Epstein

P1: PSA/spd P2: PSA/ary QC: KKK/anil T1: KKK
March 30, 1999 15:30 Annual Reviews AR082-23
Annu. Rev. Plant Physiol. Plant Mol. Biol. 1999. 50:641–64

Copyright °c 1999 by Annual Reviews. All rights reserved
SILICON
Annu. Rev. Plant. Physiol. Plant. Mol. Biol. 1999.50:641-664. Downloaded from www.annualreviews.org
Emanuel Epstein
Access provided by UNIVERSIDADE DE SAO PAULO on 05/08/23. For personal use only.
Department of Land, Air and Water Resources—Soils and Biogeochemistry,

University of California at Davis, Davis, California 95616-8627;
e-mail: eqepstein@ucdavis.edu
KEY WORDS: essentiality, growth, development, mechanical strength, stress
“. . . silicon in life here on earth is little understood as yet. There is a lot of work to be
done” RJP Williams (136).
ABSTRACT
Silicon is present in plants in amounts equivalent to those of such macronutrient
elements as calcium, magnesium, and phosphorus, and in grasses often at higher
levels than any other inorganic constituent. Yet except for certain algae, including
prominently the diatoms, and the Equisetaceae (horsetails or scouring rushes), it
is not considered an essential element for plants. As a result it is routinely omit-
ted from formulations of culture solutions and considered a nonentity in much of
plant physiological research. But silicon-deprived plants grown in conventional
nutrient solutions to which silicon has not been added are in many ways exper-
imental artifacts. They are often structurally weaker than silicon-replete plants,
abnormal in growth, development, viability, and reproduction, more susceptible
to such abiotic stresses as metal toxicities, and easier prey to disease organisms
and to herbivores ranging from phytophagous insects to mammals. Many of these
same conditions afflict plants in silicon-poor soils—and there are such. Taken
together, the evidence is overwhelming that silicon should be included among the
elements having a major bearing on plant life.
CONTENTS
INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 642
“SILICON DEFICIENCY” IN PLANT PHYSIOLOGY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 642
HISTORY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 643
ESSENTIALITY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 645
SILICON IN PLANTS: THE AMOUNTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 647
641
1040-2519/99/0601-0641$08.00
642 EPSTEIN
SILICON AND GROWTH AND DEVELOPMENT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 650

ABIOTIC STRESSES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 653
BIOTIC STRESSES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 656
BIOCHEMISTRY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 657
CONCLUSION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 659
INTRODUCTION
Silicon is a quantitatively major inorganic constituent of higher plants but ab-

sent or nearly so from even major scientific publications devoted to them. The
reason for the astonishing discrepancy between the Si content of plants and the
Si content of the enterprise of plant science is the conclusion of the developers
of the solution culture technique, in the early 1860s, that Si need not be included
in the formulation of nutrient solutions, i.e. that Si is dispensable in the growth
of plants. This conclusion subsequently was reinforced by the promulgation
of a logically flawed definition of “essentiality,” the wide acceptance of which
did nothing to dissuade plant physiologists from their disinterest in this ele-
ment. There is, however, a large body of botanical, agronomic, horticultural,
and plant pathological knowledge of Si, along with some plant physiologi-
cal experimentation, that forcibly drives home the lesson that Si cannot be
written off as a plant biological nonentity. In this review evidence is sum-
marized that will drive home the significance to plant life of this ubiquitous
element.
“SILICON DEFICIENCY” IN PLANT PHYSIOLOGY

What better place to look for the significance accorded Si than this very series,
which has chronicled the advances in our field since 1950? Except for inci-
dental brief references, mainly concerned with clay minerals, the first volume
to include a discussion of Si was Volume 5 (1954), in a chapter devoted to the
mineral nutrition of phytoplankton (81), in which the well-known requirement
of diatoms for Si was discussed. For a decade after that we find nothing but an
occasional mention of the element, mainly in connection, again, with algae.
In 1964, Clements (31) sounded a note that will recur in the present review.
In a chapter entitled “Interaction of Factors Affecting Yield,” Clements, who
worked in Hawaii on the physiology of sugarcane, reported a number of in-
stances where applications of silicates to this and other crops gave striking
yield responses. He thought that silicate eliminated some toxic agents in those
highly weathered soils. This field-oriented line of inquiry was resumed in 1966,
Volume 17, where Bollard & Butler (17), in a chapter on mineral nutrition, de-
voted a section to Si, briefly discussing some aspects of it in relation to soil/plant
SILICON 643
interactions, Si uptake and transport, and the protection it may afford plants
against insects and disease agents.
In 1969, Volume 20, Lewin & Reimann (84) published the first and till now,
only chapter with the word silicon in its title: “Silicon and Plant Growth.” The
authors gave an account of the then current knowledge of Si, in respect to both
diatoms and higher plants.
In their discussion of metal toxicity in plants, Foy et al (54) summarized
evidence regarding the protection afforded by Si against the damage wrought

plants by high concentrations of metal ions in their substrate, such as Al, Mn, and
Fe. Metal toxicity is an increasingly acute problem, and we shall have occasion
to discuss the role of Si in often mitigating its adverse effects on plant growth.
In the last reference but one to Si in this series, in 1980, Clarkson & Hanson
(30) wrote briefly about Si, concluding with the sentence: “When more is
known about cell wall biosynthesis in higher plants the need for silicic acid can
be examined.” But despite the huge body of work on the biosynthesis of cell
wall components that has since been done (the preceding volume of this series
has no less than three reviews bearing on this topic), the expectation expressed
in that sentence has proven illusory. Finally, Carpita (25) recently referred
briefly to Si in the walls of grasses.
For another instance of “Si deficiency” in the literature let us turn to a volume
specifically devoted to the plant organ directly exposed to Si, the roots (134).
This 1002-page tome devoted to plant roots has about four lines devoted to
Si. The reasons for this neglect of the element, and further instances of it, are
discussed in the next section. These cases are not given in a spirit of finger-
pointing but to document our collective, astonishing disregard of an element
present in plants in amounts equivalent to those of Ca, Mg, S, and P, and often
in excess of them.
HISTORY
Research on the plant physiology of Si depended on the advent of the solution
culture technique (47). With some notable exceptions (mainly certain highly
weathered tropical soils), soils contain high percentages of Si. In most of them,
indeed, Si is second only to oxygen as a soil constituent; the mean values are
O, 49% and Si, 31% (125). In the soil solution, the direct source of Si that plant
roots draw upon, the element, in the form of H4SiO4 over the physiological
range of pH values, is present at concentrations normally ranging from 0.1 mM
to 0.6 mM (46), roughly two orders of magnitude higher than the concentrations
of P in soil solutions, for which Tisdale et al (130) give an average (but highly
variable) value of 0.0016 mM. Plants growing in soil therefore are exposed to
Si, and control of their Si status is well-nigh impossible.
644 EPSTEIN
That difficulty, at the experimental level, changed with the introduction of

the solution culture technique around 1860. An account of the then available
knowledge of plant nutrition is given by Sachs (110), in a chapter entitled Nutri-
ents (Nährstoffe). He stressed that only with this technique can the investigator
control the composition of the inorganic substrate of plants, and determine
which elements are indispensable constituents (unentbehrliche Bestandtheile)
of plant nutrition (p. 141). He concluded that Si does not belong among the in-
dispensable elements in the same sense as K, P, etc. He did, however, comment

on the wide distribution of Si in plants.
The conclusion of Sachs as to the dispensability of Si for plant life was

subsequently given further sanction by the publication, in 1939, of a definition
of essentiality (4): “an element is not considered essential unless (a) a deficiency
of it makes it impossible for the plant to complete the vegetative or reproductive
stage of its life cycle; (b) such deficiency is specific to the element in question,
and can be prevented or corrected only by supplying this element; and (c) the
element is directly involved in the nutrition of the plant quite apart from its
possible effects in correcting some unfavorable microbiological or chemical
condition of the soil or other culture medium.”
The near-universal acceptance of this definition is puzzling in view of its
flaws. As for (a), many plants may be quite severely deficient in a nutrient
element and yet complete their life cycle; (b) is redundant, (a) already having
made reference to the element (“it”) under consideration, excluding all others;
and (c) presumes that designation of an element as essential has to entail knowl-
edge of its direct involvement in the nutrition of the plant. But at the time when
the essentiality of B was established by Katherine Warington, AL Sommer, and
CB Lipman in the 1920s (45, 47), nothing was known about its direct involve-
ment in plant nutrition. The evidence simply was that the plants failed unless
the element was supplied. The same argument applies to the discovery of the
essentiality of several other elements at the time when their essentiality was
discovered. At any rate, the promulgation of this definition and its acceptance
reinforced the impression that Si could be dispensed with. Thus the classical
compilation of the chemical composition of plants by Goodall & Gregory (59)
contains not a single value of the Si content of any crop. When, in 1966, Hewitt
(63) published the second edition of what still is by far the most comprehensive
compilation of the composition of nutrient solutions, his listing of about 140
such formulations includes Si in just 5, meant specifically for research on Si
or Si accumulator plants. Even in recent books dealing specifically with plant
nutrition (14, 103), Si is given short shrift.
A turning point in research on Si was the publication, in 1967, of a review by
Jones & Handreck (78). It remains indispensable reading for anyone interested
in Si in soils, plants, and animals. Nearly every topic on Si in plants being
SILICON 645
pursued today was discussed, or at least touched upon, in that classical paper.
A look at its lengthy list of references impresses the reader by the extent to
which they come from agricultural and crop science publications rather than
from plant physiological ones.
The further history of advances made in our knowledge of Si in plants is
recorded in a number of reviews. That by Lewin & Reimann (84) in this series
has already been mentioned. Shkolnik (120) has summarized literature not
readily available in the West, and Richter & Suntheim (108) have given early
references, many to papers in German that have not been cited often. Raven’s
review (106) deals with transport, primarily, and is required reading. Epstein
(46) has discussed the reasons for considering experimental plants grown in
conventional nutrient solutions (i.e. without inclusion of Si in their formulation)
to be experimental artifacts. Savant et al (116) have recently reviewed silicon
management and sustainable rice production. Despite its specialized topic, this
is a wide-ranging review drawing on the extensive research published in Pacific
and Far East countries, and elsewhere.
Other reviews are mentioned in various contexts, but those referred to will
give readers an entrée into the material available concerning this element. So
will study of several multi-authored volumes (13, 49, 124). The 1975 book by
Voronkov et al (132) provides a large amount of analytical data on the Si content
of living things, ranging from bacteria to humans. The authors discuss possible
functions and remark on the paucity of what is known about its transport.
Vitti et al (131) give a general account of Si in soils and plants, including some
references pertaining to Brazil in particular. In the broad sense, the subject of
Si in plants is a subset of the inorganic chemistry of life, to which the books by
Fraústo da Silva & Williams (55), Kaim & Schwederski (79), and Simkiss &
Wilbur (123) are wide-ranging introductions. For trenchant, brief discussions
of Si in biology, reference is made to Birchall (15), Exley (50), Perry & Keeling-
Tucker (101), and Williams (137).
ESSENTIALITY
At this time only two groups of plants are known to have an absolute and
quantitatively major requirement for Si: the diatoms and other members of
the yellow-brown or golden algae, the Chrysophyceae (84, 124, 135), and the
“scouring rushes,” Equisitaceae (27).
As for higher plants in general, however, on which the present review focuses,
essentiality of Si in the Arnon-Stout (4) sense has not been proven (5, 126).
However, it is difficult to purge solution cultures thoroughly of Si. Woolley
(144) has done what still remains the most painstaking experimentation on this
subject and reduced the Si content of tomato plant tops to a mere 0.0006% on
646 EPSTEIN
a dry weight basis, but the growth of the plants was not diminished compared
with that of plants supplied with Si. Work in Japan, however, has led to the
conclusion, in 1990, that further intensive research “should qualify silicon as
an essential element for higher plants in the near future” (126). Werner & Roth
(135) are more sanguine and “consider it to be an essential element,” without,
however, spelling out the criteria for this opinion.
At this point we have to come to grips with the application of the Arnon
& Stout (4) definition of essentiality, already criticized above, to Si. This is
not the place for a full-blown discussion of this subject, but a few points need
to be made nevertheless. They have to do with (a) the limitations of purging

elements from experimental cultures; (b) the genotypic variations in nutrient
requirements; and (c) quantitative considerations. As for (a), when Woolley
(144), in the research already referred to, grew tomato plants in highly purified
culture solutions and failed to find differences between their growth and that
of plants deliberately supplied with Si, he concluded, correctly, that this did
not show Si to be nonessential but only that, if it were essential, it would be
required at tissue levels of less than 0.2 µmol · g−1 dry weight, or 0.0006%.
For demonstrating the essentiality of Ni, it was necessary to purify the medium
to the point where tissue levels were reduced to less than 0.00001% (21, 22).
Clearly, it has not been possible to reduce tissue levels of Si to comparably low
values, and the possibility of a micronutrient function for Si cannot be ruled
out. This same point concerning the limitations of purification procedures has
been made repeatedly (5, 45, 46). It applies to Si with particular force, because
the uncharged H4SiO4 passes through the ion exchangers so widely used for
preparing laboratory-grade water.
The second reservation about the definition of essentiality is that it conveys an
impression of general applicability that is not warranted. Sodium is an example:
It is a micronutrient for C4 plants (23), but not known to be a micronutrient for
plants generally. For Si, equal care needs to be taken not to generalize, in view
of its essentiality for some groups of plants.
Finally, quantitative considerations. Chlorine is assuredly a micronutrient for
higher plants (24), but it may well play a macronutrient role not only in some
halophytes (133) but in some genotypes and conditions, for such a common crop
as winter wheat (44, 82). For Si, no micronutrient function has been shown;
plants unable to grow without it have a quantitatively major requirement for the
element.
Thus it is necessary to go beyond mere listings of elements as essential or
not known to be so, and to give judicious interpretations of the complexities of
the real world. In view of what follows, Si will be considered “quasi-essential”
for many of those plants for which its absolute essentiality has not been estab-
lished. An element is defined as quasi-essential if it is ubiquitous in plants,
SILICON 647
and if a deficiency of it can be severe enough to result in demonstrable ad-

verse effects or abnormalities in respect to growth, development, reproduction,
or viability. That deficient situation prevails for Si in many highly weathered
tropical soils, such as Ultisols and Oxisols (53, 115), and, as already pointed
out, in conventional, Hoagland-type solution cultures. By extension, these same
considerations apply when plants are grown in commercial (or hobby) hydro-
ponics systems (11). The addition of clay to soilless media promoted the growth
of greenhouse crops (40); might not that finding be due to Si having become
available to the plants from the clay?
SILICON IN PLANTS: THE AMOUNTS

A tabulation of the ranges of the tissue levels of mineral elements found in plants
gives the range for Si as 0.1–10% on a dry weight basis, it being understood that
both lower and higher values may be encountered (46). Comparison of these
values with those for such elements as Ca (0.1–0.6%), S (0.1–1.5%), and others
shows Si to be present in amounts equivalent to those of several macronutrient
elements, and even exceeding them at the highest levels.
Very comprehensive sets of Si analyses of plant material have been compiled
by Japanese workers who grew plants of numerous species in the same soil
and then analyzed leaves for a number of elements including Si (98, 127–129),
expressing the results in terms of percentages or ppm (mg · kg−1) on a dry weight
basis. It is impossible in this small compass to do more than single out some
facts. The lowest value for Si recorded in any of these tabulations is 0.01%,
for very few species including Sansevieria trifasciata Prain (Agavaceae) and
Lycoris radiata Herb. (Amaryllidaceae) (98). Values below 0.1% were found
but rarely. The highest value among these leaf analyses was that for rice,
6.3% (98, 127). The coefficient of variation was higher for Si than for any
other element. In the angiosperms, there was a significant negative correlation
between values for Si and B, and a positive one for Ca and B.
Figure 1 condenses the information on the Si content of 147 species of
angiosperms (98), separately for dicots (left bar), and monocots (right bar).
Clearly, quantitatively, the values for Si fall in the same range as those for the
inorganic macronutrient elements, but their variability is wide. The observation,
often made in earlier publications, that the Si content of monocots is by and
large higher than that of dicots is borne out.
As Figure 1 indicates, the Si content of plants varies vastly in different geno-
types. Such differences may occur even among genotypes of the same species,
as shown, for example, in studies of rice ecotypes (38, 143). Other factors are
important for it, as is the case for many elements, but for Si, the differences
in Si content are exceptionally large. The availability of Si in the soil or other
648 EPSTEIN
Figure 1 Box and whisker chart showing the statistics of values of maximum, minimum, upper
quartile, lower quartile, and median, indicated, respectively, by bars at the top and bottom ends
of the vertical lines, upper and lower sides of the boxes, and bars within the boxes. Left symbol
and ordinate, dicots; right, monocots. The figure is based on the analysis of 147 species of
Angiospermae. From (98), by permission.
substrate does, of course, influence the Si content of the plant. Plants themselves
alter the chemical form and availability of soil Si (80; See 14a). Other gener-
alizations emerging from the reviews already cited are that, as a rule, most of
the Si absorbed is transferred from the roots to the shoots, and that within the
shoots, its distribution is far from even. For a recent (and extreme) instance
of the latter phenomenon, Ernst et al (48) analyzed different parts of the fruits
(nutlets) of the sedge Schoenus nigricans (Cyperaceae) collected at a wet dune
slack. The highest values by far were those of the pericarp of the fruit, ranging
up to 34%. Vegetative parts of the fruiting shoots had Si values lower by more
than an order of magnitude. Such differential, sharply localized distributions
of Si have often been noted, in both roots and shoots (68–70, 99, 113, 114).
Customarily, however, it is leaf tissue that is mainly analyzed for Si, as is
the case for other mineral elements. Analyses of many plants led Jones &
Handreck (78) to propose a rough division of plants into three groups: wetland
Gramineae (paddy-grown rice) have the highest values, on the order of 10–15%
on a dry weight basis; dryland grasses such as rye and oats are intermediate at
SILICON 649
Figure 2 Distribution of Si accumulators in the plant kingdom. Modified after Figure 2 of (129),
by permission.
about 1–3%; and most dicots have less than 1%. However, in view of the large
variability already referred to (see Figure 1), all such categorizations should be
viewed with caution. Finally, attention is drawn to the fact that even relatively
“low” values for tissue Si contents, such as 0.1%, are still comparable with the
low values found for such macronutrient elements as S, P, and Mg (46) (see
also Figure 1).
Figure 2 shows the distribution of Si in the plant kingdom, with marked dif-
ferences among groups, in that such relatively primitive plants as the Bryophyta
650 EPSTEIN
are high-Si plants, as are some of the evolutionarily advanced ones, such as the
Poales. In the Pteridophytes, both high-Si and low-Si data appear, and there
are scattered high-Si groups in the dicots, and in monocots besides the Poales.
Evolution seems to have “adopted” Si as a major mineral plant constituent a
number of times, and “made do” with more moderate but by no means negligible
Si contents for much of the plant kingdom.
The processes of absorption and transport of Si resulting in the eventual Si
contents of plants and their parts are unlikely to differ fundamentally from those
of other elements, to judge by the findings of Jarvis (75), Stookey (125a), and
MM Rafi & E Epstein (unpublished results). However, because it is in the form

of an uncharged molecule, H4SiO4, and is not redistributed once deposited as
“opal” (see below), its translocation within the plant is even more markedly
affected by the transpiration stream than is the case for other elements (see
below, under Abiotic Stresses). In the marine diatom, Cylindrotheca fusiformis,
Hildebrand et al (65) recently identified a cDNA that encodes a Si transporter
(68Ge, a chemical analog of Si, was used as a tracer). The findings represent the
first identification of a Si transporter in any organism.
SILICON AND GROWTH AND DEVELOPMENT

The literature on Si in plants is replete with reports that Si promotes the growth of
plants. In many instances the growth stimulation was due to the protection that
Si afforded plants against the detrimental effects of abiotic and biotic stresses,
discussed below. Another factor to keep in mind is that when experimental plants
are grown in soil, the interaction between added silicon and soil constituents
such as P may produce effects not directly attributable to Si (89, 109, 121).
Nevertheless, an appreciable body of evidence supports the conclusion that
often Si enhances plant growth and development and does so directly, hence its
designation here as “quasi-essential.”
Since experiments with solution culture are likely to provide the most un-
equivocal evidence in this regard, several such investigations are given first.
Adatia & Besford (1) grew cucumber (Cucumis sativus cv. Corona) in recir-
culating nutrient solutions to which Si had not been deliberately added (less
than 0.17 mM Si) and solutions with Si at ten times that concentration. The
lower leaves of the high-Si plants were darker green, better disposed for light
interception, and were noticeably rougher, especially their petioles, in com-
parison with these features of the low-Si plants. Leaf thickness was greater
than in the low-Si plants, and the dry weight per unit area of the high-Si
plant leaves was also greater. The leaves of the low-Si plants were prone to
wilting, and senesced earlier than those of the high-Si plants. Other indices
of plant performance were also higher in the high-Si plants. For example,
SILICON 651
on a unit area basis, the activity of Rubisco was 50% higher in the leaves
of the high-Si plants. Fruit yield, on the other hand, was the same in the two
treatments.
Other such instances of growth and development being positively affected
by Si have been given for such diverse plants as rice, Oryza sativa (97), loblolly
pine, Pinus taeda (43), cotton, Gossypium hirsutum (85), annual brome, Bro-
mus secalinus (56), and Poinsettia, Euphorbia pulcherrima (93), among many,
all these reports referring to experiments with solution cultures. Not all such
comparisons have yielded similar results, however. Thus Wutcher (145) com-
pared the growth of young orange trees, Citrus sinensis, in solution culture
without Si addition and with deliberate inclusion of Si; the results indicated
“only a limited role of this element in citrus nutrition.”
Turning to field-grown plants, or experimental ones in soil in pots, positive
responses are not as clear-cut as is the case with plants in solution culture,
as already pointed out. Nevertheless, the evidence for effects of Si per se is
strong. As early as 1976, Lian (85a) declared that “Si nowadays is generally
considered to be an essential element agronomically (though not, in the strict
sense, physiologically) for rice growth. . .today, the application of silicate fer-
tilizers is very common in Japan and Korea. An annual consumption of over
one million ton (38) and 400,000 tons (5) has been reported in these countries”
(the numbers in parentheses refer to Lian’s references by E Takahashi and CS
Parks, respectively). The Si material is mostly derived from slags, the refuse of
iron ore smelters most prominently. By now, good effects from the application
of Si fertilizers to rice have been reported from many locations, indicating “the
agronomic essentiality of Si management for increasing and/or sustaining rice
yields” (116). Winslow et al (143) have drawn attention to the difference be-
tween lowland, Si-rich soils where rice of the indica ecotype is grown in paddy
culture, and upland, Si-deficient soils, where the japonica ecotype is cultivated
under rainfed conditions. Yamauchi & Winslow (146) and Goto & Goto (60),
among others, also stress the importance of Si for the growth of japonica rice
in low-Si, upland soils. In many areas where rice is grown, depletion of plant-
available Si in soils may be the cause of declining rice yields (115). These
authors make clear that this conclusion refers not only to upland, acidic soils
but other soils such as calcareous ones as well.
While rice takes pride of place among crops that have been shown to respond
positively to Si applications, it is not the only one. As early as 1965 and 1966,
Clements (32) and Ayres (6) reported that application of silicate slag stimulated
the growth of sugarcane, Saccharum spp., on low-Si soils. Other research has
borne out that conclusion (3, 41, 42, 121). Silicon is used to good effect as a
fertilizer for sugarcane in Brazil, Australia, South Africa, and India (LE Datnoff,
personal communication).
652 EPSTEIN
As for other species, the depletion of plant-available Si commented on above

in connection with the decline of rice yields (116) may well be a factor, but
evidence is lacking. Silicate rock (granite) powder, applied to soils of the south-
western part of Western Australia, promoted the growth of wheat (66) and of
subterranean clover, Trifolium subterraneum, and ryegrass, Lolium rigidum
(35). The authors attributed the growth responses to K+ released from the rock
powder, but the possible contribution of other elements including Si is under
investigation (P Hinsinger, personal communication). Application of rock dust

to soils as a means to improve the yield and quality of crops has been touted
in the popular press but without documentation and explanation of the putative
effects (9).
The quantitatively major “product” of plant growth and development is cell
wall, and the physical properties of plants are determined to a large extent by
those of its cell walls. Virtually all the reviews mentioned above under the
heading, History, refer to the role of Si in minimizing or preventing lodging of
plants, especially of grain crops such as rice and wheat. The effect is attributed
to the deposition of Si in the form of solid amorphous silica, SiO2 · nH2O, in the
cell walls. The reinforcement of the walls provided by these “opal phytoliths”
adds mechanical strength.
Deposition of silica in the surface cell layers, and especially the epidermis,
has a bearing as well on physical surface properties. Thus Adatia & Besford
(1), in their experiments with cucumber plants growing in circulating nutri-
ent solutions, noted that the basal leaves of the high-Si plants and the petioles
were rough, and the cucumber fruit of the Si-supplied plants in the experi-
ments of Samuels et al (112) had a dull appearance, or bloom. The leaf edges
and the awns of Si-deprived wheat plants were fairly smooth, but those of
plants grown in solutions supplied with Si were exceedingly rough (104). For
the awns, the difference between the Si-deprived and Si-replete plants was
quantified in terms of the friction force—the force needed to be overcome
if the heads of the plants were to slide down an inclined plane. Despite the
numerous reports on physical differences between low-Si and high-Si plants,
this is so far the only quantification of any such difference. It was due to the
deposition of Si in the trichomes of the awns of the Si-replete plants, as deter-
mined by X-ray microanalysis, vs its absence, i.e. undetectability, in the Si-
deprived ones. These authors (104) provide formulations of +Si and −Si culture
solutions.
This Si reinforcement of cell walls, and further effects elicited by Si, protect
the plants against the depredations of disease organisms and herbivores includ-
ing insects, as discussed below under the heading, Biotic Stresses. Indeed, we
shall see that it is in relation to abiotic and biotic stresses that the role of Si in
plant life is most patent.
SILICON 653
ABIOTIC STRESSES
In 1957, Williams & Vlamis (138) made a remarkable discovery, the repercus-
sions of which have recently become more significant than at any time before.
When they grew barley, Hordeum vulgare cv. Atlas 46, in a conventional nutri-
ent solution with 9.l µM Mn, the leaves developed dark brown necrotic spots,
the formation of which was prevented by adding Si at 357 µM (10 ppm). The
yield depression caused by the Mn toxicity was likewise eliminated by the ad-
dition of Si. Their findings (138–141) did not indicate that Si diminished the
total Mn content of the leaves but rather, that it caused Mn to be more evenly
distributed instead of being concentrated in discrete necrotic spots. The inter-
action between Mn and Si has been studied by several workers, who also give
further references (71, 72, 76).
The discovery that Si often confers on plants protection against Mn toxicity
in turn led to the finding that this role of Si applies to other metal ions as well,
particularly Al. Aluminum (Al3+) is highly toxic at the elevated concentrations
in acid soils. For this reason, considerable research has lately been devoted
to the effects of this toxic metal on plants, and the role of Si in minimizing
or eliminating them. In many of these investigations, no decision could be
reached regarding the mechanism of the alleviation of toxicity by Si. Interaction
between Si and the metals, reducing the activity of toxic metal ions in the
medium, is one possible mechanism (67). The solubility of Si in the medium is
another. The concentration of Si in soil solutions is on the order of 0.1–0.6 mM
(46), and changes little over the entire range of physiological pH values. When
culture solutions with much higher concentrations are used, concerns arise about
polymerization and precipitation. The discussion will make clear, however, that
not only processes in the substrate, but others occurring within the plant, need
to be considered in connection with the interactions between Si and metals
such as Mn and Al. There may be considerable differences in the protection Si
provides against Al toxicity, as shown, for example, for the indica and japonica
ecotypes of rice (105).
Li et al (85) found cotton, Gossypium hirsutum, to be highly sensitive to
200 µM Al in solution, the effect being mitigated by additions of Si. Purely
external interactions as discussed above may have been responsible. External
effects also seem to have predominated in experiments in which Si dimin-
ished the toxicity of Al to soybean (10). The precipitation of subcolloidal, inert
hydroxyaluminosilicate species seems to have diminished the concentration
(activity) of phytotoxic Al in solution. From experiments with corn, Zea mays,
in which Si afforded protection against the inhibition of root elongation by Al,
the authors also concluded that the formation of Al-Si complexes was respon-
sible (88).
654 EPSTEIN
This kind of explanation based entirely on extraneous processes in the so-

lution is much less likely, however, in the experiments by Barceló et al with
teosinte, a wild form of Zea mays that grows in acid soils prone to high Al levels
(7). These authors found plants exposed to toxic Al concentrations (60 µM or
120 µM) to be less inhibited in their growth when Si at the low concentration
of 4 µM was included in the culture solution. The mechanism of this effect
seemed to be mainly a decrease of Al uptake by the plants due to Si. Silicon and
Al also changed the concentrations of organic acids in the plants; most signifi-
cant were the increased concentrations of malic and formic acids in the plants
grown in the presence of added Si and Al. The same group, working with an
Al-sensitive variety of corn, Zea mays var. BR 201 F, compared root elongation
of plants that had been pretreated with Si and then exposed to Al with that of
plants not pretreated with Si. The Si pretreatment was effective in counteract-
ing the inhibition of root elongation by Al (36). The temporal separation of
the Si and Al treatments notwithstanding, the Si pretreatment diminished the
absorption of Al by the plants. Since Al and Si were not present in the medium
simultaneously a purely external interaction between the two elements is an
unlikely explanation of the results. In wheat, too, the mitigation of Al toxicity
was shown to be due to internal, physiological processes (33). Hammond et al
(61) found Al toxicity in barley to be mitigated by Si. They concluded, surely
with ample justification, that the “overall Al/Si effect is likely to be complex,”
but drew attention to their finding that Si reversed the inhibitory effect of Al on
the absorption of Ca by the roots and Ca transport to the shoots.
Readers are referred to the paper by Hodson & Evans (68) and Cocker
et al (34) for a review of Al/Si interactions. The authors (68) tabulate inves-
tigations of no less than nine species in which Al toxicity as affected by Si has
been studied. In most of them, Si did indeed ameliorate various adverse effects
of Al. Mechanisms of this amelioration are discussed. In a subsequent paper,
Cocker et al (34) have proposed that low-solubility aluminosilicates or hydrox-
yaluminosilicates (or both) are formed within the root cell wall (apoplastic)
space, thereby reducing the concentration of free, toxic Al3+ ions. The pro-
tection that Si provides against Al toxicity is a lively topic not just in plant
physiology but in research on biota generally, including fish (51, 87).
Of the potentially toxic metals Al looms largest, because of its ubiquity
in soils and its heightened solubility under acid conditions such as prevail in
heavily weathered tropical soils. A second reason for emphasizing Al is that
forest decline due to anthropogenic soil acidification (58) has become a grave
problem. Barceló et al (8) have called Al phytotoxicity a “challenge for plant
scientists,” and so it is. These matters give an impetus for research on the
mechanisms of Si/Al interactions. Hodson & Evans (68) are surely justified in
their injunction: “Workers on Al toxicity must take Si into account.” The same
SILICON 655
comment applies to many contexts besides that of Al toxicity. The frequent

failure to heed this admonition represents a triumph of plant physiological
“ideology” (the adherence to a flawed definition of essentiality) over the facts
of life, and common sense. It has to be kept in mind that plants grow in a
medium containing Si, and absorb it: the +Si situation is the norm, or “control.”
Depriving them of the element amounts often to the gratuitous imposition of
an atypical environmental stress.
Salinity is another abiotic stress that has been shown in some investigations
to be mitigated by Si. Matoh et al (91) reported such an instance for rice grown
in solution culture with additions of either NaCl, seawater, or polyethylene gly-

col at concentrations up to the equivalent of 20% seawater. Silicate at 0.89 mM
reduced the translocation of Na+ to the shoots and increased dry matter pro-
duction of the stressed plants over that of the controls. For wheat (2) and barley
(86) similar findings have been reported, namely a repression of Na+ transport
in plants growing in salinized solutions supplied with Si, with concomitant
improvement in their growth, compared with that of plants in similar solutions
to which Si had not been added. The results reported for Prosopis juliflora, a
small leguminous tree, came from an experiment in which the plants grew in a
sand-peat-vermiculite mix (20). At the highest NaCl concentration (260 mM),
Si at 0.47 mM had a positive effect on plant growth.
Because salinity has both ionic and osmotic components the question arises
as to the role of Si in plant water relations, and vice versa. Because silicon
is present over the physiological pH range as uncharged silicic acid, H4SiO4,
and absorbed as such, the idea has surfaced time and again that it is absorbed
passively along with the water and may indeed serve as an index of water use
(46, 78, 106), but the idea has lost currency (46, 92). Nevertheless, transpiration
favors translocation of Si, and Si often accumulates at the terminal sites of the
transpiration stream, at leaf edges, in trichomes, specialized bulliform cells, and
other sites of intense transpiration, but exceptions have also been noted (114).
Conversely, effects of Si on water relations have also been reported (91, 116),
but the findings so far are contradictory and inconclusive. Nevertheless, the
osmotic effect of silicic acid, H4SiO4, its proneness to formation of opal, its
intimate association with the cell wall matrix (102), and the rigidity that it
bestows on cell walls once deposited as opal (46, 104, 106) all make it likely
that Si plays a role in plant water relations—a fertile field for investigation.
Reference has already been made to the physical strength that Si bestows on
cell walls. Some of these effects also fall under the present heading of abiotic
stresses, such as resistance to lodging, where the stress is gravity. Roots cope
with a stress shoots, except emerging ones, seldom do; roots have to overcome
the physical impedance offered by the solid matrix of the soil. It is possible
that Si plays a role here, but evidence is lacking.
656 EPSTEIN
BIOTIC STRESSES
The interaction between plants and their environment takes place mainly via
plant surfaces, both below and above the ground. That statement applies in full
force to the encounter between plants and disease and pest organisms. The cell
wall with its cuticle is the foremost defense to breach. Bacteria and fungi do
so by chemical means, whereas pests including phytophagous insects utilize
mechanical means mainly though not exclusively. From early on (78), the as-
sumption has often been made that it is the physical incrustation of cell walls
by opal phytoliths that is responsible for the frequently observed protection af-
forded plants by Si against diseases and pests, and no doubt it plays a role (16).
We shall see, however, that Si is involved in very complex, as yet incompletely
understood defense mechanisms. When, as is often done for control of the
medium, the plants are grown experimentally in conventional solution cultures
to which no Si has been added, the plants’ own physiological defense mecha-
nisms are set aside: The plants are to some extent “experimental artifacts” (46).
As usual, Jones & Handreck (78) furnish a starting point. They summarized
several instances of protection provided plants by Si against fungal and insect
attack. A more recent review (46) cites more, so that the present one is confined
to just a limited number.
In the experiments by Adatia & Besford (1) on the effects of Si on cucumber
plants grown in a recirculating nutrient solution, the plants in the low-Si cultures
were attacked by the powdery mildew fungus Sphaerotheca fuliginea despite
applications of fungicide, whereas the high-Si plants remained nearly free of
disease symptoms. Bélanger et al (11, 12) have shown the effectiveness of Si in
protecting greenhouse crops against disease, including spectacular pictures (12)
of leaves of cucumber grown in solution culture, and the total protection ob-
tained when Si was supplied. For grapevines, Vitis vinifera, Si applications
proved highly successful in protecting the plants against powdery mildew, a
major fungal disease in vineyards (18, 107).
Cucumber roots are subject to attack by fungi of the genus Pythium, and their
shoots by Sphaerotheca fulginea (95, 96). Chérif et al (28, 29) examined the
effects of Si added to nutrient solutions at 1.7 mM on the responses of the plants
to infection by Pythium spp. Silicon in the infected plants elicited resistance to
fungal attack and the appearance of precursors of fungitoxic aglycones acting
possibly like phytoalexins. Other potentially protective metobolic processes and
inhibitory substances were identified in the infected, Si supplied plants. Very re-
cently, Fawe et al (52) identified the active, Si-induced agent protecting cucum-
ber against attacks by powdery mildew as a phytoalexin. These investigations
clearly show that Si elicits a veritable cascade of biochemical defense mecha-
nisms in the infected plants. Reviewing these and other findings, these authors
(52a) relate Si to one type of induced disease resistance, specifically “systemic
SILICON 657
acquired resistance (SAR).” It is characterized by accumulation of salicylic acid

and pathogenesis-related (PR) proteins. This response is typically elicited by
pathogens, but their effect can be mimicked by “activators,” of which Si is one
(52a, 117). The defense mechanisms mobilized by Si include the accumulation
of lignin, and generally, phenolic compounds, as well as chitinases and peroxi-
dases. Fawe et al (52a) make an impressive case for the significance of Si in plant
defense mechanisms against disease. Perhaps Si acts as a second messenger.
Monocots are no less immune to diseases than dicots such as cucumber.

Indeed, field experience about the role of Si in plant protection against disease
is extensive; only a few recent instances can be given here. Winslow (142)
found that rice grown on Si-poor, upland soils in West Africa had a shoot Si
content much below that recommended for rice. Silicon fertilization gave a
large increase in Si content and yield, and significantly reduced the severity of
fungal diseases. The responses varied among genotypes.
Rice grown under entirely different conditions, namely, on low-Si organic
Histosols in Florida, responded similarly: Incidence of disease was negatively
correlated with Si content in comparisons of plots receiving, with those not
receiving Si applications (39). Genotypic variations were noted in this investi-
gation as well. For a comprehensive review readers are referred to Savant et al
(116), and for a recent investigation of control by Si of rice blast, accompanied
by a comprehensive bibliography, to Seebold (118). Latin American instances
are discussed by Vitti et al (131). Silicon provides protection not only against
disease organisms but herbivores as well, be they phytophagous insects (111)
or mammals (57, 94).
As in the dicots discussed above, in monocots, too, there is a complex in-
terplay between Si (or Si deprivation) and biochemistry. Thus in oat, Avena
sativa, Carver et al (26) found that Si deprivation resulted in greater accu-
mulation of phenolic compounds in epidermal cells under attack by Blumeria
graminis, a pathogenic fungus. The activity of phenylalanine ammonia-lyase
was also enhanced by Si deprivation, even in uninocculated plants. Under Si
deprivation, the production of structural phenolic compounds may have substi-
tuted for the silicification that normally contributes to the structural strength of
cell walls. These response mechanisms are by no means uniform in different
species; readers are referred to the paper by Carver et al (26) and its references.
BIOCHEMISTRY
With apologies to Winston Churchill, the biochemistry of Si is a riddle wrapped
in a mystery inside an enigma. When Jones & Handreck (78) published their
groundbreaking review in 1967, Si biochemistry was one of the few relevant
topics they did not include: There was no evidence to discuss. And when Carpita
(25) recently referred to the incorporation of Si into cell walls of grasses, he
658 EPSTEIN
wrote, “Little has been reported on any chemical interaction with other cell wall
constituents.” The amorphous nature of cell wall silica (90, 147) might support
the idea that there is no biochemistry involved.
Various attempts to demonstrate the occurrence of organo-silicon compounds,
reported by Peggs & Bowen (100) were unsuccessful, including their own, in
which they studied Equisetum arvense (horsetail) and Thuja plicata (western
red cedar, Cupressaceae).
Nevertheless, the evidence discussed above on the role of Si in the response

of plants to pathogens indicates that Si is not biochemically inert. Because
of its prominent presence in the cell wall, Si is a candidate for association

with cell wall components and hence, with polysaccharides (19), and with cell
wall–associated proteins.
Inanaga et al (73, 74) have presented evidence that in rice, Si may be involved
in Si-aromatic ring associations between lignin and carbohydrate. In Phalaris
canariensis (canary grass) and Equisetum telmateia, low levels of proteins are
trapped inside silica bodies, along with monosaccharides (62). In the cell wall
of the diatom Cylindrotheca fusiformis, a 200-kDa protein is associated with a
specific structure of the silica scaffold (83). In this case, trapping of the protein
within the cell wall silica was specifically ruled out: The protein is located at
the surface of the silica. These workers cloned a family of cDNAs that code
for these silica-associated proteins.
Further demonstrating an intimate connection between proteins and Si,
Shimizu et al (119) recently identified three proteins that seem to control biosili-
cification in the sponge (Porifera) Tethya aurantia. They called these proteins
silicateins (for silica proteins); they belong to a single protein family. The most
abundant of these proteins, silicatein α, belongs to the cathepsin L class of
the papain-like cysteine protease superfamily. The biosilicification in these
sponges may therefore be mediated through the enzymatic activity of these
silicateins. Cha et al (26a) in subsequent in vitro experiments have extended
these exciting findings and strengthened that conclusion.
In the marine diatom Cylindrotheca fusiformis, Hildebrand et al (64) isolated
Si-responsive cDNA clones derived from Si-responsive mRNAs. One possible
role for such cDNAs might be that of encoding Si transport proteins. Indeed,
this group, associated with the late “silicophile,” BE Volcani, has recently cloned
a family of genes that code for Si transport proteins (65a). Perhaps the new
type of hydrogen bond recently discussed by Crabtree (37) may be involved in
complexes between Si and cell wall components (carbohydrates, proteins) and
possibly other Si biochemistry, but that, for now, is speculation.
These recent exciting findings from organisms other than higher plants, and
from chemistry, should spur plant physiologists, biochemists, and molecular
biologists to probe deeply into the corresponding features, metabolites, and
processes in higher plants.
SILICON 659
CONCLUSION
This survey has covered a great deal more ground than is customary in annual
reviews of very actively pursued topics in plant physiology and plant molecular
biology. As shown early on, the ubiquitous and quantitatively prominent ele-
ment, Si, has not received the attention it requires, in view of the significance of
it in plant life. It has thus been necessary to cast the net rather wide and bring
together not just strictly plant physiological evidence but to include experience
gathered by botanists, agronomists, horticulturists, plant pathologists, and still

others. There is no justification for the disregard of Si in so much of the science

of plant physiology. The evidence is overwhelming that in the real world of
plant life, Si matters.
ACKNOWLEDGMENTS
I thank members of the Davis campus for discussion: RG Burau, WH Casey,
RA Dahlgren, RJ Southard, and RJ Zasoski, all of my own department. D
Potter of the Department of Pomology assisted in the interpretation of Figure 2,
and JE Hill of the Department of Agronomy and Range Science advised me
on rice ecotypes. RR Bélanger made available unpublished material and gave
encouragement and counsel. LE Datnoff, MJ Hodson, and P Hinsinger did
likewise. J Ma directed me to material published in Japanese and helped me
with the interpretation of Figure 2, as did M Kobayashi. E Malavolta sent me
material from Brazil. None of these colleagues bears any responsibility for
shortcomings of this review. The secretarial staff’s competence and devotion
are deeply appreciated. I thank Peggy for putting up with one more of these
distractions. My own research on Si was supported by the Department of
Energy. Finally, I express my appreciation to Annual Reviews for inviting me
to write this, my third contribution to this series.
Visit the Annual Reviews home page at

http://www.AnnualReviews.org
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Annual Review of Plant Physiology and Plant Molecular Biology
Volume 50, 1999
CONTENTS
Educator and Editor, Martin Gibbs 1
Phosphate Translocators in Plastids, Ulf-Ingo Flügge 27
The 1-Deoxy-D-xylulose-5-phosphate Pathway of Isoprenoid Biosynthesis
47
in Plants, Hartmut K. Lichtenthaler
Chlorophyll Degradation, Philippe Matile, Stefan Hörtensteiner, Howard
67
Thomas
Plant Protein Serine/Threonine Kinases: Classification and Functions, D.
97
G. Hardie
Improving the Nutrient Composition of Plants to Enhance Human
133
Nutrition and Health, Michael A. Grusak, Dean DellaPenna
Gametophyte Development in Ferns, Jo Ann Banks 163
C4 Gene Expression, Jen Sheen 187

Genetic Analysis of Hormone Signaling, Peter McCourt 219
Cellulose Biosynthesis: Exciting Times for a Difficult Field of Study,
245
Deborah P. Delmer
Nitrate Reductase Structure, Function, and Regulation: Bridging the Gap
277
Between Biochemistry and Physiology, Wilbur H. Campbell
Crassulacean Acid Metabolism: Molecular Genetics, John C. Cushman,
305
Hans J. Bohnert
Photoprotection Revisited: Genetic and Molecular Approaches, Krishna
333
K. Niyogi
Molecular and Cellular Aspects of the Arbuscular Mycorrhizal Symbiosis,
361
Maria J. Harrison
Enzymes and Other Agents that Enhance Cell Wall Extensibility, Daniel
391
J. Cosgrove
Leaf Development in Angiosperms, Neelima Sinha 419
The Pressure Probe: A Versatile Tool in Plant Cell Physiology, A. Deri
447
Tomos, Roger A. Leigh
The Shikimate Pathway, Klaus M. Herrmann, Lisa M. Weaver 473
Asymmetric Cell Division in Plants, Ben Scheres, Philip N. Benfey\ 505
CO2-Concentrating Mechanisms in Photosynthetic Microorganisms,
539
Aaron Kaplan, Leonora Reinhold
Plant Cold Acclimation: Freezing Tolerance Genes and Regulatory
571
Mechanisms, Michael F. Thomashow
The Water-Water Cycle in Chloroplasts: Scavenging of Active Oxygens
601
and Dissipation of Excess Photons, Kozi Asada
Silicon, Emanuel Epstein 641
Phosphate Acquisition, K. G. Raghothama 665
Roots in Soil: Unearthing the Complexities of Roots and Their
695
Rhizospheres, Margaret E. McCully

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P1: PSA/spd P2: PSA/ary QC: KKK/anil T1: KKK

March 30, 1999 15:30 Annual Reviews AR082-23

Annu. Rev. Plant Physiol. Plant Mol. Biol. 1999. 50:641–64

Department of Land, Air and Water Resources—Soils and Biogeochemistry,

KEY WORDS: essentiality, growth, development, mechanical strength, stress

SILICON AND GROWTH AND DEVELOPMENT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 650

Silicon is a quantitatively major inorganic constituent of higher plants but ab-

“SILICON DEFICIENCY” IN PLANT PHYSIOLOGY

evidence regarding the protection afforded by Si against the damage wrought

That difficulty, at the experimental level, changed with the introduction of

dispensable elements in the same sense as K, P, etc. He did, however, comment

The conclusion of Sachs as to the dispensability of Si for plant life was

to be made nevertheless. They have to do with (a) the limitations of purging

and if a deficiency of it can be severe enough to result in demonstrable ad-

SILICON IN PLANTS: THE AMOUNTS

MM Rafi & E Epstein (unpublished results). However, because it is in the form

SILICON AND GROWTH AND DEVELOPMENT

As for other species, the depletion of plant-available Si commented on above

investigation (P Hinsinger, personal communication). Application of rock dust

This kind of explanation based entirely on extraneous processes in the so-

comment applies to many contexts besides that of Al toxicity. The frequent

in solution culture with additions of either NaCl, seawater, or polyethylene gly-

acquired resistance (SAR).” It is characterized by accumulation of salicylic acid

Monocots are no less immune to diseases than dicots such as cucumber.

Nevertheless, the evidence discussed above on the role of Si in the response

of its prominent presence in the cell wall, Si is a candidate for association

gathered by botanists, agronomists, horticulturists, plant pathologists, and still

others. There is no justification for the disregard of Si in so much of the science

Visit the Annual Reviews home page at

10. Baylis AD, Gragopoulou C, Davidson Physiol. 29:526–32

toxicity by silicon in higher plants: so- ganic solution to bioorganic essentiality.

drogen bond. Science 282:2000–1 tance?

W, Schroeder JI. 1997. A gene family of ganic Chemistry: Inorganic Elements in

biological systems. J. Chem. Soc. Chem. privation causes physical abnormalities

C4 Gene Expression, Jen Sheen 187

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