P l a n t a n d Soil X X I I I , no.

STUDIES

A u g u s t 1965

OF SILICA IN THE OAT PLANT

III. UPTAKE OF SILICA FROM SOILS BY THE PLANT

by L. H. P. J O N E S and K. A. H A N D R E C K
C.S.I.R.O., Division of Plant Industry, c/o SchooI of Agrieulture,
University of Melbourne, Australia

INTRODUCTION

It is known that plants take up different amounts and different

proportions of silicon from a soll according to their species. Thus,
the proportion of silica in gramineous species is of the order of ten
to twenty times that found in leguminous species 4 24. Not only do
species differ in uptake but plants of the one species (e.g. rice s)
absorb different amounts of silica when grown in different soils. The
proportions of silica in both gramineous and leguminous species are
decreased on raising the pH of the soil 6, and we have noted that
oats grown in different solls of the same pH contain, at maturity,
different proportions of siliea. We present here further evidence of
how the soil determines the uptake of silica b y plants.
An understanding of the chemistry of silicon in the soil solution
is of importance because the soll solution is the immediate source
of silicon for plants. However, apart from a reference 24 to work
earried out 100 years ago in which soil solutions were obtained b y
displacement with water, there are no reports concerned speciIically
with silicon in the soll solution. Other data 17 is 19 recorded for
'si]icon in soil solutions' have been obtained with aqueous extracts
of soils; this is a misuse of the term 'soil solution'. The data from
aqueous extracts cannot be directly related to the uptake of silica
b y plants and it is even diIficult to relate one extract to another
beeause the results depend on the ratio of water to soiI and the
duration of shaking. Throughout our investigations a pressure celt 22
has been used to obtain solutions from soils maintained at field
--

79

--

B0

L. H. P. J O N E S AND K. A. H A N D R E C K

capacity. By using the reduced silico-molybdate method of analysis

under prescribed conditions (see page 8) we have found that all the
silicon in the soil solution is present as monosilicic acid. This is
consistent with the Iact that silica passes into solution as monosilicic
acid at pH's below 9 and as silicate ions at pH's above 9 2 7 15. Since
monosilicic acid may be regarded as a hydrated form of SiO2,
presumably Sf(OH)4, the terms monosilicic acid and silica in solution
will hereafter be used synonomously.
The eoncentration of silica in solutions obtained from a wide
range of potted soils has been found to differ, but never to exceed
B0 ppm calculated as Sf02. One field soil was available at various
pH's in the range 5.4 to 7.2 and the concentration of sitica in solution
was found to decrease from 70 to 23 ppm SiO~ with increasing pH.
These figures contrast with the solubility of amorphous silica in
water which is of the order of 120 ppm Sf02 at 25C and which is
independent of pH over the range I to 9 2 7 15. The present studies
deal firstly with the reasons for the differences between soils and
for the undersaturation of soil solutions and secondly with the
relation between silica in the soil solution and its uptake by oats.
Although these two topics were studied concurrently and with
identical soils they will be presented under separate headings.
SILICA IN THE SOIL SOLUTION

The mechanisms responsible for the different concentrations of

silica in solution in solls and for the undersaturation oB soil solutions
are complex. There are, however, several studies which suggest that
the solubility of silica is influenced by aluminium when added as a
salt 2o or as an oxide 9. These studies led to the present work on the
effeets of fron and aluminium oxides on the solubility of silica in
both simple systems and in soils. The work with simple systems has
been reported elsewhere 10 but a brief account of it is presented
here because it is basic to this paper.

Simple systems
The simple systems were designed to study the sorption of
monosilicic acid by fron and aluminium oxides in suspensions with
pH's ranging from 1 to 12. The procedure was to shake 100 mg
samples of oxide with 100 ml of solutions of monosilicie aeid con-

S T U D I E S OF S I L I C A I N T H E OAT P L A N T

8[

taining 120 ppm Si02 and to determine the silica remaining in

solution at equilibrium. Three samples of ferric oxide were used; all
contained haematite but they ranged from practically amorphous
to poorly crystalline. Four samples of aluminium oxide were used;
all contained boehmite but they ranged from practically amorphous
to moderately well crystalline.
All oxide samples removed monosilicic acid from solution and
their adsorption curves had certain features in common. Each
sample showed a maximum adsorption close to pH 10, above which
silicate ion is formed and the solubility of silica increased sharply.
High adsorption was confined to the range of pH 4 to 10 and as pH
decreased within this range smaller amounts of monosilicic acid
were adsorbed. Below pH 4 the amounts adsorbed decreased sharply
until at about p H 2 all the monosilicic acid remained in solution.
Among the least crystalline oxides aluminium oxides were more
efIective, weight for weight, in adsorbing monosilicic acid than
ferric oxides. Thus, at pH 6 an aluminium oxide adsorbed 9.1 mg
Si02 per 100 mg and a Ierric oxide 4.0 mg SiO2 per 100 mg. The most
crystalline aluminium oxide adsorbed only one third as much as the
least crystalline whereas the ferric oxides did not differ greatly
amongst themselves in their ability to adsorb monosilicic aeid.

Soil systems
The effects of ferric and aluminium oxides on silica in the soil
solution were studied using two groups of potted soits. In the first
group, orte of the terric oxides and two of the aluminium oxides
were added to the soll from the experimental area of the School of
Agriculture, University of Melbourne. This soil is a sandy loam
containing 1.76 per cent organic carbon * and 2.1 per cent free ferric
oxide **; its mechanical, chemical, and mineralogical compositions
are shown in Tables 1,2, and 3. The ferric oxide (I) was a commercial
product from B.D.H. Ltd., England, which consisted of poorly
crystalline haematite. One of the aluminium oxides (III) was
prepared b y the method of S c h u y l e n b o r g h aa but in its preparation the final pH was not allowed to exceed 7.2; it was practically
* Determined by the method of W a l k l e y and B l a c k ( P i p e r ~l) and expresse d as
per cent by weight of the air-dry soll.
** E x t r a c t e d by the method of A g u i l e r a and J a c k s o n a; the results of analyses are
expressed as per cent by weight of the air-dry soll.

82

L. H. P. JONES

AND

K. A. HANDRECK

amorphous to X-rays but had an incipient boehmite structure. The

other sample (IV) was a chromatographic aluminium oxide (Brockmann) manufactured b y E. Merck, Germany; it consisted of moderately well crystalline boehmite. The oxides were ground to pass a
200-mesh I.M.M. sieve and were uniformly mixed with separate
samples of the University soff at the rate of 5 per cent b y weight of
the air-dry soil. These mixtures and the control soil were weighed
into polyethylene pots. All pots of soff had their pH's adjusted to
5.9 4- 0.1 ; all were brought to pF 2.0 b y adding distilled water to
make 18 per cent b y weight and maintained with this water content
for 6 months. The pH was determined from time to time and
remained at 5.9 4- 0.2.
TABLE 1
Mechanical composition of soil (%) *
Soll

Coarse sand
(0.2-2 mm)

University...
Penola . . . .
Wollongbar . ,

33.0
1.4
0.6

Fine sand
(0.02-0.2 mm)

I
i

39.2
27.5
16.8

Silt
m
Clay
(0.002-0.02 m ) (< 0.002 mm)

15.5
12.0
22.3

I
1

12.3
59.1
60,3

* Recalculated to a basis of sand + silt + clay = 100.

At various times samples were temporarily removed for studies of

silica in the soil solutions. In order to simulate the system of a soll
in which the solution is continually depleted by plants and replenished from the solid phase, the following procedure was followed.
A sample of 354 g soil (300 g oven-dry soff) was placed in a pressure
cell sealed at the bottom with a Visking cellulose membrane. A gas
pressure, applied for 9 hours, was adjusted so as to bring the soil to
pF 2.6 thereby removing 30 to 32 per cent of the soil's water. (This
loss is of the same order as the maximum daily loss due to transpiration from the soil-plant systems). At the end of the 9-hour period
the soil was removed from the pressure cell, rewetted with distilled
water equivalent in volume to that removed, mixed, and allowed
to stand in a sealed polyethylene bag for 15 hours. A further sample
of soil solution was obtained during the next 9 hours and this cycle
was repeated for eight weeks. The concentration of silica decreased
progressively in the first few cycles and levelled off b y the sixth
cycle to a steady coneentration characteristic of the particular soil.
Throughout our work the concentration of silica in the soil solution

STUDIES

83

OF SILICA IN THE OAT PLANT

is defined as this steady concentration which the soil can maintain

in a cycle consisting of alternate removal of solution and adding of
water to bring the soil back to field capacity.
TABLE 2
S i l i c a - s e s q u i o x i d e r e l a t i o n s of c l a y f r a c t i o n s
Per cent by weight
Soil

University . . .
Penola.
Wollongbar...

MoIecular ratio

SiO2

A1203

Fe20a

51.97
48.25
22.32

20.41
17.26
30.39

7.56
7.03
22.50

SiO~o

SiO~

A120a

R208

4.3
4.7
1.2

3.5
3.7
0.8

During the six-month period the concentration of silica in solution from the control soll remained constant at 48 ppm SiOs.
Ferric oxide I lowered the concentration to 27 ppm at the beginning
of the period but during the six months this increased to a constant
value of 40 ppm. The aluminium oxides had a more striking effect,
thus Sample I I I lowered the concentration at the beginning of the
period to 7 ppm Si02 and Sample IV to 12 ppm. During the six
months these concentrations increased only slightly to constant
values of 9 and 15 ppm respectively.
TABLE 3
M i n e r a l c o m p o s i t i o n of solls (per c e n t b y w e i g h t )
C o n s t i t u e n t s **
Soil

Fraction*

University

Fine sand
Silt
Clay

Penola

Fine sand
Silt
Clay

Wollongbar Fine sand

Silt
Clay

Gibbsite

Boehmite

Cxoethite

HaeMagmatite netite
1

Ilmenite

I~aolin

30
10

20
33

11
< 10

Felspar

Opal
Quartz phytoliths

<1
tr.

88
> 90

4
n.d.

35

2
tr.
65

<1
< 10
fr.

> 90
> 90
<10

7
n.d.

40

0.1
tr.
50

14
10

0.1
n.d.

0.2

23
40
33

Mica

22
I0
33

* F i n e S a n d , 0.02 to 0.2 m m ; Silt, 0 . 0 0 2 to 0.02 m m ; C l a y < 0 . 0 0 2 m m .

** T h e c o m p o s i t i o n of t h e fine s a n d f r a c t i o n s w a s d e t e r m i n e d f r o m a c o u n t of n o t less t h a n 1000 g r a i n s . T h e
c o m p o s i t i o n of b o t h t h e silt a n d c l a y f r a c t i o n s w a s d e t e r m i n e d b y X - r a y d i f f r a c t i o n ; t h e q u a n t i t a t i v e e s t i m a t e s
re 4- 1 0 % .
tr. - t r a c e ; n . d . - n o t d e t e r m i n e d .

84

L. H. P. J O N E S A N D

K. A. H A N D R E C K

The second group of potted soils comprised: the University soil

without added sesquioxides; a black clay from a reclaimed swamp
at Penola in the south-east of South Australia, which contains 5.58
per cent organic carbon and 1.4 per cent free ferric oxide; and a
krasnozem from Wollongbar near Lismore in New South Wales,
which contains 4.94 per cent organic carbon and 25.6 per cent free
ferric oxide. The Penola and Wollongbar soils were chosen because,
although they contain similarly large amounts of clay, they have
widely different amounts of free sesquioxides; their mechanical,
chemical and mineralogical compositions are shown in Tables I, 2,
and 3.
The three soils were weighed into po]yethylene pots and had
their pH's adjusted to 5.6; additional pots of the University soil
were adjusted to pH 6.8. All were brought to pF 2.0 by adding
distilled water; at this pF the Penola and Wollongbar soils contained
54 and 52 per cent water respectively. The pots were maintained
with these water contents for six months during which time samples
were temporarily removed, for studies of silica in the soil solutions.
The method used was the same as that described above except that
the gas pressure which was applied for 9 hours was adjusted to bring
the Penola and Wollongbar soils to pF 3.0, thereby removing 17 per
cent of the water from each of the soils.
The concentrations of silica in solution in these soils are shown in
Table 4. Samples of the University soil were also available from the
field with pH's ranging from 5.5 to 7.2 and the concentrations of
silica in solutions obtained from them are shown in Fig. 1, together
with the results obtained with potted University soils. The levels of
silica in solutions in the first cycle from the various samples of the
University soll were 23 to 36 per cent higher than the steady concentrations, whereas with the Penola and Wollongbar soils the levels
were 7 and t4 per cent higher respectively.
TABLE 4
T h e c o n c e n t r a t i o n of siliea in soiI s o l u t i o n s ( p p m SiO2)
Soll
University
University
Penola
, Wollongbar . . . . . . . . . .

Soil p H

Initial
concentration

Steady
concentration

5.6
6.8
5.6
5.6

72
37
72
8

54
gO
67
7

STUDIES OF SILICA IN THE OAT PLANT

'-"

85

8O

0
o

initial

concn.

stady

concn.

d.

cL

60

\ ~

\.

0
o
03

40

0
uq
C

CJ

2O

0
5"0

I
6-0

I
7.0

Soil

8.0

pH

Fig. 1. T h e r e l a t i o n b e t w e e n soil p H a n d t h e c o n c e n t r a t i o n of silica in s o l u t i o n

in t h e U n i v e r s i t y soff.

Discussion

The studies with simple systems show that both ferric and
aluminium oxides adsorb monosilicic acid from solution. The
mechanism of adsorption is obscure, whether through the formation
of a salt as suggested b y I l e r 7 (page 69) or whether through
hydrogen bonding, which is more plausible. Any explanation taust
take aceount of our observations that pH decreased during equilibration of an oxide with monosilicic acid and that adsorbed monosilicic acid m a y be desorbed on decreasing the pH of a suspension.
Out results with simple systems are in agreement with those of
two other groups 5 17 is who have recently found that monosilicic
acid is adsorbed on the surfaces of ferric and aluminium oxides.
Like ourselves, they found that adsorption depended on pH.

86

L. H. P. JONES AND K. A. HANDRECK

However, further comparisons with our work are difficult because

the other groups studied suspensions in which the ratios of solid to
solution were so high that different adsorbing capacities amongst
their oxides would be masked.
The present studies show that the concentration of silica in the
soil solution decreases with increasing pH or with increasing amounts
of sesquioxides, the more so as these oxides become more amorphous.
The contrast between the Penola and Wollongbar soils is especially
striking. One cannot be sure whether the low figure for Wollongbar
is due to its iron or aluminium oxides (Tables 2 and 3) but the earlier
work on simple systems suggests that aluminium is especially
responsible.
It appears that soils are capable of maintaining constant concentrations of monosilicic acid in solution despite repeated withdrawals.
This implies that monosilicic acid is released fairly rapidly from the
various solid sources in the soil. It has been suggested that much of
the monosilicic acid is derived from adsorbing surfaces ~. I n the
present studies the concentrations of silica in the first cycles were
always higher than the steady concentrations and it is reasonable
to assume that the excess silica is derived from loosely adsorbed
monosilicie acid of which there would be a limited amount. After
this has been removed it is probable that adsorbing surfaces continue
to be the main immediate source of monosilicic acid for the soil
solution but these surfaces must be continually replenished from
other sources. In the present studies most of the replenishment
would take place during the 15-hour equilibration periods, the
monosilicic acid coming from dissolution of the clay minerals and
both crystalline and amorphous forms of silica. (Amorphous silica
would fange from colloidal silica to opal particles of plant origin i.e.
phytoliths 1~ IS). Of these sources the amorphous silica releases
monosilicic acid at the greatest rate 11 and may be considered the
most important primary source of silica for the soil solution.
The Penola soil has a high concentration of opal phytoliths in its
fine sand fraction (Tables 1 and 3) and presumably a high concentration of colloidal silica. Because of this and the apparent absence
of free sesquioxides it is difficult to explain the lowering of the level
of silica in solution to about 70 ppm Si02 eren when monosilicic acid
is not being removed. This may be due to an additional equilibrium
involving a combination of aluminium with colloidal silica. In

STUDIES OF SILICA IN THE OAT PLANT

87

support of this it may be noted that a trace amount of aluminium

decreases the rate of depolymerization of colloidal silica 20 and that
adsorbed cations, notably aluminium, decrease the rate of dissolution of amorphous silica from diatoms 16
UPTAKE OF SILICA BY PLANTS

The uptake of silica by oats was studied in two pot experiments

conducted in successive years. These experiments were carried out
concurrently with the work on soil solutions using the same two
groups of soils. The first experiment compares the effects of fron
and aluminium oxides on silica in the soil solution and its uptake by
plants; the succeeding experiment goes further and relates the
uptake of silica to its concentration in the soil solution and the
amount of water transpired.
' A l g e r i a n ' oats were g r o w n in b o t h p o t e x p e r i m e n t s . T h e e u l t u r e m e t h o d s
were d i f f e r e n t for t h e t w o years a n d will b e d e s c r i b e d u n d e r s e p a r a t e h e a d i n g s
b u t t h e a n a l y t i c a l m e t h o d s were c o m m o n t o b o t h e x p e r i m e n t s a n d were as
follows.
(i) S i l i c a i n s o l u t i o n
i n p l a n t s . - A s u s p e n s i o n was s e p a r a t e d f r o m
h e s h p l a n t m a t e r i a l b y m e a n s of a s c r e w - t y p e juice e x t r a c t o r a n d t h i s
s u s p e n s i o n was c e n t r i f u g e d a t 20,000 g for 1 h o u r t o s e p a r a t e solid m a t e r i a l .
T h e l i q u i d o b t a i n e d b y t h i s m e t h o d is p r o b a b l y r e p r e s e n t a t i v e of t h e l i q u i d
p h a s e oI t h e p l a n t tissue since microscopic e x a m i n a t i o n of t h e c r u s h e d p l a n t
m a t e r i a l r e v e a l e d t h a t m o s t of t h e p l a n t eells were r u p t u r e d . T h e c e n t r i f u g a t e was d i l u t e d w i t h w a t e r a n d a n a l i q u o t t h e n h e a t e d in a t e I l o n b e a k e r
w i t h c h l o r i n e w a t e t t o d e s t r o y organic m a t t e r . A f t e r excess c h l o r i n e h a d b e e n
e x p e l l e d on a w a t e r - b a t h t h e l i q u i d was cooled a n d filtered i n t o a s t a n d a r d
Ilask f r o m w h i c h a n a l i q u o t was t a k e n for t h e e s t i m a t i o n of silica.
(ii) T o t a l s i l i c a i n p l a n t s . - T h e d r y p l a n t m a t e r i a l (tops) was g r o u n d
in a S e r v a l l ' O m n i - m i x e r ' . A s u b - s a m p l e was t h e n dried a t 105C Ior 15 hours,
t r a n s f e r r e d t o a p l a t i n u m d i s h a n d i g n i t e d in a n electrie m u f f l e furnace. T h e
t e m p e r a t u r e of t h e f u r n a c e was r a i s e d f r o m 200C t o 500C o v e r a p e r i o d of 6
hours, t h e n h e l d a t 500C for 10 hours. A n h y d r o u s s o d i u m c a r b o n a t e (E.
Merck, G e r m a n y ) was m i x e d w i t h t h e a s h a n d t h e m i x t u r e fused o v e r a M e k e r
b u r n e r for hour. T h e cooled c a k e was dissolved in distilled vater a n d m a d e
u p to a s t a n d a r d v o l u m e ; a n a l i q u o t was t h e n a n a l y s e d for siliea.
R o o t s were t a k e n for a n a l y s i s f r o m " n a t u r e p l a n t s a n d a l t h o u g h t h e y were
r e p e a t e d l y w a s h e d it was i m p o s s i b l e t o c o m p l e t e l y r e m o v e c o n t a m i n a t i n g
p a r t i c l e s of soil. W e h a v e n e v e r t h e l e s s f o u n d t h a t t h e a m o u n t s of silica i n
t h e r o o t s were less t h a n 2 p e r c e n t of t h e silica in t h e w h o l e p l a n t . T h e s a m e
r e l a t i o n b e t w e e n siliea in t h e roots a n d t h e w h o l e p l a n t h a s also b e e n f o u n d
i n o a t s g r o w n in w a t e r c u l t u r e s c o n t a i n i n g silicic aeid 11. W e h a v e a c c o r d i n g l y

B~

L. H. P. JONES AND K. A. HANDRECK

ignored silica in roots in the succeeding calculations and discussion.

(iii) C h e m i c a l a n a l y s i s . - Silica was estimated b y the reduced silic o m o l y b d a t e m e t h o d 14 aa under the following conditions. A suitable aliquot
(containing less t h a n 40#g SiO2) was transferred to a 25 ml s t a n d a r d flask.
Distilled w a t e r was added to bring the v o l u m e to a p p r o x i m a t e l y 15 ml, and
2 ml of a solution containing 7% (V~r/V) sodium m o l y b d a t e in N hydrochloric
acid was t h e n added. The reaction of silica w i t h m o l y b d a t e was 'Irozen' b y
adding 5.5 ml of 10 N hydrochloric acid 26 at either 2 or 10 minutes after
adding the m o l y b d a t e 1. The silicomolybdic acid was t h e n reduced w i t h 0.5
ml of a solution containing 0 . 2 % 1 : 2 : 4 a m i n o n a p h t o l s u l p h o n i c acid, 2.4%
sodium sulphite and 12% sodium metabisulphite. Ten minutes after adding
the reducing solution the absorption was read at 670 m/z on either a Hilger
' S p e k k e r ' A b s o r p t i o m e t e r or a U n i c a m S p e c t r o p h o t o m e t e r .
B y allowing t h e m o l y b d a t e to react for 10 minutes it was possible to
analyse for monosilicic and disilicic acids exclusive of higher polymers and
colloidal silica. In order to analyse specifically for monosilicie acid the
reaction was 'frozen' at 2 minutes 1

Pot experiments, I96r. The eMects o/ /erric and aluminium oxides on

uptake
The soll systems comprised the U n i v e r s i t y soll w i t h no added oxide (i.e.
control soll), and three lots of the Universi~y soll separately m i x e d with 5 per
cent b y weight of the following oxides: ferric oxide I, a l u m i n i u m oxide I I I
and a l u m i n i u m oxide IV. Samples of the control soil and the three soll-oxide
m i x t u r e s (hereaffer simply called solls) were weighed into polyethylene pots
it1 7.00 kg lots. There were sixteen pots of the control soll and six pots of each
of t h e other soils. The p H ' s of all pots were b r o u g h t to 5.9 ~ 0.1 w i t h sulphuric acid. A t the same t i m e each p o t had m i x e d into it t h e following:
KH2PO4, 0.26 g; KNO~, 1.03 g; MgSO4.7t-I20, 0.25 g; CuSO4.5H20, 0.023g;
Z n S O 4 . 7 H 2 0 , 0.023 g; M n S O 4 . 4 H 2 0 , 0.23 g. All pots were housed under a
glass c a n o p y in a birdproof erlclosure in the grounds of the School of Agriculture, U n i v e r s i t y of Melbourne.
The pots were watered to p F 2.0 and t w e n t y - f o u r seeds were sown in six
pots of eaeh of the Iour soils in m i d - A u g u s t ; the n u m b e r of seedlings was
reduced to t w e n t y soon after emergence. In addition to the pots w i t h plants
there were ten pots of the control soil in which plants were not grown. Six
of these 'fallow' pots and the six control pots in which plants were grown were
mulched w i t h opaque white p o l y e t h y l e n e pellets (approx. 2 2 5 m m ) ;
these were applied at the rate of 750 g per pot to give a layer 3.2 cm deep on
the surface of the soil. The remaining four 'fallow' pots of ttle control soil were
w i t h o u t a mulch.
T h r o u g h o u t the growing season all pots were weighed daily at a b o u t 5 p.m.
and brought back to p F 2.0 w i t h distilled watet. Plants were h a r v e s t e d for
analysis at six stages during the growing season. F o u r plants were t a k e n
from each pot at t h e first two harvests and three from each p o t at each of the
sueeeeding four harvests. N o r m a l l y at a h a r v e s t all t h e p l a n t m a t e r i a l from

S T U D I E S OF SILICA IN T H E OAT P L A N T

89

t h e o n e soll w a s b u l k e d , w a s h e d , d r i e d a t 105C, w e i g h e d , a n d g r o u n d ; s u b s a m p l e s w e r e t h e n t a k e n f o r a n a l y s i s o f t o t a l silica. H o w e v e r , a t t h e f i r s t a n d

f i I t h h a r v e s t s , p a r t of t h e p l a n t m a t e r i a l f r o m e a c h of t h e f o u r s o i l s w a s u s e d
i m m e d i a t e l y f o r t h e e s t i m a t i o n of s i l i c a in s o l u t i o n i n t h e p l a n t ; t h e r e s t of
t h e m a t e r i a l w a s a n a l y s e d for t o t a l silica.

The plants growing in the soil containing aluminium oxide III

showed symptoms of phosphate deficiency but were kept healthy
b y spraying frequently with 0.1% CaH4(P04)2. Those growing in
the soff containing aluminium oxide IV developed slight symptoms
soon after tillering but required only two sprayings with the solution
of CaH4(PO4)2.
Differenees in the amounts of silica in the plants were detected
when the plants were f e r with the hand. Thus, fewer opaline hooks la
could be felt on the surIaces of leaves of plants growing in the soils
containing added aluminium oxides than on those of plants growing
in the control soff.
The concentration of silica in solution in the plants differed
according to the soff (Table 5) and was roughly proportional to the
concentration of silica in the soil solution.
TABLE 5
Effects of sesquioxides on the concentration of silica in solution (ppm SiO2)
in oats grown in University soil
Oxide added
None (control)
Ferric oxide I
A l m n i n i u m oxide III
A l u m i n i u m oxide IV

.......

Silica in
soii solution
p p m SiO2
48
40
9
15

Days after sowing

34

77

123
87
44
58

173
140
53
76

The results of analyses for total silica in the plants (Table 6 and
Figs. 2 and 3) show that all the oxides lowered the uptake of silica.
The proportion of silica in the plants increased progressively with
age but at all stages the relation between plants growing in the four
solls was maintained. The total amounts of silica at maturity were
proportional to the concentrations of silica in solution in the solls.
Thus, fron and aluminium oxides lowered the concentration in the
soll solution and in turn reduced the uptake of silica by oats. Just
as aluminium oxides had a greater effect than fron oxides on the
concentration of siliea in soll solutions so they were more effective

90

L. H. P. J O N E S AND K. A. H A N D R E C K

in reducing the uptake of silica by oats. These results suggest that

there is a quantitative relation between the concentration of silica
in the soil solution and the amount taken up b y oats.
TABLE 6
Effects of sesquioxides on the uptake of silica by oats *
grown in University soil
Oxide added

Si02 content of plants

Silica in
soil solution
ppm SiO2

Dry mart e r
g/plant

% D.M.

mg/plant

48
40
9
15

2.90
2.80
2.42
2.94

2.81
2.44
0.49
0.89

81.5
68.3
11.9
26.2

None (control)
Ferric oxide I
Aluminium oxide I I I .
Aluminium oxide IV .
At 112 days after sowmg.

IOO

80
o c o n t r o l - no oxide added
[] ferric oxid T

1%1

aluminium oxide
aluminium oxide LV

cO
cm 6 0

Ln

tU

~. 40
.~

._v
u')

20

0
20

40

60

Days

80

after

I00

120

sowing

F i g . 2. T h e e f f e c t of s e s q u i o x i d e s o n t h e t o t a l s i l i c a i n o a t s g r o w n i n t h e
U n i v e r s i t y soff.

91

STUDIES OF SILICA IN THE OAT PLANT

Since passive absorption can account for most of the water taken
up by a plant, and the siliea of the soil solution is uncharged monosilicic acid, it is reasonable to expect a passive uptake of silica in the
transpiration stream. Knowing the concentration of silica in the
soil solution and the amount of water transpired b y the plant, it
should be possible to calculate the amount of silica in the plant. The
polyethylene pellets used as a mulch on the control pots reduced
evaporation from the soil surfaee b y about 95 per cent and allowed
the loss of water due to transpiration to be measured. Each plant
had transpired 1550 ml at maturity and with 48 ppm SiO2 in the
soll solution we should expect 74 mg SiO2 per plant. This was close
enough to the 81.5 mg SiO~ found by analysis to suggest that the
uptake of silica is passive. This hypothesis was tested in the experiments that follow.
E

3"0

__
IM

O
B

2.0

d~dded

- .

tn

,ca

~.. l.O

.-~

a uminium oxide _

~ aluminium

oxide

.-cD

. m _ _ i
"1~"

20

40

--a

i ~

(so
Days

80
after

1oo

mo

sowing

Fig. 3. The efect of sesquioxides on the concentration of silica in oats grown

in the University soll.

Pot experiments, x962. The relaHor~ o~ uptake to silica in the soil

solution and to trampiratio~z
The solls comprised the University soil without added sesquioxides, and
those from Penola and \ollongbar. They were weighed into polyethylene
pots which contained 7.00 kg of the University- and 4.75 kg of the Penola- and
*vVollongbar soils. There were eighteen po~s of the University soil and nine

92

L. H. P. JONES AND K. A. HANDRECK

pots of e a c h of t h e others. T h e p H ' s of live p o t s of e a c h of t h e t h r e e solls were

a d j u s t e d to 5.6 w i t h s u l p h u r i c a c i d ; five a d d i t i o n a l pots of t h e U n i v e r s i t y
soil were a d j u s t e d to p H 6.8 a n d t h e s e will b e r e f e r r e d t o as a f o u r t h soil.
E a c h p o t h a d m i x e d i n t o it t h e following: KH2PO4, 0.52 g; C a ( N O 3 ) 2 . 4 H 2 0 ,
1.03 g; M g S O 4 . 7 H 2 0 , 0.25 g; CuSO4.5H~O, 0.023 g; a n d d u r i n g t h e growing
season e a c h p o t received four surface a p p t i c a t i o n s of C a ( N O 3 ) 2 . 4 H 2 0 each
of 1.0 g.
T h e p o t s were w a t e r e d to p F 2.0 a n d t h i r t y seeds were sown in five p o t s
of e a c h soll in m i d - A u g u s t ; t h e n u m b e r of seedlings was r e d u c e d t o e i g h t e e n
soon a f t e r emergence. I n a d d i t i o n t o t h e p o t s w i t h p l a n t s t h e r e vere four p o t s
of e a c h soil in w h i c h p l a n t s were n o t grown. All p o t s were m u l c h e d w i t h
p o l y e t h y l e n e pellets a t t h e r a t e of 800 g p e r p o t a n d h o u s e d in a glass-house.
T h r o u g h o u t t h e g r o w i n g season t h e p o t s were weighed d a i l y a t a b o u t 5
p.m. a n d b r o u g h t b a c k t o p F 2.0 w i t h distilled w a t e t . P l a n t s were h a r v e s t e d
for a n a l y s i s a t live stages d u r i n g t h e growing season. T h e n u m b e r s of p l a n t s
t a k e n f r o m e a c h p o t a t successive h a r v e s t s were as follows: six, four, t h r e e ,
two, a n d t h r e e . A t t h e first t h r e e h a r v e s t s p a r t of t h e p l a n t m a t e r i a l f r o m
e a c h of t h e four soils was t a k e n for t h e e s t i m a t i o n of silica in s o l u t i o n in t h e
p l a n t ; t h e r e s t of t h e m a t e r i a l was r e s e r v e d for a n a l y s i s for t o t a l silica. A t
t h e last t w o h a r v e s t s all t h e p l a n t m a t e r i a l was r e s e r v e d for a n a l y s i s for t o t a l
silica.
TABLE 7
The concentration of silica in solution (ppm SiO2) in oats
grown in three solls of pH 5.6
Silica in
Days after sowing
SoiI
soil solution
44
58
ppm SiO~
University
54
78
119
l%noIa
67
148
t 53
. Wollongbar . . . . . .
7
44
68

82
130
222

70

The total silica in the plants was proportional to the concentrations of silica in solution in the four soils (Tables 8 and 9). Since
the amounts of silica in solution in these four soils differed over a
wide range, it is possible to compare more critically the uptake of
silica as found b y analysis with the uptake expected from measurements of both the watet transpired and the concentration of silica
in the soil solution. Throughout the period of growth there was
close agreement between the expected uptake of silica and the
amounts found b y analysis (Tables O and 9). These results confirm
our findings that these soils are capable of maintaining a steady
concentration of silica in solution despite repeated withdrawals.
They also show that the uptake of silica b y this variety of oats can

S T U D I E S OF SILICA IN T H E OAT P L A N T

93

be explained simply in terms of silica in the soil solution and the

water transpired; they thereby confirm our hypothesis of the passive
uptake of silica.
TABLE 8

The uptake of silica by oats grown in three soils of p H 5.6

Soil
SiO2 content of plants
Days
Dry
Watet I SiO2
(Silica in solution
after
marter
transpired expected
% D.M.
mg/plant ml/plant mg/plant
in parentheses)
sowing
g/plant
44
0.168
University
2.00
3.36
67
k62
(54 p p m SiO2)

Penola
(67 p p m SiO2)

Wollongbar
(7 p p m SiO2)

58
82
I09
124

0.410
2.12
5,80
6.40

2.36
2.69
2.77

9.39
50.0
156
177

175
910
2785
3265

).45
t9.1
150
176

44
58
82
109
124

0.212
0.507
3.23
6.18
6.92

2.69
3.04
3,40
3.80
3,96

5.70
15.4
110
235
274

86
243
1585
3350
3900

5.76
16.3
106

44
58
82
109
124

0.242
0.603
2.93
6.24
7.07

0.31
0.35
0.34
0.38
0.40

0.750
2.11
9.96
23.7
28.3

108
294
1415
3260
3860

0.756
2.06
9.90

2.29

224

261

22.8

27.0

The figures for silica in solution in the plant call for comment.
Besides differences which are related to the concentration of silica
in the soil solution, there were differences in the concentration of
silica in plants from the one soil on different days (Tables 5 and 7).
The passive uptake of silica can explain these differenees. Thus,
higher concentrations of silica in solution in the plants were found
on days of higher temperature and on these days both the transpiration rate and the rate at whieh monosilicic acid entered the plant
would increase.
TABLE 9

Effeet of pH on the uptake of silica by oats * grown in University soil

Soil
pH

Si~-cn~en-t B plan~t s |t

Silica in
solution

Dry marter
g/plant

% D.M.

6.40
6.13

2.77
1.68

Water

mg/pla~t t transpiredml/plant

p p m SiO~
5.6
6.8

54
30

* At 124 days after sowing.

177
103

3265
3250

SiO2

expected
mg/plant
176
97.5

94

L. H. P. JONES AND K. A. HANDRECK

It is likely that the uptake of silica by other gramineous species is

also passive. This suggestion is supported b y data on the composition of barley 23 which show the amount of silica in the plants to
be proportional to the water transpired. From the amounts of water
transpired and the amounts oI silica in the plants it can be calculated
that 68 to 88 ppm Si02 would have been required in the soil solution
to account for the silica found in the plants at various stages
through the growing period. It is interesting that these figures are
so consistent. However, they are probably consistently too high
because the silica in the plant material was determined as acid
insoluble ash. In this method the amount of silica is overestimated
by 10 to 20 per cent because of the alkalis, alkaline earths, and
water retained by the silica. If an allowance of this order is made,
the apparent concentration of silica in the soil solution would be
comparable to the concentrations found in our soils.
In so far as solls are capable of maintaining a constant level of
silica in solution it should be possible to estimate the amount of
water transpired b y a cereal crop growing in the field simply b y
analysing the plants for silica and determining the concentration of
silica in the soil solution.
So far this discussion has dealt with the oat and other gramineous
species which contain comparable proportions of silica. What
happens with other species may be very different. For example,
leguminous species contain relatively low proportions of silica 4 24
and taust have some mechanism for rejecting it at the root surface.
SUMMARY
Silicon in soil solutions is p r e s e n t e n t i r e l y as monosilicic acid a n d soils are
c a p a b l e of m a i n t a i n i n g a s t e a d y c o n c e n t r a t i o n in s o l u t i o n despite r e p e a t e d
w i t h d r a w a l s . This c o n c e n t r a t i o n is a l w a y s below t h e s a t u r a t i o n Iigure of
120 p p m SiO~ a n d varies a m o n g s t soils of t h e s a m e p H .
B o t h ferric a n d a l u m i n i u m oxides a d s o r b monosilicic acid b u t a m o n g
oxides of similar c r y s t a l l i n i t y , a I u m i n i u m oxide is m o r e effective t h a n ferric
oxide. W h e n a d d e d to a soil, b o t h ferric a n d a l u m i n i u m oxides lowered t h e
c o n c e n t r a t i o n of monosilieic acid in solution. A l u m i n i u m oxide a g a i n h a d t h e
g r e a t e r effect a n d one s a m p l e lowered t h e c o n c e n t r a t i o n of monosilicie acid
f r o m 48 t o 9 p p m SiO2.
I n p o t e x p e r i m e n t s oats were g r o w n in a soll t o w h i c h ferric a n d a l u m i n i u m
oxides h a d b e e n a d d e d a n d also in soils differing n a t u r a l l y in t h e i r c o n t e n t of
free sesquioxides. T h e u p t a k e of silica b y t h e p l a n t s c a n b e e x p l a i n e d s i m p l y

S T U D I E S OF SILICA IN T H E OAT P L A N T

95

i n t e r m s o f t h e c o n c e n t r a t i o n of s i l i c a in t h e soil s o l u t i o n a n d t h e a m o u n t of
watet transpired. This indicates that oats take up silica passively in the
transpiration stream.
ACKNOWLEDGEMENTS
W e a r e g r a t e f u l t o P r o f e s s o r s G. W . L e e p e r a n d S.M. W a d h a m
for their
i n t e r e s t ii1 t h i s w o r k a n d t o D r . A. A. M i l n e f o r h e r a s s i s t a n c e i n t h e m i n e r a l o g i c a l w o r k . P r o f e s s o r H . C. F o r s t e r
kindly provided laboratory and
glass-house facilities.
Received March t3, 1964
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18
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26

S T U D I E S OF SILICA IN THE OAT PLANT

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