RESEARCH EDITORIAL
this complex and humungous issue. Yet, as
doi:10.2489/jswc.70.6.329
Soil carbon sequestration and aggregation
by cover cropping
R. Lal
Agriculture is the foundation of human
civilization. Transition from hunter-gath-
erer to settled agriculture, as evidenced by
the use of crude implements for tilling and
harvesting, may have occurred in modern
day Iraq and elsewhere around 11,000 years
ago (Troeh et al. 2004). The beneficial role
of growing cover crops in improving soil
productivity had been recognized for at least
three millennia and probably longer. Use of
cover crops as green manure had been advo-
cated in most ancient cultures (e.g., Indus
Valley and Middle East) to improve soil
fertility. In China, for example, use of cover
cropping dates back to the Zhou dynasty
3,000 years ago (Lipman 1912; Pieters 1927;
Meisinger et al. 1991). Cato, a Roman philos-
opher during the 3rd century BC, advocated
the use of green manure and compost to
improve soil productivity (Winiwater 2006;
Brevik and Hartemink 2010). By the 4th
century AD, grain yields of wheat (Triticum
aestivum) in Rome declined drastically due to
the lack of cover cropping and were hardly
270 to 400 kg ha1 (4 to 6 bu ac1), and that
of barley (Hordeum vulgare) were only 220
to 320 kg ha1 (195.8 to 285.1 lb ac1; Paine
and Harrison 1993). Therefore, during 395
AD, farmers in southern Italy (Campania)
were forced to abandon as much as 134,000
ha (330,980 ac) of cropland because of low
production (Simkhovich 1937). In Japan,
the importance of soil fertility maintenance
by green manuring and growth of legumes
dates back to 9th century AD.
In the 16th century Europe, Tusser (1580;
Rasmussen 1975; Paine and Harrison 1993),
published several volumes in agriculture and
stated the disadvantages of declining fertility
and low yields when farmers continuously
grow oats (Avena sativa), rye (Secale cereale), bar-
ley, and wheat.Tusser advocated the use of peas
(Pisum sativum) and buckwheat (Fagopyrum
esculentum) for restoring soil fertility.
JOURNAL OF SOIL AND WATER CONSERVATION
In the United States, experiments with
crimson clover (Trifolium incarnatum) and
hairy vetch (Vicia villosa) were done toward
the end of the 19th century (Duggar 1898).
Thomas Jefferson advocated the use of cover
crops in Virginia to supply nitrogen (N) to
the succeeding crop (Betts 1953). Pieters
(1917, 1927) summarized the research con-
ducted in cover cropping during the early
1900s in the United States.
Modern agriculture evolved since the
industrial revolution of around 1750, but
more so since the 1950s. Major advances in
agriculture have been made since World War
II. The use of heavy machinery, input-re-
sponsive varieties grown as monoculture,
chemicals (e.g., fertilizers, pesticides, and
herbicides), and supplemental irrigation
or subsoil drainage began in the 1950s.
Consequently, food production increased
dramatically in all industrialized countries
and emerging economies. However, modern
agriculture also created some unprecedented
environmental problems, especially those
related to soil degradation, water pollution
and eutrophication, air pollution, emission
of greenhouse gases (GHGs) into the atmo-
sphere, and loss of biodiversity (Kirchmann
and Thorvaldsson 2000). Thus, there have
been some drastic human-induced alterations
in the pedosphere, atmosphere, hydrosphere,
biosphere, and the lithosphere (figure 1).
Soil degradation, by accelerated erosion and
other processes, has been extremely severe in
some regions, and has even caused extinc-
tion of some vulnerable soils (Tennesen
2014). These transformative trends are set-
in-motion because of drastic changes to the
pedosphere by land misuse and soil misman-
agement (figure 1).
Thus, a major challenge lies in revers-
ing the degradation trends while increasing
agricultural production and improving
the environment. Admittedly, there is nei-
ther a silver bullet nor a panacea to address
Thomas Tusser proposed around 1580, rather
than being greedy and practicing continu-
ous cultivation of food grains and other cash
crops, it is ethical to include fallow and cover
crops in the rotation cycle.
Therefore, the objective of this article is
to deliberate the impacts of cover cropping
on carbon (C) sequestration, improvements
in soil quality, increase in aggregation, and
reduction in the net emission of GHGs into
the atmosphere.
Addressing Challenges of Climate
Change through Cover Cropping
Climate change is a major global issue of the
21st century and must be addressed. Several
Copyright 2015 Soil and Water Conservation Society. All rights reserved.
Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org
important initiatives undertaken in 2014
indicate an emerging consensus that each and
every inhabitant of planet Earth is a perpe-
trator and victim of climate change. In June
of 2014, the US Environmental Protection
Agency (EPA) proposed the Clean Power
Plan to cut C emissions from existing power
plants by up to 30% by 2030, relative to 2005
levels (Kintisch 2014). The United States-
China accord announced in November of
2014 (The White House 2014) and the Lima
accord in December of 2014 (Davenport
2014) indicate the start of global cooperation
in reducing GHG emissions.
An important link between agriculture
and climate change cannot be ignored.
Agricultural activities, both directly and
indirectly, contribute about 30% of the total
anthropogenic emissions (IPCC 2014). Soils
of agroecosystems can be a source or a sink
of GHGs depending on land use and man-
agement. The soils can provide a large C
sink (Pacala and Socolow 2004; Socolow
and Pacala 2006), especially through regu-
lar and large in-put of biomass-C to soil by
activities such as cover cropping and conser-
vation agriculture (CA).
A cover crop is not a cash crop, but is used
to produce crops and is an integral part of
a production system (i.e., CA). It is a tran-
sition crop between two production crops,
and has its own challenges and tradeoffs (e.g.,
demand for land, water, requirement of N by
winter grasses, pest incidence, and slow soil
warming in spring). Cover crops can play an
Rattan Lal is the director at the Carbon Manage-
ment and Sequestration Center at The Ohio State
University, Columbus, Ohio.
NOV/DEC 2015VOL. 70, NO. 6 329
 Figure 1
Interaction between lithosphere, pedosphere, biosphere, and the atmosphere affecting climate change.
important role in enhancing soil C seques-
tration, off-setting some gaseous emissions
from croplands and agroecosystems, while
also providing other ecosystem services and
Climate
co-benefits (table 1).
change
Atmosphere Temperature Soil Carbon Sequestration and
Precipitation Hydrosphere
Cover Cropping

Soil C sequestration under cover cropping
ity
L Weat ers and CA is the transfer of atmospheric carbon
I each herin Soil quality iodiv s s
e
H lluvia ing g Hydrologic B Biom ystemary et dioxide (CO2 ) into the soil organic C (SOC)
M umi tion energy balance Ecos prim ity, n tiv- and soil inorganic C (SIC) pools through
G iner ficati et ctiv duc -
e a o Gaseous fluxes, N produ e pro osys ity production of plant biomass and its return to
ero olog lizati n emissions m ec iv
sio ic
n
on
Elemental bio net duct the soil such that some of the biomass-C is
transformations ity, pro
tem not reemitted into the atmosphere (Lal 2004).
Salinization
The process of soil/ecosystem C budget, and
the presumed drawdown of atmospheric

Copyright 2015 Soil and Water Conservation Society. All rights reserved.
Pedosphere

Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org

CO2 , should appropriately be assessed within
Biogeochemical Pedospheric Soil organic the same landscape unit (Olson et al. 2014).
transformations processes carbon pool World soils are an important component
of the global C cycle (figure 2) and strongly
Fo ol
ssi
l po affect atmospheric chemistry. Whereas the
car Lithospheric on
b on processes carb gross primary productivity (GPP) of ~123
res ni c Pg C y1 (135.5 109 tn C yr1) is balanced
erve rg a
s o
Lithosphere Soil in by plant and soil respiration of about 60 Pg
C y1 (66.1 109 tn C yr1) each, some of
the photosynthesized C is retained in the
biosphere as the net ecosystem produc-
Table 1 tivity (NEP) and net biome productivity
Impacts of cover crops on enhancing soil functions. (NBP; Jansson et al. 2010). Net primary pro-
Attribute Function ductivity (NPP) is the rate at which plants
incorporate atmospheric C through photo-
Soil quality High crop productivity and total production
synthesis. The NPP is estimated at 63 Pg C
Deep rooting depth and favorable edaphic conditions y1 (69.4 109 tn C yr1; Jasson et al. 2010).
High soil nutrient and water retention capacity The NEP is defined as the rate of change in
Favorable soil structure and tilth; and low compactability, ecosystem C storage over time (NEP = dC
crusting, hardsetting dt1; Abner and Melillo 1991; Janisch and
High bioturbation Harman 2002). An ecosystem is a CO2 sink
Improve workability when NEP is positive and a source when
Facilitate conservation agriculture and no-till farming it is negative. The NBP refers to the net
Climate change Large soil carbon (C) pool, more soil C sequestration change in ecosystem C pool with correction
Moderated gaseous flux (N2O, CH4)
for losses by fire, removal by harvest, and any
input of organic fertilizers. Relationships
Change in albedo
between NPP, NEP, and NBP are shown in
Favorable soil temperature and heat flux
equations 1 to 3 (Jansson et al. 2010):
Water resources High water infiltration, low surface runoff
Reduced nonpoint source pollution, hypoxia NPP = GPP PR, (1)
Improve renewability
Soil pests and pathogen Create disease-suppressive soils NEP = NPP MR, and (2)
Reduce weeds
Decrease nematodes and other soil-born pests and pathogens NBP = NEP LC, (3)
Ecosystem functions Elemental cycling
Hydrological and energy budget where PR is plant respiration; MR is soil
Habitat for biodiversity microbial respiration; and LC is losses of C by
harvesting, fire, and erosion, etc. Cover crop-
ping, in conjunction with CA, can strongly
influence NBP (figure 3) but also affect

330 NOV/DEC 2015VOL. 70, NO. 6 JOURNAL OF SOIL AND WATER CONSERVATION
 Figure 2
Impacts of cover cropping and conservation agriculture on soil carbon (C) sequestration
(redrawn and updated from Lal 2004). Also refer to equations 1 to 3 in the text.
1
y (gross primary prod
Pg C
1 23
nt respiration P
(Pla
Biota
Live: 560 Pg C
Detritus: 60 Pg C
Microbia
60 P
l res
gC
pira 1
y
tio
n(
R)
M
4,000 Pg C (2 m depth)
Potential of cropland soils
(+0.6 to 1.2 Pg C y1)
emission of nitrous oxide (N2O) and meth-
ane (CH4) from soil into the atmosphere
(figure 3). Improvements in soil structure and
tilth by cover cropping enhance soil aeration,
which may increase CH4 oxidation/uptake
and reduce N2O emission. Low O2 availabil-
ity in soil, created by unfavorable structure
and anaerobiosis, can increase emissions of
N2O and nitric oxide (NO; Zhu et al. 2013;
Panek et al. 2000). Among numerous ecosys-
tem services provisioned by cover cropping
(Schipanski et al. 2014), soil C sequestration
for climate change adaptation and mitigation
(figure 4) is timely because of the increasing
global interest in reducing the net atmo-
spheric emissions of GHGs. These services
can be broadly grouped into (1) risk manage-
ment over the short term and (2) stewardship
and ethics over the long term (figure 4). Risk
management is directly related to short-term
economic gains and profitability (Snapp et
al. 2005), whereas stewardship and ethics
JOURNAL OF SOIL AND WATER CONSERVATION
ucti
vity
6 0 Pg C y 1 ) by harvesting crop residues (corn, etc.) for
R)
Atmosphere
800 Pg C
C + 4.3 Pg C y1
g C y1
Pg C y1
.1 P
60
n1
on
sio
ati
ero
p ir
oil
S
es
il r
Soil
So
enhance the moral responsibilities through
consideration of the societal value of soil C
(Lal 2014) over longer periods of time and
indirectly. Furthermore, several disservices
of conventional agroecosystems (e.g., loss
of wildlife habitat, nutrient runoff, sedi-
mentation and nonpoint source pollution,
GHG emissions, and pesticide transport
in water) can be reduced by incorporating
cover crops in the rotation cycle (Power
2010). Increase in return of biomass-C to
the soil by cover cropping, both below- and
aboveground, can create a positive C bud-
get through reducing losses by erosion and
offsetting mineralization. There exists a
positive correlation between SOC change
and biomass-C input into the soil (Larson
et al. 1972; Rasmussen et al. 1980). Dell et
al. (2008) reported SOC sequestration rate
of 0.5 Mg C ha1 y1 (0.22 tn ac1 C yr1)
under no-tillage (NT) system. Some adverse
effects of removal of silage corn (Zea mays)
can be alleviated by cover cropping, if the
biomass-C added through cover cropping
is sufficient to offset the amount removed
biofuel feedstock (Fronning et al. 2008) and
other purposes. With adoption of recom-
mended management practices (RMPs) for
sustainable management of arable land, how-
ever, cover cropping can improve soil quality
even in a high desert farming system, such
as southern Idaho (Hunter and Falen 2012).
Choice of an appropriate cover crop
depends on site-specific conditions, and on
sufficient understanding of processes affect-
ing the biomass production and turnover
rate of organic substances in the soil. Thus,
rate of SOC sequestration by cover cropping
Copyright 2015 Soil and Water Conservation Society. All rights reserved.
Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org
varies with soil type, cover crop species, and
management (tables 2 and 3). Information
is also needed about the biomass produc-
tion rate (above- and belowground) and
about the rate of decomposition to assess
the overall ecosystem C budget and soil
carbohydrate concentration for a specific
cover crop. Kuo et al. (1997) observed that
annual ryegrass was better suited as a win-
ter cover crop to sequester SOC in Pullman,
Washington, United States, than Austrian
winter pea (Lathyrus hirsutus), hairy vetch,
or canola (Brassica napus). Efficiency of SOC
sequestration also depends on availability
of N. In the central Great Plains region of
Akron, Colorado, United States, Halvorson
et al. (2002a, 2002b) observed more seques-
tration with more input of N fertilizer and
less leaching with cereals. The rate of SOC
sequestration was 94, 140, and 182 kg C ha1
y1 (83.7,124.6, and 162.0 lb C ac1 yr1) for
45, 67, and 134 kg N ha1 (40; 60; 1,119 lb
N ac1) rate, respectively (SOC [Mg ha1] =
14.8 + 0.023x 0.00006x 2, r 2 = 0.85, where
x is the rate of N application [kg ha1], and
SOC is for the 0.15 cm [0.06 in] depth).
Efficiency of SOC sequestration, conversion
of biomass-C on the surface into SOC, also
follows a quadratic function (Efficiency [%]
= 15.3 + 0.829x 0.0037x 2, where x is
the N rate in kg ha1, r 2 = 0.8). In terms of
the leaching losses of soil N, nonleguminous
cover crops (e.g., rye) may be more effective
in reducing residual nitrate (NO3) and its
leaching than leguminous cover crops, such
as hairy vetch or crimson clover (Sainju et al.
1998; Dabney et al. 2001). Cover crops can
reduce the NO3 leaching potential in rain-
fed (Strock et al. 2004; Kladivko et al. 2004;
Kaspar et al. 2007) and irrigated systems
NOV/DEC 2015VOL. 70, NO. 6 331
 Table 2
Rate of soil organic carbon (SOC) sequestration by cover cropping.
Cropping Cover Rate
Location Region system crop Depth (cm) (Mg C ha1 y1) Reference
Pennsylvania Appalachian ridge, United States Cornsilage Rye 0 to 10 0 Dell et al. 2008
Illinois Southern, United States Cornsoybean Mixed 0 to 75 0.49 to 0.85 Olson et al. 2014
Illinois Southern, United States Cornsoybean Mixed 0 to 75 0 Olson et al. 2010
Harvesting for biofuel Rye 0 Fronning et al. 2008
Pisa Central Italy Corn Mixed 0 to 30 0.08 to 0.34 Mazzoncini et al. 2011
Nebraska Western Corn Belt, United States Cornsoybean Clover 0 to 7.6 0.135 to 0.174 Liebig et al. 2002
Fort Valley Georgia, United States Wheatsoybean Alfalfa 0 to 20 0.9 Sainju et al. 2002
Fort Valley Georgia, United States Mixed 0 to 20 0.5 to 0.6 Sainju et al. 2002
Belle Mina Alabama, United States Cottoncottoncorn Rye 0 to 20 0.53 to 1.12 Sainju et al. 2008

ha1 y1 (0.036 to 0.15 tn ac1 yr1; table 4). In

Figure 3 comparison with continuous corn, the rate
Process affecting gaseous emission and soil carbon (C) sequestration by cover cropping. of SOC sequestration in 0 to 7.6 cm (0 to 3
in) layer in soil of western Nebraska, United

Copyright 2015 Soil and Water Conservation Society. All rights reserved.
Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org
States, under cornsoybean (Glycine max)
oat rotation with clover over 15 years was
Increasing 0.123 to 0.174 Mg C ha1 y1 (0.05 to 0.078
net biome I n cr tn C ac1 yr1; table 5). The amount of crop
ual eas residue returned over the 15 year period was
sid productivity i ng
re (NBP)
e co 5.3, 4.8, 5.3, and 5.1 Mg ha1 y1 (2.36, 2.14,
e an il C sy 2.36, and 2.27 tn C ac1 yr1) for continu-
so
ti m g m

ous corn, cornsoybean, cornoatsorghum

st
em
of
sin

(Sorghum bicolor)soybean, and cornsoy-

e

Biomass production
Cb
rea

synthesis

Enhancing rhizospheric Erosion control beancornoatclover rotation, respectively.

Photo-

ud g
Inc

processes
Improving depth
Reducing fire risks A five-year study conducted in Georgia,
et

distribution of soil C Decreasing losses of United States, with and without hairy vetch
dissolved organic C
at three levels of N and for three tillage (NT,
Increasing Cover chisel plow, and moldboard plow) treatments
cropping
soil organic Stabilizing CH4 uptake Reducing CH4 indicated a strong increase in SOC pool with
impacts on GHG
and inorganic aggregates emissions from emissions a cover crop, but only for the NT system of
C pools arable lands seedbed preparation (table 6). The rate of
Increasing uptake of Improved aeration
SOC sequestration for 0 to 20 cm (0 to 7.9
available N by the Good soil structure and
in) depth was 0.9 Mg ha1 y1 (0.40 tn ac1
Uptake of
soluble N

cover crop tilth yr1). A companion experiment showed an

Mo trific
den

Moderating soil hydro- average rate of SOC sequestration of 0.50,

sis

thermal regime and

de ati

0.52, and 0.63 Mg C ha1 y1 (0.22, 0.23,

i

ne
r a on

process
tin ,

and 0.28 tn ac1 yr1) for cover cropping

ge

it
g

no

an rifica with crimson clover, hairy vetch, and rye,

n

Decreasing
ha

d et
lea tion, N2O emissions d m
ch e in cotton (Gossypium hirsutum)-based system
ing uc
Reducing Red
leaching of
NO3
(Delgado 1998, 2001). Cover crops can even There exists a strong interaction between
function as vertical filter strips to recover and NT and cover cropping. For example, in
mine NO3 from soil (Delgado 1998, 2001; Pisa, central Italy, Mazzoncini et al. (2011)
Delgado et al. 2001a, 2001b). Some cover reported that the SOC pool in 0 to 30 cm (0
crops can also increase the amount of soluble to 11.8 in) depth increased under NT at the
organic C in the soil (Zhou et al. 2010) and rate of 0.04 Mg C ha1 y1 (0.018 tn C ac1
thus exacerbate the leaching losses. yr1). With cover cropping, however, the rate
of increase ranged from 0.08 to 0.34 Mg C
respectively. The rate of SOC sequestration
with rye cover crop in Belle Mina, Alabama,
United States, for 0 to 20 cm (0 to 7.9 in)
depth was 0.53 to 0.76 Mg C ha1 y1 (0.24
to 0.34 tn C ac1 yr1), which increased to
1.12 Mg C ha1 y1 (0.50 tn C ac1 yr1) with
addition of poultry litter (table 7).
An experiment conducted in South Africa
indicated increase in soil organic matter
(SOM) in the 0 to 5 cm (0 to 1.97 in) layer
with increase in input of biomass-C (SOM
[g kg1] = 0.2x + 21.6, r 2 = 0.43, x = Mg ha1
of biomass) and fertilizer N (SOM [g kg1] =
0.15x + 28.3, r 2 = 0.77, x = kg N ha1) under
corncover crop and cornoat rotation,

332 NOV/DEC 2015VOL. 70, NO. 6 JOURNAL OF SOIL AND WATER CONSERVATION
 Figure 4
Ecosystem services provisioned by cover cropping through short- and long-term benefits.
Direct effects
Weed and pest
Erosion control
management
Mycorrhizal Water
colonization conservation
Risk management
respectively (Dube et al. 2012), by improving
biomass production. Application of N and
residue retention with NT can improve SOC
concentration (and stock) in the surface layer
up to the 30 cm (0 to 11.8 in) depth (Havlin
et al. 1990). Improvements in soil fertility,
and thus input of additional biomass-C, by
growing cover crops was reported in tropical
savanna ecosystems (Ibewiro et al. 2000; Tain
et al. 200) and in temperate regions (McVay
et al. 1989; Sainju et al. 2005). However,
empirical and modeling (Kemanian and
Stckle 2010) techniques need to be devel-
oped to assess the rate of SOC concentration.
The rates of SOC sequestration, discussed
above and summarized in tables 2 and 3, are
in accord with those reported for NT system
by West and Post (2002).
A study conducted by Olson et al. (2014)
in Illinois, United States, showed SOC
sequestration by cover copping at the rate of
0.5 to 0.9 Mg C ha1 y1 (0.22 to 0.40 tn
C ac1 yr1) to 75 cm (29.5 in) depth. Even
if only half of these rates (0.5 to 0.9 Mg C
ha1 y1) of 0.25 to 0.50 Mg C ha1 y1 (0.11 Cover Cropping and Conservation
to 0.20 tn C ac1 yr1) can be sequestered by
cover cropping on arable land at global scale,
it can make a significant impact on net C
emission from agroecosystems. For arable
land area of 1.5 billion ha (3.71 billion ac),
and the SOC sequestration rate of Olson et
al. (2014), technical or maximum potential
of SOC sequestration by cover cropping in
conjunction with NT is 0.4 to 0.8 Pg C y1
(0.41 to 0.82 109 tn C yr1). It is import-
JOURNAL OF SOIL AND WATER CONSERVATION
Cover cropping and soil quality
Impact
Indirect effects
Ecosystem services
Plant nutrients Soil structure
Retention Soil biodiversity
Fixation (N) and aggregation
Soil organic
GHG emissions Economic profit Water quality
carbon
Copyright 2015 Soil and Water Conservation Society. All rights reserved.
Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org
Stewardship and ethics
Table 3
Impact of cover cropping on rate of soil organic carbon (SOC) sequestration under the two
tillage systems under cornsoybean rotation on a typic Fraguidal in Southern Illinois (adapted
from Olson et al. [2014]).
Gain in SOC stock by Rate of SOC
12 years of cover cropping sequestration
Tillage Depth (cm) in 0 to 75 cm (Mg ha1) (Mg ha1 y1)
No-till 0 to 15 5.4 0.45
15 to 75 4.9 0.41
0 to 75 10.3 0.85
Chisel plow 0 to 15 3.4 0.28
15 to 75 2.5 0.21
0 to 75 5.9 0.49
Moldboard plow 0 to 15 0.1 0.008
15 to 75 1.1 0.09
0 to 75 1.2 0.10
ant, however, that cover cropping is practiced harvested in plants and animal produce, and
in conjunction with NT, residue mulching, (5) effective weed control by judicious com-
integrated nutrient management (INM), and bination of IPM. Environmental benefits of
integrated pest management (IPM). cover cropping increase when a mixture of
several species is used rather than one species
(Kramberger et al. 2013). Indeed, some con-
Agriculture cerns about impacts of NT on crop yields
No-till farming is a component of CA (Lal (Pittelkow et al. 2014) or limited potential
2015). Principal components of CA include of soil C sequestration (Powlson et al. 2014)
(1) minimal or no soil disturbances, (2) reten- are probably due to the lack of adoption of
tion of crop residue mulch, (3) use of complex a complete package of NT and CA com-
crop rotations including cover crops, (4) use prising of the five components listed above,
of INM by a judicious combination of inor- especially the absence of crop residue mulch
ganic and organic fertilizers and elemental and a cover cop in the rotation cycle.
recycling along with biological nutrient fix- Indeed, benefits of NT and CA are pro-
ation to replace those removed by crops or visioned through the presence of crop
NOV/DEC 2015VOL. 70, NO. 6 333
 Table 4
Effects of cover cropping on soil organic carbon (SOC) sequestration in Central Italy (recalculated from Mazzoncini et al. [2011]).
Rate of SOC
N supplied by cover SOC pool in 0 to 30 cm (Mg ha1) sequestration
Cover crop cover crop (Kg N ha ) 1993 2008 Change (Mg ha1 y1)
1

Control 45.2 46.6 1.4 0

Nonlegume 55 45.9 48.5 2.6 0.08
Low N supply legume 87 44.4 50.5 6.1 0.31
High N supply legume 110 44.7 51.2 6.5 0.34
Notes: Nonlegume = Rye, brown mustard (Brassica juncea). Low nitrogen (N) supply legume = Crimson clover (Trifolium incarnatum), squarrosum
clover (Trifolium squarrosum).

Table 5
Effect of cover crops in the rotation cycle and rate of nitrogen (N) fertilizer for 16 years on soil organic carbon (SOC) sequestration in Nebraska,
United States (recalculated from Liebig at el. [2002]).
SOC in 0 to 7.6 cm (Mg ha1) SOC sequestration rate (Mg ha1 y1)
Crop sequence Zero N Low N High N Zero N Low N High N

Copyright 2015 Soil and Water Conservation Society. All rights reserved.
Continuous corn 13.5 14.6 17.8 0 0.074 0.287

Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org

Cornsoybean 14.1 14.7 15.1 0.041 0.080 0.107
Cornoatcloversorghumsoybean 15.8 15.5 16.1 0.154 0.133 0.174
Cornsoybeansorghumoatclover 14.9 15.2 14.9 0.094 0.113 0.094
Notes: N rate = 0, 90, and 180 kg ha1 for corn and grain sorghum; and 0, 34, and 68 kg ha1 for soybean and oat and clover.

Figure 5 residue mulch and inclusion of an appropri-

Factors affecting soil structure and aggregation by cover cropping. ate cover crop in the rotation cycle. Cover
crops suppress weed growth as living mulch
and as residues after they have been killed
Soil (Teasdale et al. 2007; Ross et al. 2001) as
micro-fauna well as through allelopathic effects (Weston
on Fungi 1996; Barnes and Putnam 1983; Liebman
ati Mycorrhizal association Bio
b iliz Bacteria t ur
and Davis 2000; Kelton et al. 2012; Bais et al.
a 2003). Further, leguminous cover crops can
st b
also fix N (Sainju et al. 2000) and improve
e

at
at

ion

soil fertility.
reg
A gg

Percentages of arable land under CA ranges

from 44% to 71% in South America, 23% in
the United States, ~40% in Canada, 6% in
matic factor Soil China, 1% in India, and <1% in countries of
Cementing/ Cli s
Africa (Richards et al. 2014). By 2014, CA
binding agents microfauna
Free

was practiced on some 155 million ha (383

d dr ying

Sesquioxides Aggregation Gopher

Carbonates Earthworms
and aggregate million ac; Kassam et al. 2014) or about 10%
zing an

Clay and clay minerals Termites

Organic compounds stabilization Centipeds, millipedes of the arable land area. The area under CA is
Nematodes
an

Fungal hyphae increasing rapidly in China (Hou et al. 2011)

th Protozoa
d

Root hair awi tin

g

ng Wet and elsewhere.

Phy

Soil Aggregation and Cover Cropping

si

Understanding the impact of cover cropping

se s
ca
lc

on SOC sequestration also requires knowl-

es
by

m
he

ica
oc
pla

tr edge about its impact on soil aggregation

oo l pro
pr

Plant ic
n

ts ces roots er and structural attributes. For example, soil

s ph
Root and hairs i zos biodiversity, supported by a complex habitat
Root exudates Rh for soil biota, is enhanced by cover crop-
Depth of root ping, which is also a creative way to improve
penetration
microcommunities in soil (Wilson 2002).
Soil biodiversity, relevant to formation and
stabilization of structural aggregates, include

334 NOV/DEC 2015VOL. 70, NO. 6 JOURNAL OF SOIL AND WATER CONSERVATION

bacteria, fungi, protozoans, actinomycetes,
nematodes, mites, collembola and primitive
microarthropods (e.g., proturans, symphylans,
and dipulurans), and other mesofauna (e.g.,
enchytraeid worms) (Bruns and Byrne 2004;
Wallwork 1970). Microbial by-products (e.g.,
polysaccharides) improve aggregate stability
(Gupta and Germida 1988; Peixoto et al.
2006) as cementing agents.
Glomalin (glycoprotein) produced by
arbuscular mycorrhizal fungi (AMF) is a
strong cementing agent that enhances aggre-
gate stability (Wright et al. 1996a, 1996b)
for a wide range of soil types (Wright and
Upadhyaya 1998). Mycorrhizal fungi provide
many beneficial services to plants, including
absorption of water and mineral nutrients.
They also produce glomalin (named after
Glomales, which is the taxonomic order of
the fungi), which has a strong impact on
soil structure (see the following section).
Conversion to CA can strongly enhance
AMF inoculation, production of glomalin,
and formation and stabilization of aggregates.
In addition to the effects on other processes
(Wright et al. 1998; Wright and Upadhyaya
1999), glomalin-induced changes in soil
structure can also impact the SOC pool
(Rilling et al. 2001). Rise in CO2 by AFM
activity by cover cropping improves soil
structure (Rilling et al. 1999) and increases
mean residence time of C in soil.
Aggregation is a complex process, and
cover cropping increases aggregation (Mendes
et al. 1999) through several interacting pro-
cesses. Cover cropping impacts soil structure
(figure 4) on a long-term basis through direct
effects of vegetative cover and rooting char-
acteristics, but also through buildup of SOC
concentration. Indirectly, cover cropping also
impacts rooting characteristics and rhizo-
spheric processes, with profound impacts on
formation of organo-mineral complexes and
stabilization of structural units. Establishing
cover crops also affects soil macrofauna (e.g.,
earthworms and termites) and microfauna
(e.g., fungi), which enhance aggregation and
aggregate stabilization. Important soil micro-
organisms with strong impacts on aggregation
are mycorrhizal and saprophytic fungi and
bacteria (figure 5).
The aggregate hierarchy concept (Tisdall
and Oades 1982) is important in soils contain-
ing a higher amount of SOC (Elliot 1986).
Inorganic binding agents are relatively more
important in soils with inherently low SOC
concentration (Six et al. 2004; Fernandez-
JOURNAL OF SOIL AND WATER CONSERVATION
Table 6
Effects of hairy vetch cover crop on soil organic carbon (SOC) sequestration in Fort Valley,
Georgia, United States, over five year period (calculated from Sainju et al. [2002]).
SOC sequestration rate (Mg ha1 y1)
Tillage 0 N 90 kg N ha1 180 kg N ha1
No-till 0.9 0.9 0.88
Chisel plow 0 0.06 0.08
Moldboard plow 0.14 0.14 0.08
Notes: N = nitrogen.
Table 7
Effects of 10 years of cover cropping on soil organic carbon (SOC) sequestration rate at Belle
Mina, Alabama (adapted from Sainju et al. [2008]).
Rate of SOC
SOC pool sequestration
Rotation Fertilizer (Mg ha1) (Mg ha1 y1)
Fallow None 32.5 0
Copyright 2015 Soil and Water Conservation Society. All rights reserved.
Cottoncottoncorn None 37.8 0.53
Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org
Cottoncottoncorn NH4, NO3 38.6 0.61
Ryecottonryecottoncorn NH4, NO3 40.1 0.76
Ryecottonryecottoncorn Poultry litter 43.7 1.12
LSD (.05) 4.9
Notes: Nitrogen (N) application by ammonium (NH4), nitrate (NO3), and poultry litter = 100 kg N
ha1. LSD = least significant difference. All treatments were under no-tillage farming.
Ugalde et al. 2011). Microaggregates (20 to 250 aggregation (Tissdall 1994). Colonization by
m [0.0008 to 0.01 in]), stabilized by organic AMF and its effects on aggregation are also
substances, polyvalent cations, and sesquioxides, affected by temperature (Hetrick and Bloom
are combined into macroaggregates (>250 1984; Kabir et al. 1997b).There are also positive
m) by fungal hyphae and microbial by-prod- effects of AMF on agronomic yields and other
ucts. A better understanding of soil aggregate ecosystem services (Power 2010; Smith 2008;
formation may be realized by considering Unger 1998;Vannette and Hunter 2009).
the interactions between roots and mycorrhi- Cover crops also increase the particulate
zal fungi in relationship to plant community organic matter, or the so-called POM frac-
composition (Miller and Jastrow 1990). Cover tion (Motta et al. 2007; Ding et al. 2005),
crops affect aggregation by providing cement- which impacts soil aggregation (Dube et
ing material, especially the AMF (Schipanski et al. 2012; Verchot et al. 2010; Six et al. 2002,
al. 2014; Deguchi et al. 2007; Kabir and Koide 2004). Bioturbation caused by macrofauna
2000; White and Weil 2010), and lengthy fal- (e.g., gopher) under cover cropping can
low rotations reduce the incidence of AMF also influence inoculation potential of AMF
colonization (Thompson 1987; Boswell et (Koide and Mooney 1987).
al. 1998; Kabir et al. 1997a). Kabir and Koide It is evident, therefore, that soil aggrega-
(2000) assessed the impacts of winter wheat as tion is improved by cover crops (Robertson
a cover crop and a perennial weed dandelion et al. 1991, 1995; Hermawan and Bomke
(Taraxacum officinale) on AMF and soil aggrega- 1997; Calkins and Swanson 1998; Sainju et al.
tion under corn. Water stable aggregation was 2003; Liu et al. 2005; table 8) partly because
13%, 24%, and 28% under fallow, winter wheat, of increase in polysaccharides and other
and dandelion treatment, respectively. Positive binding agents, such as water extractable car-
impacts of AMF on soil aggregation have been bohydrates (Metzger et al. 1987; Kinsburky
widely reported (Miller and Jastrow 1992).The et al. 1989; Haynes and Swift 1990) and
AMF hyphal network can enhance aggrega- acid extractable C fractions (Haynes et al.
tion (Kabir and Koide 2000) by production of 1991; Degens 1997). Some of these organic
glomalin, which binds particles (Wright and compounds are formed in soils from micro-
Upadhaya 1996), providing a skeletal struc- bial processes (Aoyama et al. 1999). Cover
ture (Miller and Jastrow 1992) and C substrate, crop experiments conducted in Vancouver,
which enhance activity of other microor- Canada, for a short period of only eight
ganisms (Andrade et al. 1998) and improve months showed significant impacts on water
NOV/DEC 2015VOL. 70, NO. 6 335

stable aggregation (WSA) and the mean
weight diameter (MWD) of aggregates (table
7). Annual ryegrass and fall rye increased
the MWD of aggregates, and all cover crops
increased soil dilute acid extractable poly-
saccharides. Liu et al. (2005) concluded that
the dilute acid-extractable polysaccharide
fraction is an important binding agent under
short-term cover cropping. Similar results
regarding the effects of different cover crop
species on aggregate stability and aggregation
for some soils in the state of Goias, Brazil,
were reported by Nascente et al. (2014).
Cover crops increased SOC concentration
in the surface layer, which was passively cor-
related with WSA and the MWD.
Plant-derived carbohydrates are import-
Table 8
Effects of nonleguminous winter crops (August 20, 1994, to April 14, 1995) structural properties
of a soil in British Columbia, Canada (adapted from Liu et al. [2005]).
Treatment WSA (2 to 6 mm) (%) MWD (mm)
Control (bare soil) 20.5c 1.24c
Annual rye grass 42.5a 1.99a
Fall rye 33.5b 1.67ab
Spring barley 25.5bc 1.35bc
Notes: WSA = water stable aggregation. MWD = mean weight diameter. Means followed by the
same letter within the same column are not significantly different at p < 0.05.
ated with tillage methods in systems agroecoregions to assess the long-term posi-
of INM. tive effects of cover cropping on soil quality,
4. Soil aggregation: Effects of cover nutrient cycling, water holding capacity,
cropping on formation and stabiliza- infiltration rate, runoff, and nutrient pollu-
tion of microaggregates, distribution tion to the environment.
of SOC within aggregate size classes,
and mechanisms of protection of References

ant to aggregate formation in soil under
undisturbed conditions (Martins et al. 2011).
Soils under different vegetation types and
climate may have different modes of SOC
stabilization (Elliott 1986). In addition to
stabilizing SOC, improvement in soil struc-
ture by cover cropping also improves water
quality (Hoorman 2009) by reducing leach-
ing (Wilson et al. 1991) and improving
use efficiency of N and water (Reeves and
Touchton 1991).
Researchable Priorities
Despite ~3,000 years of history of cover
cropping, numerous knowledge gaps remain
in the context of global issues of the 21st
century. Whereas significant advances have
been made since 2000 (Delgado et al. 2007),
important knowledge gaps that need to be
addressed include the following:
1. Choice of cover crop: Site-specific
research is needed to identify appro-
priate cover crops and suitable
techniques of their management
along with those of crops to be grown
with them.
2. Biomass production: Research is
needed to assess the amount of bio-
mass produced, plant nutrients fixed,
shoot:root partitioning of biomass-C,
rooting characteristics, and turnover
time of SOM and different fractions.
3. Soil C sequestration:The rate of SOC
sequestration to 1 m (3.3 ft) depth
must be assessed on the basis of long-
term (>10 years) use of a rotational
sequences involving site-specific
cover crops, and the interactive effects
of cover cropping should be evalu-
planned research on principal soils and major
SOC against microbial processes
needs to be understood.
5. Water and nutrient requirements:
Field experiments are needed for
evaluating additional water and nutri-
ent requirements (footprint) of cover
cropping, along with assessment of
other managerial (herbicides and
mechanical mowing) needs. Research
is also needed on the long-term
effects of cover cropping on plant
available water holding capacity of
soil in relation to improvements in
SOC pool, aggregation, and changes
in pore size distribution.
6. Types of organic C: Effects of differ-
ent cover crops and their management
should be assessed on types of organic
substances (soluble, particulate, and
different fractions) added to the soil,
and their MRT, and impacts on soil
aggregation and other processes.
7. Incubation study: Laboratory incuba-
tion studies are needed to understand
the decomposition processes and
emission of GHGs.
8. Mixtures of cover crops: Impact of
one species versus mixture of several
species (grasses, legumes, and compo-
sitional) must be assessed on edaphic
factors and soil properties.
9. Net C gains: Net SOC sequestration
must be assessed with due consider-
ation to C costs of management.
10. Societal value of soil C: Assessing the
societal value of soil C sequestered by
cover cropping is essential to promot-
ing the adoption of this practice.
There is a strong need to conduct well-
Copyright 2015 Soil and Water Conservation Society. All rights reserved.
Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org
Andrade, G., K.L. Mihara, R.G. Linderman, and G.J.
Bethlenfalvay. 1998. Soil aggregation status and
rhizobacteria in the mycorrhizosphere. Plant and Soil
202(1):89-96.
Aoyama, M., D.A. Angers, and A. N'Dayegamiye. 1999.
Particulate and mineral-associated organic matter in
water-stable aggregates as affected by mineral fertilizer
and manure applications. Canadian Journal of Soil
Science 79(2):295-302.
Bais, H.P., R. Vepachedu, S. Gilroy, R.M. Callaway, and J.M.
Vivanco. 2003. Allelopathy and exotic plant invasion:
From molecules and genes to species interaction.
Science 301:1337-1380.
Barnes, J.P., and A.R. Putnam. 1983. Rye residues contribute
weed suppression in no-tillage copping systems. Journal
of Chemistry and Ecology 9:1057-1045.
Betts, E.M. 1953. Thomas Jefferson's farm book with
commentary and relevant extracts for other writings.
Charlottsville: American Philosophical Society,
University Press of Virginia.
Boswell, E.P., R.T. Koide, D.L. Shumway, and H.D. Addy.
1998. Winter wheat cover cropping, VA mycorrhizal
fungi and maize growth and yield. Agriculture
Ecosystems & Environment 67(1):55-65.
Brevik, E.C., and A.E. Hartemink. 2010. Early soil knowledge
and the birth and development of soil science. Catena
83(1):23-33.
Bruns, M.A, and L.B. Byrne. 2004. Scale model of a soil
aggregate and associated organisms: A teaching tool
for soil ecology. Journal of Natural Resources and Life
Science Education 33:85-91.
Calkins, J.B., and B.T. Swanson. 1998. Comparsion of
conventional and alternative nursery field management
systems: Soil physical properties. Journal of Environment
and Horticulture 16:90-97.
Dabney, S.M., J.A. Delgado, and D.W. Reeves. 2001. Use of
winter cover crops to improve soil and water quality.
Communications in Soil Science and Plant Analysis 7
and 8:1221-1250.

336 NOV/DEC 2015VOL. 70, NO. 6 JOURNAL OF SOIL AND WATER CONSERVATION
Davenport, C. 2014. A Climate Accord Based on Global Peer
Pressure. The New York Times. http://www.nytimes.
com/2014/12/15/world/americas/lima-climate-deal.html.
Degens, B.P. 1997. Macro-aggregation of soils by biological
bonding and binding mechanisms and the factors
affecting these: A review. Australian Journal of Soil
Research 35(3):431-459.
Deguchi, S., Y. Shimazaki, S. Uozumi, K. Tawaraya, H.
Kawamoto, and O. Tanaka. 2007. White clover living
mulch increases the yield of silage corn via arbuscular
mycorrhizal fungus colonization. Plant and Soil
291(1-2):291-299.
Delgado, J.A. 1998. Sequential NLEAP simulations to
examine effect of early and late planted winter cover
crops on nitrogen dynamics. Journal of Soil and Water
Conservation 53(3):241-244.
Delgado, J.A. 2001. Use of simulations for evaluation of best
management practices on irrigated cropping systems.
In Modeling Carbon and Nitrogen Dynamics for Soil
Management, ed. M.J. Shaffer, L. Ma, and S. Hansen,
355-381. Boca Raton, FL: Lewis Publishers.
Delgado, J.A., M.A. Dillon, R.T. Sparks, and S.Y.C. Essah.
2007. A decade of advances in cover crops. Journal of
Soil and Water Conservation 62(5):110A-117A.
Delgado, J.A., R.R. Riggenbach, R.T. Sparks, M.A. Dillon,
L.M. Kawanabe, and R.J. Ristau. 2001a. Evaluation of
nitrate-nitrogen transport in a potato-barley rotation.
Soil Science Society of America Journal 65:878-883.
Delgado, J.A., R.J. Ristau, M.A. Dillon, H.R. Duke, A.
Stuebe, R.F. Follett, M.J. Shaffer, R.R. Riggenbach, R.T.
Sparks, A.Thompson, L.M. Kawanabe, A. Kunugi, and K.
Thompson. 2001b. Use of innovative tools to increase
nitrogen use efficiency and protect environmental
quality in crop rotations. Communications in Soil
Science and Plant Analysis 32:1321-1354.
Dell, C.J., P.R. Salon, C.D. Franks, E.C. Benham, and
Y. Plowden. 2008. No-till and cover crop impacts on
soil carbon and associated properties on Pennsylvania
dairy farms. Journal of Soil and Water Conservation
63(3):136-142, doi:10.2489/jswc.63.3.136.
Ding, G., X. Liu, S. Herbert, J. Novak, D. Amarasiriwardena,
and B. Xing. 2005. Effect of cover crop management on
soil organic matter. Geoderma 130:229-239.
Dube, E., C. Chiduza, and P. Muchaonyerwa. 2012.
Conservation agriculture effects on soil organic matter
on a Haplic Cambisol after four years of maize-oat and
maize-grazing vetch rotations in South Africa. Soil &
Tillage Research 123:21-28.
Duggar, J.F. 1898. Experiments with crimson clover and
hairy vetch. Alabama Agricultural Experiment Station,
Bulletin No. 96.
Elliott, E.T. 1986. Aggregate structure and carbon, nitrogen,
and phosphorus in native and cultivated soils. Soil
Science Society of America Journal 50:627-633.
Fernandez-Ugalde, O., I. Virto, P. Barre, N. Gartzia-
Bengoetxea, A. Enrique, M.J. Imaz, and P. Bescansa.
2011. Effect of carbonates on the hierarchical model of
JOURNAL OF SOIL AND WATER CONSERVATION
aggregation in calcareous semi-arid Mediterranean soils.
Geoderma 164(3-4):203-214.
Fronning, B.E., K.D. Thelen, and D.H. Min. 2008. Use of
manure, compost, and cover crops to supplant crop
residue carbon in corn stover removed cropping systems.
Agronomy Journal 100(6):1703-1710.
Gupta, V.V.S.R., and J.J. Germida. 1988. Distribution
of microbial biomass and its activity in different soil
aggregate size classes as affected by cultivation. Soil
Biology & Biochemistry 20(6):777-786.
Halvorson, A.D., G.A. Peterson, and C.A. Roule. 2002a.
Tillage system and crop rotation effects on dryland
crop yields and soil carbon in the central Great Plains.
Agronomy Journal 94:1429-1436.
Halvorson, A.D., B.J. Wienhold, and A.L. Black. 2002b.
Tillage, nitrogen, and cropping system effects on soil
carbon sequestration. Soil Science Society of America
Journal 66(3):906-912.
Havlin, J.L., D.E. Kissel, L.D. Maddux, M.M. Classen, and
J.H. Long. 1990. Crop-rotation and tillage effects on soil
organic-carbon and nitrogen. Soil Science Society of
America Journal 54(2):448-452.
Haynes, R.J., and R.S. Swift. 1990. Stability of soil aggregates
in relation to organic-constituents and soil-water
content. Journal of Soil Science 41(1):73-83.
Haynes, R.J., R.S. Swift, and R.C. Stephen. 1991.
Influence of mixed cropping rotations (pasture arable)
on organic-matter content, water stable aggregation and
clod porosity in a group of soils. Soil & Tillage Research
19(1):77-87.
Hermawan, B., and A.A. Bomke. 1997. Effects of winter
cover crops and successive spring tillage on soil
aggregation. Soil & Tillage Research 44(1-2):109-120.
Hetrick, B.A.D., and J. Bloom. 1984. The influence of
temperature on colonization of winter-wheat by
vesicular-arbuscular mycorrhizal FUNGI. Mycologia
76(5):953-956.
Hoorman, J.J. 2009. Using cover crops to improve soil and
water quality. In Fact Sheet: Ohio State University.
Hou, R.X., Z. Ouyang, Y.S. Li, D.D. Tyler, F.D. Li, and G.V.
Wilson. 2012. Effects of tillage and residue management
on soil organic carbon and total nitrogen in the North
China Plain. Soil Science Society of America Journal
76(1):230-240.
Hunter, L., and C. Falen. 2012. Green manure for oils
nutrient management in a high-desert farming systems.
University of Idaho.
Ibewiro, B., N. Sanginga, B.Vanlauwe, and R. Merckx. 2000.
Nitrogen contributions from decomposing cover crop
residues to maize in a tropical derived savanna. Nutrient
Cycling in Agroecosystems 57(2):131-140.
IPCC (Intergovernmental Panel on Climate Change). 2014.
Part A: Global and Sectoral Aspects. Contribution of
Working Group II to the Fifth Assessment Report
of the Intergovernmental Panel on Climate Change.
In Climate Change 2014: Impacts, Adaptation, and
Vulnerability. Cambridge and New York: Cambridge
University Press.
Janisch, J.E., and M.E. Harmen. 2002. Successional changes
in live and dead wood carbon stores: Implications for
net ecosystem, productivity. Tree Physiology 22:77-89.
Jansson, C., S.D. Wullschleger, U.C. Kalluri, and
G.A. Tuskan. 2010. Phytosequestration: Carbon
biosequestration by plants and the prospects of genetic
engineering. Bioscience 60(9):685-696.
Kabir, Z., and R.T. Koide. 2000. The effect of dandelion or
a cover crop on mycorrhiza inoculum potential, soil
aggregation and yield of maize. Agriculture Ecosystems
& Environment 78(2):167-174.
Kabir, Z., I.P. OHalloran, and C. Hamel. 1997a.
Overwinter survival of arbuscular mycorrhizal hyphae
is favored by attachment to roots but diminished by
Copyright 2015 Soil and Water Conservation Society. All rights reserved.
disturbance. Mycorrhiza 7(4):197-200.
Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org
Kabir, Z., I.P. OHalloran, J.W. Fyles, and C. Hamel. 1997b.
Seasonal changes of arbuscular mycorrhizal fungi as
affected by tillage practices and fertilization: Hyphal
density and mycorrhizal root colonization. Plant and
Soil 192(2):285-293.
Kaspar, T.C., D.B. Jaynes, T.B. Parkin, and T.B. Moorman.
2007. Rye cover crop and gamagrass strip effects on
NO3 concentration and load in tile drainage. Journal of
Environmental Quality 36:1503-1511.
Kassam, A., T. Friedrich, F. Shaxson, H. Bartz, I. Mello, J.
Kienzle, and Pretty J. 2014. The spread of conservation
agriculture: Policy and institutional support for adoption
and uptake. Field Actions Science Reports 2014.
Kelton, J., A.J. Price, and J. Mosjidis. 2012. Allelopathis weed
suppression through the use of cover crops, 115-130.
www.intechopem.com.
Kemanian, A.R., and C.O. Stockle. 2010. C-Farm: A simple
model to evaluate the carbon balance of soil profiles.
European Journal of Agronomy 32(1):22-29.
Kintisch, E. 2014. Climate change: A bold baby step on
emissions. Science 344(6188):1070-1071.
Kirchmann, H., and G. Thorvaldsson. 2000. Challenging
targets for future agriculture. European Journal of
Agronomy 12(3-4):145-161.
Kladivko, E.J., J.R. Frankenberger, D.B. Jaynes, D.W. Meek,
B.J. Jenkinson, and N.R. Fausey. 2004. Nitrate leaching
to subsurface drains as affected by drain spacing
and changes in crop production system. Journal of
Environmental Quality 33(5):1803-1813.
Koide, R.T., and H.A. Mooney. 1987. Spatial variation in
inoculum potential of vesicular arbuscular mycorrhizal
fungi caused by formation of gopher mounds. New
Phytologist 107(1):173-182.
Kramberger, B., A. Gselman, M. Podvrsnik, J. Kristl, and
M. Lesnik. 2013. Environmental advantages of binary
mixtures of Trifolium incarnatum and Lolium multiflorum
over individual pure stands. Plant Soil and Environment
59(1):22-28.
Kuo, S., U.M. Sainju, and E.J. Jellum. 1997. Winter cover
crop effects on soil organic carbon and carbohydrate
NOV/DEC 2015VOL. 70, NO. 6 337
 in soil. Soil Science Society of America Journal
61(1):145-152.
Lal, R. 2004. Soil carbon sequestration impacts on
global climate change and food security. Science
304(5677):1623-1627
Lal, R. 2014. Societal value of soil carbon. Journal of Soil and
Water Conservation 69(6):186A-192A, doi:10.2489/
jswc.69.6.186A.
Lal, R. 2015. Sequestering carbon and increasing productivity
by conservation agriculture. Journal of Soil and Water
Conservation 70(3):55A-62A, doi:10.2489/jswc.70.3.55A.
Larson, W.E., Y.B. Morachan, C.E. Clapp, and W.H. Pierre.
1972. Effects of increasing amounts of organic residues
on continuous corn: II. Organic carbon, nitrogen,
phosphorus, and sulfur. Agronomy Journal 64(2):204-&.
Liebig, M.A., G.E. Varvel, J.W. Doran, and B.J. Wienhold.
2002. Crop sequence and nitrogen fertilization effects
on soil properties in the Western Corn Belt. Soil Science
Society of America Journal 66(2):596-601.
Liebman, M., and A.S. Davis. 2000. Integration of soil, crop
and weed management in low-external input farming
systems. Weed Research 40:27-47.
Lipman, J.G. 1912. The associative growth of legumes and
nonlegumes. New Brunswick: Bulletin No. 253, New
Jersey Agricultural Experiment Station.
Liu, A.G., B.L. Ma, and A.A. Bomke. 2005. Effects of cover
crops on soil aggregate stability, total organic carbon, and
polysaccharides. Soil Science Society of America Journal
69(6):2041-2048.
Martins, M.D., D.A. Angers, and J.E. Cora. 2012.
Carbohydrate composition and water-stable aggregation
of an oxisol as affected by crop sequence under no-till.
Soil Science Society of America Journal 76(2):475-484.
Mazzoncini, M., T.B. Sapkota, P. Barberi, D. Antichi, and
R. Risaliti. 2011. Long-term effect of tillage, nitrogen
fertilization and cover crops on soil organic carbon
and total nitrogen content. Soil & Tillage Research
114(2):165-174.
Mcvay, K.A., D.E. Radcliffe, and W.L. Hargrove. 1989.Winter
legume effects on soil properties and nitrogen-fertilizer
requirements. Soil Science Society of America Journal
53(6):1856-1862.
Meisinger, J., W.L. Hargrove, R.L. Mikkelsen, J.R.
Williams, and V.W. Benson. 1991. Effects of cover
crops on groundwater quality. In Cover Crops for
Clean Water, ed. W.L. Hargrove. Ankeny, IA: Soil and
Water Conservation Society.
Mendes, I.C., A.K. Bandick, R.P. Dick, and P.J. Bottomley.
1999. Microbial biomass and activities in soil aggregates
affected by winter cover crops. Soil Science Society of
America Journal 63(4):873-881.
Metzger, L., D. Levanon, and U. Mingelgrin. 1987. The
effect of sewage-sludge on soil structural stability
microbiological aspects. Soil Science Society of America
Journal 51(2):346-351.
Miller, R.M., and J.D Jastrow. 1992. The role of mycorrhizal
fungi caused by formation of gopher mounds, ed.
338 NOV/DEC 2015VOL. 70, NO. 6
G.J. Behtlenfalvay and R.G. Linderman. ASA Special
Publication. Madison,WI: Soil Science Society of America.
Miller, R.M., and J.D. Jastrow. 1990. Hierarchy of root and
mycorrhizal fungal interactions with soil aggregation.
Soil Biology & Biochemistry 22(5):579-584.
Motta, A.C.V., D.W. Reeves, C. Burmester, and Y. Feng.
2007. Conservation tillage, rotations, and cover crop
affecting soil quality in the Tennessee valley: Particulate
organic matter, organic matter, and microbial biomass.
Communications in Soil Science and Plant Analysis
38(19-20):2831-2847.
Nascente, A.S., C.A.C. Crusciol, and L.F. Stone. 2014. Straw
degradation and nitrogen release from cover crops under
no-tillage. Revista Caatinga 27(2):166-175.
Olson, K., S.A. Ebelhar, and J.M. Lang. 2014. Long-term
effects of cover crops on crop yields, soil organic carbon
stocks and sequestration. Open Journal of Soil Science
4:284-292.
Pacala, S., and R. Socolow. 2004. Stabilization wedges:
Solving the climate problem for the next 50 years with
current technologies. Science 305(5686):968-972.
Paine, L.K., and H. Harrison. 1993. The historical roots of
living mulch and related practices. HortTechnology
3(2):137-143.
Panek, J.A., P.A. Matson, I. Ortiz-Monasterio, and
P. Brooks. 2000. Distinguishing nitrification and
denitrification sources of N(2)O in a Mexican
wheat system using (15)N. Ecological Applications
10(2):506-514.
Peixoto, R.S., H.L.C. Coutinho, B. Madari, P.L.O.A.
Machado, N.G. Rumjanek, J.D. Van Elsas, L. Seldin,
and A.S. Rosado. 2006. Soil aggregation and bacterial
community structure as affected by tillage and cover
cropping in the Brazilian Cerrados. Soil & Tillage
Research 90(1-2):16-28.
Pieters, A.J. 1917. Green manuring: A review of the American
experiment station literature. Journal of the American
Society of Agronomy 9(2):62-82.
Pieters, A.J. 1927. Green manuring, principles and practices.
New York: Wiley.
Pittelkow, C.M., X. Liang, B.A. Linquist, K.J. van Groenigen,
J. Lee, M.E. Lundy, N. van Gestel, J. Six, R.T. Venrerea,
and C. van Kessel. 2014. Productivity limits and
potentials of the principles of conservation agriculture.
Nature 517(7534):365-8.
Power, A.G. 2010. Ecosystem services and agriculture:
tradeoffs and synergies. Philosophical Transactions
of the Royal Society B-Biological Sciences
365(1554):2959-2971.
Powlson, D.S., C.M. Stirling, M.L. Jat, B.G. Gerard, C.A.
Palm, P.A. Sanchez, and K.G. Cassman. 2014. Limited
potential of no-till agriculture for climate change
mitigation. Nature Climate Change 4(8):678-683.
Rasmussen, P.E., R.R. Allmaras, C.R. Rohde, and N.C.
Roager. 1980. Crop residue influences on soil carbon
and nitrogen in a wheat-fallow system. Soil Science
Society of America Journal 44(3):596-600.
Rasmussen, W.D. 1975. Agriculture in the United States: A
documented history.Vol. 1. New York: Random House.
Reeves, D.W., and J.T. Touchton. 1991. Influence of fall
tillage and cover crops on soil water and nitrogen use
efficiency of corn grown on a coastal plain soil. In Cover
Crops for Clean Water, ed. W.L. Hargrove. Ankeny,
IA: Soil and Water Conservation Society.
Richards, M., T. Sapkota, C. Stirling, C. Thierfelder,
N. Verhulst, T. Friedrich, and J. Kienzle. 2014.
Conservation Agriculture: Implementation guidance for
policy makers and investors. In Practice Brief, Climate-
Smart Agriculture. Food and Agriculture Organization
of the United Nations.
Rilling, M., S.F. Wright, M.F. Allen, and C.B. Field. 1999.
Rise in carbon dioxide changes soil structure. Nature
400:628.
Copyright 2015 Soil and Water Conservation Society. All rights reserved.
Rilling, M.C., S.F. Wright, K.A. Nichols, W.F. Schmidt,
Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org
and M.S. Torn. 2001. Large contribution of arbuscular
mycorrhizal fungi to soil carbon pools in tropical forest
soils. Plant Soil 233:167-177.
Robertson, E.B., S. Sarig, and M.K. Firestone. 1991.
Cover crop management of polysaccharide-mediated
aggregation in an orchard soil. Soil Science Society of
America Journal 55(3):734-739.
Robertson, E.B., S. Sarig, C. Shennan, and M.K. Firestone.
1995. Nutritional management of microbial
polysaccharide production and aggregation in an
agricultural soil. Soil Science Society of America Journal
59(6):1587-1594.
Ross, S.M., J.R. King, R.C. Izaurralde, and J.T. O'Donovan.
2001. Weed suppression by seven clover species.
Agronomy Journal 93(4):820-827.
Sainju, U.M., B.P. Singh, and W.F. Whitehead. 1998. Cover
crop root distribution and its effects on soil nitrogen
cycling. Agronomy Journal 90(4):511-518.
Sainju, U.M., B.P. Singh, and W.F. Whitehead. 2002.
Long-term effects of tillage, cover crops, and
nitrogen fertilization on organic carbon and nitrogen
concentrations in sandy loam soils in Georgia, USA. Soil
& Tillage Research 63(3-4):167-179.
Sainju, U.M., B.P. Singh, S. Rahman, and V.R. Reddy. 2000.
Tillage, cover cropping, and nitrogen fertilization
influence tomato yield and nitrogen uptake. Hortscience
35(2):217-221.
Sainju, U.M.,W.F.Whitehead, and B.P. Singh. 2005. Biculture
legume-cereal cover crops for enhanced biomass
yield and carbon and nitrogen. Agronomy Journal
97(5):1403-1412.
Sainju, U.M., W.F. Whitehead, and B.R. Singh. 2003.
Cover crops and nitrogen fertilization effects on soil
aggregation and carbon and nitrogen pools. Canadian
Journal of Soil Science 83(2):155-165.
Sainju, U.M., Z.N. Senwo, E.Z. Nyakatawa, I.A. Tazisong,
and K.C. Reddy. 2008. Soil carbon and nitrogen
sequestration as affected by long-term tillage, cropping
systems, and nitrogen fertilizer sources. Agriculture
Ecosystems & Environment 127(3-4):234-240.
JOURNAL OF SOIL AND WATER CONSERVATION
Schipanski, M.E., M. Barbercheck, M.R. Douglas, D.M.
Finney, K. Haider, J.P. Kaye, A.R. Kemanian, D.A.
Mortensen, M.R. Ryan, J. Tooker, and C. White. 2014.
A framework for evaluating ecosystem services provided
by cover crops in agroecosystems. Agricultural Systems
125:12-22.
Simkhovich, V.G. 1937. Toward an understanding of Jesus.
New York: The Macmillan Co.
Six, J., H. Bossuyt, S. Degryze, and K. Denef. 2004. A history
of research on the link between (micro) aggregates, soil
biota, and soil organic matter dynamics. Soil & Tillage
Research 79(1):7-31.
Six, J., R.T. Conant, E.A. Paul, and K. Paustian. 2002.
Stabilization mechanisms of soil organic matter:
Implications for C-saturation of soils. Plant and Soil
241(2):155-176.
Smith, S.E., and D.J. Read. 2008. Mycorrhizal Symbiosis,
3rd ed. Amsterdam: Academic.
Snapp, S.S., S.M. Swinton, R. Labarta, D. Mutch,
J.R. Black, R. Leep, J. Nyiraneza, and K. O'Neil.
2005. Evaluating cover crops for benefits, costs and
performance within cropping system niches. Agronomy
Journal 97(1):322-332.
Socolow, R.H., and S.W. Pacala. 2006. A plan to keep
carbon in check. Scientific American 295(3):50-57.
Strock, J.S., P.M. Porter, and M.P. Russelle. 2004. Cover
cropping to reduce nitrate loss through subsurface
drainage in the northern US Corn Belt. Journal of
Environmental Quality 33:1010-1016.
Teasdale, J.R., L.O. Brandsaeter, A. Calegari, F.S. Neto,
M.K. Upadhyaya, and R.E. Blackshaw. 2007. Cover
Crops and Weed Management. In Non Chemical Weed
Management: Principles, Concepts and Technology,
ed. M.K. Blackshaw and M.K. Upadhyaya, 49-64.
Vancouver: University of British Columbia.
Tennesen, M. 2014. Rare earth. Science 346(6210):692-695.
The White House. 2014. U.S.-China Joint Announcement
on Climate Change. Washington, DC: Office of
the Press Secretary. https://www.whitehouse.gov/
t he -p ress- of fice/ 2 0 1 4 / 1 1 / 1 1 / us-china -j o in t-
announcement-climate-change.
Thompson, J.P. 1987. Decline of vesicular-arbuscular
mycorrhizae in long fallow disorder of field crops
and its expression in phosphorus deficiency of
sunflower. Australian Journal of Agricultural Research
38(5):847-867.
Tian, G., G.O. Kolawole, B.T. Kang, and G. Kirchhof.
2000. Nitrogen fertilizer replacement indexes of legume
cover crops in the derived savanna of West Africa. Plant
and Soil 224(2):287-296.
Tisdall, J.M., and J.M. Oades. 1982. Organic matter and
water-stable aggregates in soils. Journal of Soil Science
33:141-163.
Troeh, F.R., J.A. Hobbs, and R.L. Donahue. 2004. Soil and
water conservation for productivity and environmental
protection, 4th ed. Upper Saddle River, NJ: Pearson
Prentice Hall.
JOURNAL OF SOIL AND WATER CONSERVATION
Unger, P.W., and M.F. Vigil. 1998. Cover crop effects on National Academy of Sciences of the United States of
soil water relationships. Journal of Soil and Water America 110(16):6328-6333.
Conservation 53(3):200-207.
Vannette, R.L., and M.D. Hunter. 2009. Mycorrhizal fungi as
mediators of defense against insect pests in agricultural
systems. Agricultural and Forest Entomology
11(4):351-358.
Verchot, L.V., L. Dutaur, K.D. Shepherd, and A.Albrecht. 2010.
Matter stabilization in soil aggregates: Understanding
the biogeochemical mechanisms that determining the
fate of carbon inputs in soils. Geoderma 161:182-193.
Wallwork, J.A. 1970. Ecology of soil animals. London:
McGraw-Hill.
West, T.O., and W.M. Post. 2002. Soil organic carbon
sequestration rates by tillage and crop rotation: A global
data analysis. Soil Science Society of America Journal
Copyright 2015 Soil and Water Conservation Society. All rights reserved.
66(6):1930-1946.
Journal of Soil and Water Conservation 70(6):329-339 www.swcs.org
Weston, L.A. 1996. Utilization of allelopathy for weed
management in agroecosystems. Agronomy Journal
88:860-86.
White, C.M., and R.R. Weil. 2010. Forage radish
and cereal rye cover crop effects on mycorrhizal
fungus colonization of maize roots. Plant and Soil
328(1-2):507-521.
Wilson, E.O. 2002.The future of life. New York: Alfred A. Knopf.
Wilson, G.V., D.D. Tyler, J. Logan, and K. Turnage. 1991.
Tillage and cover crop effects on nitrate leaching.
In Cover Crops for Clean Water, ed. W.L. Hargrove.
Ankeny, IA: Soil Water Conservation Society.
Wright, S.F., and A. Upadhyaya. 1998. A survey of soils for
aggregate stability and glomalin a glycoprotein produced
by hyphae of arbuscular mycorrhizal fungi. Plant and
Soil 198:97-107.
Wright, S.F., and A. Upadhyaya. 1999. Quantification of
arbuscular mycorrhizal fungi activity by the glomalin
concentration on hyphal traps. Mycorriza 8:283-285.
Wright, S.F., and A. Upadhyaya. 1996. Extraction of an
abundant and unusual protein from soil and comparison
with hyphal protein of arbuscular mycorrhizal fungi.
Soil Science 161(9):575-586.
Wright, S.F., A. Upadhyaya, and J.S. Buyer. 1998. Comparison
of N-linked oligosaccharides of glomalin from arbuscular
mycorrhizal fungi and soils by capillary electrophoresis.
Soil Biology and Biochemistry 30:1853-1857.
Wright, S.F., M. Franke-Snyder, J.B. Morton, and A.
Upadhyaya. 1996. Time-course study and partial
characterization of a protein on hyphae of arbuscular
mycorrhizal fungi during active colonization of roots.
Plant and Soil 181:193-203.
Zhou, X., H. Wu, Z. Xu, and C. Chen. 2010. Winter cover
crops increase soil carbon and nitrogen cycling processes
and microbial functional diversity. In 19th World
Congress of Soil Science. Brisbane, Australia.
Zhu, X., M. Burger, T.A. Doane, and W.R. Horwath.
2013. Ammonia oxidation pathways and nitrifier
denitrification are significant sources of N2O and NO
under low oxygen availability. Proceedings of the
NOV/DEC 2015VOL. 70, NO. 6 339