Research Signpost

37/661 (2), Fort P.O., Trivandrum-695 023, Kerala, India

Plant-Microbe Interactions, 2008: ISBN: 978-81-7895-314-4

Editors: E. Ait Barka and C. Clément

Enhancing biological control of

14 soilborne plant diseases by
organic soil amendments
Pervaiz A. Abbasi, George Lazarovits and Kenneth L. Conn
Agriculture and Agri-Food Canada, Southern Crop Protection and Food
Research Centre, 1391 Sandford Street, London, Ontario, N5V 4T3 Canada

Abstract
Organic amendments have been used in sustainable
agriculture primarily as biofertilizers and soil
conditioners. Such amendments provide nutrients to
crops, alter the biological, chemical, and physical
properties of soil, and can lead to the control of
soilborne plant diseases. Disease suppression occurs
through a number of mechanisms including biological
control and generation of breakdown products that
are toxic to pathogens and pests. Some amendments
may also induce resistance in plants. Biological control
of soilborne plant pathogens and pests is a natural
process wide spread in agricultural soils. Agricultural
Correspondence/Reprint request: Dr. Pervaiz A. Abbasi, Agriculture and Agri-Food Canada, Southern Crop
Protection and Food Research Centre, 1391 Sandford Street, London, Ontario, N5V 4T3 Canada
Email: abbasip@agr.gc.ca
2 Pervaiz A.Abbasi et al.

practices that favour this phenomenon can lead to healthier crops with no
adverse impacts on the environment. There is compelling evidence that
organic amendments help to maintain the abundance and diversity of soil
microbial communities. They are useful for building high populations of
resident beneficial organisms while decreasing populations or activities of
plant pathogens and nematode pests. The use of organic amendments as a
disease management strategy will increase if their field efficacy remains
consistent. In this chapter, we provide an overview of the work on organic
amendments focusing primarily on specific amendments and the diseases they
control, their modes of actions, and ways to improve their efficacy.

1. Introduction
Plant diseases caused by soilborne pathogens and plant-parasitic
nematodes are a major factor in crop losses world-wide. The primary means of
controlling such pathogens and pests has been the use of agrochemicals,
particularly that of broad spectrum biocides. However, the demand for
reduced-risk and non-chemical options of managing soilborne diseases is
growing due to increasing public concerns about the impacts of pesticides on
environment and human health. Biological control, an attractive alternative to
chemical pesticides, can be achieved either by introducing known biocontrol
agents (BCAs) or by adding substances such as organic amendments that
stimulate the activity of resident microorganisms including BCAs in the soil
(9, 64, 66, 67). Organic soil amendments from the agricultural-related industry
have been used for centuries as fertilizers, but their effects on plant diseases
were also noticed. These amendments increase the activity and diversity of the
resident microbial community by increasing the organic matter content and
nutrient levels of the amended soil (72). Changes in the overall soil
microbiology can lead to biological control of soilborne plant pathogens. Also,
it appears that the decomposition of organic amendments can result in the
generation of compounds that kill plant pathogens (64, 66, 67) and nematode
pests (9). Toxic metabolites from microorganisms associated with the
breakdown of organic matter may also play a role in reducing the pathogen and
pest inoculum. It is interesting to note that the biological control option for
managing soilborne diseases may also involve chemicals. Host plants and
biocontrol agents can respond to a pathogen attack by producing antimicrobial
molecules (18). The breakdown products of some organic amendments are also
antimicrobial chemicals.
Organic matter is one of the most important components of soil and is
relevant to plant health (73). Over the last 50 years there has been severe
depletion in soil organic matter content of many agricultural soils due to
intensive cultivation. This has resulted in degradation of soil structure. There is
Short Title 3

a need to revitalize these soils with practices such as addition of organic

amendments. Increasing the organic matter content of soils, in addition to
providing nutrients to plants, also improves soil physical, chemical, and
biological properties including the ability of soils to retain nutrients and make
them available to plants; increased soil buffering capacity; and greater biological
diversity (50). Such changes can create conditions that are suppressive to plant
pathogens (15, 32) and increase plant resistance to herbivores (13). One of the
key long-term benefits of incorporating organic amendments to soil is the
generation of natural disease suppressiveness. Agricultural soils naturally
suppressive to soilborne plant pathogens and nematode pests have been found
worldwide (10, 111, 112). The basis of suppressiveness in most cases has been
attributed to biological activity (111), but soil chemical and physical factors may
also play a role. It is possible to artificially create suppressiveness in soils but
conditions may be specific and unique for each soil.

2. Biological control of soilborne diseases

Biological control involves managing pests and pathogens such that their
populations or activities are reduced to keep crop losses below economic
thresholds. It can be accomplished by general microbial activity (general
suppression) or the activity of specific organisms (specific suppression). Both
types of biological disease suppressions and the parameters affecting them
have been extensively studied for compost-amended substrates (56, 59).
Despite the enormous research efforts on biological control of plant diseases,
the number of microbial products available commercially represents only
about 1% of agricultural chemical sales (41). The performance of BCAs under
controlled conditions in potting substrates has not been consistent (31). The
right kind of potting substrates may be critical for sustained efficacy (59). It is
likely that BCAs may do well when the disease pressure is not severe or when
the pathogen populations are low. Under field conditions, the efficacy of
microbial products can be more unpredictable since the BCAs face enormous
difficulties and challenges in establishing themselves in the rhizosphere and
maintaining the population densities required for disease suppression. Field
performance of BCAs is a key factor for successful development and adoption
of a biological product. Absence of other control measures, cost-effective mass
production and formulation, market size, and registration requirements may
also impact the development of a microbial product (40). It seems that the use
of BCAs for biological control of soilborne diseases is often not an effective
stand-alone practical control measure. The performance of a microbial product
in the field can be improved by integrating it with other disease management
practices such as chemical treatments, soil solarization or steam sterilization,
organic amendments, or other agronomic practices (95).
4 Pervaiz A.Abbasi et al.

2.1. BCAs and their activity

The most commonly known bacterial BCAs belong to the genera
Agrobacterium, Bacillus, Pseudomonas, and Streptomyces, whereas fungal or
fungal-like BCAs belong to the genera, Candida, Coniothyrium, Trichoderma,
Fusarium, Pythium, and mycorrhizal fungi. All these organisms are natural soil
inhabitants. Both pathogenic and non-pathogenic microorganisms may be
useful as BCAs and can induce plant defenses (108). The root-inhibiting
fluorescent pseudomonads have been extensively studied for their disease
suppressive (111) and growth promoting activities. The rhizobacteria at the
root zones can impact plant health, yield, and soil quality (98). The activity of
these and other soil inhabitants is affected by soil type and agricultural
practices. The population structure of Pseudomonas spp. in soil is influenced
by agricultural regimes; for example as compared to arable land, specific
antagonistic subpopulations were stimulated in grassland (46). Specific
examples of some of these BCAs and their products are discussed in various
reviews (41, 78, 85). Similarly, mycorrhizal fungi, both arbuscular mycorrhizal
(AM) and ectomycorrhizal, also have the enormous potential to act as BCAs of
soilborne plant diseases (104). They are known to provide control of a number
of plant diseases in a range of systems, however, as of yet no mycorrhizal
product has been introduced in the market for commercial use (113). The
likely reason for this is that these fungi are obligate symbionts and could not be
mass-produced on growth medium. AM fungi are ubiquitous soil microbes in
nature that form symbiotic associations with plant roots. AM fungi are
particularly important in organic or sustainable farming systems to manage
soilborne diseases (52). BCAs provide control of soilborne plant diseases
through several mechanisms. Some of the common known mechanisms
include competition, production of antibiotics, antagonism or parasitism, and
induction of systemic resistance (51, 53, 109, 110).
Biological control at some form and level is always active in natural soils
where it prevents most soilborne plant pathogens and pests from causing
diseases. This condition is described as “biological buffering capacity” of a
soil which is highly effective in suppressing most soilborne plant pathogens
and nematode pests (16). Various studies of suppressive soils suggest that
biological control could also be achieved by enhancing the natural level of
suppressiveness that exists in every soil (10). Naturally-occurring resident
antagonists play a key role in this natural soil suppressiveness. Their
populations and activities can be exploited to enhance the disease control
effect at will. Practices such as crop rotations and organic soil amendments
have been used quite often to achieve this effect by exploiting the activity of
resident antagonists (75).
Short Title 5

2.2. Impact of organic amendments on BCAs

Organic amendments can enhance resident BCAs or act as substrates for
introduced BCAs. Specific impacts of organic amendments on resident BCAs
in a soil may influence the outcome of any host-pathogen interaction.
Application of chicken litter as a soil amendment and soil solarization
stimulated a shift in the soil microbial population dynamics with a significant
increase in Bacillus spp. and fluorescent pseudomonads in the rhizosphere,
rhizoplane, and interior root tissues of tomatoes and sweet potatoes (96). These
microorganisms may have contributed to the observed increased growth
response of vegetables and disease suppression (96). The use of organic
amendments, such as chitin and other material, may stimulate populations of
the existing antagonists or growth promoting organisms in the root
environment to induce disease suppressiveness and plant growth promotion
(63). Similarly, the stimulation of antagonists and plant growth promoting
rhizobacteria by compost has been described for the tomato rhizosphere (12).
Organic soil amendments have also been reported to stimulate and enhance the
activities of microorganisms that are antagonistic to plant-parasitic nematodes
in the amended soil (9, 91, 93). Some more examples of organic amendments-
stimulated biological control of soilborne diseases are given in the next section
on mechanisms of disease reduction by organic amendments.
The use of BCAs as inoculants of composts or potting soil has a potential
application in agriculture and horticulture (88). BCAs such as Trichoderma spp.
have been used as compost inoculants to improve the consistency of control of
soilborne pathogens using composts (82). The inoculation of composts with
BCAs may improve the efficacy and reliability of disease control obtained. Paper
mill residuals compost induced resistance to Fusarium wilt of tomato and
reduced fungal colonization of root tissues (87). The disease suppression was
enhanced when suppressive compost was inoculated with a known biocontrol
agent, Pythium oligandrum (20, 87). The use of a combined approach to control
soilborne diseases is a worthwhile strategy in reducing the variability in the
performance of BCAs and improving their disease control effect. Combining
BCA, Trichoderma viride isolates L4 or S17A, with composted onion waste
completely eliminated incidence of Allium white rot caused by Sclerotium
cepivorum (25). The control of Allium white rot was also achieved with spent
mushroom compost added with T. viride S17A in glasshouse and field
experiments (30). The co-application of a known biocontrol agent, Trichoderma
hamatum 382, and a co-product of ethanol production from corn (condensed
distiller’s solubles, CDS) in a non-suppressive batch of peat-based mix enhanced
damping-off suppression of radish seedlings compared to CDS or the biocontrol
agent alone in a growth room bioassay (6). The disease control effect was
enhanced if planting was delayed for 1 week after incorporating CDS and the
biocontrol agent in the infested mix.
6 Pervaiz A.Abbasi et al.

3. Managing soilborne diseases with organic soil

amendments
Organic material from agriculture-related industry such as animal by-
products, poultry by-products, fish by-products, plant-based products,
manures, and composts have been suppressive against soilborne plant diseases
when used as soil amendment. Disease suppression has been accomplished
with fresh as well as composted organic material though results have been
inconsistent between fields. Several reviews are available on this subject (15,
16, 29, 57, 59, 60, 64, 66, 67, 84). The use of organic soil amendments such as
animal manures and composts is a common practice in organic farms. It is now
a well known fact that soilborne plant diseases are less severe in organically-
managed soils than in conventionally-managed soils (106). The most common
of the organic amendments can be classified into groups based on the source
from which these were obtained.

3.1. Animal and poultry by-products

Animal rendering and poultry industries generate large amounts of waste
that can be processed into value-added products with a use in agriculture, for
instance meat and bone meal from the animal processing industry and feather
meal from the poultry industry. Meat and bone meal (MBM) is a high
nitrogenous (7.7% total N) material that can be used as a soil amendment to
control soilborne plant pathogens. Incorporation of MBM at a broad-cast rate
of 37 t/ha (equivalent to 1.9% mass/mass soil) to two potato fields with a
history of potato scab and verticillium wilt 2 weeks before planting potatoes
reduced the severity of verticillium wilt and potato scab in both locations (65).
Reduced disease levels in the MBM plots were evident for an additional year
after the amendment application but returned to control levels in the third year.
The populations of plant-parasitic nematodes were also reduced at both
locations and remained below the unamended control levels for 3 years (65).
The high amendment rates of MBM also affected tuber yield in the first year.
In the MBM-treated plots, potato plants showed stunting and signs of
phytotoxicity which resulted in the reduction of tuber yield (65).
Poultry feathers have been processed into an organic fertilizer (Nature
Safe 10-2-8, Griffin Industries, Ltd., Cold Spring, Kentucky, USA). The
efficacy of this product to suppress soilborne diseases was demonstrated in two
commercial potato fields located in Prince Edward Island, Canada. These
fields had a history of potato scab and verticillium wilt. Soil incorporation of
this product at a broad-cast rate of 8.6 t/ha (equivalent to 0.43% mass/mass
soil) into field plots 3 weeks before planting potatoes reduced scab severity
with 90% of the marketable tubers in the year of application at both sites. The
Short Title 7

amendment effect lasted for another year, but in the third year after the
amendment incorporation, the scab severity on tubers from the treated plots
was not different from the control plots (Conn and Lazarovits, unpublished
data). The application of both MBM and Nature Safe amendments should be
made in furrows to reduce the total N on per hectare basis.

3.2. Fish by-products

Soil amendments of various by-products of fish industry are routinely used
as fertilizers for crop production. Fish meal, the dried protein obtained from
processed fish, has been used as a soil amendment with great success in
vegetable production systems (21, 43). Fish solid waste has also been known to
reduce the populations of plant-parasitic nematodes (8). Fish emulsions or fish
soluble nutrients are liquid by-products of the processing of fish into fish meal.
They have been used mainly as foliar fertilizers (14), but can also provide
disease protection against some foliar (2) and soilborne diseases (3, 5). Foliar
sprays of a diluted solution of fish emulsion could control bacterial spot of
tomato and pepper (2). Fish emulsion added as a pre-planting amendment to
potting substrates and to an organic soil controlled damping-off diseases of
cucumber and radish (3). Most recently we demonstrated that fish emulsion as
a pre-plant soil amendment can control potato scab (5). The use of fish
emulsion as a disease control strategy for soilborne diseases such as potato
scab and verticillium wilt needs to be further validated in the context of
improving soil organic matter content and for improvement of soil and crop
health in general. The broadcast rates of fish emulsion effective against potato
scab may not be economical and it is possible that banding or furrow
applications can lower costs without compromising efficacy. There is also a
need to conduct long-term studies with fish emulsion and other similar
material applied serially over time at lower rates that can be economically
feasible. For instance, applying the material yearly at lower rates for several
years may improve the efficacy over time for yield improvements and disease
suppression.

3.3. Plant-based products

Soy meal, obtained after removal of oil from soybeans, is a high
nitrogenous (7.3% total nitrogen) organic material with a potential to be used
as a soil amendment to manage soilborne diseases. Pre-plant soil amendment
of soy meal at a broad-cast rate of 37 t/ha to two commercial potato fields in
Ontario, Canada with high pathogen inoculum of Streptomyces scabies and
Verticillium dahliae reduced the severity of verticillium wilt and common scab
on potatoes in both locations (65). The disease control effect of soy meal lasted
for an additional year after application. The populations of plant parasitic
8 Pervaiz A.Abbasi et al.

nematodes were also reduced at both locations and remained below the
unamended control levels for 3 years (65). As described above for MBM, the
soy meal amendment at these high rates was also phytotoxic and reduced total
tuber yield in the year of application (65). This amendment should also be
applied in furrows.
Ammonium lignosulfonate (ALS) is a co-product of paper manufacturing.
It is produced when the liquor from the acid sulfite pulp-making process is
neutralized with ammonia. We tested ALS as a pre-plant soil amendment to
control potato scab and verticillium wilt in several commercial fields in
Ontario and Prince Edward Island, Canada. A single application of ALS
reduced the incidence of verticillium wilt and potato scab at all locations in the
year of application (94). Disease severity remained lower in subsequent years
in some locations when potatoes were re-planted into the same soils without
further addition of ALS. The reduction of potato scab and verticillium wilt was
not site- or soil-specific unlike other amendments we have tested which were
greatly influenced by organic matter content and soil pH.
Neem cake is also a high nitrogen-containing (6.4%) organic material
derived from seed of the neem tree after extracting oil. Addition of neem cake to
agricultural soils before planting can reduce plant-parasitic nematodes and
viability of microsclerotia of verticillium wilt pathogen, V. dahliae, depending
upon the rates of neem cake and soil types (4). Higher broad-cast rates of neem
cake in soil may not be economical and may be phytotoxic, unless applied in
furrows. Weekly foliar sprays of a diluted aqueous solution of neem oil reduced
severity of bacterial spot on tomato and pepper foliage and fruit (2). Reduction of
bacterial spot severity, however, was inconsistent in the following years.
Corn distillation products or condensed distiller’s solubles (CDS) is a co-
product obtained after the removal of ethyl alcohol by distillation from yeast
fermentation of corn and condensing the thin stillage fraction to a semi-solid.
CDS has no immediate use at this stage. When tested as a pre-plant
amendment to peat-based mixes or soils, CDS provided protection against
plant diseases caused by some soilborne pathogens (6). Addition of CDS to a
potato soil 2 weeks before planting eggplants reduced verticillium wilt disease.
Similarly, CDS pre-plant soil amendment reduced potato scab severity in a
potato soil with medium levels of disease pressure under greenhouse and
microplot conditions, but it was not as effective in the field. However,
marketable tubers (with < 5% surface scab) were significantly increased under
all three conditions (6). While CDS did not result in an increase in yield, it did
not reduce it either. Non-uniform mixing of the material may have been
partially responsible for this reduced efficacy in the field.
The effectiveness of soil amendment with fresh organic material such as
broccoli or grass followed by covering of the amended plots with plastic
Short Title 9

sheeting to reduce the survival of Fusarium oxysporum f. sp. asparagi,

Rhizoctonia solani, and Verticillium dahliae was demonstrated in field
experiments (22). The generation of anaerobic and reducing soil conditions as
indicated by rapid depletion of oxygen and a decrease in redox potential might
have responsible for reduced viability of the pathogen inoculum. Mishra and
Srivastava (81) found that a combination of soybean green manuring and
partially decomposed wheat straw was the most effective means of reducing
potato scab with a decrease of 57% in disease severity and 55% in disease
incidence, and increasing potato yield by 14%. An increase in disease has been
reported with soil amendment of green manuring (74, 116).

3.4. Manures
Application of manures by growers to their fields is one of the oldest
agricultural practices. This can have dramatic effects on plant diseases caused
by soilborne pathogens and plant-parasitic nematode pests and on general
populations of soil microorganisms. However, there are contradictory claims
from growers about the effectiveness of manures to reduce plant diseases.
Incorporation of chicken manure at 66 t/ha and liquid swine manure
(LSM) at 55,000 L/ha as a pre-plant soil amendment was effective in
reducing verticillium wilt, common scab, and populations of plant-parasitic
nematodes in the year of application at two commercial potato field locations
in Ontario, Canada (26). These rates of amendments for effectiveness seem
to be very high. In soil microcosms and plot studies, the addition of pig
slurry to infested soil lowered populations of Ralstonia solanacearum biovar
2, the causative agent of wilting disease of potato and other crops (49). The
number of infected and or diseased plants was also reduced by the
amendment. A combination of soil solarization and pig slurry amendment
showed an additive effect of both treatments (49). In a greenhouse study,
Riegel and Noe (89) noted that adding chicken litter (chicken manure and
pine shaving beddings) into soil artificially infested with Meloidogyne
incognita eggs reduced the population densities of root-knot nematode after
45 and 90 days of planting cotton. The decrease in M. incognita populations
was correlated with an increase in the litter rate and increase in bacterial and
fungal counts in the amended soil. The effect of manure soil amendment has
also been shown to increase the resistance of plants produced on such soils
against herbivores. Potato plants produced in the field plots receiving manure
soil amendments in combination with reduced amounts of synthetic
fertilizers showed a reduction in densities of Colorado potato beetle
compared to plots receiving full rates of synthetic fertilizer (13). It is
believed that a good plant growth due to balance nutrients in organically-
managed soils may have resulted in resistance to herbivores.
10 Pervaiz A.Abbasi et al.

3.5. Composts
Composts have been used as soil amendments for centuries mainly to
enhance soil fertility but they can also maintain or improve organic matter
level and biological and physical properties of the amended soil. Their wide
spread availability and use in sustainable agriculture as environmentally safe
biofertilizers and soil conditioners is increasing all over the world.
Composts are also known to influence plant diseases caused by soilborne
pathogens (44, 54, 58, 80, 115) as well as foliar pathogens (1, 87). The effects of
composts on diseases in the field have been generally more variable than in
greenhouse or growth room experiments (82). The degree of stabilization of
composts can have a positive effect on it disease suppressing effect (47). Several
studies under controlled conditions have demonstrated a suppressive effect of
composts on soilborne diseases such as damping-off, root rots, and wilts (57, 58,
59). Disease suppression with compost-amended substrates and parameters
affecting their efficacy has been extensively studied. Addition of poultry litter or
poultry litter compost in combination with a sorghum sudangrass rotation was
most effective in reducing populations of Pratylenchus penetrans, although
efficacy varied across years (39). In a greenhouse study, potato plants produced
in soils amended with vegetable compost and wood-chip polyacrylamide cores
showed significantly lower infection rates of V. dahliae compared to in control
and manure and vegetable composts alone (38). In a recent study, Darby et al.
(33) showed that composted and fresh dairy manure solids can reduce damping-
off of cucumber and root rots of snap bean and sweet corn. The amended soils
were no longer suppressive after 12 months, however, suppressiveness was
restored after re-amendment the following year (33). In a field study utilizing
organic soil fertility amendments, Bulluck and Ristaino (23) demonstrated that
composted cotton-gin trash reduced the incidence of southern blight in
processing tomatoes and enhanced populations of beneficial microbes such as
antagonistic soil fungi in the genus Trichoderma and fluorescent pseudomonads.

4. Mechanisms of disease reduction by organic

amendments
Organic materials added to infested soils affect plant diseases but the
mechanism(s) of disease reduction have been for most part ignored. However,
in order to optimize their efficacy, such information is vital. As described
below, many mechanisms have been proposed including stimulation of
resident microbes with biocontrol activity, induction of systemic resistance in
plants, generation of toxic chemicals, and in some amendments presence of
toxic substances such as volatile fatty acids (VFAs). It is likely that more than
one mechanism is involved but which ones play a dominant role is likely to be
Short Title 11

soil specific. Most, if not all, of the organic amendments enhance microbial
activity but not all lead to reduced disease. The effects of organic amendments,
suggests that both chemical and biological components of compost-amended
soils can contribute to disease suppression (1, 23, 118). However, a further
characterization and understanding of the different mechanisms by which
organic amendments reduce plant diseases may help improve the disease
control effect and reduce variability (59). It will also increase the spectrum of
soils that can be treated.

4.1. Stimulation of BCAs

Organic amendments generally enhance total microbial numbers in the
amended soils including BCAs. Organic matter can act as a substrate for growth
of antagonists, but it also can lead to enhance enzyme activities required for
biological control (35, 114). The higher microbial activity in organically-
managed sandy soils has been described as partly responsible for take-all
suppression (55). The higher soil bacterial populations were evident with LSM
and chicken manure amendments (26). These amendments can reduce
populations of the pathogen and hence disease by more than one mechanism.
Some of the longer term reductions seen may be derived from the increased
microbial activity (26). Both enhanced activity and diversity of microbial
population in the composted swine waste-amended potting mixes were
associated with suppressiveness to Rhizoctonia damping-off of impatiens (34).
The increased in populations of beneficial microbes such as Trichoderma and
fluorescent pseudomonads may be responsible for reduction in the incidence of
southern blight in processing tomatoes with soil amendment of composted
cotton-gin trash (23). Pérez-Piqueres et al. (86) reported that differences in soil
suppressiveness to Rhizoctonia solani diseases were related to differences in
chemical composition, nutrients availability, and microbial composition due to
both incorporation and stimulation of microorganisms by the compost
amendments. These authors also noticed that soil characteristics of the amended
soil differed with the nature of the composts and the soil types. Fatty acid
metabolizing and seed colonizing microbial communities were found associated
with the leaf compost suppressive to Pythium damping-off of cotton (77).
The reduction in the incidence of verticillium wilt and potato scab in
commercial potato fields amended with ALS also may be due to stimulated
BCAs. ALS, which was not toxic to Verticillium dahliae microsclerotia or to S.
scabies, caused an increase in total microbial populations and diversity,
particularly that of fungi (94). Verticillium dahliae colonies derived from
microsclerotia recovered from the ALS plots were much smaller and heavily
infested with other competing fungi. This was not seen with microsclerotia
from untreated plots. Thus, increased populations of fungi which may act as
12 Pervaiz A.Abbasi et al.

BCAs may be a mechanism by which ALS reduced verticillium wilt and potato
scab. Disease reduction by fish emulsion may also be due to more than one
mechanism depending on rates and types of soils or substrates used. In peat-
based mix, fish emulsion does not appear to have direct toxicity to damping-
off pathogens but appears to create a biological climate that is suppressive to
disease initiation (3). The delayed improvement of disease control with time
corresponded with increased microbial activity in the amended mix.
Furthermore, in some batches of peat-based mixes, there was no control
possibly due to low microbiological activity in the matrix. Seven days after
incorporation of fish emulsion to peat-based mix, the populations of culturable
fungi and bacteria had increased. This incubation time also corresponded to the
initiation of the disease suppression by the amended peat-based mix. Fish
emulsion as an amendment to muck soil was shown to favour known
biocontrol and growth-promoting agents (Hill, Abbasi, Lazarovits, and
Hemmingsen, unpublished data). The microbial communities of the fish
emulsion-amended muck soil were characterized based on DNA analysis of the
chaperonin-60 sequence and high-throughput sequencing. The fish emulsion-
amended muck soil was also suppressive to damping-off of cucumber.

4.2. Induction of resistance in plants

Organic soil amendments can stimulate microorganisms such as
rhizobacteria and other non-pathogenic organisms that may induce resistance
in plants. Microbial metabolites have been involved as inducers of systemic
resistance in plants in rhizobacteria-mediated biological control of plant
diseases (17). Induced resistance has been implicated in suppressive soil
systems. Non-pathogenic isolates of Fusarium oxysporum induced systemic
resistance in watermelons against Fusarium wilt (68). Paper mill residuals
compost induced resistance to Fusarium wilt of tomato. The suppression was
associated with reduced fungal colonization of root tissues by the pathogen due
to an increase in physical barriers to fungal penetration (87). The disease
suppression and physical protection was enhanced when suppressive compost
was inoculated with a known biocontrol agent, Pythium oligandrum (20, 87).
Similarly, plants grown in composted pine bark mix inoculated with
Trichoderma hamatum 382 provided protection against bacterial leaf spots of
tomato, radish, and lettuce (11). Compost-induced systemic resistance has been
implicated in reduction of these bacterial leaf spots and other root and foliage
diseases of crop plants (117, 118).

4.3. Generation of toxic chemicals

The breakdown products of some organic amendments are chemicals but
they are produced through natural processes by the action of microbial activity
Short Title 13

in the soil. These should also fall under the category of biological control.
Nitrogen transformation products, such as ammonia and nitrous acid, can be
lethal to some pathogens and pests (19, 70, 79, 99, 101, 105). These toxicants
are produced after addition of higher rates (2-10% by weight) of high nitrogen-
containing organic amendments such as chicken manure, MBM, chitin and
chitosan, and soy meal to soil. Ammonia accumulation occurs most readily in
soils with low rates of nitrification, low cation exchange capacity, moisture and
organic carbon content, and high specific density and sand content (102, 103).
The nitrous acid toxicity seems to occur in soils of neutral to acidic pH and with
high rates of nitrification. The use of high rates of nitrogen containing
amendments for controlling soilborne pathogens and pests may be impractical.
The high rates can cause phytotoxicity and lead to surface and groundwater
contamination. Since much lower rates are needed in soils having low organic
carbon and high sand contents, they may find some use at such sites. To be
practical, the amendments should be applied in soils which are most responsive
in producing ammonia or nitrous acid.
Production of other toxic chemicals such as sulfur-containing compounds,
isothiocyanates which are released from hydrolysis of glucosinolates, and
others has been described for Brassicaceaous soil amendments (45, 69). Some
volatile toxic compounds can also be produced by creating anaerobic and
reducing conditions in the soil by incorporating fresh organic material such as
broccoli or grass followed by covering of the amended plots with plastic
sheeting (22). Although these toxicants are produced in the soil after
amendment incorporation through natural processes, they may also affect
beneficial and non-target organisms depending upon the concentrations. Most
of these amendments have also been described to enhance microbial activity in
the soil (26, 65, 76). The release of toxic ammonia was considered responsible
for stimulating fungal parasites of nematodes (90).

4.4. Presence of VFAs

Some organic amendments such as LSM and fish emulsion contain VFAs
which can be toxic to pathogens in low pH soils. As described above, both
these amendments also stimulate the microbial activity in the soil. Only the
protonated forms of these VFAs (e.g. acetic acid and not acetate) are toxic to
V. dahliae (100). VFAs from LSM are toxic to V. dahliae microsclerotia in
acid soils within a day after application (28). A concentration of about 3 mM
(mass/mass soil water) of the nonionized form of VFAs from LSM in soil is
needed to kill V. dahliae (28). Within a few days after addition of LSM to soil
the VFAs disappear, most likely due to degradation by microorganisms. The
total VFA concentration in LSM can reach 400 mM and acetic, propionic, and
isobutyric acids are the major VFAs while n-butyric, n-valeric, isovaleric, and
14 Pervaiz A.Abbasi et al.

n-caproic acids are present in lesser amounts (100). In some acid soils, LSM
addition results in toxicity from VFAs as well as nitrous acid.
Fish emulsion also contains VFAs (190 mM total VFAs) with acetic acid
being the major VFA. It is possible that VFA toxicity may be one of the
mechanisms of disease reduction by fish emulsion in the low pH soils (5)
although populations of Trichoderma spp. are also known to increase in acid
soils. The fact that 20% of the VFAs in fish emulsion are formic acid increases
the toxicity of the VFAs in fish emulsion compared to LSM which does not
contain formic acid. Formic acid is 7 times more toxic to V. dahliae compared
to acetic acid (100). LSM can be formulated to enhance its VFA contents with
VFA-producing bacteria.

5. Additional benefits of organic soil amendments

Besides providing control of plant diseases, incorporation of organic
amendments into soils can have other favourable effects. Some of these effects
which also may be directly or indirectly related to disease control are described
below.
• Fertility effect. Organic soil amendments are known to influence plant
growth by reducing diseases caused by soilborne pathogens. Most of the
examples in the literature, as described above, have cited a reduction in
disease, but in some cases an increase in disease has been reported (74, 116).
Organic amendments can also improve plant growth by directly supplying
nutrients. Organic amendments may also enhance soil fauna which are known
to influence plant growth and pathogen survival and dispersal (42). The use of
organic sources of fertilizer or waste products have been shown to result in
very satisfactory yields over extended periods of crop cultivation (62, 92).
• Development of suppressive soils. Suppressive soils are defined as soils
which allow only a low level of disease development on a susceptible host
even though a virulent pathogen is present in the soil. One of the long-term
benefits of using organic soil amendments is the generation of conditions
suppressive to soilborne pathogens and pests leading to plant growth
promotion. Modification of biological, chemical, and physical properties of
soil through cultural practices and inputs is required to generate suppressive
conditions (97). Plant genotypes should also be considered in exploiting the
potential of this natural disease suppression system as they affect the activity
of resident soil microbial communities (76). Soil incorporation of summer and
winter cover crops and barley straw, sorghum residues, and native grass hay as
mulch layers topped with poultry litter and dolomite to stimulate rapid decay
and raised the pH and Ca contents, led to suppressiveness against Phytophthora
root rot of avocado (reviewed in 97). This was a successful attempt of creating
suppressive conditions by an Australian avocado grower.
Short Title 15

• Improved soil properties. Organic amendments improved soil physical,

chemical, and biological properties. Besides providing nutrients to plants,
organic amendments increase organic matter content of the amended soils (50).
High organic matter content of a soil can retain nutrients and make them
available to plants, increase soil buffering capacity and biological diversity.
Application of compost made from potato, sawdust, and manure at volume
ratios of 3:3:1 in an intensive 3-year potato rotation provided benefits for
potato productivity and in soil physical and biological properties (24). Organic
farming systems in The Netherlands resulted in lower levels of both nitrate and
total soluble nitrogen in the soil, increase in number and diversity of bacteria,
and species richness in both bacteria and nematode communities (107).
• Healthy impact on environment. Recycling of agricultural waste
material can bring about a favourable effect on the health of environment. The
use of organic soil amendments for disease management can also lead to a
gradual reduction in the use of harmful agrochemicals. A reduction in mineral
sources of fertilizer is also possible in some cases. Growing healthy crops by
improving soil health is by itself a healthy impact on environment.
• Substrate for BCAs. Microorganisms are a major component of soil and
contribute significantly to soil quality and productivity (7). Organic
amendments generally cause a significant increase in the size of resident soil
microbial communities including BCAs. They also provide an ideal matrix for
delivery of BCAs. For instance, fish emulsion and other organic materials used
as an amendment to control soilborne diseases are known to enhance total
fungal and bacterial soil populations. Fish emulsion has also been reported as a
good substrate for delivery of plant growth promoting rhizobacteria (37).
Similarly, straw mulch could be used to deliver BCAs in protected crops such
as ginseng and strawberry (114). In a cropping system under natural
conditions, indigenous soil microorganisms generally contribute to disease
suppression by reducing and competing with pathogen populations. These
indigenous microorganisms have been adapted to the specific environmental
conditions in any particular soil. Organic amendments can influence these
indigenous microorganisms and enhance their biological control effectiveness.
• Compatibility with other control strategies. Organic soil amendments
can be compatible with any other disease control strategy and be an integral
component of integrated disease management system. Combining organic
amendments with other disease control approaches such as soil solarization can
also provide improved control of soilborne plant diseases. The synergistic
effect of solarization and nitrogenous amendments on pathogen control can
serve as a means to lower application rates and shorten times for solarization.
Solarization, in combination with chicken manure soil amendment at 18.8 t/ha,
was effective in near removal of disease incidence of the sweet potato crop and
16 Pervaiz A.Abbasi et al.

its stored roots (96). Stevens et al. (96) also showed that soil solarization and
chicken litter soil amendment stimulated a shift in the soil microbial
population dynamics. Field solarization of soil amended with composted
chicken manure gave better control of pathogens and higher yield of lettuce
and tomato than either treatment alone (44, 45). Disease control in solarized
and amended soil is attributed to killing of the pathogen by a combination of
increased temperatures and enhanced generation of biotoxic volatile
compounds. The increased soil temperature due to solarization can have
profound effect on physical and biological properties of the soil amended with
organic amendment (48). Gelsomino and Cacco (48) found that a combination
of farmyard manure and solarization resulted in an increase in soil temperature,
pH, and electrical conductivity, and an altered bacterial community which
remained stable over time. Soil amendment with mustard pod residues in
conjunction with soil solarization reduced viable propagules of Macrophomina
phaseolina and Fusarium oxysporum f.sp. cumini (61).

6. Constraints of using organic amendments

The conditions described below are not really limitations or constraints,
but are factors that need to be considered while selecting a particular soil
amendment to manage soilborne plant diseases. These factors were identified
based on the use of specific amendments under specific soil conditions.
• Soil specificity. Organic soil amendments such as MBM, soymeal, and
LSM can effectively reduce soilborne diseases at specific soil conditions. The
pH of soil and of amendment can affect the efficacy of these and other
amendments (27, 102). By selecting the soils of appropriate pH or by adjusting
the pH of the amendments, the disease reducing efficacy of high nitrogen- or
VFA-containing amendments can be improved. Acidification of LSM may
expand the range of soils where it can be effective against potato scab, but it
may also reduce the yield. Acidified LSM may not be economical for high pH
and/or buffered soils but maybe ideal for soils with low buffering capacities.
• Less effectiveness in high disease pressure soil. Organic soil amendments
in general are effective for disease control only in soils with a medium level of
disease pressure (5). In a high disease pressure soil, the disease reduction may
not be impressively obvious. However, a reduction in pathogen inoculum over
time is expected and that may lead to less disease. Also, pathogen inoculum in
such soils may be reduced by adopting a long-term integrated disease
management strategy. It is not likely that organic amendments completely
eliminate the pathogen inoculum from the low disease pressure soils.
• Method and timing of applications. Non-uniform incorporation of the
organic material into the soil is one of the factors contributing to inconsistent
Short Title 17

field efficacy of the amendments. The growers need to have an access to the
proper equipment to apply the material where it needed the most for any desired
results. The amendments effective only at high rates can be applied in bands or
in-furrows to lower costs but not efficacy. The timing of application also varies
and depends on the organic materials for maximum disease control. Applying the
high nitrogen- or VFA-containing amendments in the fall immediately after
harvesting a crop when the soil pH is the lowest may provide better control than
application in the spring, prior to the crop. There is also a need to examine the
optimal time to apply the organic material in a crop rotation cycle. Generation of
anaerobic or reducing soil conditions following application of the organic
material and tarping can result in fermentation products such as VFAs which are
known to reduce pathogen inoculum (22, 71). The combination of these
technologies may be practical under certain conditions.
• High rates of application. Organic amendments need to be applied at
quite high rates for any effectiveness against soilborne diseases. The 1%
(20,000 litres/ha) broadcast rate of fish emulsion effective against potato scab
(5) can not be economical unless applied in furrows or bands. Similarly
effective rates of LSM are also quite high. Field application of higher rates of
solid material may not be economically viable and physically practical. Lower
rates applied more frequently may provide longer lasting disease suppression
and that may be economically feasible. This does add to the cost as application
and handling can be expensive, but it seems to create a disease suppressive
condition in fields where the practice has been used for years. An organic soil
amendment applied once at higher rate may be economical if its disease
control and yield effects last for more than 2 years. There is also real concern
about pollution and ground water contamination with high rates of application.
• Other issues. Inconsistent field efficacy, small market size, and lack of
equipment for applying organic amendments into soil are some of the factors
preventing the wide spread use of this technology. Currently, most of these
products are looked on as a disposal issue rather than an underutilized energy
resource. More than a billion tons of organic and inorganic agricultural
recyclable by-products are generated every year in the United States (36).
These include crop residue, cattle manure, poultry and swine manure,
municipal solid wastes, and pulp and paper industry wastes. If a value-added
benefit for these products can be demonstrated, they may be recycled for
agricultural use with great potential benefit. The odourless, safe, and pathogen-
free liquid organic products may be adopted more rapidly as they can be
readily added to the planting rows by manure injectors or even by irrigation
systems. In order for organic amendments to be adopted, their performance
should be eye catching to growers and in some cases comparable to
agrochemicals.
18 Pervaiz A.Abbasi et al.

7. Concluding remarks and future work

The demand for the use of organic amendments for managing soilborne
diseases and enhancing soil fertility will continue to expand as an alternative to
agrochemicals. The use of this technology may still play a meaningful role in
specialty and high-value crops and in organic farming. Consistency in field
efficacy of organic soil amendments may stimulate conventional growers to
adopt and integrate such practices as well. The integrated approach using the
combinations of technologies such as organic amendments with BCAs or
solarization or cultural practices is the way forward to manage diseases with
improved efficacy. This approach can provide superior disease control against
a broad range of pathogens and pests than either control strategy alone even
though there may be additive costs attached to it. There is a great need to focus
on the soil management practices that improve soil quality and health by
maintaining the abundance and diversity of resident soil microbial
communities, high population of beneficial organisms, and low population of
plant pathogens and pests. Future understanding and research targeting the
constraints of using organic soil amendments for disease management may
further improve the efficacy leading to large market size.

8. References
1. Abbasi, P. A., Al-Dahmani, J., Sahin, F., Hoitink, H. A. J., and S.A. Miller. 2002.
Effect of compost amendments on disease severity and yield of tomato in
conventional and organic production systems. Plant Dis. 86: 156-161.
2. Abbasi, P.A., Cuppels, D.A., and G. Lazarovits. 2003. Effect of foliar applications
of neem oil and fish emulsion on bacterial spot and yield of tomatoes and peppers.
Can. J. Plant Pathol. 25: 41-48.
3. Abbasi, P.A., Conn, K.L., and G. Lazarovits. 2004. Suppression of Rhizoctonia
and Pythium damping-off of radish and cucumber seedlings by addition of fish
emulsion to peat mix or soil. Can. J. Plant Pathol. 26: 177-187.
4. Abbasi, P.A., Riga, E., Conn, K.L., and G. Lazarovits. 2005. Effect of neem cake soil
amendment on reduction of damping-off severity and population densities of plant-
parasitic nematodes and soilborne plant pathogens. Can. J. Plant Pathol. 27: 38-45.
5. Abbasi, P.A., Conn, K.L., and G. Lazarovits. 2006. Effect of fish emulsion used as
a pre-plant soil amendment on verticillium wilt, common scab, and tuber yield of
potato. Can. J. Plant Pathol. 28: 509-518.
6. Abbasi, P.A., Conn, K.L., and G. Lazarovits. 2007. Managing soilborne diseases of
vegetable crops with a pre-plant soil or substrate amendment of a corn distillation
product. Biocont. Sci. Technol. 17: 331-344.
7. Abawi, G.S., and T.L. Widmer. 2000. Impact of soil health management practices
on soilborne pathogens, nematodes and root diseases of vegetable crops. Appl. Soil
Ecol. 15: 37-47.
8. Akhtar, M., and I. Mahmood. 1995. Suppression of nematode populations with
animal by-products. Biores. Technol. 51: 269-271.
Short Title 19

9. Akhtar, M., and A. Malik. 2000. Roles of organic soil amendments and soil
organisms in the biological control of plant-parasitic nematodes: a review.
Bioresource Technol. 74: 35-47.
10. Alabouvette, C. 1999. Fusarium wilt suppressive soils: an example of disease-
suppressive soils. Australas. Plant Pathol. 28: 57-64.
11. Aldahmani, J.H., Abbasi, P.A., Sahin, F., Hoitink, H.A.J., and S.A. Miller. 2005.
Reduction of bacterial leaf spot severity on radish, lettuce and tomato plants grown
in compost-amended potting mixes. Can. J. Plant Pathol. 27: 86-193.
12. Alvarez De Brito, M.A., Gagne, S., and H. Antoun. 1995. Effect of compost on
rhizosphere microflora of the tomato and on the incidence of plant growth-
promoting rhizobacteria. Appl. Environ. Microbiol. 61: 194-199.
13. Alyokhin, A., Porter, G., Groden, E., and F. Drummond. 2005. Colorado potato
beetle response to soil amendments: a case in support of the mineral balance
hypothesis? Agric. Ecosys. Environ. 109: 234-244.
14. Aung, L.H., Flick, G.J., Bluss, G.R., Aycock, H.S., Keefer, R.F., Singh, R.,
Brandon, D.M., Griffin, J.L., Hovermale, C.H., and C.A. Stutte. 1984. Growth
responses of crop plants to fish soluble nutrients fertilization. Virginia Polytechnic
Institute and University, Blacksburg, Va. Bulletin 84-9.
15. Bailey, K.L., and G. Lazarovits. 2003. Suppressing soil-borne diseases with
residue management and organic amendments. Soil Tillage Res. 72: 169-180.
16. Baker, K.F., and R.J. Cook. 1974. Biological Control of Plant Pathogens. Freeman,
San Francisco CA. (Reprinted in 1982, American Phytopathological Society, St.
Paul,MN)
17. Bakker, P.A.H.M., Ran, L.X., Pieterse, C.M.J., L.C. van Loon. 2003.
Understanding the involvement of rhizobacteria-mediated induction of systemic
resistance in biocontrol of plant diseases. Can. J. Plant Pathol. 25: 5-9.
18. Bélanger, R.R. 2006. Controlling diseases without fungicides: a new chemical
warfare. Can. J. Plant Pathol. 28 (Suppl.): S233-S238.
19. Bell, A.A., Hubbard, J.C., Liu, L., Davis, R.M., and K.V. Subbarao. 1998. Effects
of chitin and chitosan on the incidence and severity of Fusarium yellows of celery.
Plant Dis. 82: 322-328.
20. Benhamou, N., Rey, P., Picard, K., and Y. Tirilly. 1999. Ultrastructural and
cytochemical aspects of the interaction between the mycoparasite Pythium
oligandrum and soilborne plant pathogens. Phytopathology 89: 506-517.
21. Blatt, C.R., and K.B. McRae. 1998. Comparison of four organic amendments with
a chemical fertilizer applied to three vegetables in rotation. Can. J. Plant Sci. 78:
641-646.
22. Blok, W.J., Lamers, J.G., Termorshuizen, A.J., and G.J. Bollen. 2000. Control of
soilborne plant pathogens by incorporating fresh organic amendments followed by
tarping. Phytopathology 90: 253-259.
23. Bulluck, L.R. III, and J.B. Ristaino. 2002. Effect of synthetic and organic fertility
amendments on southern blight, soil microbial communities, and yield of
processing tomatoes. Phytopathology 92: 181-189.
24. Carter, M.R., Sanderson, J.B., and J.A. MacLeod. 2004. Influence of compost on
the physical properties and organic matter fractions of a fine sandy loam
throughout the cycle of a potato rotation. Can. J. Soil Sci. 84: 211-218.
20 Pervaiz A.Abbasi et al.

25. Clarkson, J.P., Scruby, A., Mead, A., Wright, C., Smith, B., and J.M. Whipps.
2006. Integrated control of Allium white rot with Trichoderma viride, tebuconazole
and composted onion waste. Plant Pathol. 55: 375-386.
26. Conn, K.L., and G. Lazarovits. 1999. Impact of animal manures on verticillium
wilt, potato scab, and soil microbial populations. Can. J. Plant Pathol. 21: 81-92.
27. Conn, K.L., and G. Lazarovits. 2000. Soil factors influencing the efficacy of liquid
swine manure added to soil to kill Verticillium dahliae. Can. J. Plant Pathol. 22:
400-406.
28. Conn, K.L., Tenuta, M., and G. Lazarovits. 2005. Liquid swine manure can kill
Verticillium dahliae microsclerotia in soil by volatile fatty acid, nitrous acid, and
ammonia toxicity. Phytopathology 95: 28-35.
29. Cook, R.J., and K.F. Baker. 1983. The Nature and Practice of Biological Control of
Plant Pathogens. St. Paul, MN, APS. 433 pp.
30. Coventry, E., Noble, R., Mead, A., Marin, F.R., Perez, J.A., and J.M. Whipps.
2006. Allium white rot suppression with composts and Trichoderma viride in
relation to sclerotia viability. Phytopathology 96: 1009-1020.
31. Daughtrey, M.L., and D.M. Benson. 2005. Principles of plant health management
for ornamental plants. Annu. Rev. Phytopathol. 43: 141-169.
32. Davis, J.R., Huisman, O.C., Everson D.O., and A.T. Schneider. 2001. Verticillium
wilt of potato: a model of key factors related to disease severity and tuber yield in
southeastern Idaho. Amer. J. Potato Res. 78: 291-300.
33. Darby, H.M., Stone, A.G., and R.P. Dick. 2006. Compost and manure mediated
impacts on soilborne pathogens and soil quality. Soil Sci. Soc. Amer. J. 70: 347-
358.
34. Diab, H.G., Hu, S., and D.M. Benson. 2003. Suppression of Rhizoctonia solani on
impatiens by enhanced microbial activity in composted swine waste-amended
potting mixes. Phytopathology 93: 1115-1123.
35. Downer, A.J., Menge, J.A., and E. Pond. 2001. Association of cellulytic enzyme
activities in eucalyptus mulches with biological control of Phytophthora
cinnamomi. Phytopathology 91: 847-855.
36. Edwards, J.H. and A.V. Someshwar. 2000. Chemical, physical, and biological
characterisitics of agricultural and forest by-products for land applications. In Land
Application of Agricultural, Industrial, and Municipal By-Products, ed J.M.
Bartels, and W.A. Dick, Soil Society of America Series Book 6: 1-62.
37. El-Tarabily, K.A., Nassar, A.H., Hardy, G.E.St.J., and K. Sivasithamparam. 2003.
Fish emulsion as a food base for rhizobacteria promoting growth of radish
(Raphanus sativus L. var. sativus) in a sandy soil. Plant Soil 252: 397-411.
38. Entry, J.A., Strausbaugh, C.A., and R.E. Sojka. 2005. Compost amendments
decrease Verticillium dahliae infection on potato. Compost Sci. Utiliz. 13: 43-49.
39. Everts, K.L., Sardanelli, S., Kratochvil, R.J., Armentrout, D.K., and L.E.
Gallagher. 2006. Root-knot and root-lesion nematode suppression by cover crops,
poultry litter, and poultry litter compost. Plant Dis. 90: 487-492.
40. Fokkema, N.J. 1996. Biological control of fungal plant diseases. Entomophaga 41:
333-342.
41. Fravel, D. 2005. Commercialization and implementation of biocontrol. Annu. Rev.
Phytopathol. 43: 337-359.
Short Title 21

42. Friberg, H., Lagerlöf, J., and B. Rämert. 2005. Influence of soil fauna on fungal plant
pathogens in agricultural and horticultural systems. Biocont. Sci. Technol. 15: 641-658.
43. Gagnon, B., and S. Berrouard. 1994. Effects of several organic fertilizers on
growth of greenhouse tomato transplants. Can. J. Plant Sci. 74: 167-168.
44. Gamliel, A., and J.J. Stapleton. 1993. Effect of chicken compost or ammonium
phosphate and solarization on pathogen control, rhizosphere microorganisms, and
lettuce growth. Plant Dis. 77: 886-891.
45. Gamliel, A., and J.J. Stapleton. 1997. Improvement of soil solarization with volatile
compounds generated from organic amendments. Phytoparasitica 25(Suppl.): S31-S38.
46. Garbeva, P., van Veen, J.A., and J.D. van Elsas. 2004. Assessment of the diversity,
and antagonism towards Rhizoctonia solani AG3, of Pseudomonas species in soil
from different agricultural regimes. FEMS Microbiol. Ecol. 47: 51-64.
47. Garcia, C., Pascual, J.A., Mena, E., and T. Hernández. 2004. Influence of the
stabilisation of organic materials on their biopesticide effect in soils. Bioresource
Technol. 95: 215-221.
48. Gelsomino, A., and G. Cacco. 2006. Compositional shifts of bacterial groups in a
solarized and amended soil as determined by denaturing gradient gel
electrophoresis. Soil Biol. Biochem. 38: 91-102.
49. Gorissen, A., van Overbeek, I.S., and J.D. van Elsas. 2004. Pig slurry reduces the
survival of Ralstonia solanacearum biovar 2 in soil. Can. J. Microbiol. 50: 587-593.
50. Grandy, A.S., Porter, G.A., and M.S. Erich. 2002. Organic amendment and rotation
crop effects on the recovery of soil organic matter and aggregation in potato
cropping systems. Soil Sci. Soc. Am. J. 66: 1311-1319.
51. Handelsman, J., and E.V. Stabb. 1996. Biocontrol of soilborne plant pathogens.
Plant Cell 8: 1855-1869.
52. Harrier, L.A., and C.A. Watson. 2004. The potential role of arbuscular mycorrhizal
(AM) fungi in the bioprotection of plants against soil-borne pathogens in organic
and/or other sustainable farming systems. Pest Manage. Sci. 60: 149-157.
53. Haas, D., and G. Défago. 2005. Biological control of soil-borne pathogens by
fluorescent pseudomonads. Nature Rev. Microbiol. 3: 307-319.
54. Hadar, Y., and R. Mandelbaum. 1992. Suppressive composts for bio-control of
soilborne plant pathogens. Phytoparasitica 20(Suppl.): S113-S116.
55. Hiddink, G.A., van Bruggen, A.H.C., Termorshuizen, A.J., Raaijmakers, J.M., and
A.V. Semenov. 2005. Effect of organic management of soils on suppressiveness to
Gaeumannomyces graminis var. tritici and its antagonist, Pseudomonas
fluorescens. Eur. J. Plant Pathol. 113: 417-435.
56. Hoitink, H.A.J., and P.C. Fay. 1986. Basis for the control of soilborne plant
pathogens with composts. Annu. Rev. Phytopathol.24: 93-114.
57. Hoitink, H.A.J., Inbar, Y., and M.J. Boehm. 1991. Status of compost-amended
potting mixes naturally suppressive to soilborne diseases of floricultural crops.
Plant Dis. 75: 869-873.
58. Hoitink, H.A.J., Stone, A.G., and D.Y. Han. 1997. Suppression of plant diseases by
composts. HortScience 32: 184-187.
59. Hoitink, H.A.J., and M.J. Boehm. 1999. Biocontrol within the context of soil
microbial communities: a substrate dependent phenomenon. Annu. Rev.
Phytopathol. 37: 427-446.
22 Pervaiz A.Abbasi et al.

60. Huber, D.M., and R.D. Watson. 1970. Effect of organic amendments on soil-borne
plant pathogens. Phytopathology 60: 22-26.
61. Israel, S., Mawar, R., and S. Lodha. 2005. Soil solarisation, amendments and bio-
control agents for the control of Macrophomina phaseolina and Fusarium
oxysporum f.sp. cumini in aridisols. Annals Appl. Biol. 146: 481-491.
62. Kelman, A. and R.J. Cook. 1977. Plant pathology in the People’s Republic of
China. Annu. Rev. Phytopathol. 15: 409-429.
63. Kloepper, J.W., Rodríguez-Kábana, R., Zehnder, G.W., Murphy, J.F., Sikora, E.,
and C. Fernández. 1999. Plant root-bacterial interactions in biological control of
soilborne diseases and potential extension to systemic and foliar
diseases. Australas. Plant Pathol. 28: 21-26.
64. Lazarovits, G. 2001. Management of soilborne plant pathogens with organic soil
amendments: a disease control strategy salvaged from the past. Can. J. Plant
Pathol. 23: 1-7.
65. Lazarovits, G., Conn, K.L., and J. Potter. 1999. Reduction of potato scab,
verticillium wilt, and nematodes by soymeal and meat and bone meal in two
Ontario potato fields. Can. J. Plant Pathol. 21: 345-353.
66. Lazarovits, G., Tenuta, M., and K.L. Conn. 2001. Organic amendments as a
disease control strategy for soilborne diseases of high-value agricultural crops.
Australas. Plant Pathol. 30: 111-117.
67. Lazarovits, G., Conn, K.L., Abbasi, P.A., and M. Tenuta. 2005. Understanding
the mode of action of organic soil amendments provides the way for improved
management of soilborne plant pathogens. Acta Hort. 698: 215-225.
68. Larkin, R.P., Hopkins, D.L., and F.N. Martin. 1996. Suppression of Fusarium
wilt of watermelon by nonpathogenic Fusarium oxysporum and other
microorganisms recovered from a disease-suppressive soil. Phytopathology 86:
812-819.
69. Lewis, J.A., and G.C. Papavizas. 1971. Effect of sulfur-containing volatile
compounds and vapors from cabbage decomposition on Aphanomyces euteiches.
Phytopathology 61: 208-214.
70. Loffler, H.J.M., Cohen, E.B., Oolbekkink, G.T., and B. Schippers. 1986. Nitrite as
a factor in the decline of Fusarium oxysporum f. sp. dianthi in soil supplemented
with urea or ammonium chloride. Neth. J. Plant Pathol. 92: 153-162.
71. Lynch, J.M. 1978. Production and phytotoxicity of acetic acid in anaerobic soils
containing plant residues. Soil Biol. Biochem. 10: 131-135.
72. Mäder, P., Fliessbach, A., Dubois, D., Gunst, L., Fried, P., and U. Niggli. 2002.
Soil fertility and biodiversity in organic farming. Science 296: 1694-1697.
73. Magdoff, F. and R.R. Weil. 2004. Soil organic matter management strategies. In
Soil Organic Matter in Sustainable Agriculture, ed F. Magdoff, and R.R. Weil,
pp. 45-65. CRC Press, Boca Raton, Florida.
74. Manici, L.M., Caputo, F., and V. Babini. 2004. Effect of green manure on Pythium
spp. population and microbial communities in intensive cropping systems. Plant
Soil 263: 133-142.
75. Mathre, D.E., Cook, R.J., and N.W. Calla. 1999. From discovery to use: traversing
the world of commercializing biocontrol agents for plant disease control. Plant Dis.
83: 972-983.
Short Title 23

76. Mazzola, M. 2004. Assessment and management of soil microbial community

structure for disease suppression. Annu. Rev. Phytopathol. 42: 35-59.
77. McKellar, M.E., and E.B. Nelson. 2003. Compost-induced suppression of Pythium
damping-off is mediated by fatty-acid-metabolizing seed-colonizing microbial
communities. Appl. Environ. Microbiol. 69: 452-460.
78. McSpadden Gardener, B. B., and D.R. Fravel. 2002. Biological control of plant
pathogens: research, commercialization, and application in the USA. Online. Plant
Health Progress doi:10.1094/PHP-2002-0510-01-RV.
79. Michel, V.V., and T.W. Mew. 1998. Effect of a soil amendment on the survival of
Ralstonia solanacearum in different soils. Phytopathology 88: 300-305.
80. Millner, P.D., Ringer, C.E., and J.L. Maas. 2004. Suppression of strawberry root
disease with animal manure composts. Compost Sci. Utiliz. 12: 298-307.
81. Mishra, K.K., and J.S. Srivastava. 2004. Soil amendments to control common scab
of potato. Potato Res. 47: 101-109.
82. Noble, R., and E. Coventry. 2005. Suppression of soil-borne plant diseases with
composts: A review. Biocontrol Sci. Technol. 15: 3-20.
83. Papavizas, G.C., and R.D. Lumsden. 1980. Biological control of soilborne fungal
propagules. Annu. Rev. Phytopathol. 18: 389-413.
84. Patrick, Z.A., and T.A. Toussoun. 1965. Plant residues and organic amendments in
relation to biological control. In Ecology of Soil-borne Plant Pathogens, ed K.F.
Baker, and W.C. Snyder. University of California Press, Berkeley, Los Angeles,
CA. pp. 440-459.
85. Paulitz, T.C., and R.R. Bélanger. 2001. Biological control in greenhouse system.
Annu. Rev. Phytopathol. 39: 103-133.
86. Pérez-Piqueres, A., Edel-Hermann, V., Alabouvette, C., and C. Steinberg. 2006.
Response of soil microbial communities to compost amendments. Soil Biol.
Biochem. 38: 460-470.
87. Pharand, B., Carisse, O., and N. Benhamou. 2002. Cytological aspects of compost-
mediated induced resistance against Fusarium crown and root rot in tomato.
Phytopathology 92: 424-438.
88. Postma, J., Montanari, M., and P.H.J.F. van den Boogert. 2003. Microbial
enrichment to enhance the disease suppressive activity of compost. Eur. J. Soil
Biol. 39: 157-163.
89. Riegel, C., and J.P. Noe. 2000. Chicken litter soil amendment effects on soilborne
microbes and Meloidogyne incognita on cotton. Plant Dis. 84: 1275-1281.
90. Rodriguez-Kabana, R., King, P.S., and M.H. Pope. 1981. Combination of
anhydrous ammonia and ethylene dibromide for control of nematodes parasitic on
soybeans. Nematropica 11: 27-41.
91. Rodriguez-Kabana, R., and G. Morgan-Jones. 1987. Biological control of
nematodes: soil amendments and microbial antagonists. Plant Soil 100: 237-247.
92. Shen, D. 1997. Microbial diversity and application of microbial products for
agricultural purposes. Agric. Ecosyst. Environ. 62: 237-245.
93. Sikora, R.A. 1992. Management of the antagonistic potential in agricultural
ecosystems for the biological control of plant parasitic nematodes. Annu. Rev.
Phytopathol. 30: 245-270.
24 Pervaiz A.Abbasi et al.

94. Soltani, N., Conn, K.L., Abbasi, P.A., and G. Lazarovits. 2002. Reduction of
potato scab and verticillium wilt with ammonium lignosulfonate soil amendment in
four Ontario potato fields. Can. J. Plant Pathol. 24: 332-339.
95. Spadaro, D., and M.L. Gullino. 2005. Improving the efficacy of biocontrol agents
against soilborne pathogens. Crop Prot. 24: 601-613.
96. Stevens C., Khan V.A., Rodriguez-Kabana R., Ploper L.D., Backman P.A., Collins
D.J., Brown J.E., Wilson M.A., and E.C.K. Igwegbe. 2003. Integration of soil
solarization with chemical, biological and cultural control for the management of
soilborne diseases of vegetables. Plant Soil. 253: 493-506.
97. Stone, A.S., Scheurell, S.J., and H.M. Darby. 2004. Suppression of soilborne
diseases in field agriculture systems: organic matter management, cover cropping,
and other cultural practices. In Soil Organic Matter in Sustainable Agriculture, ed
F. Magdoff, and R.R. Weil, pp. 131-177. CRC Press, Boca Raton, Florida.
98. Sturz, A.V., and B.R. Christie. 2003. Beneficial microbial allelopathies in the root
zone: the management of soil quality and plant disease with rhizobacteria. Soil
Tillage Res. 72: 107-123.
99. Tenuta, M. 2001. Nitrogen transformation products from nitrogenous amendments
and the soil properties affecting their toxicity to Verticillium dahliae Kleb. Ph.D.
Dissertation University of Western Ontario, London, Canada.
100. Tenuta, M., Conn, K.L., and G. Lazarovits. 2002. Volatile fatty acids in liquid
swine manure can kill microsclerotia of Verticillium dahliae. Phytopathology 92:
548-552.
101. Tenuta, M. and G. Lazarovits. 2002a. Ammonia and nitrous acid from nitrogenous
amendments kill microsclerotia of Verticillium dahliae. Phytopathology 92: 255-
264.
102. Tenuta, M., and G. Lazarovits. 2002b. Identification of specific soil properties that
affect the accumulation and toxicity of ammonia to Verticillium dahliae. Can. J.
Plant Pathol. 24: 219-229.
103. Tenuta, M., and G. Lazarovits. 2004. Soil properties associated with the variable
effectiveness of meat and bone meal to kill microsclerotia of Verticillium dahliae.
Applied Soil. Ecol. 25: 219-236.
104. Traquair, J.A. 1994. Fungal biocontrol of root diseases: endomycorrhizal
suppression of cylindrocarpon root rot. Can. J. Bot. 73(Suppl. 1): S89-S95.
105. Tsao, P.H., and J.J. Oster. 1981. Relation of ammonia and nitrous acid to
suppression of Phytophthora in soils amended with nitrogenous organic
substances. Phytopathology 71: 53-59.
106. van Bruggen, A.H.C., and A.J. Termorshuizen. 2003. Integrated approaches to root
disease management in organic farming systems. Australas. Plant Pathol. 32: 141-156.
107. van Diepeningen, A.D., de Vos, O.J., Korthals, G.W., and A.H.C. van Bruggen.
2006. Effects of organic versus conventional management on chemical and
biological parameters in agricultural soils. Appl. Soil Ecol. 31: 120-135.
108. van Loon, L.C., Bakker, P.A.H.M., and C.M.J. Pietrse. 1998. Systemic resistance
induced by rhizosphere bacteria. Ann. Rev. Phytopathol. 36: 453-483.
109. van Loon, L.C., and P.A.H.M. Bakker. 2005. Induced systemic resistance as a
mechanism of disease suppression by rhizobacteria. In PGPR: Biocontrol and
Biofertilization, ed Z.A. Siddique. Springer, Dordrecht, The Netherlands.
Short Title 25

110. Weller, D.M. 1998. Biological control of soilborne plant pathogens in the
rhizosphere with bacteria. Annu. Rev. Phytopathol. 26: 379-407.
111. Weller, D.M., Raaijmakers, J.M., McSpadden Gardener, B.B., and L.S.
Thomashow. 2002. Microbial populations responsible for specific soil
suppressiveness to plant pathogens. Annu. Rev. Phytopathol. 40: 309-348.
112. Westphal, A. 2005. Detection and description of soils with specific nematode
suppressiveness. J. Nematol. 37: 121-130.
113. Whipps, J.M. 2004. Prospects and limitations for mycorrhizas in biocontrol of root
pathogens. Can. J. Bot. 82: 1198-1227.
114. White, G.J., and J.A. Traquair. 2006. Necrotrophic mycoparasitism of Botrytis
cinerea by cellulolytic and ligninocellulolytic Basidiomycetes. Can. J. Microbiol.
52: 508-518.
115. Workneh, F., and A.H.C. van Bruggen. 1994. Suppression of corky root of
tomatoes in organically managed soil associated with soil microbial activity and
nitrogen status of soil and tomato tissue. Phytopathology 81: 688-694.
116. Yulianti, T., Sivasithampram, K., and D.W. Turner. 2006. Saprophytic growth of
Rhizoctonia solani Kühn AG2-1 (ZG5) in soil amended with fresh green manures
affects the severity of damping-off in canola. Soil Biol. Biochem. 38: 923-930.
117. Zhang, W., Hoitink, H.A.J., and W.A. Dick. 1996. Compost-induced systemic
acquired resistance in cucumber to Pythium root rot and anthracnose.
Phytopathology 86: 1066-1070.
118. Zhang, W., Han, D.Y., Dick, W.A., Davis, K.R., and H.A.J. Hoitink. 1998.
Compost and compost water extract-induced systemic acquired resistance in
cucumber and Arabidopsis. Phytopathology 88: 450-455.