Physiological and Molecular Plant Pathology 64 (2004) 189–199

www.elsevier.com/locate/pmpp

Powdery mildew of Arabidopsis thaliana: a pathosystem for exploring

the role of silicon in plant–microbe interactions
Dalila Ghanmia, David J. McNallya, Nicole Benhamoua,b, Jim G. Menziesc,
Richard R. Bélangera,*
a
Centre de Recherche en Horticulture, Sainte-Foy, Qué., Quebec, Canada G1K 7P4
b
Recherche en Sciences de la Vie et de la Santé, Pavillon Charles-Eugène Marchand, Université Laval, Qué. Quebec, Canada G1K 7P4
c
Agriculture and Agri-Food Canada, 195 Dafoe Road, Winnipeg, Manitoba, Canada R3T 2M9
Accepted 30 July 2004

Abstract
Silicon (Si) has been used in agriculture to protect plants against disease for hundreds of years and its prophylactic effects in monocots and
dicots are well documented. The mechanisms by which Si exerts its protective effects in planta, however, are uncertain and presently the
subject of debate. In this study, we sought to determine if Arabidopsis thaliana could be used to clarify the role of Si in plant–pathogen
interactions. Accordingly, X-ray microanalysis mapping, light microscopy, scanning and transmission electron microscope techniques were
used to examine the leaves of SiK fed A. thaliana plants inoculated with the powdery mildew fungus, Erysiphe cichoracearum. The results
of this study demonstrate for the first time, that A. thaliana is a species that absorbs Si and that the incidence of powdery mildew disease for
SiK fed plants is significantly lower compared to control plants. In particular, treatment with Si appeared to induce the production of an
electron-dense, fungitoxic substance that accumulated within and around the collapsed fungal haustoria of infected epidermal cells within the
leaves of disease-resistant plants. These results with Arabidopsis corroborate recent observations in other non-related species and support the
emerging theory that the mechanisms by which Si imparts resistance to plants are complex and are not entirely explained by the traditionally
proposed role of Si as a reinforcer of mechanical resistance. Collectively, the findings of the present study have established the Arabidopsis
thaliana-Erysiphe cichoracearum pathosystem as a valid model to investigate the role of Si in plant–microbe interactions.
q 2004 Elsevier Ltd. All rights reserved.

Keywords: Silicon; Arabidopsis thaliana; Erysiphe cichoracearum; Powdery mildew; Plant–pathogen interactions; Induced resistance; Defense reactions;
X-ray microanalysis mapping

1. Introduction have highlighted the advantages of treating plants with Si

The beneficial effects of silicon (Si) have been
demonstrated in several plant species, however, it is not
considered essential for plant nutrition [16,32]. Although
consistent evidence of direct positive effects on yield
have been traditionally hard to obtain, most experiments
have demonstrated the advantages of Si application in
terms of disease control. Research efforts during the last
few years have focused on examining the role of Si in
the outcome of plant–microbe interactions and several
studies originating from Europe, North America and Asia
* Corresponding author. Tel.: 418 656 2758; fax: 418 656 7856.
E-mail address: richard.belanger@plg.ulaval.ca (R.R. Bélanger).
0885-5765/$ - see front matter q 2004 Elsevier Ltd. All rights reserved.
doi:10.1016/j.pmpp.2004.07.005
to control plant diseases [18]. In that context, treatment
with Si was recently shown to be a safe and effective
means of protecting wheat and barley from Blumeria
graminis [25,52], rice from Pyricularia oryzae [42] and
cucumber from Pythium ultimum [5,10,11] and Podo-
sphaera xanthii[34].
As a result of the accumulating evidence demonstrat-
ing the protective effects of Si in different pathosystems,
the role of this element in plant defense has gained
acceptance. By contrast, the mechanisms by which Si
exerts its protective effects in planta are presently the
subject of ongoing debate and controversy. Traditionally,
polymerized Si deposited within the epidermal cells
of plants was thought to act as a physical barrier,
190 D. Ghanmi et al. / Physiological and Molecular Plant Pathology 64 (2004) 189–199
reinforcing mechanical resistance against pathogenic
penetration [23,39]. Evidence supporting this school of
thought has largely been provided by studies correlating
Si accumulation with increased resistance against fungal
pathogens in cereal species [15,49]. For example, using
barley Carver et al. [9] reported that powdery mildew-
resistant plants presented an early accumulation of Si
compared to vulnerable plants. Based on these obser-
vations, the authors argued that the accumulation of Si in
the apoplast effectively reduced penetration of the
pathogen resulting in a greater level of resistance
among these plants.
The role of Si as a reinforcer of mechanical resistance
was questioned a few years later when Samuels et al. [44]
demonstrated that treatment with Si decreased powdery
mildew disease in cucumber and Menzies et al. [35] showed
that attempted sites of fungal penetration were surrounded
by a phenolic-like material. These preliminary observations
were later corroborated by Fawe et al. [17] who showed that
treatment with Si induced defense reactions in cucumber,
including the production of fungitoxic flavonoid com-
pounds. Recently, studies involving wheat and rice have
supported an active role for Si as a mediator of defense
reactions. Indeed, treatment with Si was shown to induce
defense mechanisms in wheat against Blumeria graminis f.
sp. tritici which were linked to the accumulation of an
electron-dense, phenolic material surrounding fungal haus-
toria within the epidermal cells of these plants [4].
Moreover, Si was shown to trigger the production of
antifungal momilactone phytoalexins in rice effectively
conveying increased resistance against blast disease caused
by Magnaporthe grisea[41].
In light of these recent studies demonstrating the
active role of Si in plant disease resistance, it appears
that the mechanisms by which Si exerts its protective
effects are more complex than currently acknowledged
and are not entirely explained by the traditional role of
this element as a component of mechanical resistance. In
the present study, we sought to determine whether
Arabidopsis thaliana, a model species that is being
increasingly used to explore plant–pathogen interactions
[3,37], could be exploited to clarify the role of Si in
plant defense. Arabidopsis thaliana is particularly well-
suited for this type of investigation given the substantial
amount of information available for this species and its
compatibility with powdery mildew, a disease known to
be controlled by treatment with Si [7,9]. To our
knowledge, the role of Si, in either a physiological
or pathological context, has never been examined for
A. thaliana. Consequently, it is not known whether
A. thaliana actively absorbs Si, since plant species differ
considerably in their ability to absorb Si [38], or whether
the application of Si influences how this species interacts
with powdery mildew. In this context, three objectives
were established: (1) to determine if A. thaliana absorbs
Si supplied in the form of potassium silicate; (2) to
determine if the application of Si reduces the level of
disease on A. thaliana plants inoculated with the
powdery mildew fungus Erisyphe cichoracearumand;
(3) to examine the ultrastructure of E. cichoracearum—
A. thaliana interactions on the leaves of plants treated
with Si.
2. Materials and methods
2.1. Plant material and application of Si
Arabidopsis thaliana, accession Colombia (Col-0), was
used because of its reported compatibility with the
powdery mildew fungus Erysiphe cichoracearum [1].
This accession was obtained from the ABRC (Arabidop-
sis Biological Resource Center, Ohio State University,
Columbus OH). A. thaliana seeds were germinated in
0.2% agar media and incubated at 4 8C for 3 days
according to Frye and Innes [19], prior to being
transplanted into pots containing Pro-Mixw (Premier
Horticulture, Rivière du Loup, Qué.) at a density of 5
plants per pot. A. thaliana plants were then placed in
environment-controlled growth chambers (16 h light,
23 8C/8 h dark, 18 8C; 60–70% humidity) and covered
with plastic sheets (to ensure a high level of humidity)
which were later removed following the emergence of
the first real leaves. For the purpose of this study, two
treatments were used: (1) A. thaliana plants watered with
a nutrient solution containing soluble silicon (SiC) and;
(2) A. thaliana plants watered with a nutrient solution
without soluble silicon (SiK). The SiC nutrient solution
contained an optimized concentration of Si (100 ppm) in
the form of potassium silicate as outlined by Chérif et al.
[12]. A. thaliana plants were then watered with either the
SiC or SiK nutrient solution four times a week starting
from the first leaf stage for the duration of
the experiment.
2.2. Inoculation of plants and evaluation of disease severity
Erysiphe cichoracearum, designate UCSC (University
of California at Santa Cruz), obtained from Dr Somerville,
was maintained on squash plants grown under the same
conditions as A. thaliana plants. Inoculation with
E. cichoracearum was performed at the sixth leaf stage
(3–4 weeks after germination) by brushing conidia from
the infected leaves of squash plants onto the healthy leaves
of A. thalianaplants according to Xiao et al. [51]. Disease
severity was quantified eight days after inoculation and was
based on an arbitrary scale of 0–4 (where 0 represented no
powdery mildew and a score of 4 meant that approximately
80% of leaf surfaces were covered with powdery mildew)
as described by Adam et al. [2]. A total of 20 plants
were used for each treatment, and the experiment was
repeated twice.
 D. Ghanmi et al. / Physiological and Molecular Plant Pathology 64 (2004) 189–199 191

2.3. X-ray microanalysis mapping and scanning electron
microscopy (SEM)
Scanning electron microscopy was used to observe the
development of E. cichoracearum and X-ray microanalysis
mapping was used to determine the Si concentration within
the leaves of SiC and SiK A. thaliana plants. For each
treatment, a mininum of three leaf samples from different
A. thaliana plants harvested 8 days after inoculation were
prepared for X-ray microanalysis mapping and SEM
analyses as outlined by Chérif et al. [12]. Samples were
then analyzed using a CAMECA SX-100 Universal EPMT
microscope (Cameca instruments Inc., Trumbull, USA)
operating at a voltage of 15 kV and a current of 20 nA.
2.4. Transmission electron microscopy (TEM)
and light microscopy
For microscopy analyses, a minimum of ten leaf samples
from SiC and SiK A. thaliana plants harvested 8 days after
inoculation were fixed in 3% gluteraldehyde and then rinsed
with a 0.1 M sodium cacodylate buffer (pH 7.2) prior to
being dehydrated by soaking in an ethanol series containing:
30, 50, 70, 80, 95 and 100% ethanol. Leaf samples from
SiC and SiK plants were then post-fixed for 1 h at room
temperature in 1% osmium tetroxide (pH 7.2) before being
fixed in Epon. Resin-embedded leaf samples were then
cross-sectioned into 3 mm ultra thin sections using a
diamond knife and visualized using a JEOL 1200 EX
transmission electron microscope (JEOL, Tokyo, Japon)
operating at 80 kV.
For light microscopy, thin sections from resin-embedded
leaf samples harvested from SiC and SiK plants were
stained with 1% aqueous toluidine blue prior to being
analyzed with an Olympus (Shinjuku-Kv, Tokyo) model
BH2-RFCA bifocal microscope. Light microscope images
were recorded using a Coolsnap-Prow Media Cyberneticsw
(Silver spring, MD) color camera (1 s exposure) and
analyzed using the Image-Prow Plus version 4.5.1 software
by Media Cyberneticsw (Silver Spring, MD).

Fig. 1. X-ray microanalysis mapping images showing trichomes on the surfaces of leaves harvested from Arabidopsis thaliana plants treated with: (a) a nutrient
solution without silicon (SiK) or; (b) a nutrient solution containing 100 ppm of silicon (SiC). The concentration of silicon is indicated by color (see inset),
where white represents the highest concentration of silicon and black indicates no silicon. Observations are representative of analyses on a minimum of three
samples.
192 D. Ghanmi et al. / Physiological and Molecular Plant Pathology 64 (2004) 189–199
3. Results
3.1. X-ray microanalysis mapping
X-ray microanalysis mapping of Si distribution in
A. thaliana plants revealed no detectable levels of Si within
the leaves of SiK plants (Fig. 1a). By contrast, X-ray
microanalysis mapping of SiC leaves showed a uniform
pattern of Si deposition with a strong deposition of
the element on and around trichomes (Fig. 1b).
3.2. Evaluation of disease severity and scanning electron
microscope (SEM) analysis
SiC and SiK plants inoculated with E. cichoracearum
displayed a different level of infection when examined 8
days after inoculation. SiK plants exhibited obvious
symptoms of the disease with fungal colonies extending
from leaf margins to mid-rib sections (Fig. 2a) whereas in
SiC plants, fungal development was limited and rarely
observed (Fig. 2b). In a similar manner, the disease rating
Fig. 2. The leaves of Arabidopsis thaliana plants treated with: (a) a nutrient
solution without silicon (SiK) and; (b) a nutrient solution containing
100 ppm of silicon (SiC). Leaves from both treatments were harvested
8 days after inoculation with the powdery mildew fungus Erysiphe
cichoracearum.
for SiK plants was systematically higher and averaged
1.5G0.2 whereas it was significantly lower for SiC plants
at 0.2G0.1. These results were corroborated by scanning
electron microscope observations where dense mycelial
mats and conidial colonies were observed on the leaves of
SiK plants (Fig. 3a), while mycelial development at the leaf
surface of SiC plants was reduced to a loosened network of
fine hyphal filaments (Fig. 3b).
3.3. Light and transmission electron microscopy
(TEM) analysis
Preliminary observations using light microscopy
revealed that with SiK plants, hyphae developed abun-
dantly at the leaf surface (Fig. 4a) and formed turgescent
haustoria in the underlying epidermal cells (Fig. 4b and c).
Even at low magnification, haustorial digitations, charac-
teristic of a successful infection, were readily visible. In the
case of SiC plants however, haustoria appeared to be
collapsed and encased in a dark material whereby no
structures were discernible (Fig. 4d and e).
For SiK leaf samples, the cytology of infection with
E. cichoracearum resembled that previously described in
a number of plant species infected by biotrophic fungi
[4,21]. For instance, hyphae of the pathogen growing at
the leaf surface produced germ tubes that formed
appressoria which adhered to the host cell surface
(Fig. 5a). Successful epidermal penetration was associ-
ated with the formation of narrow penetration pegs that
crossed the cuticle and the underlying epidermal cells.
Enlargement of the infection pegs in epidermal cells
resulted in the elaboration of multilobed structures, the
haustorial bodies (Fig. 5b). In transverse sections, the
multi-shaped haustorial lobes appeared disconnected from
each other (Fig. 5b). The haustorial cytoplasm contained
numerous organelles including several mitochondria. In
all cases, the haustorial body was surrounded by a
circumvoluted membrane, the extrahaustorial membrane,
which is derived from the host’s plasma membrane
(Fig.5b). Interestingly, some samples revealed the host
plant’s ability to form hemispherical protuberances
resembling papilla at the sites of attempted fungal
penetration (Fig. 5c and d). In most cases however, the
fungus escaped this barrier produced by the host and
successfully colonized epidermal cells as evidenced by
the presence of turgescent haustoria with typical ultra-
structure (Fig. 5d).
Although the pathogen was able to penetrate the
epidermis of SiC leaf samples, marked changes in the
morphological and structural features of haustorial bodies
were noticed (Fig. 6). For instance, most haustorial
bodies were severely damaged and haustorial lobes were
frequently distorted. Degradation events were mainly
characterized by a necrotization of the fungal lobes
which appeared as aggregated, polymorphic structures
lying in an amorphous matrix (Fig. 6a and b). A dense
 D. Ghanmi et al. / Physiological and Molecular Plant Pathology 64 (2004) 189–199 193

Fig. 3. Scanning electron micrographs of leaf surfaces of Arabidopsis thaliana plants treated with: (a) a nutrient solution without silicon (SiK) and;
(b) a nutrient solution containing 100 ppm of silicon (SiC). Leaves from both treatments were harvested 8 days after inoculation with the powdery mildew
fungus Erysiphe cichoracearum. Observations are representative of analyses on a minimum of three samples.

granular material was generally found to accumulate in an electron-dense material (Fig. 6c and d) which was
the space between the aggregated haustoria and itself surrounded by multi-textured appositions that
the extrahaustorial membrane (Fig. 6a and b). In appeared to have been sequentially deposited as judged
addition, penetration pegs appeared totally encased in by their lamellar aspect (Fig. 6c).
194 D. Ghanmi et al. / Physiological and Molecular Plant Pathology 64 (2004) 189–199

Fig. 4. Light micrographs of toluidine blue O-stained semi-thin sections of leaves harvested from Arabidopsis thaliana plants treated with: (a–c) a nutrient
solution containing no silicon (SiK) and; (d and e) a nutrient solution containing 100 ppm of silicon (SiC). Leaf samples from all treatments were harvested
8 days post inoculation with the powdery mildew fungus Erysiphe cichoracearum. (a, amyloplast; ch, collapsed haustorium; dm, dense material; e, epidermal
cell; fm, fungal mycelium; ,h, haustorium). Observations are representative of analyses on a minimum of ten samples.

4. Discussion
In recent times, A. thaliana has emerged as a promising
model for investigating plant–microbe interactions. Several
factors make A. thaliana particularly interesting for this type
of research including the extensive amount of publicly
available data for this species, the ongoing development of
molecular tools and resources facilitating the rapid analysis
of its genome [8,13,20,28,47], and the recent identification
of compatible isolates of powdery mildew fungi [1,26,
40,51]. Although the role of Si had never been examined in
A. thaliana, evidence is provided from the present study that
the A. thaliana–E. cichoracearum pathosystem is a reliable
model to be used for clarifying the controversial role of this
element in the defense strategy of plants.
X-ray microanalysis mapping of the leaves of A. thaliana
plants submitted to exogenously supplied Si demonstrated
that the plant could absorb soluble Si. This finding was
pivotal to the continuation of this work since higher plants
differ considerably in their capacity to absorb Si. For
example, Jones and Handrick [24] have classified plants
into two groups depending on their content of polymerized
Si (% of dry weight). Species that accumulate Si include
members of the Equisetaceae and Cyperaceae (10–15%),
and more recently cucumber [18] and wheat [4]. Examples
of non-accumulating plants include most grasses (1–3%)
and dicots such as tomato [38]. The observed concentra-
tion of polymerized Si on the trichomes of SiK fed
A. thaliana plants was typical of plants that accumulate Si
and is explained by the fact that soluble silicon is
absorbed by plants as uncharged silicic acid (Si(OH)4).
Once absorbed, Si then moves from the roots to the leaves in
plantaby following the transpiration stream and eventually
polymerizes in the extracellular spaces and walls of
epidermal cells at sites of strong evapotranspiration and
basal cells of trichomes [4,44].
The most striking difference between the leaves of SiC
and SiK A. thaliana plants inoculated with E. cichora-
cearum was the extent of fungal colonization. This
observation is noteworthy because it demonstrates that
although E. cichoracearum was able to successfully
penetrate and infect the epidermal cells of SiC plants, its
rate of development was altered. Closer inspection of leaf
surfaces with scanning electron microscopy corroborated
these observations as it revealed that the leaves of SiC
plants were systematically less colonized compared to SiK
plants, whose leaves supported dense mycelial mats. The
ability of Si amendments to reduce the incidence of
powdery mildew disease in A. thaliana supports similar
findings in non-related species such as cucumber [11,12,17],
muskmelon and zucchini squash [36], rose and strawberry
[48], and more recently wheat [4] and rice [41,42].
Several lines of evidence lead us to suggest that the
ability of SiC A. thaliana plants to resist infection with
E. cichoracearum is not entirely explained by the
traditionally proposed role of Si as a reinforcer of
mechanical resistance. Support for this concept is provided
by the extensive deposition of a substance with high affinity
for toluidine blue O stain within and around the collapsed
fungal haustoria, including along the epidermal cell wall
and upon the extrahaustorial membrane of the pathogen
within infected SiC epidermal cells. Although the toluidine
 D. Ghanmi et al. / Physiological and Molecular Plant Pathology 64 (2004) 189–199 195

Fig. 5. Transmission electron micrographs of ultra-thin sections of leaf samples harvested from Arabidopsis thaliana plants treated with a nutrient solution
without silicon (SiK). (a, b, d) infected epidermal leaf cells containing fully developed, multi-lobed haustoria with fungal mycelia immediately outside these
cells. (c) a fungal penetration peg embedded within an epidermal cell papilla that successfully stopped the development of a haustorium. All leaf samples were
harvested 8 days following inoculation with the powdery mildew fungus Erysiphe cichoracearum. (a, !13000; b, !6000; c, !8000; d, !8000). Cu, cuticule;
EHM, exhahaustorial membrane; Ep, epidermis; F, fungus; HB, haustorial body; P, papilla; Pe, penetration peg; L, Lobe. Observations are representative of
analyses on a minimum of ten samples.
196 D. Ghanmi et al. / Physiological and Molecular Plant Pathology 64 (2004) 189–199

Fig. 6. Transmission electron micrographs of ultra-thin sections of leaf samples harvested from Arabidopsis thaliana plants treated with a nutrient solution
containing 100 ppm of silicon (SiC). (a and b) Infected epidermal cells containing collapsed electron-dense fungal haustoria that are detached from the
extrahaustorial membrane and surrounded by a dark osmiophilic substance and; (c), infected epidermal cells showing the electron-dense granular material
surrounding the haustorial neck of the fungus. (d) close-up view of collapsed haustorium and lobes embedded within a granular and electron-dense deposit.
All leaf samples were harvested 8 days following inoculation with the powdery mildew fungus Erysiphe cichoracearum. (a, !5000; b, !8000; c, !16000;
d, !13000). Em, electron-dense material; GM, granular material; HB, haustorial body; Pe, penetration peg; WA, wall apposition. Observations are
representative of analyses on a minimum of ten samples.

blue O reaction cannot be considered highly specific, it is
known to stain phenolic compounds a deep blue color
[6,46]. The intense blue coloration observed around
the collapsed haustoria exclusively within the infected
epidermal cells of SiC A. thaliana plants may well
correspond to the accumulation of phenolic compounds.
Previous studies, albeit using different systems, have
reported similar findings. For instance, using powdery
mildew-infected cucumber plants treated with Milsanaw,
a reported elicitor of induced resistance in this species [14],
Wurms et al. [50] observed collapsed fungal haustoria
encased in a similar material with a high affinity
for toluidine blue O stain. Phytochemical analyses of
elicited, powdery mildew-infected cucumber leaves later
corroborated these findings when it was revealed
that following elicitation, autofluorescent C-glycosyl
 D. Ghanmi et al. / Physiological and Molecular Plant Pathology 64 (2004) 189–199
flavonoid phytoalexins rapidly accumulated around and
within the fungal haustoria of infected epidermal cells prior
to the collapse of the pathogen [33].
The formation of papilla and cell wall appositions has
been reported as an effective induced defense mechanism
within some cereal species [9,29,30]. By studying papilla
formation in cereals, it was shown that mature papilla are
mostly heterogeneous in composition and often contain
proteins, phenolic compounds such as lignin, complex
polysaccharides such as callose and pectin [43,45] and
polymerized Si thereby explaining their refractive, glass-
like appearance [30,31,53]. As a first active defense
reaction, papilla relate to powdery mildew fungal pen-
etration in two ways: in some instances, penetration fails
when papilla are present and alternatively, penetration may
succeed and the papilla becomes a collar for the haustorial
neck [52]. Using transmission electron microscopy, both
outcomes were observed for SiC and SiK A. thaliana
plants during this study, strongly suggesting that papilla
formation may not be sufficient by itself to explain the level
of protection observed for SiC plants. These results
corroborate the findings of Chérif et al. [12] who reported
that the increased resistance of cucumber to P. ultimum
induced by exogenously supplied Si was not associated with
the accumulation of insoluble Si at sites of pathogen
penetration, regardless of the plant organ examined. Studies
examining the concentration of Si near penetration sites of
fungi in wheat, barley, cucumber and morning glory have
also reached similar conclusions [9,29].
In previous ultrastructural studies of cereal powdery
mildews, hypersensitive response (HR) was reported as a
common phenomenon observed in incompatible inter-
actions [22]. Using the Blumeria graminis f. sp. hordei-
barley pathosystem, Koga et al. [27] proposed that during
HR, phenolic compounds were released and accumulated on
the cell walls of epidermal cells undergoing fungal ingress.
The authors proposed that soluble Si, if present, would form
insoluble complexes with phenolic compounds thereby
strengthening the mechanical resistance of epidermal cells
against penetration by the pathogen. In such cases, it was
suggested that Si accumulation was secondary to HR and
largely played a passive and mechanical role in the defense
response of the plant. Although degenerated epidermal
cells were at times observed within the leaves of SiC
A. thaliana plants using TEM during this study, the degree
to which HR response was responsible for the increased
resistance of SiC plants to E. cichoracearum remains
difficult to establish. Rather, the results of this study militate
strongly in favor of the induction of other active defense
responses. In line with this hypothesis, the observation that
E. cichoracearum frequently penetrated and produced
mature haustoria within the epidermal cells of SiC A.
thalianaplants suggests that Si is not a key mechanical
deterrent to fungal penetration. Moreover, the granular
electron-dense nature of the material that accumulated
within the infected epidermal cells of SiC A. thaliana
197
plants is highly characteristic of phenolic compounds [12]
and bore a striking resemblance to the ultrastructural
observations recently reported for powdery mildew-infected
cucumber and wheat plants treated with Si [4,50]. Finally,
the fact that this electron-dense material, likely enriched in
phenolic substances, was consistently observed around the
collapsed haustoria in SiC A. thalianaplants speaks in favor
of a direct toxic effect on the pathogen.
In conclusion, this study provides the first evidence that
A. thalianahas the ability to absorb soluble Si which, in turn
protects it against infection with the powdery mildew
fungus E. cichoracearum. Although the precise mechanisms
by which Si exerts its protective effects in A. thaliana are
not fully understood, some lines of evidence indicate that
they likely do not involve Si behaving exclusively as a
reinforcer of mechanical resistance. Instead, powdery
mildew resistance observed for SiC A. thaliana plants
appears to be the result of induced defense reactions,
including the production of an electron-dense material
likely enriched in phenolic compounds with an apparent
toxic effect on the pathogen. Accordingly, these results with
A. thaliana corroborate recent observations in other species
and support the theory that once absorbed by the plant, Si
operates as a mediator of defense reactions in planta. The
results of this study combined with the extensive resources
readily available for A. thaliana make the Arabidopsis
thaliana-Erysiphe cichoracearum pathosystem a reliable
model with which to investigate the role of Si in plant–
microbe interactions.
Acknowledgements
This work was supported by grants from the Natural
Sciences and Engineering Research Council of Canada
(NSERC) and the Canada Research Chairs program to
R.R. Bélanger. The authors thank Dr Somerville for
generously providing E. cichoracearum as well as
D. Auclair and A. Goulet for excellent technical assistance.
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