Applied Clay Science 64 (2012) 12–17

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Applied Clay Science

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / c l a y

Research paper

Weathering of biotite in the presence of arbuscular mycorrhizae in selected

agricultural crops
J.M. Arocena a,⁎, B. Velde b, S.J. Robertson a
a
University of Northern British Columbia, Prince George, BC, Canada V2N4Z9
b
UMR 8538 CNRS Ecole Normale Supérieure, 24 rue Lhomond 75231 Paris, France

a r t i c l e i n f o a b s t r a c t

Article history: The mutual association of roots and fungus, mycorrhizae, benefits the general ecosystem by producing soil clays from
Received 17 October 2010 primary phyllosilicates. In laboratory experiments, X-ray diffraction analyses of clay materials showed that the
Received in revised form 17 June 2011 fungus Glomus when inoculated onto barley and canola produced various types of less K-containing phyllosilicates
Accepted 19 June 2011
(e.g., illites, smectites and hydroxy-interlayered vermiculites) from biotite and left some biotite (the K-rich mineral)
intact. Non-inoculated plants left no intact biotite. Both non-inoculated and inoculated alfalfa produced only some
Keywords:
Mycorrhizae
illites from biotite. We propose that mycorrhizae inoculated samples exhibit selective K-extraction from some biotite
Agricultural crops particles to benefit plant growth and at the same time leaving unaltered biotite for future extractions. However,
PCR results from molecular DNA analysis (i.e., PCR) of fungal population, X-ray absorption near edge spectroscopic
Clay minerals analysis of the oxidation of Fe2+ to Fe3+ and K-uptake by plants seems to indicate that the K-extraction process from
biotite is by no means simple. The transformation of biotite and K (and Mg)-uptake depend on the plant–fungus
symbiotic system, and the formation of clay minerals will then be a function of plant type and symbiotic relations in
the soils.
© 2011 Elsevier B.V. All rights reserved.

1. Introduction Newly-formed HIV (often called soil vermiculites) are commonly

The root zone of an estimated 95% of all plant species is home to
fungal–root symbiosis (or mycorrhizae) (Sylvia, 1998). Our conven-
tional knowledge is that mutual benefits result when host plants
provide fungi with photosynthates and the latter facilitate acquisition of
essential nutrients and water from the soil by the plants. Access of plants
to soil resources is increased by 2 or 3 orders of magnitude due to wide
extension of the root system through the prolific growth of fungal
hyphae. Mycorrhizal association is known to improve the nutrient and
water supplying capabilities of the soil, hence increase soil fertility
(Sylvia, 1998). However, we believe that mycorrhizal association is
more than the immediate nutritional benefits between plant and fungus
because new clay minerals had been observed in several root–soil
interface (or rhizosphere soils). The uptake of K by plants, such as
ryegrass, has been reported to convert phlogopite (Mg and K-rich mica)
into vermiculite, a high-charged hydroxy-interlayered vermiculite
(HIV) mineral (Hinsinger et al., 2006; Mortland et al., 1956). Further,
the biologically-mediated removal of Mg from chlorite (Mg-rich high
temperature phyllosilicate mineral) leaves behind HIV (Arocena and
Velde, 2009). Negative charges associated with surfaces of these
high-charged clays are effective adsorption sites to store essential
cations (e.g., Ca, K and Mg) and water, thus improve the natural fertility
of soils.
⁎ Corresponding author. Tel.: + 1 250 960 5811; fax: + 1 250 960 5539.
E-mail address: arocenaj@unbc.ca (J.M. Arocena).
observed in many rhizosphere soils (e.g., April and Keller, 1990; Barré
et al., 2008; Hinsinger et al., 2006). However, the identity of fungi
associated with the specific synthesis of new clay minerals in
rhizosphere soils is limited to our work on ectomycorrhizal (or outside
the root) associations in spruce and fir forest ecosystems (Glowa et al.,
2004). In the Canadian sub-boreal forests, the symbiosis between
Piloderma (an ectomycorrhizal fungus) and white spruce roots
transformed soil mica and chlorite to HIV with subsequent improve-
ment in soil fertility arising from increased cation exchange capacity
(CEC) and exchangeable K and Mg (Arocena et al., 1999; Glowa et al.,
2004). The removal of K and Mg through uptake by white spruce trees
developed some negatively charged clays from high temperature
biotite and chlorite, and consequently the synthesis of HIV and other
high-charged phyllosilicates in rhizosphere soils can be traced in part
to plant interactions in the soil zones of alteration sequences.
Fungal species in lichens have been reported to produce
secondary phyllosilicates also. Prieto et al. (1997) reported the
transformation of mica in granitic rocks into vermiculite under the
influence of Tephromela atra (Huds.). In addition, we observed the
formation of 2:1 clay minerals under colonies of Aspicilia caesioci-
nerea (Nyl.ex Malbr.) Arnold on granitic rocks in Kunlun Shan,
Qinghai (Tibetan) plateau in China (Arocena et al., 2003). In basalt
deposits in eastern Turkey, 2:1 type clays were reported under the
colonies of the following crustose, foliose and pathogenic lichen
species including Lecidella meiococca (Nyl.) Leuckert and Hertel,
Buellia disciformis, Tephromela atra (Huds.) Hafellner, Caloplaca

0169-1317/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.clay.2011.06.013
 J.M. Arocena et al. / Applied Clay Science 64 (2012) 12–17
flavorubescens, Scoliciosporum umbrinum (Ach.) Arnold, Candelariella
coralliza (Arocena et al., 2007).
In agricultural crops where fungal associations are mostly
arbuscular mycorrhizae, AM (or inside the root), we are not aware
of any report regarding the identity of fungi directly associated with
any secondary clay mineral formation, thus the need for studies
reported in this paper.
The objective of this study was to investigate the breakdown of
biotite to understand the role of Glomus, a major genus of arbuscular
mycorrhizal fungi, in the formation of secondary low-K content clay
minerals in rhizosphere soils of three agricultural crops: barley
(Hordeum vulgare), alfalfa (Medicago sativa), and canola (Brassica
napus). We conducted the study in a growth chamber for controlled
optimal growth conditions for plants and fungi.
2. Materials and methods
2.1. Growth chamber experiment and K-uptake
We grew three common agricultural crops (i.e., barley, canola and
alfalfa) in a growth chamber maintained at 20 °C, 30% RH, and day
\night cycle of 12/12 h to ensure optimal plant growth conditions. The
growth medium was a mixture of 20–100 mesh Ottawa sand (691 g)
and b200 mesh biotite (9 g) where the essential nutrients (i.e., N, P,
Ca, Na, Mg, B, Zn, Cu, Mo, Fe) except K were provided to the plants
through a modified Hoagland solution (e.g., Hoagland and Snyder,
1933). One group of seedlings (IN) was inoculated with MycoApply
Seed Inoculant (Fort Myers, Florida) and maintained in a growth
chamber while the second set in another growth chamber was left
non-inoculated (NIN). The plants were grown to 100 days and then
harvested. Some pots were thinned during the experiment to
primarily minimize space competition within the growth chamber.
Each treatment was replicated three times.
Plant samples (i.e., below and above-ground) were collected at the
end of the experiment and oven dried at 70 °C prior to determination
of elemental uptake. Total K and Mg contents taken up by crops were
estimated through the dry ash procedure using HNO3 and 5 M HCl
(Kalra and Maynard, 1991) was determined by ICP-MS. Our emphasis
was on mineralogical changes in the biotite rather than plant growth
thus, total K and Mg contents were presented as concentrations in
plants.
2.2. Biotite composition
We used ground specimen biotite (catalog # 46-1193) supplied by
Ward's Natural Science (Rochester, NY USA). The mean (and standard
deviation, n = 8) elemental composition of the biotite using micro-
probe at the Ecole Normale Supérieure (Paris, France) were as follows
(%): Na2O — 0.63 (0.17); MgO — 13.7 (0.46); Al2O3 — 10.92 (0.26);
SiO2 — 39.89 (0.70); K2O — 8.46 (0.37); TiO2 — 2.19 (0.11); MnO —
0.63 (0.21); FeO — 17.59 (1.16); and F — 7.14 (1.21).
3. Methods
3.1. AM fungal DNA analysis
Total DNA was extracted from the MycoApply Seed Inoculant (Fort
Myers, Florida) and root samples of non-inoculated (NIN) and
inoculated (IN) alfalfa, barley and canola (n = 3) using a 2X CTAB
(hexadecyl trimethyl ammonium bromide) protocol for plants
(Palumbi, 1996). The NS31 and D3-labeled AM1 primers were used
in PCR amplification of the internal transcribed spacer (ITS) region of
ribosomal DNA of the three traditional AM fungal families (Glomaceae,
Acaulosporaceae, and Gigasporaceae) (Helgason et al., 1998; Mummey
et al., 2005). Thermocycler conditions were an initial denaturation
step for 4 min at 94 °C, annealing for 1 min at 55 °C, and extension at
13
72 °C for 2 min, followed by 34 cycles of denaturation (94 °C for 30 s),
annealing (50 °C for 40 s) and extension (72 °C for 1 min) and final
extension at 72 °C for 6.5 min. PCR products were run on a high
resolution 2.5% agarose gel and digital images were saved using the
Gel Print 2000I photographic system (BioPhotonic Corp.).
PCR products were loaded into a CEQTM 8000 automated sequencer
(version 6.02) and run at 60 °C separation temperature, 4 kV voltage,
and 120 min separation time. Analysis was performed using the
amplicon fragment length polymorphism (AFLP) program of the
CEQTM 8000 sequencer and the quartic model for size standard with
the minimum relative peak height set at 1% and a bin width of 3 bp. The
fluorescent signal strength of each fragment peak was normalized to the
total peak area within each sample (Osborne et al., 2006). The relative
abundance of different DNA fragment lengths in each sample were
compared using Nonmetric Multidimensional Scaling (NMS), which
was calculated on the basis of a Sørensen (Bray–Curtis) distance
measure with 250 runs with real and randomized data (PC-ORD 5.0,
McCune and Mefford, 1999). Pairwise differences between treatment
(IN, NIN and AM inoculum) and plant groups were tested statistically
using Multi-Response Permutation Procedures (MRPP) with the
Sørensen distance measure (McCune and Grace, 2002).
3.2. X-ray diffraction analyses
Clay fractions were separated from rhizosphere (i.e., soils clinging
to roots) soils collected from IN and NIN samples. Calcium- and
K-saturated air dried and glycol treated oriented clay samples (b0.5 g)
were analyzed using a Bruker D8 with GADDS® X-ray diffractometer
using Co-Kα radiation generated at 40 kV and 20 mA. The low amount
of clay (b0.5 g) was sufficient for the 850 μm collimator and the
efficient area detector of the XRD diffractometer. X-ray diffractograms
were subjected to deconvolution analysis (Lanson, 1997) to estimate
the relative proportions of various secondary clay minerals from
calculated areas under each XRD peak.
3.3. Fe-X-ray absorption near edge spectroscopy
X-ray absorption near edge spectroscopy (XANES) was conducted to
determine the oxidation of Fe2+ in biotite to Fe3+. Selected clay samples
were ground to b50 μm using a mortar and pestle and mounted on
Capton™ tape and oriented at 45° prior to X-ray beam exposure at the
VESPERS beamline in the Canadian Light Source (CLS), University of
Saskatchewan synchrotron facilities. X-ray absorption spectra at 100 eV
range to include the Fe-edge (7112 eV) were recorded using a
multi-element detector. We collected at least three spectra from each
selected clay samples (contrasting samples based on XRD results) and
the reference samples (e.g., biotite and hematite) used in the study. The
estimates of the Fe2+ and Fe3+ were conducted by comparing the area
under the corresponding absorption peaks in the pre-edge part of the
spectra after least square fitting or deconvolution (Lanson, 1997; Wilke
et al., 2009).
4. Results
4.1. Clay mineralogy and K-uptake
Deconvolution analysis (Lanson, 1997) of X-ray diffraction patterns
of clay minerals collected from rhizosphere soils showed that various
types of less-K clays were produced from the K-rich high temperature
biotite in both non-inoculated (NIN) and Glomus-inoculated (IN)
barley, canola and alfalfa (Fig. 1A–E). In Fig. 1F, the biotite in the
control treatments (both IN and NIN) retained its original structure
with a sharp 1.0 nm peak indicating no significant structural
modification.
The oxidation of Fe 2+ in biotite to Fe 3+ is shown in the X-ray
absorption at lower energy (Fe 2+) in untreated biotite compared to
14 J.M. Arocena et al. / Applied Clay Science 64 (2012) 12–17

(A) Barley IN (B) Barley NIN

ML
(72%)
M
ML
(40%)
I

HIV I
(26%)

(C) Canola IN (D) Canola NIN

I

ML ML
HIV M
(40%) (80%)
(16%)
I

(E) Alfalfa IN (F) Control - No plant M

Alfalfa NIN* - I (50%) M No Plant - NIN

I
(60%)
46-1193 (Biotite)

No Plant - IN

6 8 10 12 8 10 12
CoKαο2θ

Ca, Al-OH Ca, Al-OH

K K
NH4+, Mg-OH NH4+, Mg-OH

Fig. 1. Representative spectra for X-ray diffraction analyses of biotite extracted from the experiments after plant growth. The spectra have been background substracted and the
intensities decomposed into fundamental components of mica, illite mixed layered mineral and hydroxy interlayered mineral (Lanson, 1997). Legend: = observed XRD
pattern; = fitted line; = HIV; = mixed layer; = illite, and = biotite.

higher energy (Fe 3+) in clay samples collected from the rhizosphere
of common agricultural crops with (IN) or without (NIN) Glomus
inoculation (Fig. 2). The estimates of the conversion of Fe 2+ into Fe 3+
ranges from 50% in alfalfa-IN to 100% in barley-NIN. Clay minerals
produced by the alteration of biotite by NIN plants were dominated by
a mixed layer (ML) mineral (mixed layering of K-rich micaceous
mineral and a low K mineral, smectite indicated as such by a peak shift
from 11.5 to 12.6 A upon glycol saturation in the barley IN sample of
Fig. 1), and illite (Fig. 1B, D and F). The presence of these new mineral
phases was indicated by X-ray diffraction analysis using the following
criteria:
Illite, a phase near 1.0 nm (~ 1.05 nm) with a more intense width at half
height (WHH) of ~ 0.8°2θ compared to the initial biotite which had a
WHH of 1.3°2. Careful work by Fordham (1990), (Fig. 1, Table 1)
indicates that altered biotites in soils have similar characteristics.
Detailed microprobe analyses of altered biotite, called illite by the
author, indicate a slight loss of potassium, and an increase in water
content as indicated by a lower oxide total in the microprobe analysis.
These characteristics indicate a parallel between altered trioctahedral
mica the dioctahedral illite minerals. We thus use the term illite below
to describe the presence of a trioctahedral, less potassic and more
hydrous mica-like mineral.
Mixed layered (ML) mineral (smectite/mica) had a peak position
N1.08 nm up to N1.2 nm indicating a mixing of illite and smectite of up
to 30% smectite assuming a bihydrated cation with two water layers.
These phases were seen to partially expand after glycol solvation.
 J.M. Arocena et al. / Applied Clay Science 64 (2012) 12–17 15

2+
Fe ~ 50%
3+
Fe ~ 50%
Alfalfa-IN

2+
Fe ~ 40%
3+
Fe ~ 60% Alfalfa-NIN

2+
Fe ~ 20%
3+ Barley-IN
Fe ~ 80%
3+
Hematite (Fe )
normalized absorption

2+
Fe ~ 0%
3+
Fe ~ 100% Barley-NIN

3+
Alfalfa-IN
Hematite (Fe )
Alfalfa-NIN

2+
Biotite (Fe )

Barley-NIN
7104 7106 7108 7110 7112 7114
energy (eV) 2+
Biotite (Fe )
Observed spectrum - solid line
Fitted line - dash dot line
2+
Fe line - dash line
3+
Fe line - dot line Barley-IN

IN-Glomus-inoculated; NIN - non inoculated

7060 7080 7100 7120 7140 7160

energy (eV)

Fig. 2. Iron X-ray absorption near edge spectroscopy (Fe-XANES) of clay samples and reference Fe2+ (biotite) and Fe3+ (hematite). Inset shows shift from lower (Fe2+) to higher
energy (Fe3+) absorption and the relative estimates of Fe2+ and Fe3+ in various clay samples.

HI mineral or HIV with a peak position at 1.42 nm and a narrow peak
(WHH of 0.8°2 θ) and unchanged by glycol treatment.
The relationships between K-uptake by crops and the formation of
illite, mixed layers and HIV are shown in Fig. 3A–D. K-rich mineral (or
biotite) with a peak at near 10.2°2 θ (Co-Kα radiation) was still
present in IN samples, while in all NIN samples, biotite appeared to
have been altered to less K-minerals except for alfalfa IN (Fig. 3A).
Both IN and NIN plants produced illite with contents ranged from low
15% (alfalfa NIN) to as high as 83% in barley NIN (Fig. 3B). Similarly,
mixed layer clays were observed in all treatments other than alfalfa IN
and NIN treatments although canola-IN samples exhibit both ML and
no ML formation (Fig. 3C). Estimates of HIV content were 18 and 12%
for barley IN and canola IN, respectively, and the production of HIV
was limited to these two treatments (Fig. 3D). We did not observe
significant differences in K and Mg concentrations in plants in both IN
Table 1
Mean⁎ (and standard error) uptake of K and Mg by inoculated (IN) and non-inoculated
(NIN) selected agricultural crops (mg K or Mg g−1 plant biomass) (n* = 3).
K uptake (mg K g−1)
and NIN treatments but K and Mg in biomass of barley were
significantly lower and higher, respectively than alfalfa and canola
(Table 1).
4.2. AM fungal community analyses
The number of DNA fragments (representing AM fungal units) was
greater in IN compared to NIN samples of alfalfa and canola, but lower
in IN compared to NIN samples of barley. The relative abundance of
DNA fragments of the IN and NIN groups did not vary significantly
from one another overall or within plant groups. Only NIN community
profiles varied significantly from the profile generated from the AMF
inoculant. The IN groups shared 10 DNA fragments with the inoculant
that were not present in the NIN group. Five AM fungal units were
shared by the inoculant and each of the alfalfa-IN and canola-IN
groups; only one AM fungal unit was present in both the inoculant
and barley-IN groups. Hence, some DNA fragments were present in
the NIN samples at the end of the experiment, but IN samples mainly
contained the same AM fungi as the original inoculant.
5. Discussion
Mg uptake (mg K g−1)

Canola-NIN 5.57a (0.52) 12.0ab (0.18) Information on the alteration of fresh biotite in alteration sequences
Canola-IN 5.08a (0.25) 14.1b (0.43)
is given by Jeong et al. (2006). This study investigated the weathering
Alfalfa-NIN 6.56a (1.22) 10.3a (0.37)
Alfalfa-IN 5.14a (0.10) 10.6a (0.15) reaction effects in natural systems where no plants were involved in the
Barley-NIN 3.98b (0.26) 20.2c (2.62) change of mineralogy, samples coming from deeper than 60 cm. Deep
Barley-IN 4.22b (0.22) 17.8c (1.02) weathering produced essentially two types of phases: hydro-biotite
⁎ Means followed by similar letters are not significantly different (p N 0.5) using the which is a regularly ordered smectite–biotite alternate structure,
Fisher's Least Square Difference test; n — number of replicates. hydroxyl-interlayered (HI) mineral and a structurally-modified mica.
16 J.M. Arocena et al. / Applied Clay Science 64 (2012) 12–17
100 (A)Biotite (B) Illite
80
60
40
Relative Content (%)
20
0 K (and Mg) from biotite. The synthesis of HIV in rhizosphere can also be
100
(C) Mixed Layer (D) Hydroxy-Interlayered
Vermiculite (HIV)
80
60
40
20
0
40 60 80 100 40 60
K- uptake (mg K per pot)
Fig. 3. Plot of estimated potassium extraction by plants and the relative contents (peak
area%) of the major clay mineral phases present for the experiments. (A) — Biotite,
(B) — Illite, (C) — Mixed Layer, (D) — Hydroxy-Interlayered Vermiculite.
The authors insist on the effect of oxidation which produces the mineral
changes. Overall there is a loss of potassium from the structure and the
oxidation state of iron is modified strongly. However examinations of
their data suggest that there are different mechanisms of K extraction,
especially in the HI samples where the loss of K is greater than the
oxidation effect. Here, the HI mineral is likely to have been produced by
a direct Al(OH)2+ exchange for K +. We will use Jeong et al. (2006) as
background information for the interpretation of the results of the
laboratory studies reported in this paper.
The influence of Glomus, a genus of AM fungi used to improve the
yield in cultivated crops (Powell, 1981), when inoculated to barley,
canola and alfalfa in the production of several types of less K-content
minerals from the initial biotite was not straight forward. Establishment
of mycorrhizal association seems to be very much influenced by the
symbiosis between the fungus and the agricultural crops as indicated by
the variable number of DNA fragments among IN plants. For barley, the
presence of less K minerals (i.e., smectite, HIV) in the IN than in NIN
samples indicates a more efficient K (and Mg)-extraction process from
individual biotite grains when Glomus and other organisms are present in
the system. The AM fungi represented by the 10 DNA fragments shared
between the inoculant and IN treatment groups may belong to the
Glomus genus and play a role in the changes in clay minerals observed.
Although the majority of these AM fungi were associated with alfalfa and
canola seeds, it is not the number but differences in the functional
attributes of different AM fungi colonizing a root system may increase
resource acquisition and plant survival (Helgason and Fitter, 2005).
Secondary clay minerals synthesized from biotite by the crop-Glomus
sp association have decreasing K content from 1.0 to 0.0 K atom per
formula unit and represents a continuum of clay minerals from biotite to
illite to mixed layer to HIV (see Fig. 1). Although some NIN plants have
higher K-uptake than IN plants, the synthesis of less-K secondary clays
(e.g., ML and illite) increases with K uptake by crops whether or not
initiated by inoculation treatment. This supports earlier claim that the
complex mechanisms of K removal from biotite involved more than
Glomus inoculation but also the symbiotic relationships between crops
and fungus. Nevertheless, electrical charges on ML and HIV result from
the lowering of layer charge by oxidation of Fe2+ and/or removal of K in
biotite. The extraction of K (and Mg) is commonly compensated by
cations (e.g., Ca2+, NH4+, Mg2) in the soil system many of which is
essential to maintain vigorous growth of plants and organisms and can
improve the long term fertility of soils. We interpret the shift in the Fe
absorption band to higher energy in the Fe-XANES data reported in this
paper to indicate that there two divalent Fe positions in the biotite, one
of which is more easily oxidized than the other. The Fe oxidation may
have been facilitated by the action of Glomus and enhanced the release of
due to a direct exchange of hydroxyl–aluminum cations for K. The effects
on plant fertility due to the presence of exchangeable cations (such as
Al–OH) is questionable, but the presence of exchanged K for the
hydroxyl interlayer ion is a very important source of K for plant growth
Alfalfa IN
Barley IN
(e.g., April and Keller,1990; Carnicelli et al., 1997; Fichter et al., 1998;
Canola IN Zanelli et al., 2006).
Barley NIN
Canola NIN The presence of unaltered biotite in the barley-IN samples suggests
Alfalfa NIN that there was a selective extraction of K (and Mg) from some biotite
grains by mycorrhizal associations in the barley-IN system leaving
others biotite intact. This may suggests a greater proximity of these
grains to the mycorrhizal hyphae. In the barley-NIN samples, no
biotite is left intact and all material is altered but less in the extreme
cases than in the IN samples. Thus, mycorrhizal fungi (in case of
80 100 barley-Glomus) selectively extract K to immediately benefit plant
growth and at the same time leave unaltered biotite for future
extractions. Further in the barley-NIN treatment, all biotite grains
were transformed with the implication that much of the K released by
root–mineral interaction can be lost from the plant growth system
through leaching into the ground water.
It is interesting to note that the extent of alteration of the biotite
grains is not only dependent on the mycorrhizal treatment, but also
on the type of plant used. Both IN and NIN alfalfa produced illite which
indicates only minor destabilization of the biotite structure due to
minimal removal of K showing that plant species is an important
factor in plant–soil mineral breakdown. Canola and barley have
similar effects in the IN experiments but barley shows a greater effect
(shift to lower 2θ position) than in the NIN experiment. This indicates,
as one might expect, somewhat different responses of extraction
efficiency depending upon plant type. However, canola extracts large
amounts of K in both IN and NIN treatments even though the
extraction mechanism is different oxidation to produce smectite
layers or hydroxy-aluminum K exchange. It is known that lupin, a
nitrogen fixing plant as is alfalfa, tends to be very weakly mycorrhizal
(see review by Hodge et al., 2009) which explains the small effect
observed in the experiments reported here.
A fascinating study was made by Balogh-Brunstad et al. (2008) on
biotite weathering in the laboratory using ectomycorrhizal fungi and
biotite without the presence of plants. They observed that the effects
of alteration were very similar to those of organic acid attack alone,
hence no specific effect was noted when the fungi were present alone.
This coupled with the observations in the present study that NIN and
IN control plots have no detectable effects on the transformation of
biotite indicates strongly that the plant–mineral reactions are strongly
influenced by symbiotic processes which enhance the efficiency of K
release for plant growth.
Coming back to the background study of Jeong et al. (2006) our clay
minerals derived from biotite appear to be of a different origin from
those formed by water rock interaction under oxidizing weathering
conditions although the mechanisms can be similar. The biotite in our
untreated soils only slightly altered and hence the change in mineralogy
in the plant experiments would be due to bio-interactions exclusively.
Further, the key mineral in the weathering of biotite, the formation of
hydro-biotite, a regularly interlayered mixed layered mineral with
sharp (narrow) peaks was not observed in our experimental products.
 J.M. Arocena et al. / Applied Clay Science 64 (2012) 12–17
Instead an ML mineral of random mixed layering characteristic was
formed. In both water–rock and biologically driven alterations one can
find a hydroxy-interlayer mineral formed by hydroxy ion–potassium
exchange. Thus, biological processes appear to be much more rapid than
those of mineral chemical processes but probably affected by the same
chemical processes.
6. Conclusion
We can conclude that the symbiotic plant–fungus system maybe
more efficient through the selective extraction of K in some biotite
grains, probably those in contact with the fungi, conserving K
resources for future use in other grains outside the immediate contact
zones in the soil. It seems that conservation of biotite is achieved by a
more complete extraction of K (and Mg) and oxidation of Fe thus the
formation of HIV. Roots of barley and canola without mycorrhizal
inoculation apparently alter all biotite grains in the soil, releasing K
more indiscriminately. Our results provide useful information to the
important role of fungus–root symbiosis in the maintenance of
fertility and long term productivity of agricultural (and forest) soils.
They also underline the varied response of minerals to interactions
with different plants and symbiotic systems.
Acknowledgments
We acknowledge the financial support from the Natural Sciences
and Engineering Research Council, and the Canada Research Chair
Program. We thank S. Brown, A. Esler, R. Miller and M. Thompson for
assistance in the experiment and laboratory analyses. Part of the
research described in this paper was performed at the Canadian Light
Source, which is supported by the Natural Sciences and Engineering
Research Council of Canada, the National Research Council Canada, the
Canadian Institutes of Health Research, the Province of Saskatchewan,
Western Economic Diversification Canada, and the University of
Saskatchewan.
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