J. Phytopathology 156, 458–463 (2008) doi: 10.1111/j.1439-0434.2007.01392.

x
 2008 The Authors
Journal compilation  2008 Blackwell Verlag, Berlin

Cell and Molecular Biology Laboratory, University of São Paulo, USP-Piracicaba, Brazil

The Potential Use of a Silicon Source as a Component of an Ecological

Management of Coffee Plants
J. C. Martinati1, R. Harakava2, S. D. Guzzo2 and S. M. Tsai1
AuthorsÕaddresses: 1Cell and Molecular Biology Laboratory of Agriculture Nuclear Energy Center (CENA ⁄ USP), Av.
Centenário, n. 303, PO Box 96, 13416-00 Piracicaba, SP, Brazil; 2Phytopathology Biochemistry Laboratory, Biological
Institute, São Paulo, SP, Brazil (correspondence to J. C. Martinati. E-mail: jumarti@cena.usp.br)
Received August 31, 2007; accepted November 20, 2007

Keywords: coffee, coffee leaf scorch, silicon, resistance, defence, Brazil

Abstract perennial crop is subject to high losses in potential

Coffee is one of the most important agricultural export
commodities in the world and it represents the main
export from some developing countries. Therefore, the
development of new methods of coffee management
that improves production without causing any damage
to the environment is an attractive alternative for pro-
ducers. Much effort has been invested towards under-
standing the mode of action of compounds that can
induce resistance against several pathogens without
injuring the environment. Many researches have con-
sidered silicon efficient in avoiding plant pathogen pen-
etration and development. Our aim was to verify the
effect of potassium silicate and calcium ⁄ magnesium sili-
cate in the development of coffee seedlings (Coffea
arabica cv. Mundo Novo) as well as to evaluate the inci-
dence of coffee leaf rust development under greenhouse
conditions. The experiment was a completely random-
ized design with 12 treatments with 10 plants per treat-
ment. The treatments were 0, 0.25, 1.25, 2.5, 4 and
5 lm of Si for each source of silicon incorporated into
the soil. The seedlings were inoculated with a uredini-
ospores suspension of Hemileia vastatrix (2 mg ⁄ ml) at
the seventh month after planting (six pair of leaves).
Evaluations were performed by counting the number of
lesions per leaf. The statistical analysis showed that the
number of lesions reduced by up to 66% at the highest
silicon dose when compared to the number of lesions in
control plants. Infected plants were found to have a lin-
ear decrease of lesions with the increase of silicate con-
centration. The lowest number of lesions per leaf area
was observed in plants that received 5 lm of Si from
potassium silicate. This result indicates the use of
silicon as an alternative for an ecological management
system for coffee disease protection.
Introduction
Coffea arabica is one of the most important agricul-
tural export products in developing countries. This
production due to pests and diseases. The most
destructive disease of coffee plants is the coffee leaf
rust caused by the fungus Hemileia vastatrix (Berkekey
and Broome) and the yield losses in Brazil have been
estimated at 45% if no control methods are taken
(Matiello, 1991).
Fungicides provide efficient protection, but their
applicability can be jeopardized by adverse environ-
mental effects and by the raising of pathogen-resistant
strains. Several chemical control strategies for enhanc-
ing plant resistance to pathogens have been under-
taken. Such measures provide ecological management
allied to the priority of sustainable coffee production.
The improvement of plant resistance based on induc-
tion of host defence becomes an important option to
control the rust disease in coffee plants.
Silicon sources have increasingly been used. Silicon
(Si) has long been known to reduce the incidence of
fungal diseases in a number of pathosystems (Bélanger
et al., 2003, 2003; Rodrigues et al., 2003; Botelho
et al., 2005; Fauteux et al., 2005), but its mode of
action in plants remains unclear, although it seems
that Si acts on general mechanisms common to most
plant species.
It has been shown that Si could act as an enhancer
of plant defence responses or as an activator of strate-
gic signalling proteins. Considered to be biologically
active, Si triggers a faster and more extensive plant
defence by interacting with several key components of
plant stress signalling systems ultimately leading to
induced resistance against pathogenic fungi (Fauteux
et al., 2005).
Based on their observations with cucumber, Fawe
et al. (2001) suggested that the Si bioactivity is com-
pared with the action of secondary messengers of sys-
temic acquired resistance (SAR). Thus, Si would act as
a modulator influencing the timing and extent of plant
defence responses. The effects of Si on secondary

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Coffee Leaf Rust Control
metabolism are achieved only after elicitation as well
as through the known activators. Silicon was shown to
be involved in the increased resistance of cucumber to
Podosphaera xanthii by enhancing antifungal activity
within the plant, and this was attributed to the pres-
ence of a phytoalexin identified as flavonol aglyconerh-
amnetin (Fawe et al., 1998, 2001).
Coffee defence genes against H. vastatrix have been
isolated and identified in resistant plants (Fernadez
et al., 2006; Guzzo, 2004). These studies were carried
out in different periods of postfungus inoculation and
both of them could demonstrate that the expression of
defence genes occurs from the early stages of infection
to 72 h after infection. Those genes were also identified
in susceptible coffee plants treated with the elicitor
BTH (benzo [1, 2, 3] thiadazole-7-carbothioic acis
S-methylester). With these results, we could observe
that even susceptible coffee plants can react against
infection process by enhancing the defence mechanisms
through elicitation.
Our aim was to verify the efficacy of silicon sources in
reducing coffee leaf rust in susceptible plants to establish
an economical and sustainable coffee production and
may offer a promising alternative to chemical control.
Materials and Methods
Coffee plant
The seeds of C. arabica cv. Mundo Novo (IAC – 388-
17-1), susceptible to race II of H. vastatrix used in this
study were donated by Cooperativa Garcafé, Garça
SP, Brazil. These seeds were surface sterilized with
2–5% sodium hypochlorite for 15 min and then
washed three times in sterile distilled water. Germina-
tion occurred in seedling containers filled with sand
until cotyledonary leaves were fully developed, approx-
imately 60 days after planting. Following leaf develop-
ment, the seedlings were transplanted to plastic vases
and kept in greenhouse. All fertilization recommended
procedures were performed.
Silicon treatments
Immediately after transplanting, the coffee seedlings
were divided into two groups, according to fertilization
treatments: (i) fertilized with calcium ⁄ magnesium sili-
cate; and (ii) fertilized with potassium silicate. Both
groups were completely randomized in a total of 12
treatments with 10 plants per treatment. The treat-
ments were 0, 0.25, 1.25, 2.5, 4 and 5 lm of Si for each
source of silicon incorporated into soil every
2 months.
Inoculum and inoculation procedure
Urediniospores of H. vastatrix race II were used in the
experiments. They were collected from field-infected
leaves of cv. Mundo Novo at Fazenda Santa Elisa (SP,
Brazil). The collection procedure involved scraping off
young rust pustules, drizzling the material and wrapping
up the spores in polypropylene tubes. The tubes
containing the spores were then stored in a constant
temperature chamber at )80C until the determination
459
of germination rate. Spore batches with less than 15%
of germination were not used for inoculation proce-
dures.
To prepare the inoculation, the selected urediniosp-
ores were activated by placing the tube in a water bath
at 40C for 10 min. Spores were suspended in distilled
water at a concentration of 2 mg ⁄ ml and the suspen-
sion was maintained homogeneous during the time of
inoculation. Coffee plant seedlings at 7 months of ger-
mination were inoculated with the solution described
above. This solution was sprayed on the seedling
leaves so that it would entirely cover the abaxial sur-
face with droplets. After the inoculation, the plants
were placed in a humid dark room at 25C for 48 h.
Analysis of development variables
For the measurement of the development variables,
height of coffee plants, number of leaves per plant and
the average area of coffee plants leaves were consid-
ered. For the calculation of leaf area, the same param-
eters cited by Favarin et al. (2002) were used.
Disease assessment
The level of the resistance induced in coffee plants
against the coffee leaf rust was evaluated by evaluating
the symptoms by counting the number of lesions per
leaf area of each leaf inoculated. Plant growth was
assessed by recording the stem length and the total
number of leaves per plant.
Statistics
Twelve treatments and 10 replicates were used. Regres-
sion analysis was used to investigate whether silicon
concentration and sources affected the number of
lesions. In all regression analyses, the percentage vari-
ance accounted for (R2) was used to assess goodness
of fit. Analyses were carried out using sas ver. 8.2
(SAS Systems, Cary, NC, USA).
Results
Effect of dose ⁄ source of silicon on plant growth and
development
The statistical analysis for the average height of the
coffee plants showed significant difference on the coffee
plant growth when the source used was potassium sili-
cate. No statistical difference was observed on the
average height for coffee plants amended with Ca ⁄ Mg
silicate (Table 1).
Table 1
Linear regression analysis for plant height
Ca ⁄ Mg silicate Potassium silicate
Regression curve 8.284 + 0.0229 · dose 9.8809 + 0.0415 · dose
P>F 0.0162 <0.0001
VC 18.73% 20.51%
R2 0.973 0.973
VC, variance coefficient.
The data show a comparison between the same source at different
dosage. The linear equation expresses the height values with the
doses enrichment. P > F values lower than 0.0005 are significant.
460 Martinati et al.

The coffee plants amended with potassium silicate
showed significant statistical difference among the
average height of the coffee plants (P > F < 0.0001).
As the doses increases (in mm), the plant height
increases (in cm) at a ratio of 0.0415 cm to each mm
of Si incorporated into the soil. The coffee plants trea-
ted with Ca ⁄ Mg silicate exhibited growth as high as
the coffee plants treated with potassium silicate, how-
ever, it does not show significant statistical difference
among Si doses of Ca ⁄ Mg silicate (P > F = 0.0162).
In the same way, leaf area and number of leaves per
plant showed significant statistical difference for the
various Si doses only when the source used was potas-
sium silicate (P > F < 0.0001, Table 2). The average
leaf area of coffee plants treated with potassium sili-
cate increased at a ratio of 0.1840 cm2 ⁄ mm of Si incor-
porated into the soil. When the source used was
Ca ⁄ Mg silicate, the average leaf area do not differ sig-
nificantly as Si doses increases (P > F = 0.0285).
However, the average leaf area of coffee plants treated
with Ca ⁄ Mg silicate was higher than the average of
the coffee plants treated with potassium silicate. This
is expected as the coffee plants treated with Ca ⁄ Mg sil-
icate exhibited increased heights. The same situation
occurred with the number of leaves per plant.
The Si content in coffee leaves was also evaluated.
The statistical analysis demonstrated no correlation
between dose ⁄ source of silicate for Si content in the
coffee leaves of the Si-treated plants (Table 3). The
coffee leaves collected for this analysis were divided
into three samples to investigate the Si content in dif-
ferent parts of the coffee plants. The coffee branches
were divided into 3 parts to better analyse the silicon
content. Near to substract, at the middle and at the
top of the branch. Similarly in all the pair of leaves,
the Si content was statistically the same independent
of the leaves localization (near or far from the sub-
strate according to the division cited).
Effect of dose ⁄ source of silicon on coffee leaf rust
The symptoms of coffee leaf rust could be observed
60 days after fungus inoculation in all inoculated
plants. These symptoms were characterized by yellow-
ish chlorotic lesions on the abaxial surface of coffee
leaves. The statistical analysis showed that the number
of lesions reduced up to 66% in the highest silicon dose
of potassium silicate when compared to the number of
lesions in control plants (Table 4). Infected plants were
found to have a linear decrease of lesions with the
increase of potassium silicate concentration. The lowest
number of lesions per leaf area was observed in plants
that received 5 mm of Si from potassium silicate
(Figs 1 and 2). The number of lesions per leaf area in
coffee plants treated with potassium silicate were signif-
icantly reduced (P > F > 0.0001) with the increase of
Si doses with a ratio of 0.00293 lesions for each mm of
Si added. The same did not occur in the coffee plants

Table 4
Table 2 Linear regression analysis for the number of lesions per leaf area
Linear regression analysis for leaf area of the coffee plants
Ca ⁄ Mg silicate Potassium silicate
Ca ⁄ Mg silicate Potassium silicate
Regression curve 0.9184 + 0.0001 · dose 1.4117 ) 0.00293 · dose
Regression curve 65.24 + 0.116 · dose 25.59 + 0.1840 · dose P>F 0.8272 <0.0001
Pr > F 0.0285 <0.0001 VC 21.16% 25.08%
2
VC 28.60% 28.98% R 0.975 0.973
R2 0.976 0.978
VC, variance coefficient.
VC, variance coefficient. The second, third and fourth pair of each inoculated coffee plant
The data show a comparison between the same source at different was analyzed. The data showed a comparison between the same
dosages. The linear equation expresses leaf area values with the doses source at different dosages. P > F values lower than 0.0005 are sig-
increase. P > F values lower than 0.0005 are significant. nificant.

Table 3 1.2 Ca/Mg silicate

Linear regression analysis for the silicon content in coffee leaves
1
Number of lesions

Ca ⁄ Mg silicate Potassium silicate

0.8
1 % Si 1.4607 ) 0.066 · dose 1.5053 ) 0.279 · dose
P>F 0.1550 0.3472 0.6
VC 12.86% 7.80%
2 % Si 1.3393 ) 0.099 · dose 1.2207 ) 0.356 · dose 0.4
P>F 0.1590 0.3707
VC 24.15% 13.46% 0.2
3 % Si 0.7378 ) 0.0089 · dose 0.6133 + 0.1549 · dose
P>F 0.8709 0.1514 0
0 1 2 3 4 5 6 7 8
VC 22.64% 31.67%
Si doses (mM)
VC, variance coefficient.
The data show a comparison between the same source at different Fig. 1 Regression analysis of the average of total number of lesion
dosages. The numbers in the first column indicates: 1, upper third; 2, per leaf area in coffee plants treated with calcium ⁄ magnesium sili-
the medium third and 3, lower third pair of leaves of the coffee cate. Regression curve: 0.9184 + 0.0001 · dose. F-value = 0.8272;
branch. P > F values lower than 0.0005 are significant. R2 = 0.987
Coffee Leaf Rust Control
1.2 Potassium silicate
1
Number of lesions
0.8
0.6
0.4
0.2
0 proteins that act as active transporters in accumulated
0 1 2 3 4 5 6 7
Si doses (mM)
Fig. 2 Regression analysis of the average of total number of lesion
per leaf area in coffee plants treated with potassium silicate. Regres-
sion curve: 1.4117 ) 0.00293 · dose. F-value = 0.0001; R2 = 0.995
treated with Ca ⁄ Mg silicate, where linear regression
coefficient does not differ statistically from zero and
the number of lesions in the different Si doses has no
statistical meaning.
Results from this study provide strong evidence that
silicon amendments mediate an effective protection
against H. vastatrix in coffee plants. This study con-
firms the numerous observations on the beneficial role
of Si in several plants. However, there is no study
regarding the identification of defence genes expressed
in coffee plants treated with Si. In a second step of this
work, we will identify the defence genes involved in
the interaction Coffee – silicon – H. vastatrix.
Discussion
Many reports have demonstrated the efficiency of sili-
con compounds in controlling plant disease caused by
pathogenic fungi (Datnoff et al., 1991; Chérif et al.,
1994a; Epstein, 1994; Bélanger et al., 1995, 2003; Fau-
teux et al., 2005). These data show that the application
of Si directly in the substrate protected the coffee plant
against one of the most serious strains of H. vastatrix
that is known to cause severe necrosis and yield losses.
The lack of any effect of silicon on plant growth is
in accordance with previous studies in which the main
focus was to show the efficiency of silicon in triggering
defence responses in coffee plants.
It was also possible to observe the deficiency of cof-
fee plant in translocated Si from the root to aerial part
of the plant. Current data also point that there is no
efficient translocation of potassium silicate in coffee
plants even when the Si is applied directly in the leaves
by spraying the Si solution and thus could not control
efficiently the coffee leaf rust (Rodrigues et al., 2005).
In this study, there was also a similar situation regard-
ing the Si translocation in coffee plants, however, it
was possible to observe a correlation between the
potassium silicate doses and the decrease in the num-
ber of lesions caused by H. vastatrix.
The Si concentrations found in the coffee plants cat-
egorizes them in the group of non-accumulated plants.
Miyake and Takahashi (1985) characterized the plant
into three types according to their Si accumulation:
461
accumulated plant – >1% of Si, non-accumulated
plants – <1% of Si and the intermediated plants –
values ranging to 0.5–1% of Si. There are several
hypotheses regarding the role of Si in non-accumulated
plants (Hodson et al., 2005). One of them includes its
participation on nutritional balance, plant structure
and stress resistance. This could be converted in better
growth and ⁄ or income of cultures.
Some researches suggest the presence of membrane
8 plant roots (Ma et al., 2002; Liang et al., 2005). Proba-
bly, this could be a point to be investigated concerning
the reason of why Si is not actively translocated from
root to aerial part of coffee plants and why it is not
well absorbed when applied by spraying the Si solution
in leaves. Possibly, the fact of Si being found in lower
levels in coffee plants is because, the coffee plants does
not have a membrane protein that act as a Si trans-
porter. Some non-accumulated plant species, such as
strawberry and tomato had a better development when
amended with silicon.
Some studies describes that as potassium doses
increases, calcium concentration decreases, this being
one of the predominant factor to a higher susceptibil-
ity to plant diseases (Marschner, 1995; Junior et al.,
2003). Pozza et al. (2001) and Junior et al. (2003) also
observed that the highest severity of Ôbrown eye spotÕ
in coffee plants was observed with the increase of
potassium doses. Our results agree with those of Mor-
aes et al. (1976) who reported that the increase in per-
centage of coffee leaf rust incidence was significantly
associated with higher doses of potassium. In this
study, when a Si source containing potassium was used
(potassium silicate), the number of leaf lesions was sig-
nificantly reduced, suggesting that the active element
was Si.
Several authors describe the action of Ca in the
reduction of severity of the symptoms of diseases in
plants. According to Elad and Kirshner (1992), the
severity of Botrytis cenerea in ruscus plants could be
reduced with the application of Ca. By the results
obtained in this study, this element was not an impor-
tant factor for the reduction of coffee leaf rust where
there was no significant results regarding the decrease
of the lesions with the increase of Ca ⁄ Mg silicate
concentration.
Similar effects were observed by Botelho et al.
(2005) in coffee plant inoculated with Cercospora cof-
feicola. There was a reduction of approximately 10%
of the leaf lesion with the increase of Si dose. Between
the mechanisms by which Si can confer resistance to a
determined disease, structural barriers are cited (Koga
et al., 1988; Bowen et al., 1992; Epstein, 1999; Rodri-
gues et al., 2003), or by the activation of chemical and
biochemical plant barriers (Heath and Stumpf, 1986;
Bélanger et al., 2003; Rodrigues et al., 2003). In coffee
plants, this resistance mechanism is not fully eluci-
dated, as it does not apparently absorb the element
effectively. One of the probable mechanisms would be
the formation of complex Si-hydroxyl group provided
462
by the amino acids residues in protein kinase. When
this complex is formed, Si can act as an activator of
these proteins that would activate the nuclear tran-
scription of defence proteins. As this complex is stable,
plausibly it is not possible to detect the presence of Si
by the quantitative analysis tests which would justify
its low amount in coffee plants. The bioactivity of Si is
compared to the activation of secondary messengers of
SAR, acting as a modulator which would influence the
defence response time (Fawe et al., 2001).
However, the induction of defence mechanisms
through the action of elicitors before the occurrence of
infection changes the energetic metabolism of the plant
which could result in yield losses. On the other hand,
Si amendment seems to induce the defence mechanism
only in response to the pathogen attack. This induc-
tion is expressed through a chain reaction of several
associated biochemical changes, hence characterized by
a quick and prolonged defensive response. This char-
acteristic explains the non-specificity of the Si-induced
resistance to several uncorrelated pathogens (Chérif
et al., 1994a,b).
It is well documented that susceptibility or resistance
is determined not just by the presence or absence of
genes for resistance, but by the rapidity and magnitude
with which the genetic information is expressed (Kuc,
1990; Pan et al., 1992). This is supported by the data
from the present study showing genotypical differences
in disease resistance, with the resistant cultivar having
significantly lower disease index values than the
susceptible one, regardless of Si application.
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