al., 2002). In contrast, the manured
soils have strong aggregates when wet and weak when
dry. The differences between the soils when dry seem
to be related to differences in concentration of dispersible clay, whereas the differences when wet are
related to differences in the amount of organic binding
and bonding material. Micro- and macroaggregates
have higher carbohydrate levels in manured soils than
in those receiving chemical fertilizers (Debosz et al.,
2002a). Increased ionic concentration can be a concern in manured soil, because of increase in susceptibility to dispersion and decrease in aggregate
stability. Manuring increases concentrations of Na+,
K+ and Mg2 + (Haynes and Naidu, 1998; Hao and
Chang, 2002). Manure applications increase dispersion of large macroaggregates (Whalen and Chang,
2002), and also affect soil pH and P availability
(Whalen et al., 2000).
4.1.4. Compost
Compost additions to soil improve soil structure
and lower bulk density. Composting materials can
increase macroaggregation and rhizospheric aggregate stability (de Leon-Gonzalez et al., 2000; Caravaca et al., 2002). Soil properties and environmental
conditions determine the impact of compost on
aggregation; drought can limit the effectiveness of
compost in aggregation (de Leon-Gonzalez et al.,
2000). The effects of compost additions on soil
structure may be short-lived although effects are
generally positive on structural properties (Debosz
et al., 2002b).
4.1.5. Fertilizer and nutrients
The complexities of the chemical and physical
influences of fertilizers result in variable effects of
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
fertilization on aggregation. Fertilizer applications
generally improve soil aggregation (Haynes and
Naidu, 1998). However, under some conditions fertilizers may also decrease SOC concentration, reduce
aggregation, and reduce microbial communities compared to manured soils. Yet, using chemical fertilizers
often improves soil structure in comparison to unfertilized soils (Munkholm et al., 2002). The primary
effect of improved nutrient management is on increasing plant productivity, SOC and biological activity
(Haynes and Naidu, 1998). Increase in SOC by
fertilizer use increases aggregation and MWD (Subbian et al., 2000). Fertilizer use also improves residue
quality and quantity, but this does not necessarily
increase SOC pool (Halvorson et al., 2002). Fertilizer
applications alter soil pH and the electrolyte concentrations in soil, which can have adverse effects on soil
structure (Haynes and Naidu, 1998). The beneficial
effects of fertilizer applications generally offset any
adverse affects of fertilization. Increases in plant
residues and below-ground plant growth increase
CL, microbial activity and Cmic, which improve aggregate stability. Increases in N availability from
fertilizer treatments increase the rate of decomposition
in LF CL, while further stabilizing CI (Neff et al.,
2002).
The physical and chemical effects of fertilizers are
related to the type of fertilizer and soil. The use of
phosphetic fertilizers enhances Al3 + and Ca2 + bonding (Haynes and Naidu, 1998). Phosphoric acid promotes soil aggregation through lowered pH, Al3 +
mobilization and subsequent precipitation of Al phosphate as a cementing agent to form stable aggregates
(Haynes and Naidu, 1998). Ammonium fertilizers can
disperse clays, having an adverse effect on soil
aggregation (Haynes and Naidu, 1998). The dispersive affects of NH4+ are temporary and diminish as
NH4+ is nitrified to NO3 (Haynes and Naidu, 1998).
4.2. Crop management
Cultivation decreases total and microbial SOC pool
and soil fauna but increases metabolic CO2 (Saggar et
al., 2001). Management to increase SOC and aggregation includes fertilization, grazing management,
conversion from cultivation to native vegetation, inclusion of cover crops, legumes and grasses, earthworm inoculation and irrigation.
17
4.2.1. Crop rotations and cover crops
The aggregate dynamics vary among different
crops, crop rotations and cover crops (Jarecki and
Lal, 2003). The effect of different crops tends to
reflect the crop chemical composition (Martens,
2000), rooting structure and ability to alter the chemical and biological properties of the soil (Chan and
Heenan, 1996). These effects tend to be short-lived
under conventional tillage regimes (Chan and Heenan,
1996). In some soils, rotations may not affect aggregate stability (Filho et al., 2002). Cover crops increase
C input to the soil, reduce erosion, increase CEC,
increase aggregate stability, increase water infiltration,
and recycle nutrients. Cover crop residues may enhance microbial biomass, respiration, and N mineralization and shift microbial community (Schutter and
Dick, 2002).
4.2.2. Agroforestry
The inclusion of leguminous trees in agricultural
systems reduces soil erosion and improves soil productivity (Craswell et al., 1997; Buresh and Tian,
1997). Agroforestry influences SOM, which in turn
influences aggregation in comparison with conventional systems (Mugendi et al., 1999; Atsivor et al.,
2001).
5. Research priorities
Aggregation is correlated with root mass, root
morphology, fungal mass and use of organic matter
amendments such as mulch, manure and green manure. While these factors contribute to aggregation in
soils, a comprehensive understanding of how this
occurs is lacking. Cover crops in no-till agricultural
systems offer great potential to enhance aggregation.
Little is known about why some cover and agronomic
crops are more effective in structural development
than others. Additional information on the biochemistry and decomposition of plant residues and the
interaction with soil types and environmental conditions to promote aggregation would allow for identification of more effective management practices. A
better understanding of how aggregates are formed
and stabilized, and how they are affected by different
agricultural management and planting patterns is
necessary in order to identify agricultural practices
18
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
for sustainable crop production. Primary aggregation
mechanisms differ in different soil types; identification of practices to increase aggregation based on soil
types and dominant development processes need to be
better identified. The interaction between different
SOC sources, cations and clays are important for
understanding the development of stable soil structure. The role of secondary carbonate formation and
SOC interactions in arid and semi-arid regions is not
clear. Little is known about the impact of using
insecticides and fungicides on aggregation.
Soil structure impacts water availability, nutrient
uptake and leaching thereby affecting ground and
surface water supplies through sedimentation and
chemical contamination. It is important to develop
improved practices for the judicious use of agricultural amendments to maximize their effectiveness
while minimizing unintended and off-site impacts.
Similarly, management practices that reduce erosion,
nutrient loss and release of greenhouse gases need to
be assessed.
6. Conclusions
Soil structure holds a vital, but often overlooked
role in sustainable food production and the well being
of society. A more holistic approach to land use and
management is needed to cope with increased pressure on soil resources for sustainable food and fiber
production while reducing the adverse off-site environmental impacts of agricultural practices. The impact of soil structure ranges from a global to a highly
localized scale. Improved C sequestration in soil
aggregates can reduce the rate of increase in CO2
concentration in the atmosphere and associated global
warming. Improved soil structure enhances nutrient
recycling, water availability and biodiversity while
reducing water and wind erosion, and improving
surface and ground water quality.
Processes and mechanisms involved in soil
aggre-gation are complex with intricate feedback
mechanisms. Soil aggregation can be improved by
management practices that decrease agro-ecosystem
disturbances, improve soil fertility, increase organic
inputs, increase plant cover, and decrease SOC
decomposition rate. The SOC decomposition can
be decreased in soil by isolating labile C products
within aggregates and increasing chemical protection through input of CR compounds or transformation of CL to CR and CI compounds. Soil structure
can be enhanced through the use of crops and crop
management practices that promote aggregation
such as the use of crops with high CR and high
biomass production, the return of crop residues and
incorporation of cover crops. Aggregation also tends
to increase with increasing root length density;
extensive fibrous roots produce highest levels of
macroaggregation. Enhancing the diversity and
quantity of soil flora and fauna are important in
improving soil structure. Activity of soil fauna is
important in the formation of organo-mineral complexes and aggregation.
References
Al-Houty, W., 1998. Influence of termites on desert soil properties
in Kuwait. Arab Gulf J. Sci. Res. 16, 393 – 401.
Amezketa, E., 1999. Soil aggregate stability: a review. J. Sustain.
Agric. 14, 83 – 151.
Angers, D.A., Caron, J., 1998. Plant-induced changes in soil structure: processes and feedbacks. Biogeochemistry 42, 55 – 72.
Armstrong, A.S.B., Tanton, T.W., 1992. Gypsum applications to
aggregated saline sodic clay topsoils. J. Soil Sci. 43, 249 – 260.
Atsivor, L., Dowuona, G.N., Adiku, S.G.K., 2001. Farming systeminduced variability of some soil properties in a sub-humid zone
of Ghana. Plant Soil 236, 83 – 90.
Attou, F., Bruand, A., 1998. Experimental study of ‘fragipans’
formation in soils. Role of both clay dispersion and wetting –
drying cycles. C. R. Acad. Sci., Ser. 2, Fascicule II. Sci. Terre
Planetes 326, 545 – 552.
Attou, F., Bruand, A., Le Bissonnais, Y., 1998. Effect of clay content and silt – clay fabric on stability of artificial aggregates. Eur.
J. Soil Sci. 49, 569 – 577.
Baldock, J.A., Aoyama, M., Oades, J.M., Susanto, R.H., Grant,
C.D., 1994. Structural amelioration of a south Australian redbrown earth using calcium and organic amendments. Aust. J.
Soil Res. 32, 571 – 594.
Barral, M.T., Arias, M., Guerif, J., 1998. Effects of iron and organic
matter on the porosity and structural stability of soil aggregates.
Soil Tillage Res. 46, 261 – 272.
Batra, L., Kumar, A., Manna, M.C., Chhabra, R., 1997. Microbiological and chemical amelioration of alkaline soil by growing
Karnal grass and gypsum application. Exp. Agric. 33, 389 – 397.
Beare, M.H., Hendrix, P.F., Coleman, D.C., 1994. Water-stable
aggregates and organic matter fractions in conventional- and
no-tillage soils. Soil Sci. Soc. Am. J. 58, 777 – 786.
Boettinger, J.L., Southard, R.J., 1995. Pediment, western Mojave
Desert. Soil Sci. Soc. Am. J. 59, 1189 – 1198.
Boix-Fayos, C., Calvo-Cases, A., Imeson, A.C., Soriano-Soto,
M.D., Tiemessen, I.R., 1998. Spatial and short-term temporal
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
variations in runoff, soil aggregation and other soil properties
along a Mediterranean climatological gradient. Catena 33,
123 – 138.
Boix-Fayos, C., Calvo-Cases, A., Imeson, A.C., 2001. Influence of
soil properties on the aggregation of some Mediterranean soils
and the use of aggregate size and stability as land degradation
indicators. Catena 44, 47 – 67.
Brauman, A., 2000. Effect of gut transit and mound deposit on soil
organic matter transformations in the soil feeding termite: a
review. Eur. J. Soil Biol. 36, 117 – 125.
Brown, G.G., Barois, I., Lavelle, P., 2000. Regulation of soil organic matter dynamics and microbial activity in the drilosphere
and the role of interactions with other edaphic functional
domains. Eur. J. Soil Biol. 36, 177 – 198.
Buresh, R.J., Tian, G., 1997. Soil improvement by trees in subSaharan Africa. Agrofor. Syst. 38, 51 – 76.
Caesar-TonThat, T.C., 2002. Soil binding properties of mucilage
produced by a basidiomycete fungus in a model system. Mycol.
Res. 106, 930 – 937.
Caravaca, F., Hernandez, T., Garcia, C., Roldan, A., 2002. Improvement of rhizosphere aggregate stability of afforested semiarid
plant species subjected to mycorrhizal inoculation and compost
addition. Geoderma 108, 133 – 144.
Chan, K.Y., Heenan, D.P., 1996. The influence of crop rotation on
soil structure and soil physical properties under conventional
tillage. Soil Tillage Res. 37, 113 – 125.
Chan, K.Y., Heenan, D.P., 1999a. Lime-induced loss of soil organic
carbon and effect on aggregate stability. Soil Sci. Soc. Am. J.
63, 1841 – 1844.
Chan, K.Y., Heenan, D.P., 1999b. Microbial-induced soil aggregate
stability under different crop rotations. Biol. Fertil. Soils 30,
29 – 32.
Chan, K.Y., Heenan, D.P., So, H.B., 2003. Sequestration of carbon
and changes in soil quality under conservation tillage on lighttextured soils in Australia: a review. Aust. J. Exp. Agric. 43,
325 – 334.
Chenu, C., Le Bissonnais, Y., Arrouays, D., 2000. Organic matter
influence on clay wettability and soil aggregate stability. Soil
Sci. Soc. Am. J. 64, 1479 – 1486.
Chorom, M., Regasamy, P., Murray, R.S., 1994. Clay dispersion as
influenced by pH and net particle charge of sodic soils. Aust. J.
Soil Res. 32, 1243 – 1252.
Christensen, B.T., 2001. Physical fractionation of soil and structural
and functional complexity in organic matter turnover. Eur. J.
Soil Sci. 52, 345 – 353.
Clough, A., Skjemstad, J.O., 2000. Physical and chemical protection of soil organic carbon in three agricultural soils with different contents of calcium carbonate. Aust. J. Soil Res. 38,
1005 – 1016.
Craswell, E.T., Sajjapongse, A., Howlett, D.J.B., Dowling, A.J.,
1997. Agroforestry in the management of sloping lands in Asia
and the Pacific. Agrofor. Syst. 38, 121 – 137.
Czarnes, S., Hallett, P.D., Bengough, A.G., Young, I.M., 2000.
Root- and microbial-derived mucilages affect soil structure
and water transport. Eur. J. Soil Sci. 51, 435 – 443.
D’Acqui, L.P., Kaniele, E., Fornasier, F., Radaelli, L., Ristori,
G.G., 1998. Interaction between clay microstructure, decom-
19
position of plant residues and humificaion. Eur. J. Soil Sci.
49, 579 – 587.
D’Acqui, L.P., Churchman, G.J., Janik, L.J., Ristori, G.G., Weissmann, D.A., 1999. Effect of organic matter removal by lowtemperature ashing on dispersion of undisturbed aggregates
from a tropical crusting soil. Geoderma 93, 311 – 324.
Dalal, R.C., Bridge, B.J., 1996. Aggregation and organic matter
storage in sub-humid and semi-arid soils. In: Carter, M.R.,
Stewart, B.A. (Eds.), Structure and Organic Matter Storage in
Agricultural Soils. CRC Press, Boca Raton, FL, pp. 263 – 307.
Debosz, K., Vognsen, L., Labouriau, R., 2002a. Carbohydrates in
hot water extracts of soil aggregates as influenced by long-term
management. Commun. Soil Sci. Plant Anal. 33, 623 – 634.
Debosz, K., Petersen, S.O., Kure, L.K., Ambus, P., 2002b. Evaluating effects of sewage sludge and household compost on soil
physical, chemical and microbiological properties. Appl. Soil
Ecol. 19, 237 – 248.
De Coninck, F., 1980. Major mechanisms in formation of spodic
horizons. Geoderma 24, 101 – 128.
de Leon-Gonzalez, F., Hernandez-Serrano, M.M., Etchevers, J.D.,
Payan-Zelaya, F., Ordaz-Chaparro, V., 2000. Short-term compost effect on macroaggregation in a sandy soil under low rainfall in the valley of Mexico. Soil Tillage Res. 56, 213 – 217.
Denef, K., Six, J., Bossuyt, H., Frey, S.D., Elliott, E.T., Merckx,
R., Paustian, K., 2001. Influence of dry – wet cycles on the
interrelationship between aggregate, particulate organic matter,
and microbial community dynamics. Soil Biol. Biochem. 33,
1599 – 1611.
Denef, K., Six, J., Merckx, R., Paustian, K., 2002. Short-term
effects of biological and physical forces on aggregate formation
in soils with different clay mineralogy. Plant Soil 246, 185 – 200.
Dimoyiannis, D.G., Tsadilas, C.D., Valmis, S., 1998. Factors affecting aggregate instability of Greek agricultural soils. Commun.
Soil Sci. Plant Anal. 29, 1239 – 1251.
Dinel, H., Righi, D., Hardy, M., Jambu, P., 1997. Neutral lipids and
structural stability of physically degraded soils. Agrochimica 41,
97 – 108.
Duiker, S.W., Lal, R., 1999. Crop residue and tillage effects on
carbon sequestration in a Luvisol in central Ohio. Soil Tillage
Res. 52, 73 – 81.
Duiker, S.W., Rhoton, F.E., Torrent, J., Smeck, N.E., Lal, R., 2003.
Iron (hydr)oxide crystallinity effects on soil aggregation. Soil
Sci. Soc. Am. J. 67, 606 – 611.
Dukes, J.S., Hungate, B.A., 2002. Elevated carbon dioxide and
litter decomposition in California annual grasslands: which
mechanisms matter? Ecosystems 5, 171 – 183.
Edwards, A.P., Bremner, J.M., 1967. Microaggregates in soils.
J. Soil Sci. 18, 64 – 73.
Elhottova, D., Triska, J., Santruckova, H., Kveton, J., Santrucek, J.,
Simkova, M., 1997. Rhizosphere microflora of winter wheat
plants cultivated under elevated CO2. Plant Soil 197, 251 – 259.
Facelli, E., Facelli, J.M., 2002. Soil phosphorus heterogeneity and
mycorrhizal symbiosis regulate plant intra-specific competition
and size distribution. Oecologia 133, 54 – 61.
Fall, S., Brauman, A., Chotte, J.L., 2001. Comparative distribution
of organic matter in particle and aggregate size fractions in the
mounds of termites with different feeding habits in Senegal:
20
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
Cubitermes niokoloensis and Macrotermes bellicosus. Appl.
Soil Ecol. 17, 131 – 140.
Filho, C.C., Lourenco, A., Guimaraes, M.D.F., Fonseca, I.C.B.,
2002. Aggregate stability under different soil management systems in a red Latosol in the state of Parana, Brazil. Soil Tillage
Res. 65, 45 – 51.
Flegel, M., Schrader, S., 2000. Importance of food quality on selected enzyme activities in earthworm casts (Dendrobaena
octaedra, Lumbricidae). Soil Biol. Biochem. 32, 1191 – 1196.
Franzluebbers, A.J., 2002. Water infiltration and soil structure related to organic matter and its stratification with depth. Soil
Tillage Res. 66, 197 – 205.
Franzluebbers, A.J., Haney, R.L., Honeycutt, C.W., Arshad, M.A.,
Schomberg, H.H., Hons, F.M., 2001. Climatic influences on
active fractions of soil organic matter. Soil Biol. Biochem. 33,
1103 – 1111.
Guggenberger, G., Kaiser, K., 2003. Dissolved organic matter in
soil: challenging the paradigm of sorptive preservation. Geoderma 113, 293 – 310.
Hallett, P.D., Baumgartl, T., Young, I.M., 2001. Subcritical water
repellency of aggregates from a range of soil management practices. Soil Sci. Soc. Am. J. 65, 184 – 190.
Halvorson, A.D., Wienhold, B.J., Black, A.L., 2002. Tillage, nitrogen, and cropping system effects on soil carbon sequestration.
Soil Sci. Soc. Am. J. 66, 906 – 912.
Hansen, S., Engelstad, F., 1999. Earthworm populations in a cool
and wet district as affected by tractor traffic and fertilization.
Appl. Soil Ecol. 13, 237 – 250.
Hao, X.Y., Chang, C., 2002. Effect of 25 annual cattle manure
applications on soluble and exchangeable cations in soil. Soil
Sci. 167, 126 – 134.
Harris, R.F., Chesters, G., Allen, O.N., 1966. Dynamics of soil
aggregation. Adv. Agron. 18, 108 – 169.
Haynes, R.J., Beare, M.H., 1997. Influence of six crop species on
aggregate stability and some labile organic matter fractions. Soil
Biol. Biochem. 29, 1647 – 1653.
Haynes, R.J., Naidu, R., 1998. Influence of lime, fertilizer and
manure applications on soil organic matter content and soil
physical conditions: a review. Nutr. Cycl. Agroecosyst. 51,
123 – 137.
Hu, S., Coleman, D.C., Hendrix, P.F., Beare, M.H., 1995.
Biotic manipulation effects on soil carbohydrates and microbial biomass in a cultivated soil. Soil Biol. Biochem. 27,
1127 – 1135.
Hu, S.J., Firestone, M.K., Chapin, F.S., 1999. Soil microbial feedbacks to atmospheric CO2 enrichment. Trends Ecol. Evol. 14,
433 – 437.
Hyodo, F., Tayasu, I., Azuma, J.I., Kirtibutr, N., Abe, T., 2001.
Effect of the soil-feeding termite, Dicuspiditermes makhamensis, on soil carbon structure in a seasonal tropical forest as
revealed by CP/MAS C-13 NMR. Sociobiology 38, 487 – 493.
Jacinthe, P.A., Lal, R., Kimble, J.M., 2002a. Carbon dioxide
evolution in runoff from simulated rainfall on long-term notill and plowed soils in southwestern Ohio. Soil Tillage Res.
66, 23 – 33.
Jacinthe, P.A., Lal, R., Kimble, J.M., 2002b. Carbon budget and
seasonal carbon dioxide emission from a central Ohio Luvisol as
influenced by wheat residue amendment. Soil Tillage Res. 67,
147 – 157.
Jansen, B., Klaas, G.J., Verstraten, J.M., 2003. Mobility of Fe(II),
Fe(III) and Al in acidic forest soils mediated by dissolved organic matter: influence of solution pH and metal/organic carbon
ratios. Geoderma 113, 323 – 340.
Jarecki, M., Lal, R., 2003. Crop management for soil carbon sequestration. Crit. Rev. Plant Sci. 22, 1 – 32.
Jastrow, J.D., 1996. Soil aggregate formation and the accrual of
particulate and mineral-associated organic matter. Soil Biol.
Biochem. 28, 665 – 676.
Jastrow, J.D., Miller, R.M., Lussenhop, J., 1998. Contributions of
interacting biological mechanisms to soil aggregate stabilization
in restored prairie. Soil Biol. Biochem. 30, 905 – 916.
Jegou, D., Schrader, S., Diestel, H., Cluzeau, D., 2001. Morphological, physical and biochemical characteristics of burrow walls
formed by earthworms. Appl. Soil Ecol. 17, 165 – 174.
Kay, B.D., 1998. Soil structure and organic carbon: a review. In:
Lal, R., Kimble, J.M., Follett, R.F., Stewart, B.A. (Eds.), Soil
Processes and the Carbon Cycle. CRC Press, Boca Raton, FL,
pp. 169 – 197.
Kiem, R., Kandeler, E., 1997. Stabilization of aggregates by the
microbial biomass as affected by soil texture and type. Appl.
Soil Ecol. 5, 221 – 230.
Lal, R., 1991. Soil structure and sustainability. J. Sustain. Agric. 1,
67 – 92.
Layton, J.B., Skidmore, E.L., Thompson, C.A., 1993. Winter-associated changes in dry-soil aggregation as influenced by management. Soil Sci. Soc. Am. J. 57, 1568 – 1572.
Leinweber, P., Schulten, H.R., Jancke, H., 1999. New evidence for
the molecular composition of soil organic matter in Vertisols.
Soil Sci. 164, 857 – 870.
Logan, T.J., Lal, R., Dick, W.A., 1991. Tillage systems and soil
properties in North-America. Soil Tillage Res. 20, 241 – 270.
Lupwayi, N.Z., Arshad, M.A., Rice, W.A., Clayton, G.W., 2001.
Bacterial diversity in water-stable aggregates of soils under conventional and zero tillage management. Appl. Soil Ecol. 16,
251 – 261.
Magill, A.H., Aber, J.D., 1998. Long-term effects of experimental
nitrogen additions on foliar litter decay and humus formation in
forest ecosystems. Plant Soil 203, 301 – 311.
Martens, D.A., 2000. Plant residue biochemistry regulates soil carbon cycling and carbon sequestration. Soil Biol. Biochem. 32,
361 – 369.
Martens, D.A., Frankenberger Jr., W.T., 1992. Modification of infiltration rates in an organic-amended irrigated soil. J. Agron.
84, 707 – 717.
Mendes, I.C., Bandick, A.K., Dick, R.P., Bottomley, P.J., 1999.
Microbial biomass and activities in soil aggregates affected by
winter cover crops. Soil Sci. Soc. Am. J. 63, 873 – 881.
Molina, N.C., Caceres, M.R., Pietroboni, A.M., 2001. Factors affecting aggregate stability and water dispersible clay of recently
cultivated semiarid soils of Argentina. Arid Land Res. Manag.
15, 77 – 87.
Mugendi, D.N., Nair, P.K.R., Mugwe, J.N., O’Neill, M.K.,
Woomer, P., 1999. Alley cropping of maize with calliandra
and leucaena in the subhumid highlands of Kenya: Part 1.
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
Soil-fertility changes and maize yield. Agrofor. Syst. 46,
39 – 50.
Munkholm, L.J., Schjonning, P., Debosz, K., Jensen, H.E., Christensen, B.T., 2002. Aggregate strength and mechanical behaviour of a sandy loam soil under long-term fertilization treatments.
Eur. J. Soil Sci. 53, 129 – 137.
Munyanziza, E., Kehri, H.K., Bagyaraj, D.J., 1997. Agricultural
intensification, soil biodiversity and agro-ecosystem function
in the tropics: the role of mycorrhiza in crops and trees. Appl.
Soil Ecol. 6, 77 – 85.
Neff, J.C., Townsend, A.R., Gleixner, G., Lehman, S.J., Turnbull, J.,
Bowman, W.D., 2002. Variable effects of nitrogen additions on
the stability and turnover of soil carbon. Nature 419, 915 – 917.
Nelson, P.N., Baldock, J.A., Clarke, P., Oades, J.M., Churchman,
G.J., 1999. Dispersed clay and organic matter in soil: their nature
and associations. Aust. J. Soil Res. 37, 289 – 315.
Niklaus, P.A., Wohlfender, M., Siegwolf, R., Korner, C., 2001.
Effects of six years atmospheric CO2 enrichment on plant, soil,
and soil microbial C of a calcareous grassland. Plant Soil 233,
189 – 202.
Nissen, T.M., Wander, M.M., 2003. Management and soil-quality
effects on fertilizer-use efficiency and leaching. Soil Sci. Soc.
Am. J. 67, 1524 – 1532.
Oades, J.M., Waters, A.G., 1991. Aggregate hierarchy in soils.
Aust. J. Soil Res. 29, 815 – 828.
Pachepsky, Y.A., Rawls, W.J., 2003. Soil structure and pedotransfer
functions. Eur. J. Soil Sci. 54, 443 – 451.
Palm, C.A., Rowland, A.P., 1997. Minimum dataset for characterization of plant quality for decomposition. In: Cadisch, G., Giller,
K.E. (Eds.), Driven by Nature: Plant Litter Quality and Decomposition. CAB International, Wallingford, UK, pp. 379 – 392.
Pardo, A., Amato, M., Chiaranda, F.Q., 2000. Relationships between soil structure, root distribution and water uptake of chickpea (Cicer arietinum L.). Plant growth and water distribution.
Eur. J. Agron. 13, 39 – 45.
Pare, T., Dinel, H., Moulin, A.P., Townley-Smith, L., 1999. Organic
matter quality and structural stability of a Black Chernozemic
soil under different manure and tillage practices. Geoderma 91,
311 – 326.
Piccolo, A., Pietramellara, G., Mbagwu, J.S.C., 1997. Use of humic
substances as soil conditioners to increase aggregate stability.
Geoderma 75, 267 – 277.
Plante, A.F., McGill, W.B., 2002a. Soil aggregate dynamics and the
retention of organic matter in laboratory-incubated soil with
differing simulated tillage frequencies. Soil Tillage Res. 66,
79 – 92.
Plante, A.F., McGill, W.B., 2002b. Intraseasonal soil macroaggregate dynamics in two contrasting field soils using labeled tracer
spheres. Soil Sci. Soc. Am. J. 66, 1285 – 1295.
Poulenard, J., Podwojewski, P., Janeau, J.L., Collinet, J., 2001.
Runoff and soil erosion under rainfall simulation of Andisols
from the Ecuadorian Paramo: effect of tillage and burning. Catena 45, 185 – 207.
Powers, J.S., Schlesinger, W.H., 2002. Relationships among soil
carbon distributions and biophysical factors at nested spatial
scales in rain forests of northeastern Costa Rica. Geoderma
109, 165 – 190.
21
Qadir, M., Oster, J.D., 2002. Vegetative bioremediation of calcareous
sodic soils: history, mechanisms, and evaluation. Irrig. Sci. 21,
91 – 101.
Raimbault, B.A., Vyn, T.J., 1991. Crop rotation and tillage
effects on corn growth and soil structural stability. Agron.
J. 83, 979 – 985.
Rampazzo, N., Blum, W.E.H., Wimmer, B., 1998. Assessment of
soil structure parameters and functions in agricultural soils. Bodenkultur 49, 69 – 84.
Rillig, M.C., Wright, S.F., Kimball, B.A., Pinter, P.J., Wall, G.W.,
Ottman, M.J., Leavitt, S.W., 2001. Elevated carbon dioxide and
irrigation effects on water stable aggregates in a Sorghum field:
a possible role for arbuscular mycorrhizal fungi. Glob. Chang.
Biol. 7, 333 – 337.
Rillig, M.C., Wright, S.F., Eviner, V.T., 2002. The role of arbuscular mycorrhizal fungi and glomalin in soil aggregation:
comparing effects of five plant species. Plant Soil 238,
325 – 333.
Saggar, S., Yeates, G.W., Shepherd, T.G., 2001. Cultivation effects
on soil biological properties, microfauna and organic matter
dynamics in Eutric Gleysol and Gleyic Luvisol soils in New
Zealand. Soil Tillage Res. 58, 55 – 68.
Santos, D., Murphy, S.L.S., Taubner, H., Smucker, A.J.M., Horn,
R., 1997. Uniform separation of concentric surface layers from
soil aggregates. Soil Sci. Soc. Am. J. 61, 720 – 724.
Schack-Kirchner, H., Wilpert, K.V., Hildebrand, E.E., 2000. The
spatial distribution of soil hyphae in structured spruce-forest
soils. Plant Soil 224, 195 – 205.
Schaefer, C.E.R., 2001. Brazilian latosols and their B horizon microstructure as long-term biotic constructs. Aust. J. Soil Res. 39,
909 – 926.
Schrader, S., Zhang, H.Q., 1997. Earthworm casting: stabilization
or destabilization of soil structure? Soil Biol. Biochem. 29,
469 – 475.
Schulten, H.R., Leinweber, P., 2000. New insights into organic –
mineral particles: composition, properties and models of molecular structure. Biol. Fertil. Soils 30, 399 – 432.
Schutter, M.E., Dick, R.P., 2002. Microbial community profiles and
activities among aggregates of winter fallow and cover-cropped
soil. Soil Sci. Soc. Am. J. 66, 142 – 153.
Schwartzman, D.W., Shore, S.N., Volk, T., McMenamin, M., 1994.
Self-organization of the earths biosphere—geochemical or geophysiological. Orig. Life Evol. Biosph. 24, 435 – 450.
Seta, A.K., Karathanasis, A.D., 1996. Water dispersible colloids
and factors influencing their dispersibility from soil aggregates.
Geoderma 74, 255 – 266.
Sharma, P.K., Acharya, C.L., 2000. Carry-over of residual soil
moisture with mulching and conservation tillage practices for
sowing of rainfed wheat (Triticum aestivum L.) in north-west
India. Soil Tillage Res. 57, 43 – 52.
Shepherd, T.G., Saggar, S., Newman, R.H., Ross, C.W., Dando,
J.L., 2001. Tillage-induced changes to soil structure and organic
carbon fractions in New Zealand soils. Aust. J. Soil Res. 39,
465 – 489.
Singer, M.J., Southard, R.J., Warrington, D.J., Janitzky, P., 1992.
Stability of synthetic sand clay aggregates after wetting and
drying cycles. Soil Sci. Soc. Am. J. 56, 1843 – 1848.
22
C.J. Bronick, R. Lal / Geoderma 124 (2005) 3–22
Six, J., Elliott, E.T., Paustian, K., 1999. Aggregate and soil organic
matter dynamics under conventional and no-tillage systems. Soil
Sci. Soc. Am. J. 63, 1350 – 1358.
Six, J., Elliott, E.T., Paustian, K., 2000. Soil structure and soil
organic matter: II. A normalized stability index and the effect
of mineralogy. Soil Sci. Soc. Am. J. 64, 1042 – 1049.
Sollins, P., Homann, P., Caldwell, B.A., 1996. Stabilization and
destabilization of soil organic matter: mechanisms and controls.
Geoderma 74, 65 – 105.
Studdert, G.A., Echeverria, H.E., 2000. Crop rotations and nitrogen
fertilization to manage soil organic carbon dynamics. Soil Sci.
Soc. Am. J. 64, 1496 – 1503.
Subbian, P., Lal, R., Akala, V., 2000. Long-term effects of cropping
systems and fertilizers on soil physical properties. J. Sustain.
Agric. 16, 89 – 100.
Sumner, M.E., 1993. Sodic soils—new perspectives. Aust. J. Soil
Res. 31, 683 – 750.
Swift, R.S., 2001. Sequestration of carbon by soil. Soil Sci. 166,
858 – 871.
Tarchitzky, J., Hatcher, P.G., Chen, Y., 2000. Properties and distribution of humic substances and inorganic structure-stabilizing
components in particle-size fractions of cultivated Mediterranean soils. Soil Sci. 165, 328 – 342.
Ternan, J.L., Williams, A.G., Elmes, A., Hartley, R., 1996. Aggregate stability of soils in central Spain and the role of land management. Earth Surf. Processes Landf. 21, 181 – 193.
Thomsen, I.K., Schjonning, P., Olesen, J.E., Christensen, B.T.,
1999. Turnover of organic matter in differently textured soils:
II. Microbial activity as influenced by soil water regimes. Geoderma 89, 199 – 218.
Thomsen, I.K., Schjonning, P., Olesen, J.E., Christensen, B.T.,
2003. C and N turnover in structurally intact soils of different
texture. Soil Biol. Biochem. 35, 765 – 774.
Tisdall, J.M., 1996. Formation of soil aggregates and accumulation
of soil organic matter. In: Carter, M.R., Stewart, B.A. (Eds.),
Structure and Organic Matter Storage in Agricultural Soils. CRC
Press, Boca Raton, FL, pp. 57 – 96.
Tisdall, J.M., Oades, J.M., 1982. Organic matter and water-stable
aggregates in soils. J. Soil Sci. 33, 141 – 163.
Torn, M.S., Trumbore, S.E., Chadwick, O.A., Vitousek, P.M., Hendricks, D.M., 1997. Mineral control of soil organic carbon storage and turnover. Nature 389, 170 – 173.
Vanlauwe, G., Diels, J., Sanginga, N., Merckx, R., 1997. Residue
quality and decomposition: an unsteady relationship? In:
Cadisch, G., Giller, K.E. (Eds.), Driven by Nature: Plant Litter
Quality and Decomposition. CAB International, Wallingford,
UK, pp. 157 – 166.
Warkentin, B.P., 2001. The tillage effect in sustaining soil functions.
Journal of Plant Nutrition and Soil Science (Zeitschrift fur
Pflanzenernahrung und Bodenkunde) 164, 345 – 350.
Whalen, J.K., Chang, C., 2002. Macroaggregate characteristics in
cultivated soils after 25 annual manure applications. Soil Sci.
Soc. Am. J. 66, 1637 – 1647.
Whalen, J.K., Chang, C., Clayton, G.W., Carefoot, J.P., 2000. Cattle
manure amendments can increase the pH of acid soils. Soil Sci.
Soc. Am. J. 64, 962 – 966.
Winsome, T., McColl, J.G., 1998. Changes in chemistry and aggregation of a California forest soil worked by the earthworm Argilophilus papillifer Eisen (Megascolecidae). Soil Biol. Biochem.
30, 1677 – 1687.
Wolters, V., 2000. Invertebrate control of soil organic matter stability. Biol. Fertil. Soils 31, 1 – 19.
Wright, S.F., Upadhyaya, A., 1998. A survey of soils for aggregate
stability and glomalin, a glycoprotein produced by hyphae of
arbuscular mycorrhizal fungi. Plant Soil 198, 97 – 107.
Zhang, B., Horn, R., 2001. Mechanisms of aggregate stabilization
in Ultisols from subtropical China. Geoderma 99, 123 – 145.
Zhang, X.C., Norton, L.D., 2002. Effect of exchangeable Mg on
saturated hydraulic conductivity, disaggregation and clay dispersion of disturbed soils. J. Hydrol. 260, 194 – 205.