Mineral Nutrition and Suppression of Plant Disease

WH Elmer, Connecticut Agricultural Experiment Station, New Haven, CT, USA

LE Datnoff, Louisiana State University Agricultural Center, Baton Rouge, LA, USA
r 2014 Elsevier Inc. All rights reserved.

Glossary Rhizosphere The soil near a living root.

Oomycota A fungal-like chromistan that produces Systemic acquired resistance Systemically activated
oospores; a water mold related to algae. resistance after primary infection with a necrotizing
Pathogenesis-related proteins Groups of proteins with pathogen accompanied by increased levels of salicylic acid
different chemical properties produced in a cell within and pathogenesis-related proteins.
minutes or hours following inoculation but all being more
or less toxic to pathogens.
Phytoalexin A substance that inhibits the development of
a fungus on hypersensitive tissue formed when host plant
cells come in contact with the parasite.

Introduction molecules. Many of these pathways use enzymes that require

the micronutrients manganese (Mn), copper (Cu), zinc (Zn),
One of the fundamental strategies for maintaining plant molybdenum (Mo), or boron (B) as cofactors or activators.
health is to manage the nutritional status of the plant (Datnoff Other elements such as potassium (K) and chlorine (Cl) in-
et al., 2007a). Nutrition can often govern the fine line between fluence osmotic relations, water cycling, and root exudation,
a crop's susceptibility and resistance to a plant disease. Com- which, in turn, influence beneficial microbes. Indirect mech-
plete and balanced nutrition should always be the first line of anisms include effects on soil medium nitrification, rooting
defense against the development of a plant disease. One major medium pH, and chemical transformation of micronutrients.
problem in providing proper nutrition is that many agronomic Below, each element's effect on plant disease is discussed,
and horticultural crops vary in their nutritional requirements but caution must be exercised in that nutrition must be viewed
and those different nutrients may affect different diseases in holistically, because all nutrients affect the uptake and func-
different ways. Furthermore, the fertilization regimes necessary tion of other elements and can ultimately increase or decrease
to maximize plant health when a plant pathogen is present can plant susceptibility to disease.
differ from optimal fertilization when the pathogen pressure is
absent. This article provides an overview of the governing role
of nutrition on crop health as related to resistance and sus- Nitrogen
ceptibility to disease.
In many cases, the amount of a nutrient needed to suppress Nitrogen is the fourth most abundant element in plants and is
a plant disease may far exceed a healthy plant's nutritional an essential component of amino acids, enzymes, hormones,
requirement for that nutrient, suggesting that many nutrients phenolics, phytoalexins, and proteins, all of which can have
may participate in multiple mechanisms for plant disease direct effects on disease development. The amount of N ap-
suppression. Chemical interactions with the rooting medium, plied can have immediate effects on a plant disease, relative to
pH, and specific communities of microorganisms can, in turn, the nutritional requirement of the plant (Table 1). The greatest
influence disease development. For example, the form of ni- responses are generally in the range from deficiency to
trogen (N) can have striking effects on plant disease through physiological sufficiency. Nitrogen-deficient plants may not
root-mediated changes in pH, microbial profile in the root provide the nutrient environment necessary for obligate
zone, and alterations in the availability and function of pathogens, whereas excess N may inhibit the production of
micronutrients. Moreover, calcium (Ca) composes only ap- defense responses to other pathogens (Huber, 1980; Huber
proximately 0.5% of the dry weight of most plants, yet Ca is and Watson, 1974).
routinely applied in great quantities to container mixes to af- The time of application can also have striking effects on
fect root medium pH and suppress certain plant diseases. disease development. Side-dressing nitrogen can compensate
These observations suggest that these inorganic elements are for early-season losses caused by leaching and denitrification
influencing disease directly through physiological mechanisms but, more importantly, can avoid the early-season seedling
and indirectly through soil interactions that, in turn, affect the diseases that accompany nitrogen application, such as damp-
activity of plant pathogens. ing-off caused by species of Rhizoctonia or Pythium. However,
Both essential and beneficial elements can affect disease caution should be exercised as mechanical damage made to
defense mechanisms in plants. These nutrients play important the roots from side dressing may increase root rots. Many
roles in metabolic pathways that lead to the production times the choice as to when to apply nitrogen can be influ-
of lignin, phenols, phytoalexins, and other defense-related enced by disease pressure. For example, delaying N application

Encyclopedia of Agriculture and Food Systems, Volume 4 doi:10.1016/B978-0-444-52512-3.00251-5 231

232 Mineral Nutrition and Suppression of Plant Disease

Table 1 Influence of nitrogen form on plant disease

Disease or disorder Pathogen or causal agent Host NH4 NO3

Aflatoxin Aspergillus flavus group Corn and peanut D –

Anthracnose Gnomonia leptostyla Black walnut D D
Black root rot Thielaviopsis basicola Tobacco D I
Black root rot Rhizoctonia fragariae Strawberry D I
Blast Magnaporthe grisea Rice D I
Blossom-end rot Physiological disorder Tomato I –
Brown patch Rhizoctonia solani Ryegrass D –
Club Root Plasmodiophora brassicae Cabbage D –
Corky root Pyrenochaeta lycopersici Tomato I I
Crown rot Rhizoctonia solani Beets I D
Crown and root rot Fusarium oxysporum Tomato I D
Eyespot Pseudocercosporella herpotrichoides Wheat I D
Leaf blight Erwinia chrysanthemi Chrysanthemum I I
Leaf spot Alternaria macrospora Cotton D –
Lesion nematode Pratylenchus penetrans Strawberry D I
Mummy berry Monilinia vaccinii-corymbosi Blueberry D –
Nematodes Criconemella spp. Turfgrass – I
Root rot Fusarium oxysporum and Fusarium proliferatum Asparagus I D
Root rot Phytophthora parasitica Tomato – I
Root rot Fusarium spp. Pea I D
Root rot Thielaviopsis basicola Pansy D I
Root and stem rots Sclerotium rolfsii Many plants D D

Abbreviations: D, decrease; I, increase; − no effect.

Source: Adapted from Huber, D.M., Thompson, I.A., 2007. Nitrogen and plant disease. In: Datnoff, L.E., Elmer, W.H., Huber, D.M. (Eds.), Mineral Nutrition and Plant Disease. St. Paul,
MN: American Phytopathological Society, pp. 31−44.

in winter wheat until spring can result in early N deficiency Many of the underlying mechanisms governing disease
and increase susceptibility of plants to take-all root rot caused suppression with different nitrogen forms relate to soil pH
by Gaeumannomyces graminis (Sacc.) Arx & D. Olivier var. tritici (Smiley and Cook, 1973). Nitrate N increases the pH of the
(J. Walker) (Huber and McCay-Buis, 1993). However, a fall rooting medium and NH4-N lowers pH, making both the bulk
application can impose excess N availability during cold, wet soil and rhizosphere more acidic. These effects occur through
winter periods and increase susceptibility to Rhizoctonia root microbial and chemical reactions in the soil as well as root-
rot (Huber, 1989). mediated ion exchanges near the roots. Acidification of the
The rate of nitrogen also affects disease susceptibility. Ex- root zone also solubilizes oxides of zinc, copper, manganese,
cessive nitrogen is well documented to increase the severity of and iron, which increases their availability for uptake. As dis-
Botrytis blight on numerous greenhouse plants (Dik and cussed below, these micronutrients can play pivotal roles in
Wubben, 2007; Yermiyahu et al., 2006). Root diseases are disease suppression. When the nitrogen form is applied as
similarly exacerbated under high nitrogen rates. Root rot nitrate, Fusarium wilts and root rots of asparagus, basil, beets,
of wheat caused by Fusarium culmorum (Papendick and carnation, chrysanthemum, cyclamen, gladiolus, tobacco,
Cook, 1974) and Pythium root of geraniums (Gladstone and tomatoes, and watermelon are reported less severe than when
Moorman, 1989) are more severe under high nitrogen re- applied as NH4-N (Figure 1; Elmer, 1997; Huber and
gimes. Although growers pay close attention to rates of ni- Thompson, 2007). Our understanding of this phenomenon
trogen applied, most do not recognize the role that form of relates to the effect on rhizosphere pH, root exudation, and the
nitrogen may play in enhancing or suppressing disease. Ni- availability of Fe to microbes (Jones and Woltz, 1970).
trogen is the only nutrient that is available as both a negatively The opposite is true for diseases caused by Gaeumannomyces
charged ion, nitrate (NO3
) and a positively charged ion, spp., Thielaviopsis spp., and Verticillium spp. where NH4-N
ammonium (NH4 þ ). Nitrate N and NH4-N are metabolized suppresses disease and NO3-N favors it. Take-all root rot of
differently in plants and can have opposite effects on diseases. wheat caused by G. graminis can be suppressed in the field by
In fact, many of the conflicting research reports regarding the supplying nitrogen in the ammoniacal form, such as NH3,
role of nitrogen in plant disease may be due to a failure to (NH4)2SO4, NH4Cl, and (NH4)2PO4 (Huber, 1989; Reis et al.,
recognize and report the form of nitrogen used in the experi- 1982). Ammonium sulfate suppressed take-all patch of turf-
ments. Interestingly, the form of nitrogen can have striking grass but was more effective when combined with other cul-
effects on the effectiveness of biological control. Borrero et al. tural practices for disease control (Dernoeden, 1987). The
(2012) observed suppression of Fusarium wilt of tomato with effects of NH4-N occur through chemical reactions in the soil
Trichoderma spp. but only under increasing concentrations of as well as root-mediated ion exchanges near the roots that, in
NH4-N. Suppression from Trichoderma spp. was minimal turn, increase the availability and uptake of micronutrients
under the nitrate regime. and promote beneficial microbes in the rhizosphere.
 Mineral Nutrition and Suppression of Plant Disease 233

pH 5.5 pH 6.5
100 100

90 90

80 80

70 70

60 60

50 50

40 40

30 30

80% CaNO3 80% NH4SO4

Figure 1 Effect of N form and soil pH on percentage of tomatoes with Fusarium wilt. Data from Jones, J.P., Woltz, J.P., 1970. Fusarium wilt of
tomato: Interaction of soil liming and micronutrient amendments on disease development. Phytopathology 60, 812–813.

Managing soil acidity with NH4-N has become an important P reacts with Ca and reduces P availability. Therefore, main-
strategy for managing black root rot caused by Thielaviopsis tenance of soil pH is very important to maximize availability
basicola on tobacco and ornamentals. On eggplant, Verticillium of P and promote root health.
wilt was suppressed and yields were increased by 20% when Another important consideration is the role of beneficial
NO3-N was replaced with NH4-N (Elmer and Ferrandino, mycorrhizal fungi in P nutrition and disease development.
1994). In strawberry, the black root rot complex caused by These fungi form a symbiotic relationship with the plant's
Rhizoctonia fragariae and the lesion nematode, Pratylenchus roots known as mycorrhizae that improve plant health and
penetrans, was reduced by the application of ammonium sul- reduce disease damage. Many crops have mycorrhizal associ-
fate, but not with calcium nitrate fertilizer (Elmer and ations that are likely interrupted during propagation cycles. If
LaMondia, 1999a,b). Plants treated with ammonium sulfate mycorrhizae are present or have been established on roots,
had more yield, more leaf area, more runners, and higher there may be a need to alter the P applications because
tissue levels of K, S, Mn, and Zn than strawberries treated with mycorrhizal associations can be inhibited by increased P
calcium nitrate. availability. Commercial applications of mycorrhizal fungi are
It is very common for N to interact with other nutrients. available and may have value in management of soilborne
Potassium increases uptake of NO3 and promotes the syn- disease. Their use may allow other nutrients to be more
thesis of organic N compounds, whereas phosphorus and available as added P could lead to the precipitation of other
chlorine decrease NO3 uptake and enhance the uptake of NH4. elements.
Ammonium and Cl increase the uptake of manganese (Mn). Foliar applications of phosphorous acid (H3PO3), or its
Manganese is required for NO3 assimilation and protein syn- salts (also referred to as phosphonate, phosphate, and phos-
thesis. Molybdenum (Mo), magnesium (Mg), iron (Fe), as- phonic acid), may inhibit plant pathogens from the fungal-like
corbic acid, and energy are required for the reduction of NO3 Oomycota, i.e., species of Phytophthora and Pythium (Brunings
to NH4. Growers should also pay attention to the ion that et al., 2012). The inhibiting effect from phosphorous acid is
accompanies the nitrogen salt. probably due to either an inhibition of the metabolic activity
of the pathogen or a direct toxic effect on the pathogen.
Phosphorous acid has been demonstrated to control a number
Phosphorus of plant diseases, including downy mildew, late blight of po-
tato and tomato, foliar blights on peppers and cucurbits, root
Phosphorus (P) is the second component listed in the analysis and stem rots, and damping off.
of fertilizers due to its vital role in cell division, energy trans-
fers, and its regulatory role for transport of sugars within the
plant. Although there are exceptions, most reports on P and Potassium
plant disease suggest that increasing phosphorus above that
necessary for proper growth may be associated with increased Potassium (K) plays many essential roles in plant nutrition.
disease. In fact, those cases where P reduced disease may have Although K is not structurally bound in the plant, it increases
been in situations where the element was deficient. Cases root growth, improves water and nutrient uptake, increases
where the disease became worse following P application might cellulose and protein content, reduces lodging, enhances and
have been in soils where excess P decreased the availability of regulates at least 60 different plant growth enzymes, and can
other elements that, in turn, increased the plant's suscepti- affect the occurrence of a plant disease (Prabhu et al., 2007).
bility. In acidic soils, P reacts with Fe, Al, and Mn to form Potassium alone or its combination with N, P, and other
insoluble products, making P less available. In alkaline soils, nutrients can alter the disease severity of many soilborne and
234 Mineral Nutrition and Suppression of Plant Disease

Table 2 Effect of potassium on plant disease development

Pathogen Group Number of articles showing the effect of potassium on diseasea

Decreased Increased No effect Total

Fungal 89 33 8 130
Bacterial 19 5 * 24
Viruses 9 5 3 17
Nematodes 3 6 1 10
a
Data adapted with permission from Huber, D.M., Arny, D.C., 1985. Interactions of potassium with plant diseases. In: Munson, R.D. (Ed.), Potassium in Agriculture. Madison,
WI: American Society of Agronomy, pp. 467−488.
*Data not available.

foliar plant pathogens. These have been extensively reviewed resistance. Enhanced silicification of cell walls by K is one
before for a number of crops, diseases, and pathogens (Prabhu explanation for increased disease resistance. A deficiency of K
et al., 2007; Huber, 1980). Even though no generalization can decreases Si accumulation in epidermal cells and increases the
be made on the effect of K on disease development, this susceptibility of rice to leaf blast. Potassium, in combination
element is reported to decrease the intensity of many diseases with P, induces the development of thicker cuticles and cell
caused by pathogenic fungi, bacteria, viruses, and nematodes walls that function as mechanical barriers to infection by
(Table 2). Unfortunately, in many of these studies, investi- several pathogens or limit the growth of some pathogens as a
gators have failed to give adequate consideration to the com- result of a higher proportion of sclerenchyma tissue (Huber,
panion anions, nutrient balance, and nutrient status in order 1980).
to determine a definitive role of K (Prabhu et al., 2007). Thus, Fertilizer sources of K include various salts from which the
there may be a greater response to K in deficient than fully anion may also manifest an effect on disease in combination
sufficient plants or with an excess of K beyond that required with or independent of the K ion. The more common sources
for nutrient sufficiency. This suggests that K may affect host of K fertilizers include the chloride (muriate) or sulfate salts;
resistance more as opposed to a direct effect on the pathogen. however, carbonate, nitrate, phosphate (mono- and dibasic),
Some rice genotypes are more efficient than others in K silicon, and various organic salts are also used. It is difficult to
uptake, and this may contribute to their increased disease re- distinguish between the effect of the K and the anion in the
sistance and higher grain yield (Fageria et al., 1990). Several fertilizer salt used for disease management unless many
diseases increase with an increase in N and the effect of K can combinations are studied. It is well known that the Cl anion
be directly correlated with the level of N. The critical factor can affect various diseases independent of K (Elmer, 2007) and
affecting rice blast caused by Pyricularia oryzae is the N:K ratio that K influences plant uptake of nitrate, Ca, and other mineral
in leaves. Blast severity is low when there is a high tissue K:N elements. Similar interactions are observed with phosphate
ratio, whereas a high N:K ratio increases blast. When the salts of K (Lopez and Lucas, 2002). The level of K in plants
concentration of N is low, the addition of K suppresses rice depends on the availability of Mg and Ca. Calcium alters the
blast. However, when the level of N is high, the addition of K Ca:K ratio and interacts with other elements. The function of K
increases blast. The K and N ratio greatly influences plant in cellular organization and permeability is complimented by
growth as well as disease development, and the ratio may large Ca reserves in mature plant tissue. Calcium enhances K
change with growth of the plant. availability in neutral soils, but not in acidic soils. An inter-
Although the physiological function of K in disease resist- action between K and Mg was demonstrated on rice leaf blast;
ance mechanisms is not well understood, nutritional factors blast increased when K was applied alone but could be re-
favoring host plant resistance have been attributed to alter- duced by adding Mg.
ations in protein or amino acid availability, decreased cell
permeability, or decreased susceptibility of tissue to macer-
ation and penetration (Prabhu et al., 2007). Arginine increases Calcium
as the level of K increases and resistance of potato plants to
late blight (Phytophthora infestans) increases as the levels of K Calcium (Ca) is the third most abundant nutrient in most
increases due to the accumulation of fungistatic levels of ar- plant species. Calcium concentrations in plant tissues are
ginine in leaves. Exudates of arginine inhibit germination of usually twice that of magnesium. Because both nutrients are
sporangia of P. infestans and exudation is lowest in the absence cations with a double positive charge, they often antagonize
of K when N and P are high. The ability of various soil-applied and interact with each other in nutrient management
or foliarly applied K salts, such as potassium phosphonate, programs.
K2HPO4, KH2PO4, and KNO3, to stimulate systemic acquired Calcium is extremely important in normal cell growth
resistance in susceptible plants against powdery mildews, an- where it forms the glue between cell walls, i.e., Ca-poly-
thracnose, rust, and several other pathogens suggests that galacturonates are needed for the middle lamella for cell wall
fundamental changes in physiological pathways are probably stabilization. This element may act as a secondary messenger;
important in disease resistance (Reuveni et al., 2000; Becot consequently, it plays a role in many enzymatic reactions in-
et al., 2000; Orober et al., 2002). Anatomical changes affected volved in early plant defense mechanisms (Huber et al., 2012).
by K nutrition may also play an important role in disease Calcium is commonly applied as CaCl2, CaSO4, or Ca(NO3)2.
 Mineral Nutrition and Suppression of Plant Disease 235

Many previous studies investigating the role of Ca(NO3)2 examples on ornamentals may illustrate this point. The add-
credited all of the plant disease suppression to the NO3 ition of extra magnesium to potting soil was associated with
component and failed to recognize the calcium ion. Unfortu- increased damping-off of calendula. However, when carna-
nately, most studies do not examine different sources of Ca tions were grown on calcareous soils, applications of Mg
and its companion anion to demonstrate the contribution of suppressed Fusarium wilt. The latter study was likely a result of
calcium to plant disease control. A study on roses found that the added magnesium correcting a deficiency due to the high
foliar applications of CaCl2 reduced the incidence of gray Ca concentration.
mold (Botrytis cinerea). Calcium applications to carrot infected
with Sclerotium rolfsii reduced damage and increased yield.
Many times the influence of Ca on soil pH is more important Chlorine
than the actual concentration of Ca in soil. For example, a
reduction in the incidence of cavity spot of carrot by Pythium Chlorine has been routinely applied to crops as chloride (Cl),
coloratum was achieved with lime, whereas gypsum (CaSO4), which served as a companion ion for NH4-N, K, and Ca fer-
which does not affect pH, applied to the same soil at the same tilizers. Chloride was thought to have little value in improving
rate had no effect on the incidence of disease. The role of pH plant growth because Cl was thought to be highly available in
and Ca in suppressing clubroot of cabbage caused by Plasmo- soils and not essential. Yet reports of chloride deficiency in
diophora brassicae was recognized in 1878, and liming of the more than 11 economic crops have been published (Engel
soil for control of clubroot has been practiced for more than et al., 1997; Fixen, 1993; Gausman et al., 1958). The role of Cl
200 years. The severity of Fusarium wilt of tomato caused by in plant disease management has also long been misunder-
Fusarium oxysporum f. sp. lycopersici is also reduced with calcium stood (Elmer, 2007). The benefits of Cl are still mistakenly
applications. Greenhouse and field experiments in Florida being ascribed to the accompanying cation. For example, al-
showed that amending Fusarium-infested soil with gypsum though it is well documented that proper K nutrition will
(CaSO4) did not increase the soil pH and did not reduce the suppress some plant diseases, many subsequent studies that
occurrence of tomato Fusarium wilt, whereas the Ca content of examined different forms of K found that the ameliorating
tissues of plants grown in soil amended with hydrated lime effects on disease were restricted to KCl amendments, sug-
(Ca(OH)2) was increased and did reduce the occurrence gesting that Cl was the active ion.
of wilt. Nutritionally, Cl is regarded as a micronutrient, yet, as in
Spraying calcium salts, such as CaCl2 and Ca(NO3)2, on the case of Ca, benefits are achieved with rates that far exceed
tomatoes reduced powdery mildew colonies. A study on roses the plant's nutritional requirements. Monocots tend to be
found that foliar applications of CaCl2 reduced the incidence more tolerant of high amounts of chloride. High rates of Cl
of gray mold (B. cinerea). The same study found that increasing salts have marked effects on inhibiting soil nitrification, en-
Ca concentrations in the soil improved the shelf life of the hancing availability of Mn and other micronutrients, and on
flower. Calcium may also influence the spread of Phytophthora increasing beneficial microorganisms.
spp. in water; vinca plants that were flood irrigated and then As an element, Cl
is the only inorganic anion that is not
infected with Phytophthora spp. were healthier when the com- structurally bound to a metabolite. Its major role is to serve as
plete fertilizer solution was amended with Ca(NO3)2. Other a charge-balancing ion. When a cell absorbs Cl
, it accumu-
studies demonstrated that calcium applied as either CaCl2 or lates in the cell vacuole and lowers the cell water potential
Ca(NO3)2 in water or in Ca-free soluble fertilizer solutions below that of the medium surrounding the cell. Water then
suppressed the release and the motility of the swimming flows into the cell and increases hydrostatic cell pressure so
spore. These results demonstrate the urgent need to under- that it maintains a pressure that exceeds the force exerted by
stand more about how calcium amendments interfere with the plasmalemma. The cells remain turgid and are able to grow
Phytophthora in recirculating irrigation systems. even when drought conditions prevail. Theoretically, plants
that get root diseases might be protected by supplying suf-
ficient Cl so that they could still grow normally in soils with
Magnesium greater moisture deficits. Studies are needed to validate this
theory. Chloride ions also alter the quantity and quality of
Magnesium (Mg) is usually applied as MgSO4 or MgCl2 and organic solutes that are exuded into the rhizosphere, thus re-
has been associated with both increased disease and sup- ducing the germination of and infection by root pathogens.
pression of plant disease (Jones and Huber, 2007). Of 46 Another major role of Cl in disease suppression comes from its
studies investigating the role of magnesium on plant disease, effect on increasing the availability of Mn, which has direct
22 found that this nutrient decreased disease, 18 found that it effects on host resistance.
increased disease, and 6 found that there was little or no dif- Most reports that demonstrate disease suppression with
ference. As the data are so conflicting, no real patterns can be chloride fertilization have been conducted on monocots, but
discerned. As a consequence, there is a paucity of information there are many notable studies where crop health was ad-
regarding the direct effects of Mg on plant pathogenesis. It may vanced on dicots (Table 3; Maas, 1986; Fixen, 1993). Plants
be that some of these studies corrected a magnesium de- such as asparagus, barley, beets, celery, and coconut evolved
ficiency and the result was a healthier plant with more vigor near coastlines (Bailey and Bailey, 1976) and acquired toler-
and disease resistance. When Mg was applied in excess of what ance or even preference for chloride in their nutrition. It ap-
is required for normal growth, a nutritional imbalance de- pears that chloride nutrition may have its greatest benefits to
veloped that promoted plant stress and more disease. The few the host in providing tolerance to stresses such as drought and
236 Mineral Nutrition and Suppression of Plant Disease

Table 3 Host plants, diseases, and pathogens suppressed by chloride salts

Host plant Disease Pathogen(s)

Asparagus Fusarium crown and root rot Fusarium oxysporum

Fusarium proliferatum
Barley Common root rot Bipolaris sorokiniana Fusarium culmorum
Powdery Mildew Erysiphe graminis f. sp. hordei
Rust Puccinia hordei
Beets Rhizoctonia root and crown rot Rhizoctonia solani
Celery Fusarium yellows Fusarium oxysporum f. sp. apii
Coconut Leaf spot Bipolaris incurvata
Corn Stalk rot Gibberella zeae
Gibberella fujikuroi
Corn Smut Ustilago maydis
Cyclamen Fusarium wilt Fusarium oxysporum f. sp. cyclaminis
Date Palm Fusarium wilt Fusarium oxysporum f. sp. albedinis
Pearl Millet Downy Mildew Sclerospora graminicola
Sorghum Stalk rots Gibberella thapsinum
Gibberella zeae and Macrophomina phaseolina
Soybeans Soybean cyst nematode Heterodera glycines
Sudden death Fusarium solani f. sp. glycines
Wheat Take-all Gaeumannomyces graminis var. tritici
Root rot Fusarium culmorum
Gibberella zeae
Bipolaris sorokinian
Stripe (yellow) rust Puccinia striiformis
Leaf Rust Puccinia recondite f. sp. tritici
Powdery mildew Erysiphe graminis f. sp. tritici
Tanspot Pyrenophora tritici-repentis
Glume blotch Septoria tritici
Stagonospora blotch Phaeosphaeria avenaria

Source: Adapted from Elmer, W.H., 2007. Chlorine and plant disease. In: Datnoff, L.E., Elmer, W.H., Huber, D.M. (Eds.), Mineral Nutrition and Plant Disease. St. Paul, MN: American
Phytopathological Society, pp. 189−202.

disease. For asparagus, NaCl applications increased fern vigor and Shahzad, 2001; Suleman et al., 2001). Other processes
and spear yields by 15–30% in plots affected by F. oxysporum f. that act through the soil environment, host physiology, or
sp. asparagi and Fusarium proliferatum, but there was no evi- microbial community are more probable mechanisms for
dence that NaCl improved growth in healthy plantings (Reid disease suppression. Chloride affects nitrification (Christensen
et al., 2001). Barley had less common root rot caused by et al., 1986), manganese availability (Krishnamurti and
Bipolaris sorokiniana (Goos et al., 1989; Timm et al., 1986) Huang, 1992), and osmoregulation (Kettlewell et al., 2000;
when plants were fertilized with chloride. Kettlewell et al. Schneider, 1990). It has also been associated with enhancing
(1990) observed suppression of powdery mildew of wheat biological control (Amir et al., 1996; Elmer, 1995). Scientists at
caused by Erysiphe graminis f. sp. tritici and brown rust of barley Oregon State University suspected that chloride was affecting
caused by Puccinia hordei, when plants were sprayed with KCl. water relations in wheat (Christensen et al., 1981; Powelson
As adequate K was found in the soil, it was concluded that a and Jackson, 1978). Using pressure chambers to measure leaf
potassium deficiency was unlikely. water potential (ψ leaf) and thermocouple psychrometry to
Elmer (1997) found that applications of NaCl, KCl, CaCl2, measure osmotic potential (ψ osmotic) in cell sap, they ob-
and MgCl2 were equal in suppressing Rhizoctonia root rot of served lower leaf water potentials and lower osmotic poten-
table beets, caused by Rhizoctonia solani. Yields were increased tials in leaves treated with NH4Cl than in leaves fertilized with
in infested soil when NaCl (560 kg ha–1) was applied in an (NH4)2SO4. Turgor potential (ψ pressure), calculated from the
NH4-N regime, but not in an NO3-N regime (Elmer, 1997). formula ψ leaf ¼ ψ osmotic þ ψ pressure, was often greater in
Alternatively, Schneider (1985) found that the effectiveness of Cl-treated plants. Elmer and LaMondia (1999a,b) similarly
chloride in suppressing Fusarium yellows of celery was en- monitored the water potential of ferns of NaCl-treated as-
hanced in an NO3-N regime and in the presence of adequate paragus plants and found that NaCl applications significantly
potassium. Both studies demonstrate the complexities of lowered the predawn fern water potential and increased
chloride nutrition and its interaction with other elements. the estimated turgor potential. Kettlewell and colleagues
Chloride fertilization contributes to a range of physio- (Kettlewell et al., 2000; Mann et al., 2004) also found that KCl
logical and soil chemical responses. Although Cl applied at applications reduced the osmotic potential of the wheat leaves
high concentrations may have a minor toxic effect on the and prevented the germination of spores of E. graminis. These
pathogen, most soilborne pathogens grew better in culture as studies implicated chloride in reducing the osmotic poten-
the chloride concentration increased by 0.5–1.0% (Firdous tial of cell sap. The optimal water potential for most
 Mineral Nutrition and Suppression of Plant Disease 237

plant-infecting fungi is less than
0.5 mPa, and the growth following the drastic decrease of SO2 emissions from coal-
rates of most of them are reduced by 50% when the water burning facilities (Haneklaus et al., 2007). Most growers do
potential decreases to
3.0 mPa. There are exceptions. For not consider S nutrition as a component in their fertility
example, F. culmorum prefers an osmotic potential of
0.8 to programs. However, it is frequently applied as NH4(SO4)2,

1.2 mPa. Complete inhibition of many pathogens occurs at Ca2SO4, or MgSO4. These salts are almost exclusively applied

5.0 to
10 mPa (Cook and Papendick, 1972). The reduced with the aim of supplying the cations N, Ca, or Mg. When
growth of the pathogen before the onset of pathogenesis manipulation of pH is the sole objective, elemental S and
would play an important role in reducing disease. Schneider AlSO4 are often used. On field crops, several studies have
(1990) advanced a theory that chloride ions are used by plants shown that proper sulfur nutrition can induce resistance to
to tolerate the osmotic up-shocks that plant cells encounter plant pathogens through its effect on the function and avail-
when plant water potential is abruptly altered. In the absence ability of S-containing amino acids. Under controlled con-
of chloride ions, root cells manufacture organic ions, such as ditions in a greenhouse experiment, Wang et al. (2003) found
organic acids and amino acids, to maintain charge balance. that disease severity of oilseed rape infected with Sclerotinia
Schneider (1990) argued that when plant water potential is sclerotiorum, corn infected with Bipolaris maydis, and winter
altered by irrigation or changes in irradiation, organic solutes wheat infected with Rhizoctonia cerealis was significantly de-
are quickly exported to the root apoplast to maintain a pres- creased when the S supply was increased to young plants.
sure potential within the root cells. These organic compounds Under the conditions of S deficiency in field experimentation,
would, in turn, act as nutrition for root pathogens before soil-applied S fertilization significantly reduced the infection
pathogenesis. When chloride was applied, plants would need of oilseed rape with light leaf spot (Pyrenopeziza brassicae), of
to produce less organic and amino acids for charge balance. grapes with powdery mildew (Uncinula necator), and of potato
Roots would export chiefly chloride ions of low nutritional with stem canker (R. solani; Schnug et al., 1995; Klikocka et al.,
value in their exudates. The concept of altered root exudation, 2003). Soil-applied S reduced the infection rate and disease
as proposed by Schneider (1990), may explain changes in severity of R. solani on potato tubers by 41% and 29%, re-
microbial communities in the rhizosphere. The possible role spectively (Klikocka et al., 2003).
of beneficial microbes in chloride-treated wheat plants was Sulfur-induced resistance runs parallel to other defense
proposed by Halsey and Powelson (1980) and Christensen mechanisms, which are activated after fungal attack so that a
et al. (1981). Chloride fertilization of neutral soils has also clear differentiation is difficult, particularly under field con-
been associated with changes in the microbial community ditions. The S-containing amino acids cysteine, methionine,
(Amir et al., 1996; Elmer, 1995, 2003). Elmer (1995) isolated and glutathione play roles in protein synthesis of many
bacteria from NaCl-treated asparagus plants and found greater defense products.
densities of beneficial fluorescent pseudomonads. It is rea- In addition, salicylic acid as well as H2O2 initiates and
sonable to assume that the effect of chloride in reducing os- maintains systemic resistance, and the accumulation of sali-
motic potential also affects water cycling and root exudations. cylic acid is linked to S metabolism (Haneklaus et al., 2007).
In later split root studies, Elmer (2003) reported that NaCl As air quality improves and atmospheric S decreases further,
application to asparagus decreased root exudation of malic growers will need to be more attentive to S nutrition.
acid and amino acids, increased Cl ions, and increased fluor-
escent pseudomonads on the nontreated side of the root sys-
tem. Thus, chloride had a root-mediated effect on the Micronutrients
microbial community.
The ability to increase Mn also could increase host resist- The metals iron, manganese, zinc, copper, molybdenum,
ance by favoring the deposition of ligneous defense barriers boron, and nickel have diverse but essential roles in plants,
following infection. The synthesis of these products requires functioning as cofactors or activators of enzyme systems, as
many reactions in the shikimic acid pathway that contain previously mentioned in the section on Cl. Many of these
enzymes activated by manganese. enzyme systems play pivotal roles in disease resistance
through the production of defense products, including key
roles in phenol metabolism and lignin biosynthesis. In gen-
Sulfur eral, the concentrations that correct visual deficiency symp-
toms in plants are often far below the levels needed to ensure
Sulfur (S) is an essential element and a component of proteins proper plant health and defense against a plant disease. Most
and is the oldest recorded fungicide in use on vegetables and micronutrients become less available as the pH rises so
other crops. The fungicidal effect of elemental S was dis- growers should be aware that although a crop species may
covered in the early 1800s and has been widely used in agri- favor alkaline soil and may not show deficiencies, it may be
cultural production since the end of the nineteenth century more susceptible to attack from plant pathogens. One quick
(Hoy, 1987). Elemental S proved to be most efficient against method to correct aboveground deficiency symptoms and
rust and powdery mildew (Coleno, 1987; Cook, 1987; Hoy, boost resistance to foliar diseases is to apply a foliar appli-
1987) but was also successfully used against other diseases, cation of the needed micronutrient(s). However, as micro-
such as downy mildew in cereals (Hoy, 1987) and common nutrients are not translocated to the roots, this application
scab of potato (Vlitos and Hooker, 1951). However, the sig- would not suppress a root disease. Below is a more com-
nificance of soil-applied S on plant disease only recently be- prehensive discussion of each element's role in plant disease
came evident when S deficiency symptoms were noted suppression.
238 Mineral Nutrition and Suppression of Plant Disease

Iron At soil pH below 5.2 or above 7.8, Mn becomes chemically

available or unavailable, respectively. Its availability between
Iron (Fe) nutrition not only boosts plant vigor and health, it these ranges is largely influenced by the Mn-transforming
indirectly affects disease development in the root zone where microbes. For this reason, soil microbes, especially bacteria
its unavailability may limit the growth of pathogens (Expert, and fungi, have a profound effect on the global cycling of Mn
2007). One of the major roles where Fe influences plant dis- and its availability for plant uptake (Elmer, 1995; Erhlich,
ease is through siderophores, which are synthesized and ex- 1996; Ghiorse, 1988; Tebo et al., 1997). It has been demon-
creted by certain microorganisms to acquire their iron and strated, for example, that Mn-oxidizing microbes can decrease
thereby rob other microorganisms of Fe in the rhizosphere Mn availability by up to five orders of magnitude (Brouwers
(Buyer et al., 1994; Crowley and Gries, 1994; Marschner, et al., 2000; Tebo et al., 1997).
1995). Siderophores are low molecular weight molecules It has long been noted that certain environmental con-
(o1500 Da) that possess a high affinity for Fe3 þ . They spe- ditions that affect Mn availability has the same effects on
cifically capture Fe3 þ ions and transport them back into the disease, such as pH, N form, organic matter, moisture status,
microorganism that produces them. However, the side- soil temperature, and microbial activity (Huber and Wilhelm,
rophores can also release their Fe3 þ ions to iron-deficient 1988). One obstacle to increasing Mn levels in plants is its lack
plants. Therefore, it seems possible that microbial side- of mobility. Foliarly applied Mn is not basipetally translocated
rophores present in soils increase Fe availability on the root to roots. Furthermore, soil-applied Mn can be rapidly immo-
surface, depending on their different chemical, physical, and bilized by oxidizing soil organisms. It is not always possible to
absorptive characteristics. Mechanisms involved in these pro- override the biological and environmental factors affecting Mn
cesses have, however, been poorly defined. availability and utilization (Graham and Webb, 1991).
In addition to exploitation of microbial siderophores by Potential mechanisms of controlling disease with Mn have
plant roots, there are other beneficial effects of siderophore- not been totally resolved. Manganese affects the production of
producing rhizobacteria on plant growth. Specific Pseudomonas many host defense products. Manganese functions primarily as
strains called plant growth promoting rhizobacteria (PGPR) an activator of enzymes, including dehydrogenases, tranferases,
are able to enhance the growth of plants. Application of these hydroxylases, and decarboxylases (Burnell, 1988). Funda-
microbes can result in significant yield increases and were at- mentally, Mn serves as an activator of several enzymes in the
tributed entirely or in part to competition for Fe with patho- shikimic acid pathway for production of important secondary
genic microorganisms (Schroth and Hancock, 1982). In fact, metabolites that lead to the production of phenolics, cyano-
several strains of Pseudomonas fluorescens are effective in bio- genic glycosides, and the deposition of ligneous defense barriers
control of plant diseases due to their ability to deny available (Burnell, 1988). The role of Mn in carbohydrate, nitrogen, and
Fe to pathogenic bacteria or fungi. However, siderophore- secondary metabolism indirectly affects disease (Graham, 1983;
mediated biocontrol activity of rhizosphere Pseudomonas spe- Huber, 1989). More directly, Mn can reduce growth of the
cies is affected by the availability of Fe in the rhizosphere. The pathogen, which may explain why these organisms evolved
rhizosphere needs to be Fe limiting to maximize the bio- mechanisms for oxidizing Mn to unavailable forms.
control potential that, in turn, may limit its overall use in
biocontrol. Another mechanism of biocontrol by rhizobacteria
is induced systemic resistance (ISR). Siderophores produced by Zinc
bacteria might induce ISR. Leeman et al. (1996) and Buysens
et al. (1996) both found that ISR against plant disease was Zinc (Zn) nutrition is associated with important plant defense
governed by determinants regulated by Fe availability. pathways against fungal, bacterial, viral, and nematode
Another critical aspect concerns the signaling role of Fe in pathogens (Siddiqui et al., 2002). Zinc protects plant cells
plants and plant-associated microorganisms. The cellular and from toxic oxygen radicals and plays an important role in the
molecular regulatory mechanisms controlling the genes in- production of disease resistance signaling proteins. Zinc has
volved in Fe uptake, trafficking, and storage are still unclear. direct effects on fungal growth and secondary metabolism and
Elucidating these issues may eventually help to develop strat- indirect effects on host susceptibility. Zinc strictly modulates
egies in plant disease management. Nevertheless, it should be fungal accumulation of citric acid, stimulates glucose uptake,
recognized that biotechnological approaches targeting isolated and pigment and antibiotic production by microbes (Duffy,
components within complicated metabolic pathways are lim- 2007).
ited in their application. Zinc nutrition has pronounced effects on root diseases.
Tomato diseases caused by Macrophomina phaseolina, Fusarium
solani, and R. solani and charcoal rot of maize caused by M.
Manganese phaseolina were all reduced by increasing soil concentrations of
Zn (Siddiqui et al., 2002; Pareek and Pareek, 1999). In-furrow
A rich body of information exists on the role of manganese treatment with a mixture of zinc oxide (ZnO) and zinc sulfate
(Mn) nutrition in suppressing a plant disease (Thompson and (ZnSO4) reduced the incidence of powdery scab of potato,
Huber, 2007). Manganese in plants is absorbed in the reduced caused by Spongospora subterranea f. sp. subterranea (Falloon
Mn2 þ form and is poorly mobile in mature leaves but slightly et al., 1996). Zinc soil amendments reduced the severity of
mobile in root and stem tissue. However, the availability maize smut, caused by Ustilago maydis, and the effect was in-
of Mn in soil is influenced most strongly by soil pH and mi- creased with nitrogen fertilizers (Kostandi et al., 1997). Maize
crobial activity (Gilkes and McKenzie, 1988; Norvel, 1988). grain yields were correspondingly increased.
 Mineral Nutrition and Suppression of Plant Disease 239

Zinc influences the biocontrol activity of many bacteria. Copper is an essential micronutrient for pathogens as
Amendment of Zn soil amendments improved the activity well. Copper deficiencies in Colletotrichum lindemuthianum can
of Pseudomonas aeruginosa against M. phaseolina, F. solani, and result in the pathogen becoming avirulent to bean. Copper-
Meloidogyne javanica on tomato, even though they exhibited containing laccases can be important virulence factors for
little direct effect on root disease or galling and zinc-deficient many plant-pathogenic fungi, enabling them to degrade phys-
soils supported higher populations of P. aeruginosa (Siddiqui ical barriers and detoxify phytoalexins and tannins (Mayer and
et al., 2002). The role of Zn in microbial interactions involves Staples, 2002). For example, Heterobasidion annosum declined
biocontrol of Fusarium with Pseudomonas. Duffy and Défago in aggressiveness as laccase levels decreased (Johansson et al.,
(1997) showed that the biocontrol activity of P. fluorescens 1999). Integrated research involving Cu with other defense-
CHA0 against Fusarium root rot of tomatoes was enhanced stimulating approaches could enhance the effectiveness of
in the presence of Zn with increased production of the anti- many cultural management strategies for disease control.
biotic 2,4-diacetylphloroglucinol. The effect was mediated by
a Zn-induced inhibition of fusaric acid by the pathogen that
can, in turn, inhibit the antibiotic. Another example where Molybdenum
Zn indirectly influences biological control is for pathogenic
strains of R. solani that produce phenylacetic acid (Siddiqui Molybdenum (Mo) is not associated with influencing plant
and Shaukat, 2003, 2005). This metabolite can inhibit disease like other elements. A few studies show an increase in
the production of nematicidal compounds by P. fluorescens disease, but an equal numbers show no effect (Graham and
when damaged by root knot nematode (Meloidogyne incognita) Stangoulis, 2007). Suppression of Verticillium wilt of tomato
and is greater when R. solani is present. However, when by treatment of roots with molybdenum has been reported
sufficient Zn was present, phenylacetic acid production by (Dutta and Bremner, 1981) along with a study by de Jesus
R. solani was decreased, which relieved the repression of et al. (2004), who reported that application of Mo 25 days
nematicidal compounds and resulted in enhanced biocontrol after sowing beans suppressed angular leaf spot, caused by
of the nematode. Phaeoisariopsis griseola.
The best-known role of Mo involves nitrogen metabolism
and its role in activating the nitrate reductase enzyme, which
Copper may then influence disease development. In addition, heavy
metals like Mo are effective in deactivating viruses by de-
The recognition of the inhibitory effect that copper (Cu) has naturing protein coats (Verma and Verma, 1967), killing
on microorganisms has made it a common component of zoospores of Phytophthora sp. (Halsall, 1977) and inhibiting
many pesticides, whereas its role in plant physiological and nematodes (Haque and Mukhopadhyaya, 1983). However, at
host defense reactions that influence disease resistance or this point, manipulating Mo nutrition has not been demon-
susceptibility has received much less attention (Graham, 1983; strated to be an effective strategy for disease management.
Graham and Webb, 1991). The reduction of foliar diseases
with soil-applied Cu and benefits from Cu applied at sub-
biocidal rates are evidence of increased plant resistance rather Boron
than a direct effect on the pathogen (Graham, 1983; Graham
and Webb, 1991; Shabranskii et al., 1984). Copper-activated The role of boron (B) nutrition in plant disease is through its
physiological processes also increased the resistance of hops to wide ranging effects on structural and metabolic processes that
downy mildew (Shabranskii et al., 1984). The system where control plant defense (Stangoulis and Grahm, 2007). In gen-
Cu nutrition has made the striking difference is in barley. Low eral, B deficiency leads to host susceptibility by compromising
levels or deficiency of Cu causes pollen sterility, which reduces the plant's ability to manufacture lignin and phenolic com-
yield and predisposes plants to ergot (Azouaou and Souvré, pounds that are necessary for inhibiting pathogen invasion.
1993; Graham, 1975) by causing the normally closed flowers When B fertilizers are used in these situations, plant health and
of wheat and barley to open, exposing them to infection by disease suppression are increased. Much of the role of B in
Claviceps purpurea (Mantle and Swan, 1995). disease suppression is yet to be investigated.
Copper affects plant disease by three known mechanisms.
Direct toxicity as a bactericide or fungicide is the most com-
mon one. However, the effects of Cu on plant resistance and Nickel
pathogen virulence can also operate simultaneously. For ex-
ample, Cu is an essential cofactor in various enzyme systems The role of nickel (Ni) as an essential mineral element for
involved in plant defense against infection, the production of higher plants is a relatively recent discovery (Brown et al.,
antimicrobial compounds, and general disease resistance 1987). Its low requirement in plants qualifies it as a piconu-
(Graham, 1983; Graham and Nambiar, 1981; Graham and trient rather than a micronutrient, yet deficiencies are observed
Webb, 1991; Lebeda et al., 2001). Copper chloride induces the on a wide variety of plants (Wood and Reilly, 2007). Nickel
activity of chalcone synthase, a key enzyme in the biosynthesis deficiency is increased when root knot nematodes are present.
of diverse flavonoids involved in plant disease resistance These observations indicated an adverse influence of nema-
(Harker et al., 1990). Amino oxidases involved in H2O2 pro- todes on Ni nutrition of plants. Given that a wide range of
duction in cell walls in response to attack of pea by Ascochyta physiological processes are disrupted by insufficient Ni nu-
rabiei (Pass.) Labrousse required Cu (Laurenzi et al., 2001). trition, Ni may be a missing link in certain recalcitrant
240 Mineral Nutrition and Suppression of Plant Disease

disorders affecting several crop species that suffer from replant when subjected to acid or β-glucosidase hydrolysis displayed
disorders associated with pathogenic nematodes or fungi. strong fungistatic activity (Cherif et al., 1994). More recently,
flavonoids and momilactone phytoalexins, low molecular
weight compounds that have antifungal properties, were
Beneficial Elements (Silicon and Aluminum) found to be produced in both dicots and monocots, respect-
ively, fertilized with Si and challenged inoculated by the
As stated above, not all elements are viewed as essential, but a pathogen in comparison with nonfertilized plants also chal-
growing body of research has shown that many nonessential lenged inoculated by the pathogen (Fawe et al., 1998;
elements can suppress a number of plant diseases. It is inter- Rodrigues et al., 2004). These antifungal compounds appear
esting that the ameliorating effects of these elements are fre- to be playing an active role in plant disease suppression. In
quently only realized when the plant is under a biotic stress, addition, an increase in superoxide (O
) generation was ob-
such as a plant disease that, once again, implicates their in- served in rice leaves of plants treated with silicon 15 min after
fluence in host defense mechanisms. inoculation with Magnaporthe grisea (Maekawa et al., 2002).
Plants that accumulate silicon (Si) have shown enhanced Rodrigues et al. (2005) showed that β 1–3 glucanase, per-
resistance to a number of foliar and root diseases in both oxidase, and PR-1 proteins are associated with rice blast sup-
dicots and monocots (Datnoff et al., 2007c). Silicon is ab- pression. Both these studies suggest that other additional
sorbed by roots as monosilicic acid, Si(OH)4, and most plants mechanisms may be involved in silicon-mediated resistance to
accumulate this element in great quantities (between 1% and plant diseases. Recently, genome-wide studies for tomato, rice,
10% of their dry weight). The availability of silicon to plants and wheat amended with silicon and compared with non-
varies depending on the soil. Tropical/subtropical soils highly amended control plants have shown a differential expression
weathered, low in base saturation, and acidic are known to be of a large number of genes, and these genes are known to be
low in plant-available silicon. Organic and sandy soils also involved in host plant defense mechanisms or metabolism
contain low levels of silicon. Further, soilless mixes are silicon (Brunings et al., 2009; Chain et al., 2009; Ghareeb et al., 2010).
deficient. Consequently, plants grown on these types of root- Given the low silicon content now being found in a
ing media may benefit from a silicon application. Many number of soil types and its role in enhancing disease resist-
agronomic and horticultural crop plants supplied with silicon ance, it seems prudent to consider soluble silicon sup-
may gain protection from pathogenic fungi, bacteria, nema- plementation as amendments as simple, inexpensive methods
todes, and viruses (Table 4). Silicon appears to affect a number to reduce plant diseases.
of components of host plant resistance that includes delaying Recent research has shown that aluminum (Al) is similarly
the incubation and latent period and reducing lesion expan- taken up in plants at low levels and may be important to plant
sion rates, lesion size, and lesion number (Seebold et al., health (Shrew et al., 2007). It has been a long-held practice to
2001). Subsequently, disease progress and final disease se- lower root medium pH with AlSO4, which is often accom-
verity is dramatically reduced, and the resistance of susceptible panied by reductions for many plant diseases caused by Ver-
cultivars is augmented to almost the same level as those with ticillium and Thielaviopsis. However, evidence may suggest that
complete or partial resistance (Seebold et al., 2000; Rodrigues the aluminum ion is partially responsible for suppression of
et al., 2001). Silicon may even suppress plant disease as ef- disease through its effect on the germination of fungal spores
fectively as a fungicide (Resende et al., 2013; Seebold et al., in soil. Applications of different forms of Al to peat soils at pH
2004). As the silicon concentration (insoluble or soluble) in- 4 and 6 were followed by reductions in the population
creases in plant tissue, plant disease suppression greatly densities of Phytophthora parasitica. Strategies for incorporating
improves. Al in disease management need to be developed.
The underlying mechanisms that govern disease protection
are still not well understood; however, the effect of silicon on
plant resistance to disease is considered to be due to either an Summary
accumulation of absorbed Si in the epidermal tissue or an
expression of pathogenesis-mediated host defense responses There is no question that proper nutrient management has
(Datnoff et al., 2007c). Accumulated monosilicic acid poly- marked effects on plant diseases. Nutrients influence the plant
merizes into polysilicic acid and then transforms to amorph- pathogen both directly and indirectly by controlling the
ous silica, which forms a thickened Si cellulose membrane. By pathogen's rate of germination, penetration, colonization, and
this means, a double-cuticular layer protects and mechanically reproduction; affecting the infection rate or the progress of
strengthens plants. Silicon also might form complexes with disease; and having a toxic effect (Datnoff et al., 2007b). In
organic compounds in the cell walls of epidermal cells, thus addition, nutrients promote plant vigor, aiding recovery from
increasing their resistance to degradation by enzymes released infection, and change the anatomical or physiological plant
by fungi. state by thickening the cuticle or changing biochemical re-
Research also points to the role of silicon in planta as being actions. Hence, these nutritional practices often serve as the
active, and this suggests that the element might be a signal for first line of defense against the occurrence of plant disease.
mediating defense reactions to plant diseases (Belanger et al., However, the strategy of using fertilization practices as a
2003; Rodrigues et al., 2003). Silicon has been demonstrated management strategy against disease needs to be developed for
to stimulate chitinase activity and rapid activation of perox- each crop species.
idases and polyphenoxidases after fungal infection. Glycosi- This article highlights a few examples where manipulating
dically bound phenolics extracted from Si-amended plants fertilization reduced disease. Another more important goal
 Mineral Nutrition and Suppression of Plant Disease 241

Table 4 Host, disease, and plant pathogen response to silicon applications

Host Disease Pathogen Effecta

Monocots
Rice Leaf and neck blast Magnaporthe grisea "
Brown spot Cochliobolus miyabeanus "
Sheath blight Thanathephorus cucumeris "
Leaf scald Monographela albescens "
Stem rot Magnaporthe salvinii "
Grain discoloration Many fungal species "
Bacterial leaf blight Xanthomonas oryzae pv. oryzae "
Root knot nematode Meloidogyne spp. "
Wheat Powdery mildew Blumeria graminis "
Leaf blotch Septoria nodorum "
Foot rot Fusarium spp. "
Leaf spot Phaeosphaeria nodorum "
Eye spot Oculimacula yallundae "
Brown foot rot Fusarium culmorum ∅
Brown rust Puccinia recondita ∅
Sorghum Anthracnose Colletotrichum graminicola "
Barley Black point Alternaria spp. "
Powdery mildew Erysiphe graminis f.sp. hordei "
Rye Powdery mildew Erysiphe graminis "
Sugarcane Brown Rust Puccinia melanocephala ∅
Ring spot Leptosphaeria sacchari "
Zoysiagrass Leaf blight Rhizoctonia solani "
Creeping bentgrass Root rot Pythium aphanidermatum "
Brown patch Rhizoctonia solani "
Dollar spot Sclerotinia homoeocarpa "
Kentucky bluegrass Powdery mildew Sphaerotheca fuliginea "
Bermudagrass Leaf spot Bipolaris cynodontis "
St. Augustine-grass Gray leaf spot Magnaporthe grisea "
Corn Stalk rot Pythium aphanidermatum and Fusarium moniliforme "
Corn smut Ustilago maydis "
Dicots
Tomato Fusarium wilt Fusarium oxysporum f.sp. lycopersici (races 1 and 2) ∅
Fusarium crown and root rot Fusarium oxysporum f.sp. radicis-lycopersici "/∅b
Powdery mildew Oidiopsis sicula "/∅b
Bacterial wilt Ralstonia solanacearum "
Cucumber Powdery mildew Sphaerotheca xanthii "
Anthracnose Colletotrichum orbiculare "
Colletotrichum lagenarium ∅
Leaf spot Corynespora citrullina "
Crown and root rot Pythium ultimum and Pythium aphanidermatum "
Gray mold rot Botrytis cinerea "
Black rot Didymella bryoniae "
Fusarium wilt Fusarium oxysporum f.sp. cucumerinum "
Lettuce Pythium root rot Pythium spp. ∅
Muskmelon Powdery mildew Sphaerotheca xanthii "
Zucchini Powdery mildew Erysiphe cichoracearum "
Pumpkin Powdery mildew Sphaerotheca xanthii "
Rose Powdery mildew Sphaerotheca pannosa "
Black spot Diplocarpon rosae "
Morning glory Anthracnose Colletotrichum gloeosporioides "
Pea Leaf spot Mycosphaerella pinodes "
Paper daisies Anthracnose Colletotrichum gloeosporioides "
Grape Powdery mildew Uncinula necator "/∅b
Soybean Stem canker Diaporthe phaseolorum f.sp. meridionalis "
Rust Phakopsora pachyrhizi "c
Strawberry Powdery mildew Sphaerotheca macularis f.sp. macularis "
Tobacco BdMV Belladonna mottle virus #
a
Silicon can decrease ("), increase (#), or has no effect (∅) on disease intensity.
b
Silicon decreases disease intensity if sprayed onto the leaves but has no effect on disease if added to the nutrient solution.
c
Lemes et al., 2011.
Source: Adapted from Datnoff, L.E., Rodrigues, F.A., Seebold, K.W., 2007c. Silicon and plant disease. In: Datnoff, L.E., Elmer, W.H., Huber, D.M. (Eds.), Mineral Nutrition and Plant
Disease. St. Paul, MN: APS Press, pp. 233−246.
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was to underscore how nutrient management could advance a Burnell, J.N., 1988. The biochemistry of manganese in plants. In: Graham, R.D.,
major strategy for disease control and how that could have an Hannam, R.J., Uren, N.C. (Eds.), Manganese in Soils and Plants. Dordrecht, The
Netherlands: Kluwer Academic Publishers, pp. 125–137.
immediate impact on agriculture and food systems. Unfortu-
Buyer, J.S., Kratze, M.G., Sikora, L.J., 1994. Microbial siderophores and rhizosphere
nately, the fractionation of the data has hindered development ecology. In: Manthay, J.A., Crowley, D.E., Luster, D.G. (Eds.), Biochemistry of
of a working plan for disease management. Most experimental Metal Micronutrients in the Rhizosphere. Boca Raton, FL: CRC Press, pp. 67–80.
studies tend to focus on one plant, one element, and one Buysens, S., Heungens, K., Poppe, J., Hofte, M., 1996. Involvement of pyochelin
disease in isolation of other variables, so most research pro- and pyoverdin in suppression of Pythium-induced damping-off of tomato by
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vides only ‘snapshots’ of information. The dearth of experi- 865–871.
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agronomic and horticultural crop species whose fertility re- comprehensive transcriptomic analysis of the effect of silicon on wheat plants
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