chapter 5

Soil Ecology
Kate M. Scow and Matthew R. Werner

Overview of Soil Organisms,

T he community of organisms
that lives in soil plays many
important roles in the successful
becoming more aware of the
belowground community. This
chapter describes how cover
69
functioning of agricultural eco- crops affect soil organisms and
Microorganisms, 69
systems. This community con- provides an overview of other
Earthworms and Other
Soil Fauna, 72
sists of bacteria, fungi, protozoa, management practices that favor
nematodes (predators of micro- beneficial soil organisms.
Effect of Cover Crops on the organisms and pathogens of
Soil Community, 73 plants), earthworms, arthropods,
The Living Cover Crop, 73
Cover Crop Residues, 73
and other organisms. Although Overview of
certain species are harmful to
Practical Implications of crops, most are beneficial and
Soil Organisms
Interactions between Soil even essential for the well-being
Biota and Cover Crops for of plants. There are striking Microorganisms
Vineyards, 77 similarities between the roles of
Soil Structure, 77
microorganisms in the human Although invisible to the naked
Nutrient Management
body and in the soil. In spite of eye, soil microorganisms are
and Microbial
Communities, 77
the substantial attention paid to an important part of the below-
Mycorrhizal Fungi, 78 soilborne pests, the beneficial ground community in farm soils,
Disease Suppression, 78 species far outnumber harmful and they are a potentially valu-
Enhancing Earthworm ones in the soil. able asset to the grower. Their
Communities, 78 value lies in the roles they play
The importance of soil organ- in the decomposition of organic
Bibliography, 78
isms is often ignored and may be matter, improvement of soil struc-
masked and diminished in tilled ture, cycling of nutrients, and as
farming systems that rely on syn- a living reservoir of nutrients.
thetic fertilizers, high levels of The microbial community is most
pesticides, and minimal inputs beneficial to the grower when it
of carbon. Farming systems that is diverse, abundant, and active.
include organic amendments as Microbial populations play active
nutrient sources, on the other and passive roles in soil fertility.
hand, are particularly depen-
dent on biological activity to Microorganisms as agents of
decompose the amendments. change. The major groups of
Consequently, growers, advis- soil microorganisms are bacteria
ers, and researchers interested (including actinomycetes), fungi,
in cover crops may benefit by

69
and protozoa (fig. 5-1). Because these microor- nitrogen not only more available for some plants
ganisms are best characterized by the roles they but also directly susceptible to leaching and denitri-
play rather than by their individual species, they fication. Another group of organisms is most active
are often categorized in functional groups. One under poorly aerated conditions, such as when soils
important example is the bacteria and fungi that are flooded or poorly drained or when the demand
are responsible for the conversion of organic com- for oxygen is greater than what can be supplied
pounds to their mineral components, a process by diffusion through air-filled pores. Denitrifying
called mineralization. The mineralizers include bacteria convert nitrate to forms of nitrogen that
microbes that split complex and large plant mol- are lost to the atmosphere as nitrogen gas or nitrous
ecules (e.g., cellulose, hemicellulose, lignin) into oxide. Fermenters such as yeasts and certain bac-
smaller molecules, as well as microbes that convert teria decompose organic materials under anaerobic
the smaller molecules (e.g., sugars, amino acids, conditions, often forming foul-smelling substances.
aromatics, aliphatics) into simpler mineral forms There are groups of bacteria that oxidize (often
(e.g., ammonium- and nitrate-nitrogen, carbon diox- when oxygen is present) or reduce (usually when
ide, water, sulfate). oxygen is not present) many elements that are plant
nutrients. These microbial reactions can result in,
Many other agriculturally important functions depending on the element, the conversion of iron,
are carried out by soil microorganisms (table 5-1). sulfur, manganese, and some trace elements into
Excellent overviews of soil organisms and their forms that are more or less available to crops. Other
functions are provided in Killham (1994) and groups of organisms, such as mycorrhizal fungi,
Paul and Clark (1989). Important groups include make phosphorus more accessible to plants, either
the nitrifying bacteria (Nitrosolobus, Nitrobacter, by dissolving complex phosphorus-bearing sub-
Nitrosomonas) that are involved in the conversion stances or by effectively expanding the surface area
of ammonium to nitrate. This conversion makes the of plant roots.

hairs
SIZES-FORMS
section
septate
coccus FUNGI
single-celled (yeast)
BACTERIA
1 mm
stele cortex
spirillum

rod FUNGI elongation zone

filament
meristem
non septate
cap

CLAY CRYSTALS
SILT
sand

medium

PROTOZOA & ALGAE

50µm 1mm
1µm
1000µm

Figure 5-1. Sizes and forms of soil biota in relation to particle sizes. Source: Singer and Munns 1991, p. 145.

70 • Chapter 5
Table 5-1. Beneficial activities of microorganisms in soil

Function Type of Organism

Decomposition of plant residues Decomposers (bacteria, fungi, micro- and macrofauna).

Control of nitrogen availability to plants

Conversion of organic to mineral forms Mineralizers (most microorganisms)
Conversion of ammonium to nitrate Nitrifying bacteria
Denitrification of nitrate to nitrogen gas Denitrifying bacteria
(facultative and obligate anaerobes)

Conversion of atmospheric nitrogen gas to organic nitrogen Nitrogen-fixing bacteria (symbiotic and free-living)

Increase availability of phosphorus, iron, sulfur, Many organisms, mycorrhizal fungi

and other elements

Formation of soil humus Fungi

Soil aggregate formation and stability Polysaccharide-producing fungi and bacteria

Production of plant growth-promoting substances Primarily bacteria

Suppression of plant pathogens General microbial community, producers of toxins,

producers of iron chelators

Breakdown of pesticides and other toxic chemicals Biodegrading bacteria and fungi

Nitrogen-fixing bacteria, such as symbionts in cal binding of organic matter and clay with hyphae
the genus Rhizobium and the free-living Azotobacter (e.g., fungi and actinomycetes). Another important
and Azospirillum, convert elemental nitrogen gas function is the ability of certain microorganisms to
(N2) in the atmosphere to ammonia (NH3) that can decompose organic pesticides into harmless prod-
be readily used by crops. Rhizobia living in sym- ucts. If microorganisms did not degrade these sub-
biosis with legumes provide nitrogen for growth of stances, pesticide concentrations would eventually
these crops, reducing or eliminating the need for build up to hazardous levels in agricultural soils.
nitrogen fertilizer. If the cover crop is incorporated
into soil as a green manure, a large portion of the One group of microorganisms deserving special
nitrogen fixed by the symbiotic relationship can attention because of their importance in vineyards
fertilize the subsequent crops. Because of the large is the mycorrhizal fungi. Mycorrhizal fungi form
amount of energy required for nitrogen fixation, the beneficial symbiotic relationships with plant roots.
amount of nitrogen fixed by free-living bacteria is The association of fungus and root is called a
vastly lower than that fixed by symbiotic bacteria. mycorrhiza, which literally means “fungus root.”
This is because free-living organisms must compete Mycorrhizae are found on almost every type of
with many other soil microorganisms for the organ- plant; one exception is brassicas such as mustards
ic compounds that provide energy, whereas the and radishes (Bethlenfalvey and Lindermann 1992).
symbiotic organisms obtain the compounds directly The particular type of mycorrhiza formed on grapes
from the plant. and nonbrassica cover crops is called a vesicular-
arbuscular mycorrhiza (VAM) (Lindermann 1988).
Other functions of soil microorganisms affect VAM fungi enter the plant root cells and grow
soil physical properties. Their transformation of hyphae that extend out into the surrounding soil.
plant material into the more stable forms of organic Hyphae are long, thin strands that form the main
compounds that make up the humus is important body of fungi. The VAM hyphae effectively increase
to the maintenance of the organic fraction of soil. the surface area of the plant root system and help
Many organisms promote soil aggregate formation the plant to mine nutrients from areas that the plant
and stability through production of extracellular roots cannot reach or to obtain nutrients that diffuse
polysaccharides (e.g., bacteria and fungi) and physi- very slowly through the soil solution (e.g., phospho-

Soil Ecology • 71
rus). In exchange, VAM fungi receive carbohydrates ter, fumigated, or treated with fungicide or insec-
that the plant has produced through photosyn- ticides (Lee 1985). Also, there may be localized,
thesis. The plant benefits from improved nutrient short-term negative effects of ammonia-based fer-
uptake and because roots colonized by VAM are tilizers on earthworms. Cover-cropped, minimum-
often more resistant to attack by fungal pathogens tilled vineyards with limited or no chemical inputs
and parasitic nematodes (Perrin 1990; Hussey and can provide ideal habitats for earthworms.
Roncadori 1982). Separate plant individuals and
species may actually be connected below ground Because there are many different earthworms, it
by a bridge of VAM hyphae, with phosphorus being is not useful for a grower to learn to recognize the
transferred from one plant to another through this individual species. However, there are three main
underground network (Chiariello, Hickman, and types of earthworms that are defined by where the
Mooney 1982). earthworms live, what they eat, and what they look
like, and knowing the three types can be very use-
Microorganisms as a pool of nutrients. Nutrient ful (table 5-2). Most agroecosystems in California
cycling and energy flow in soil ecosystems is tied contain only endogeic earthworms. This type of
to the decomposition of organic matter. Soil organic earthworm lives and feeds below ground, where it
matter consists of broad groups of substances, often is somewhat protected from chemical and physical
called pools, that vary in their rates of decomposi- disturbances.
tion and functions. At one extreme, organic matter
includes readily decomposable materials that have Epigeic earthworms live in surface accumula-
not yet been transformed or still closely resemble tions of organic matter, and they usually disappear
their plant and animal origins. At the other extreme, in soils that are regularly cultivated or plowed
organic matter consists of humic substances that are because the surface layer of organic matter is bur-
virtually resistant to further decomposition. One of ied. Epigeic species such as the redworm (Eisenia
the most important pools of organic matter is the foetida) are used in vermicompost. Anecic earth-
microbial biomass (the mass of microorganisms). worms can play a very important role by incorpo-
The microbial biomass is a relatively available res- rating organic matter from the soil surface into the
ervoir of plant nutrients such as nitrogen and phos- deeper horizons and creating deep, continuous,
phorus (Marumoto, Anderson, and Domsch 1982). stable channels for aeration and infiltration. In
Although the size of the microbial biomass is rela- deciduous forests, this type of earthworm is capa-
tively small (e.g., its nitrogen content constitutes ble of burying the entire autumn leaf-fall within
only 1 to 5 percent of the total organic nitrogen in a couple of months (Knollenberg, Merritt, and
soil), the nutrients within this pool are recycled Lawson 1985). Werner (forthcoming) introduced
rapidly within the soil profile, perhaps 8 to 10 anecic earthworms in an apple orchard in coastal
times per year (Coleman, Reid, and Cole 1983). The California and saw apple litter incorporation rates
amount of nitrogen in microbial biomass in agricul- during the winter increase from 20 to 80 percent. In
tural soils ranges from 36 to 344 pounds per acre California, anecic earthworms are not widespread,
(40 to 385 kg/ha) (Paul and Voroney 1984). perhaps due to the relatively arid climate. In the
Pacific Northwest, in contrast, anecic earthworms
are especially abundant in perennial crops and
Earthworms and Other Soil Fauna are commonly harvested for the bait-worm market,
where they are known as nightcrawlers. Anecic
Earthworms, when present, stimulate the decom- earthworms could potentially be introduced into
position of cover crop residues. Their feeding and California vineyards, though their longevity is
burrowing activities incorporate residues and other uncertain and would partly depend on soil manage-
amendments into the soil, enhancing organic matter ment practices.
decomposition, humus formation, nutrient cycling,
and the development of soil structure (Werner There are many other types of invertebrate ani-
1993, 1994). Earthworm burrows can persist even mals that live in the soil, but because of their small
after the worms responsible for building them are size and cryptic habits they are seldom seen. They
gone, providing pathways for rapid root growth, are thought to play important roles in the decom-
water infiltration, and gas exchange. Deep-burrow- position of cover crops and in the functioning of
ing species can burrow through compacted soil and ecosystems in general. Soil mites and Collembola
penetrate plow pans. Earthworm numbers generally (springtails) number up to a million per square
decrease when soils are tilled, left bare in the win- meter of soil. Another important group of organ-

72 • Chapter 5
Table 5-2. Three types of earthworms and their characteristics

Type Appearance Habitat Food Examples

Endogeic Variable size; Continuous burrows Mixture of buried Allolobophora chlorotica;

light or no in soil; generally feed organic matter and Aporrectodea trapezoides;
pigmentation; and defecate mineral soil, Aporrectodea caliginosa
slow moving belowground decaying roots

Epigeic Small; dark red or Areas of high Large proportion Lumbricus rubellus; Eisenia
brown color; organic matter; of diet is foetida (red worm,
fast growing; litter layer, organic matter manure worm)
move quickly manure piles,
cool compost
piles

Anecic Large and very muscular; Build permanent, Feed by pulling Aporrectodea longa;
wedge-shaped tail; very deep organic matter Lumbricus terrestris
color on front end, vertical burrows; from surface (nightcrawler)
less on tail end; raised midden of down into burrow
fast moving but castings and residue before ingesting
slow growing marks burrow entrance

isms are the nematodes (see chapter 10). Although isms, particularly mycorrhizal fungi. The physical
the plant parasitic species receive more attention cover on the soil surface also moderates soil tem-
for their destructive activities on crops, nematodes perature and moisture changes, creating a generally
that prey on bacteria, fungi, and each other make up more hospitable habitat for soil organisms.
the majority of the nematodes present in soil.
As a general rule, soil with vegetation supports
Many mites, springtails, and nematodes feed higher microbial populations than does fallow
on the microbes that decompose organic matter. soil. Plant roots exude compounds such as amino
Because many of them carry microbial spores on acids, simple sugars, and organic acids, and they
their body, they inoculate organic matter with the slough off cells containing polysaccharides. These
very microbes that they feed on. Springtails feed on compounds provide a continuous energy supply to
fungi and have profound effects on fungi popula- microorganisms living in the root zone (the rhizo-
tion dynamics. Some mites are predators, as are sphere). Studies have shown that the size of the
pseudoscorpions, spiders, rove beetles, and centi- microbial biomass fluctuates seasonally in response
pedes. These creatures help to maintain balance by to the growth of crops such as wheat due to the rhi-
preying on the springtails and other decomposers zosphere effect (Lynch and Panting 1980).
that would quickly become overabundant without
regulation.
Cover Crop Residues

Effect of Cover Crops on the Soil Changes in soil habitat with crop incorporation.
One of the major effects of a cover crop on the soil
Community community is the increased input of organic mat-
ter contributed by cover crop residues. The large
The Living Cover Crop sources of carbon and energy for microbial com-
munities that could be provided by cash crop resi-
Cover crops directly influence the soil community dues are often removed from agricultural systems
by their root growth and plant cover over the win- by current management practices. Vineyards yield
ter. The majority of organisms found in soil are relatively little biomass that is returned to the field.
associated with the plant roots that provide them Vines will typically produce 1,500 to 2,500 pounds
with carbon and other nutrients. Long periods of per acre (1,700 to 2,800 kg/ha) of prunings (fresh
bare fallow may disrupt the community structure weight) and 800 to 1,500 pounds per acre (900 to
and reduce the numbers and activity of soil organ- 1,700 kg/ha) of leaves (dry weight). This is in con-

Soil Ecology • 73
trast to a typical corn crop biomass residue of 8,000 nematode populations in tomato plots (Temple et al.
to 12,000 pounds per acre (8,900 to 13,500 kg/ha). 1994; Scow et al. 1994). Sources of plant nutrients
In vineyards, leaves and early prunings usually stay to the organic system are cover crops (oat-vetch) and
on-site. Much of the residue associated with vines manure, whereas the low-input system receives cover
is somewhat woody and not as easily decomposed crops and mineral fertilizer. At the beginning of the
as cover crop residues. Even with minimal removal study, there was little difference in the size of the
of vine residues, a dryland cover crop producing microbial biomass among the different farming sys-
1,800 to 3,600 pounds per acre (2,000 to 4,000 tems. Only immediately after cover crop incorpora-
kg/ha) of dry matter could contribute a significant tion was the biomass higher in organic and low-input
amount of biomass to vineyards. than in conventional tomatoes. Since the third year
of the study, however, the microbial biomass and its
Growers often consider only mineral forms of activity has almost always been higher in the organic
essential plant nutrients (nitrogen, phosphorus, and low-input than conventional tomatoes. This 3-
sulfur). However, soil organic matter and plant resi- year period corresponds to the transition period often
dues are other important sources of these nutrients. observed by growers converting from conventional
Therefore, a pool of potentially available nitrogen to organic farming practices. Soil fertility can be
and other nutrients in organic forms is associated problematic during this period. Suprisingly, there is
with the cover crop biomass. The amount of nitro- little difference between the microbial populations in
gen contributed by cover crops is much greater if the organic and low-input systems, even though the
the cover crop is a legume and fixes nitrogen than organic system receives poultry manure in addition
if it is a grass crop. Organic forms of phosphorus to cover crops. Microbial populations in all farming
and sulfur are also associated with cover crops. systems appear to be sensitive to soil moisture and
Although little research has been conducted on temperature and often decline sharply in midsummer
the availability of organic phosphorus and sulfur, after irrigation has been reduced and when tempera-
it is likely that the organic forms are an important tures are quite high.
source of nutrients under certain soil chemical con-
ditions (e.g., at low and high pH). Measurement of the microbial biomass or popu-
lation density is a complicated and imperfect science
Changes in microbial populations. Studies have even for soil microbiologists. A common method for
shown that fields receiving organic amendments biomass determination involves fumigation of a soil
regularly for many years (e.g., in the form of green or sample with chloroform, extraction with potassium
animal manure) generally have larger and more active sulfate solution, and quantification of carbon in the
microbial populations than fields receiving synthetic extract. Population density is also determined by
fertilizers (Scow et al. 1994). Because microbes are fluorescent microscopy after staining a soil suspen-
often limited by available carbon in agricultural sion. Both methods require specialized equipment;
soils, fresh organic material added to soil stimulates however, there are institutions that provide these
microbial activity. Increases in activity and biomass measurements as a service. The anaerobic incuba-
cease when the input is used up and microbes die off. tion method for potentially mineralizable nitrogen
These boom and bust cycles of microbial activity may (PMN) is a simple method that does not require
be quite rapid. Buchanan and King (1992) observed complex equipment and provides an estimate of the
weekly fluctuations in microbial activity in both no- amount of organic nitrogen contained in microbial
till and reduced chemical input systems of continu- biomass (Doran 1987). The PMN is calculated by
ous maize and maize-wheat-soybean rotations. measuring ammonium before and after incubation of
soil that has been submerged in water for one week
A long-term study at the Sustainable Agriculture at a high temperature. Ammonium analyses can be
Farming Systems (SAFS) Project in Davis, California, performed by many soil analytical laboratories. The
has provided information on the effect of cover crops, SAFS project has found that PMN is strongly cor-
as well as other sustainable agricultural practices, on related with microbial biomass carbon measured by
soil communities in row crops in a Mediterranean fumigation extraction and that PMN is significantly
climate. Although the study is of row crops, some higher in organic than conventional soils. During the
of the findings are indicative of what would occur 1993 growing season, values of PMN ranged from 10
with use of cover crops in vineyards. The SAFS to 30 µg NH4–N per gram soil in the conventional
study compares the effect of four farming systems tomatoes and from 20 to 50 µg NH4–N per gram soil
(conventional 2-year rotation, conventional 4-year in organic tomatoes (Gunapala and Scow 1997).
rotation, organic, and low-input) on microbial and

74 • Chapter 5
 Measuring the diversity of microbial communi- Impact of cover crops on the soil community. Cover
ties is much more complicated than the measure- crops provide food for soil organisms throughout
ment of microbial biomass. Diversity analysis, a the growing cycle of the crop. Figure 5-2 is a simple
relatively new activity, has been made possible by model of the complex food webs that are based on
recent advances in biotechnology. Techniques used inputs of organic matter to the soil. While the cover
previously for identifying species, such as dilu- crop is alive and growing, the roots release sugars,
tion plate counts, are not capable of detecting from dead cells, mucilage, and other materials into the
90 to 98 percent of the microorganisms actually soil as root exudates that fuel the decomposition
present in soil and are therefore poor methods for process. If the cover crop is mowed, the cut por-
representing the soil community. Methods such as tions of the plant provide habitat, shade, and food
microscopic counts can at best quantify cells and at the soil surface. If the cover crop is incorporated
hyphae of bacteria and fungi but do not provide into the soil, all parts of the plant enter the organic
taxonomic information. Newer methods include matter pool and become a food source. The model
molecular biological approaches similar to those contains several feedback loops of nutrient flow.
used in criminological or paternity investigations For example, feces form a nutrient reservoir that
that analyze the DNA and RNA extracted from the continues to decompose after defecation through
soil microbial community. Another method analyz- enzymatic and microbial activity. Soil animals
es the phospholipid fatty acid (PLFA) profile that commonly reingest feces, a behavior known as
can be extracted directly from the membranes of coprophagy (McBrayer 1973).
bacterial and fungal cells present in soil (Bossio and
Scow 1998). In the near future, as these new meth- Ecological succession describes the changes
ods are increasingly applied in different studies, and developments that occur in ecosystems over
we will know far more about the diversity of soil time. Successional changes in the biological com-
microbial communities and how farming practices munity that spring up after cover crop incorpora-
affect this diversity. tion are a response to the availability of a new,
energy-rich food resource. The microcommunity
Changes in soil fauna. Although earthworms are that quickly develops in and around this resource
not particularly picky about what they eat, they do is distinct from the fauna of the surrounding soil
show preference for food that is high in nitrogen. (Dindal 1973). With decomposing straw, three phas-
This is partly because earthworms are very inef- es are discernible; an initial bacterial phase, a phase
ficient at assimilating the nitrogen contained in characterized by mites that eat bacteria, and a final
their food into their body tissues; however, their phase including springtails, other mites, and other
loss is our gain. Because earthworms are made up microarthropods (Naglitsch 1966). Successional
of 60 to 70 percent protein (dry weight), they need changes in the microarthropod community are
to ingest large quantities of organic matter to meet probably caused by changes in the food source
their nitrogen requirements. Therefore, leguminous brought about by microbial activity (Harding and
cover crops that are cut before they set seed (thus Stuttard 1974). There is a peak in species diversity
high in nitrogen) provide one of the best food mate- when decomposition is most active. However, if not
rials for earthworms. Cover crop residues ingested renewed regularly, substrates such as cover crop
by earthworm’s are ground up in the earthworm’s residues become exhausted. As this occurs, the
gut, inoculated with bacteria (some of which are opportunistic decomposer community is gradually
nitrogen-fixers), bathed in enzymes, and excreted as replaced by the species indigenous to the soils of
biologically active and nutrient-rich casts. Castings that area. The decomposer community is in essence
may contain 75 percent of the nitrogen found in the self-annihilating and nonreplenishing without peri-
ingested organic matter. The high levels of residual odic organic matter additions. Decomposer organ-
nutrients are largely available to plants. Earthworms isms have evolved many unique adaptations to
also release the nitrogen that is assimilated into allow them to locate and exploit short-lived food
their body tissue as ammonium-N and amino-N in sources. For example, in the absence of fresh
urine, mucoproteins, other secretions, and through organic matter, low numbers of fungi may persist
mortality (Lee 1985). in dormant forms (e.g., as spores). With the input
of organic matter, these fungi quickly grow to high
densities and contribute to the decomposition of
the plant residue.

Soil Ecology • 75
Figure 5-2. Food web of the compost pile. Energy flows in the direction of the arrows. 1 = First-level consumers;
2 = Second-level consumers; 3 = Third-level consumers. Source: Redrawn from Dindal n.d., pp. 6–7.

76 • Chapter 5
 Practical Implications of soil aggregates are broken apart into a liquid slurry
that mixes mineral soil with organic material and
Interactions between Soil Biota binding agents in the earthworm’s gut. The excreted
and Cover Crops for Vineyards casts become stable after drying and contribute to
soil structure. Dindal, Theoret, and Moreau (1978)
There are several questions growers might raise observed that the size of soil aggregates (casts) is
about the use of cover crops in vineyards. In con- related to the size of the earthworm species produc-
verting from conventional production to the use ing them. Lumbricus terrestris (nightcrawler) popu-
of organic inputs, how long will it take for the lations are positively correlated with the presence of
decomposer organisms to effectively break down 1/6-inch (4 mm) water-stable aggregates. Their verti-
cover crop residues and cycle nutrients? Are micro- cal burrows form large channels that allow rapid
bial inoculants necessary or worth the money? infiltration of water, exchange of gases deep into the
Are decomposer communities more diverse in soil, and penetration into soil by plant roots.
cover-cropped systems? Does this diversity improve
nutrient cycling or suppress pathogens? Does the
enhancement of decomposer organisms affect grape Nutrient Management and Microbial
yield or quality? Although it is not possible to Communities
answer all these questions at this time, we can dis-
cuss some of the known benefits resulting from the Nitrogen-rich cover crops (e.g., legumes) are a poten-
influence of cover crops on the soil community. tial source of nutrients if they are incorporated into
the soil during periods of crop demand. A potential
negative effect of using cover crops in grapes, how-
Soil Structure ever, is the possibility of nitrogen immobilization
after incorporation of a cover crop containing a high
Soil structure and stability are particularly important C/N ratio (e.g., grasses). Such effects are usually
in vineyards, which are often located on hillsides. temporary and do not result in long-term influences
Various forces cause the arrangement of primary soil on plant yields. Also, the effects can be minimized
particles (sand, silt, clay, and organic matter) into by proper timing of incorporation. An advantage of
soil aggregates. The arrangement and stability of cover crops with high C/N ratios is their ability to
particles, aggregates, and voids is soil structure. The reduce nitrate leaching by direct uptake of nitrogen
void or pore size distribution of soils is determined or by promoting immobilization of soil nitrate by
by the shape, size, and stability of aggregates. From microorganisms during periods when there is a high
an agricultural point of view, a well-structured soil potential for leaching. This characteristic can be
has low bulk density, is well aerated, absorbs rain- especially important in sandy soils.
fall and irrigation rapidly, and is easily penetrated
by plant roots and soil animals. The growth of fine The development of the large and active micro-
plant roots and fungal hyphae knits soil particles bial community that results from organic inputs is
and small aggregates together into larger units called desirable because of the multiple roles it plays in soil
macroaggregates. Grasses have a greater effect than do fertility. Large microbial populations may also sup-
other plants on stable aggregate production because port a greater diversity in higher trophic levels (e.g.,
a large proportion of plant biomass is maintained soil fauna) and thus increase the potential for the soil
underground in the root system. Production of poly- to sustain crop growth in the face of perturbations
saccharide gums by rhizosphere microorganisms and such as drought stress. Inoculation with biological
plant roots enhances the formation and stabiliza- additives containing microorganisms may result in
tion of macroaggregates because the materials act as short-term, but probably minor and not cost-effec-
glues to bind particles together. Roberson, Sarig, and tive, stimulation of processes regulating soil fertility.
Firestone (1991) found improved aggregate stability in Any benefit of inoculation, however, will almost
cover-cropped soils compared with conventional soils. always be short-lived because of the inability of the
Cover crops with a higher C/N ratio (e.g., mixtures con- foreign organisms to compete with the locals. The
taining grasses) may promote greater polysaccharide foundation of a healthy and thriving microbial com-
production and thus greater aggregate stability than do munity is good nutrition and protection from major
cover crops with lower C/N ratios (e.g., legumes). stresses. If the conditions in soil are good, there will
be plenty of microorganisms. If conditions are poor,
Earthworms also influence soil structure. As microorganisms will not survive or be active, no
earthworms burrow they ingest soil. The ingested matter how many organisms are added to soil.

Soil Ecology • 77
Mycorrhizal Fungi encourage diversification of earthworm commu-
nities by managing systems to include anecic or
Vesicular-arbuscular mycorrhizal fungi (VAM) pop- epigeic species or both. Managing the habitat to
ulations in vineyards are reduced by clean cultiva- encourage the increase of earthworm populations
tion of the vineyard floor, as well as by preplant would include minimizing physical disturbance of
fumigants and fungicides. Methyl bromide is very the soil (especially in the winter when earthworms
toxic to VAM, and this toxicity can result in stunt- are most active); limiting chemical use (especially
ing of grapevines (Menge 1982, 1983). The recovery fumigants, certain fungicides, and ammonium-
time of fungi is slower than that of bacteria, and it based fertilizers); providing a near-neutral pH; and
can be on the order of years for VAM (Liebman and providing a food source. A perennial or winter
Daar 1995). VAM inoculants are available commer- annual cover crop that is periodically mowed, with
cially in the form of fungal spores that are applied the residue left on the surface, is an excellent way
to the soil in an aqueous suspension. Another way to provide these conditions.
to increase VAM is to grow a cover crop. The dense
root growth of cover crops boosts VAM growth and In some cases it may be desirable to inoculate a
increases the likelihood of grape roots being colo- soil with anecic earthworms, either through direct
nized because of the close proximity between the inoculation with nightcrawlers purchased from a
root systems of cover crops and grapevines. This bait dealer or by transferring blocks of soil (1 cubic
was shown in an apple orchard, where VAM colo- foot [28 cm3] each) from an area with an established
nization and leaf phosphorus content was greater in anecic earthworm population. Another method is to
plots with a grass cover than in plots that had been set aside a small portion of a vineyard to be man-
treated with a herbicide (Atkinson 1983). aged intensively as an earthworm reservoir. This
method is especially useful in perennial crops like
grapes. If needed, the soil could be limed to bring it
Disease Suppression near pH 7, fertilized, and a cover crop established
and cut periodically to provide an organic mulch
In some cases, cover crops may reduce plant disease as food and physical cover. In this area of the vine-
through the enhancement of beneficial microorgan- yard, a population of the desired species could be
isms. Large and diverse microbial communities, introduced and built up. From this reservoir blocks
especially in the plant rhizosphere, may actually could periodically be taken and introduced into
suppress pathogens. Although the evidence for this the field. This might be done each year in the fall
suppression is not strong, it is highly suggestive. when earthworm activity is increasing. Remember
Both cover crops and manure have been observed to provide an organic mulch. The rate of spread
to be associated with suppressiveness. Mechanisms would vary with species and conditions in the field.
reported to be responsible for disease control with An individual nightcrawler is capable of traveling
the use of cover crop residues include an increased at least 63 feet (19.2 m) on the soil surface in the
course of one evening foray. However, the spread
resistance of crop plants through supplementation
and establishment of an entire earthworm colony
of nutrients or other factors; altered virulence or
will take longer.
growth of the pathogen directly (such as by alle-
lopathic effects of the cover crop); an increase in
activity of beneficial soil organisms resulting in
their shutting out pathogens through antagonism
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78 • Chapter 5
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Soil Ecology • 79