Abstract Arbuscular mycorrhizal (AM) fungi differ in
their response to soil pH. Thus, change in soil pH may in-
fluence the relative abundance of mycorrhizal fungi inside
roots. Root colonization by two AM fungi was studied in
relation to addition of lime (CaCO
3)
, quantity of inoculum
and inoculum placement. Addition of CaCO
3
to an acid
soil decreased the colonization of roots by Acaulospora
laevis but increased colonization by Glomus invermaium
when both fungi were present. In acid soil (pH 4.7), almost
all roots were colonized by A. laevis, while G. invermaium
was dominant when soil pH was increased to pH 7.3. This
occurred regardless of whether the inoculum was banded
or mixed throughout the soil. There was no effect of
CaCO
3
on the relative abundance of fungi inside roots at
intermediate rates of CaCO
3
application (pH 5.3–6.3)
when both fungi were inoculated together. In this experi-
ment, both fungi colonized roots at all levels of CaCO
3
when inoculated alone, except for A. laevis at the highest
level of CaCO
3
. We conclude that soil pH affects the com-
petitive ability of these two AM fungi during mycorrhiza
formation primarily by affecting hyphae growth in soil and
thus the relative abundance of hyphae at the root surface
and subsequently inside the root.
Keywords Subterranean clover · pH ·
Glomus invermaium · Acaulospora laevis · Interactions
Introduction
Field soils contain diverse communities of arbuscular
mycorrhizal (AM) fungi (Smith and Read 1997). Al-
though root systems usually become colonized by more
than one AM species (Daft and Nicolson 1974; Abbott
and Robson 1982; Merryweather and Fitter 1998;
Jacquot et al. 2000), little is known of the interactions
between species during mycorrhiza formation (Juniper et
al. 1998). The quantity of mycorrhizas formed by spe-
cies of AM fungi in field soils will depend on both the
amount of infective hyphae and their relative competi-
tive ability (Abbott and Robson 1984). The length of
root colonized by AM fungi at early stages of root
growth is related to inoculum density (Carling et al.
1979; Abbott and Robson 1984; Wilson 1984;
Giovannetti and Avio 1986) and species of AM fungi
differ in their competitive ability (Wilson and Trinick
1983; Abbott and Robson 1984; Wilson 1984; Lopez-
Aguillon and Mosse 1987; Smith et al. 2000). The dif-
ferences in competitive ability to form mycorrhizas
may be based on antagonism (Hepper et al. 1988), de-
pletion of nutrients within roots (Wilson and Trinick
1983) and carbon demand (Pearson et al. 1993). The
compounding effects on colonization of both inoculum
level (Abbott and Robson 1984) and rate of hyphae
growth from inoculum to root surface have not been
clarified.
AM fungi clearly differ in their response to soil pH
(see review by Robson and Abbott 1989). Increasing the
pH of an acid soil may result in marked changes in the
species of AM fungi present as assessed by spores in the
soil (Siqueira et al. 1990). Acaulospora laevis, for exam-
ple, is generally found more in acid than in neutral soils
(Davis et al. 1985; Porter et al. 1987a; Young et al. 1985)
and germination of spores was limited at >pH 6 (Hepper
1984; Porter 1987b). On the other hand, an isolate of
Glomus invermaium appeared less affected by soil pH,
colonizing roots of subterranean clover (Trifolium sub-
terraneum L.) over the soil pH range of 5.3 to 7.5 (Abbott
and Robson 1985).
We investigated the role of soil pH and inoculum
quantity on the relative competitive ability of A. laevis
and G. invermaium. In one experiment, we varied soil
pH by adding CaCO
3
and the soil was inoculated
throughout with propagules of A. laevis. Subsequently,
S.M. Sano · L.K. Abbott (
✉
) · M.Z. Solaiman · A.D. Robson
Soil Science and Plant Nutrition, Faculty of Natural
and Agricultural Sciences, The University of Western Australia,
Crawley, WA 6009, Australia
e-mail: labbott@cyllene.uwa.edu.au
Tel.: +61-8-93802499, Fax: +61-8-93801050
Present address:
S.M. Sano, Embrapa CPAC, Caixa Postal 08223,
Planaltina-DF, CEP 73301-970 Brazil
Mycorrhiza (2002) 12:285–290
DOI 10.1007/s00572-002-0185-6
ORIGINAL PAPER
S. M. Sano · L. K. Abbott · M. Z. Solaiman
A. D. Robson
Influence of liming, inoculum level and inoculum placement
on root colonization of subterranean clover
Received: 4 December 2001 / Accepted: 21 May 2002 / Published online: 20 July 2002
© Springer-Verlag 2002