Arbuscular mycorrhizal fungi in field crop production:

Potential and new direction

Chantal Hamel1 and Désiré-Georges Strullu2
1Agriculture and Agri-Food Canada, Semiarid Prairie Agricultural Research Centre, Box 1030 Airport Rd.,
Swift Current Saskatchewan, Canada S9H 3X2 (e-mail: hamelc@agr.gc.ca); and 2UFR Sciences,
Université d’Angers, 2 boul. Lavoisier, 49045 Angers Cedex, France. Received 13 May 2005,
accepted 23 March 2006.

Hamel, C. and Strullu, D-G. 2006. Arbuscular mycorrhizal fungi in field crop production: Potential and new direction. Can.
J. Plant Sci. 86: 941–950. Arbuscular mycorrhizal fungi (AMF) are multipurpose organisms with complex ecological ramifications
in the soil system that have been difficult to study and understand. The phytocentric concept of AMF that has prevailed since the
naming of these organisms is being replaced by a holistic vision recognizing that AMF are a key element of soil functioning and
health rather than a plant root component. Recent advances in knowledge brought about by new techniques for soil microbiology
research open the way to AMF management in crop production. Arbuscular mycorrhizal fungi may influence crop development,
even in phosphorus-rich soils. However, growing crops in soil with lower fertility would optimize the expression of the multiple
beneficial effects of AMF in agro-ecosystem and reduce nutrient seepage to the environment. The consideration of the soil myc-
orrhizal potential within the framework of soil testing and fertilization recommendations, the development of improved inoculants
and signal molecules to manipulate AMF and the development of cultivars with improved symbiotic qualities would insure the
production of good crop yields while improving agroecosystems’ sustainability.

Key words: Arbuscular mycorrhizal fungi management, field crop production, agriculture, soil quality, arbuscular mycorrhizal
effect

Hamel, C. et Strullu, D-G. 2006. Les mycorhizes à arbuscules en agriculture : nouvelles directions et orientations possibles.
Can. J. Plant Sci. 86: 941–950. Les mycorhizes à arbuscules (MA) sont des champignons aux ramifications écologiques complexes
dans le sol. Ils ont de multiples rôles mais sont difficiles à étudier et à comprendre. La théorie phytocentrique qui prévaut depuis
que ces organismes ont reçu leur nom est en train de céder la place à une vision holistique en vertu de laquelle les MA ne seraient
pas une simple composante des racines des plantes mais joueraient plutôt un rôle déterminant dans le fonctionnement et la vital-
ité du sol. De nouvelles techniques de recherche en microbiologie du sol ont récemment fait progresser nos connaissances et
ouvrent la porte à la gestion des MA en agriculture. Les mycorhizes à arbuscules pourraient influer sur le développement des cul-
tures, même dans les sols riches en phosphore. Toutefois, on optimiserait les nombreux effets bénéfiques des MA dans les écosys-
tèmes agricoles et réduirait la perte des éléments nutritifs dans l’environnement en cultivant les plantes sur des sols peu fertiles.
En tenant compte du potentiel des mycorhizes lors de l’analyse du sol et de la formulation de recommandations sur la fertilisation,
en mettant au point de meilleurs inoculants et des molécules signal pour modifier les MA, et en créant des cultivars aux plus
grandes qualités symbiotiques, on garantirait l’obtention d’un bon rendement des cultures tout en favorisant la pérennité des
écosystèmes agricoles.

Mots clés: Gestion des mycorhizes à arbuscules, grandes cultures, agriculture, qualité du sol, incidence des mycorhizes à
arbuscules

In a book on modern agronomy, Lambert et al. (1994)
defined arbuscular mycorrhizae as mixed organs resulting
from the association between fungal hyphae and roots. They
explain that the mycorrhizal hyphae improve plant absorp-
tion efficacy, in particular for the uptake of water and nutri-
ents of low mobility in soil such as phosphorus (P), as they
reach soil volumes well beyond the P depletion zone devel-
oping around roots when the root uptake rate exceeds the
rate of nutrient diffusion in soil. The concept embedded in
the above definition of arbuscular mycorrhizae is inherited
from the 1880s, when Frank created the name “mycorhiza”
i.e., “fungal root” (in Koide and Mosse 2004). Fortunately,
Frank’s concept, which reduces the arbuscular mycorrhizal
fungi (AMF) to a root component, is finally becoming obso-
lete. While the description of arbuscular mycorrhizae by
941
Lambert et al. (1994) is true, it is too restrictive. A funda-
mental aspect of the picture is overlooked: the soil.
AMF are so important in soil that a geocentric vision of
the arbuscular mycorrhizal symbiosis would have probably
been adopted at the onset, if we had been soil dwellers.
Since we do not live in the soil, our dependence on crop
plants has seemingly led us to adopt the phytocentric vision
that has been conveyed in textbooks and scientific articles
from the time of Frank up to very recently. The phytocentric
vision of the arbuscular mycorrhizal symbiosis that pre-
Abbreviatiions: AMF, arbuscular mycorrhizal fungi;
FAME, fatty acid methyl esters; PCR, polymerase chain
reaction; PGPR, plant growth promoting rhizobacteria
942 CANADIAN JOURNAL OF PLANT SCIENCE
vailed in scientific societies was concurrent to the general
exclusion of mycorrhizal research from soil science-related
disciplines. As a result, some 50 yr after Barbara Mosse cre-
ated mycorrhizal pot cultures of apple seedlings, initiating
the field of arbuscular mycorrhizal research, which yields an
average of 216 scientific publications each year since 1972
(CAB Abstract), agronomists have deserted the field of
mycorrhizal research on the premise that AMF can be
replaced by fertilizers.
AMF are multipurpose organisms with complex ecologi-
cal ramifications in the soil system, and their purpose has
been misunderstood by the public and scientists alike. This
may explain why AMF have raised relatively little interest
in the agronomic world. However, the management of AMF
in crop production is bound to become common practice in
the near future. After an era characterized by liberal use of
mineral fertilizers, new developments in crop production
techniques and policies aiming at ensuring environmental
protection are expected to lead to the reduction of fertilizer
use. Microbial inoculants are proposed as a “clean” option
to reduce crop production dependence on mineral fertilizers
(e.g., P solubilizers and AMF) and to enhance crop yield by
ways unrelated to mineral nutrition such as biochemical
plant reprogramming (Varma et al. 1999; Gray and Smith
2005; Waller et al. 2005) or bioprotection (Dehne 1982; St-
Arnaud et al. 1995). Public concerns are leading to the
development of policies to reduce the environmental
impacts of agriculture and to improve the agri-food system’s
sustainability in Canada (Stonehouse 2004) and elsewhere
(Johansson et al. 2004; Karlen et al. 2004). Precision agri-
culture technologies are already widely used to improve the
efficiency of fertilizers, and to reduce waste and seepage of
nutrients to the environment, thus, improving the sustain-
ability of crop production. The next important improvement
will be to produce more food with less agro-chemical inputs.
In a context of production system sustainability and of
environmental protection, AMF’s role as an important fac-
tor deserves more attention. The objective of this article is to
present the real function of AMF in the plant-soil system,
and to point at the technological developments needed for
AMF management in agricultural fields.
THE FUNCTION OF AMF
There are four broad groups of soil microorganisms: soil
animals, prokaryotes, fungal saprotrophs and endophytes.
Animals are involved in predation and comminution of soil
organic matter. Prokaryotes are a very diverse and versatile
group. They are responsible for much of the oxido-reduction
reactions taking place in the soil and are also largely
involved in soil organic matter mineralization, along with
fungal saprotrophs. In contrast to these later groups, AMF
are biotrophic endophytes that do not use soil organic mat-
ter as a source of carbon and energy but, rather, depend on
a host plant for their supply. In return, AMF stimulate plant
growth, improve the physical quality of their soil environ-
ment, and protect them against soil-borne pathogens.
Arbuscular mycorrhizal fungi form extensive hyphal net-
works in the top soil layer onto which most plants of an
ecosystem are connected (Read and Birch 1988). As much
as 30 m of AMF hyphae can be found per gram of soil
(Leake et al. 2004). In nature, AMF networks are involved
in the distribution of nutrients in the top layer of the soil and
among plants within a community (Marschner and Dell
1994; George et al. 1995; He et al. 2003; Neumann and
George 2004; Simard and Durall 2004), and influence plant
community structure (Hartnett and Wilson 1999; Stampe
and Daehler 2003; Urcelay and Diaz 2003).
The nature of AMF networks has been well described
recently (Giovannetti et al. 2004; de la Providencia et al.
2005). These networks are involved in the distribution of
photosynthesis-derived carbon in soil (Staddon et al. 2003a;
Zhu and Miller 2003). Arbuscular mycorrhizal hyphae,
which may account for 3–20% of root weight (Smith and
Read 1997), have a turnover of approximately 6–7 d
(Staddon et al. 2003b). Thus, considerable amounts of car-
bon are distributed in the soil via AMF networks, potential-
ly influencing soil microorganisms (Marschner et al. 2001;
Soderberg et al. 2002; Marschner and Baumann 2003) and
the accumulation of soil carbon (Rillig et al. 2001; Lovelock
et al. 2004). Our understanding of the soil systems has
improved and it has become clear that AMF are multipur-
pose key components of soil functioning and sustainability
(Leake et al. 2004). Plants, AMF and the soil matrix are
essential parts of the soil system (Fig. 1), and all these com-
ponents must be considered to understand the soil systems.
Harris and Paul (1987) estimated that 40–50% of photo-
synthesis-derived carbon is channelled to AMF; more con-
servative values of 10–20% were reported by Jakobsen et al.
(2002). Although the carbon cost of AMF to plants varies
with the organisms involved and the environmental condi-
tions, it is certainly an appreciable drain on a host plant. The
carbon cost related to AMF maintenance is offset by the
positive effects AMF have on plant growth and soil quality.
This beneficial impact of AMF is well documented (Smith
and Read 1997). The AMF effect on plant growth promotion
is most often attributed to improved uptake of water and
nutrients, in particular P, which has low mobility in soil
(Bieleski 1973) and which is required in large amount by
plants. Better nutrition, improved water relations and
increased carbon sink size have been proposed to explain
the high photosynthetic activity of AMF colonized plants, of
which the symbiotic condition would oscillate between
mutualistic and parasitic, depending on nutrient availability
(Bethlenfalvay et al. 1987). Although they are most often
beneficial, AMF have reduced plant productivity in nutri-
ent-rich agricultural soils (Ryan and Graham 2002; Stewart
et al. 2005).
Arbuscular mycorrhizal fungi are associated with, but
clearly distinct from plants and appear as the backbone of
soil ecosystems rather than as a plant part of foreign origin.
Evidence for the key role of AMF in soil systems comes
from the revelation of the very ancient origin of AMF.
Fossil records indicate that the Glomites, which existed 400
million years ago, were very similar to modern AMF, and
experts believe that the association of plant and AMF may
have helped land colonization by plants (Pirozinsky and
Dalpé 1992; Taylor et al. 1995; Schussler et al. 2001). The
fact that AMF have remained largely unchanged throughout
 HAMEL AND STRULLU — AMF IN FIELD CROP PRODUCTION
Solar
energy
Chloroplast
M
M
Fig. 1. Graphic representation of interactions in the photosynthesis powered plant-AMF-soil system. M, mineral nutrients; e, exchange of
carbon and mineral nutrients at the symbiotic interface.
the co-evolution of plants and soil, testifies to their impor-
tance to soil functioning and quality. The soil is an integrat-
ed and complex evolutive system with physical and
biological components (Crawford et al. 2005). Since evolu-
tion proceeds through the replacement of less performant
systems with performant ones, AMF, which have persisted
through time, appear as a performant multipurpose compo-
nent in soil ecosystems. As exposed by Barea et al. (2002),
AMF help plants produce more biomass with lower levels of
soil available nutrients and at the same time are photosyn-
thesis-driven soil quality builders.
AMF are involved in the maintenance of soil quality
(Barea et al. 2002). The physical soil entrapping effect and
contribution to soil organic matter of AMF hyphae directly
influence soil aggregation and structural stability (Jastrow et
al. 1998; Wright and Anderson 2000). Arbuscular mycor-
rhizal mycelium and spores produce a cell surface glyco-
protein, glomalin, which improves soil physical quality
(Rillig 2004). Glomalin was found to be closely related to
stable soil aggregate formation (Wright and Upadhyaya
1998), except in high carbonate soils where soil aggregate
stability depends on carbonates rather than on soil organic
materials (Franzluebbers et al. 2000).
Glomalin and its degradation products appear to be recal-
citrant to decomposition in soil, where they accumulate, as
determined by their long-lasting detection with a monoclon-
al antibody. Glomalin is often (Franzluebbers et al. 2000)
but not always related to soil organic matter (Rillig et al.
2001). It appears, however, that glomalin is an important
source of organic matter in soil. These fungi form abundant
943
Root hair
e
mycelial networks, have a fast turnover rate, and the gloma-
lin they produce seems to by-pass the microbial processing
imposed on fresh organic matter, thus contributing directly
to the stable soil organic matter pool. The Bradford-reac-
tive-soil-protein (BRSP) pool extracted from tropical soils
was shown to have a minimum residence time of 6 to 42 yr,
according to carbon dating, and the abundance of
immunoreactive glomalin has seemingly reached 3–10 mg
cm–3 in their A and O horizons (Rillig et al. 2001). All this
suggests that glomalin-related soil proteins’ contribution to
the stable soil organic matter pool is important. Thus, AMF
hyphal networks possess the ability to modify the physical
quality of plant habitats through their important contribution
to soil organic matter build-up and stabilization of soil
aggregates.
Extraradical AMF hyphae also influence the soil biologi-
cal environment and, hence, may influence soil biochemical
processes and the incidence of disease outbreaks.
Arbuscular mycorrhizal fungi-soil microbial interactions are
complex and cannot be generalized. The zone of soil imme-
diately surrounding AMF hyphae, the hyphosphere, hosts
selected rhizosphere bacteria (Vancura et al. 1990). These
bacteria may require amino acid and growth factors provid-
ed by AMF, and their populations may fluctuate as AMF
hyphae turnover throughout the soil volume. The AMF
impact on the overall microbial community has varied.
AMF were also shown to have little effect (Olsson et al.
1996) or negative impact (Christensen and Jakobsen 1993)
on the number and activity of total soil bacteria. Marschner
and Baumann (2003) observed a significant effect of AMF
944 CANADIAN JOURNAL OF PLANT SCIENCE
on the structure of the soil bacterial community. Numerous
researchers report AMF-related changes in the quality of the
soil microbial community (Marschner and Baumann 2003)
and variation in the size of specific microbial populations
(St-Arnaud et al. 1997; Amora-Lazcano et al. 1998;
Andrade et al. 1998a, b; Edwards et al. 1998; Filion et al.
1999; Green et al. 1999), but no general trend for an AMF
effect on soil microorganisms emerges, most likely due to
the complexity and wide biodiversity of the soil
environment.
The AMF cytoplasm may also host bacterial endophytes,
in particular plant growth promoting rhizobacteria (PGPR)
(Ruiz-Lozano and Bonfante 2000; Minerdi et al. 2002;
Bianciotto et al. 2004). The role of the endophytes living
within AMF spores and hyphae still needs to be clarified,
but some evidence suggests that they could be involved in
nutrient exchange between the partners of this consequently
tri-partite symbiosis (Minerdi et al. 2002) and stimulate
AMF spore germination (Bianciotto et al. 2004).
Important advances are being made in the field of AMF
ecology with molecular tools such as polymerase chain
reaction (PCR) (Kowalchuk et al. 2002; Vandenkoornhuyse
et al. 2003; Gollotte et al. 2004; Hunt et al. 2004; Rosendahl
and Stukenbrock 2004) and fatty acid methyl ester (FAME)
(Olsson 1999; Olsson et al. 1999; Balser et al. 2005; Nilsson
et al. 2005) based techniques that now allow us to track
these fungi in plants and soils. AMF ecology has recently
become an active field of research, which should provide
important knowledge for the management of AMF in agri-
cultural fields.
THE “MYCORRHIZAL EFFECTS”
AMF influence crop production in commercial farms
although it is not always recognized, this influence being
confounded with that of other soil factors. The creation of
non-mycorrhizal controls through the chemical destruction
of native AMF (Buttery et al. 1988; Liu et al. 2002, 2003;
Gazey et al. 2004) or their repression using non-host crops
(Vestberg et al. 2005), fallow (Abu-Zeyad et al. 1999) in
rotation, or deep tillage (Miller et al. 1995; Kabir et al. 1998;
Drijber et al. 2000; Miller 2000) have revealed that AMF
influence crop development. Considering the profound
influence of AMF on many aspects of plant physiology and
the complexity of the soil system, it is virtually impossible
to pinpoint the mechanisms responsible for the AMF effect
in any particular case. However, we know that the expres-
sion of this AMF effect in crop growth depends on three fac-
tors: environmental conditions including soil type, and plant
and fungal genotypes.
AMF were considered as a monolithic group of non-spe-
cific fungi. We now recognize that considerable variation in
plant growth response can be triggered by different AMF
isolates (Louis and Lim 1988; Castelli and Casper 2003;
Klironomos 2003; Sylvia et al. 2003; van der Heijden et al.
2003; Stewart et al. 2005). The symbiosis is more complex
than it was first thought and there is no such thing as an all-
purpose AMF isolate; the effect of an AMF is plant (Hart
and Klironomos 2002) and soil (Hamel et al. 1997; Rivera
and Fernández 2003; Rivera et al. 2007) dependant.
Considerable interaction between the AMF and plant
species isolated from a woodland site illustrated well the
ability of different AMF to enhance P uptake and growth in
different plant species and to colonize their roots (Helgason
et al. 2002). In Cuba, inoculants contain one of a few AMF
strains identified as effective, in monospecific AMF inocu-
lants. The different strains formulated are recommended for
use in different soil types (Rivera et al. 2007). Thus, the
mycorrhizal effects depend on the plant and AMF genotypes
interaction, as well as on soil conditions.
Plant genotype is a determinant of the AMF effect in two
ways. First, plant species have a specific influence on AMF
development and it appears that plant species determine, to
a large extent, the composition of AMF populations flour-
ishing in a soil (Eom et al. 2000). Some AMF are even
denied access in some plant species, even if they are good
colonizers on other species (Helgason et al. 2002; Sanders
2003), and cannot reproduce. This specificity suggests that
crop species in rotations may influence the quality of the
AMF population in the soil of a following crop. Second, the
genotype of a plant can also influence the response of this
plant to specific AMF. For example, Liu et al. (2000)
demonstrated the differential response of three maize
hybrids to inoculation with one AMF. Stewart et al. (2005),
working with different strawberry cultivars in a P-rich field,
also found large intraspecific variation in plant response to
mycorrhizal inoculation, with the growth of some cultivars
being largely decreased by inoculation with a given AMF
inoculant, while the growth of another was largely
increased. It makes no doubt that plant and AMF genotypes
influence the effect of AMF on crop development in agri-
cultural fields.
Most often, AMF effects were sought and found in plant
development. It has been overlooked, but mycorrhizal
effects are also related to soil quality. Plant genotype could
indirectly modify soil physical quality or the risk of disease
outbreak, through it effects on AMF development or popu-
lation composition. Plants, AMF and soil are three interact-
ing components of a system. In this system, plant genotype
could have an important effect on soil quality both through
direct and indirect modifications of the soil environment.
Vegetation is an important soil-forming factor (Brady and
Weil 2001). Plants directly influence soils in their quality as
the main source of metabolically active and soil organic
matter C, as well as through their influence on soil water.
Plants indirectly modify the soil environment, as they are
the determinant of the AMF networks development and,
thus, of their influence in soil.
It is well known that high soil P decreases AMF develop-
ment (Smith and Read 1997; Gazey et al. 2004; Linderman
and Davis 2004a). This sometimes led to the conclusion that
AMF cannot enhance plant growth in high P soils. However,
soil fertility is not the only determinant of AMF develop-
ment and effectiveness. The AMF effect depends also on the
plant and AMF genotypes, and AMF-related growth
enhancement at high soil P levels has occurred (Singh et al.
2002). Morin et al. (1994) demonstrated increased apple tree
biomass and leaf surface area in response to AMF inocula-
tion in P-rich soil (288 mg kg–1 Bray extractible P), while
 HAMEL AND STRULLU — AMF IN FIELD CROP PRODUCTION
Stewart et al. (2005) observed 50% increase in stolon pro-
duction per inoculated strawberry mother plant in a field soil
containing 222 mg kg–1 Mehlich-3 extractible P. In a study
to define in which field soils leek plants would benefit from
AMF inoculation, the factors associated with a positive
response to inoculation varied with soil available P level
(Hamel et al. 1997). In soils with more than 200 mg kg–1
Mehlich-3 extractible P, the soil mycorrhizal potential was
the major determinant along with the species composition of
the AMF populations indigenous to these soils. Considering
that Bray and Mehlich-3 extractible P levels of 25–30 mg
kg–1 and 45–50 mg kg–1, respectively, are considered opti-
mum for plant growth (Pierzynsky 2000), these studies indi-
cate that AMF may indeed stimulate plant growth, even in
soil very rich in available P.
Arbuscular mycorrhizal fungi are present in agricultural
fields, even in soils under intensive management. Intensive
management decreases AMF biodiversity and selects for
slow colonizing – fast sporulating species (Oehl et al. 2003),
but the examination of AMF biodiversity throughout the soil
profile revealed that genotypes presumably less fit in inten-
sively managed soil were found at greater depths (50–70
cm) in a soil zone unaffected by cultural practices (Oehl et
al. 2005). Arbuscular mycorrhizal fungi at greater depths
can be seen as a bank of biodiversity that can feed the top
soil inhabiting population when the conditions change in
surface soil, improving the adaptability of the AMF popula-
tion of cultivated soils. Furthermore, poor taxonomic AMF
diversity in intensively cropped systems may not be indica-
tive of poor AMF functional biodiversity, as pointed out by
Munkvold et al. (2004) who found a large intraspecific func-
tional diversity in AMF. Thus, the management of AMF
native to agricultural soils appears possible even in soil
modified by a history of intensive crop management.
ENHANCING MYCORRHIZAL EFFECTS IN FIELD
CROPS
It is clear that AMF effects are less prominent in soil with
high or excessive P fertility. The build up of excessive soil
P fertility levels is not desirable as P loss to the environment
may reduce water quality in rural areas (Beauchemin and
Simard 1999) and excessively rich soils do not produce
higher yields than well-managed P poor, sufficient or rich
soils. The best scenario for both farmers and society is the
one where the efficiency of P fertilizer applied at lower rates
is enhanced by AMF. In this way, soil P fertility would not
diminish potential beneficial AMF effects other than plant P
nutrition, and yields would be optimized.
In soils excessively rich in P, following repeated applica-
tion of high rates of P fertilizers, manure or compost, man-
ufactured signal molecules may be able to stimulate AMF
development and optimize AMF contribution to crop pro-
duction and soil quality. Different molecules were found to
stimulate AMF development (Tsai and Phillips 1991;
Bécard et al. 1992; Cruz et al. 2004; Dong and Zhao 2004).
Manufactured signal molecules stimulating AMF are cur-
rently being tested and will soon enter the market. These
new biotechnological products could be advantageously
integrated with known practices optimizing AMF contribu-
945
tion to crop productivity, such as cover cropping (Sorensen
et al. 2005), crop rotation (Johnson et al. 1992) reduced
tillage (Miller et al. 1995), and moderate fertilization (Miller
2000). Such tools could be also useful to enhance AMF
effects in crop species less receptive to AMF, and allow the
expression of AMF-related benefits on soil quality, and on
the development of a possibly mycorrhizae-dependent sub-
sequent crop.
Current P fertilization recommendations are very impre-
cise. Most soil test P used only estimate the available P in
the mineral fraction of the soil and ignore P potentially
available in the organic fraction of soils and the mycorrhizal
potential of soils. In order to implement strategies of
reduced fertilization and AMF management in agricultural
soils with no undue risk of yield loss, P fertilization recom-
mendations could be based on a soil test estimating both the
amount of soil P potentially available to a crop and the soil
mycorrhizal potential, which represents the potential contri-
bution of indigenous AMF populations to the recovery of
this P. A few methods have been proposed to evaluate the
status of AMF populations of soils; the most probable num-
ber or MPN (Porter 1979), the mycorrhizal soil infectivity
(MSI) (Plenchette et al. 1989) and the undisturbed core
(Brundrett et al. 1994) methods were the most favourably
considered by the scientific community. These methods all
generate estimations of the extent of the soil ability to pro-
duce mycorrhizal root colonization, but provide no informa-
tion on the quality of the AMF populations. Furthermore, all
these methods are bioassays involving growing trap plants
for weeks, which cannot be used for routine soil testing. It
may be possible to develop a convenient soil test to estimate
the potential contribution of AMF to crop nutrition based on
molecular methods. Theoretically, DNA analysis could pro-
duce estimates of the quality (Kowalchuk et al. 2002; de
Souza et al. 2004) and amounts of AMF in a soil. Real-time
PCR was used to quantify AMF (Filion et al. 2003; Alkan et
al. 2004), and PCR probes were used to enumerate AMF
isolates in ecosystems (Kowalchuk et al. 2002; Gollotte et
al. 2004). Although the analysis of phospholipids FAME
extracted from soil gives no information on AMF biodiver-
sity, it can estimate AMF abundance in soil (Balser et al.
2005; Nilsson et al. 2005). These analytical techniques
could be used in soil-testing laboratories to improve the
accuracy of soil P supply power estimates and fertilization
recommendations. The results of the test would be entered
into models specific to the crop to be grown, after calibration.
Fertilization using such an approach would maximize the
expression of AMF benefits to soils and crops, reduce agri-
cultural reliance on fertilizers, and reduce the risk of nutri-
ent loss to the environment. Attempts are currently being
made to develop such analytical techniques. The cost of
microbial identification through FAME profiling in a com-
mercial laboratory is about US$50 per sample; routine quan-
tification of AMF indicators through FAME analysis in a
soil test laboratory should cost considerably less. Soil myc-
orrhizal potential may well end up being modelled if we
finally achieve development of a practical methodology to
measure it and develop models; this would considerably
reduce the cost of AMF monitoring. A DNA-based analysis
946 CANADIAN JOURNAL OF PLANT SCIENCE
of biodiversity assessment would also be much less costly
than FAME analysis. Estimation of potential AMF contri-
bution to crop nutrition through soil testing appears feasible.
The use of manufactured bioproducts could enhance the
AMF effect. Inoculation technologies for AMF already exist
and can be very effective in containerized production
(Linderman and Davis 2004a) or in disinfested field soil
(Wininger et al. 2003). Due to the high cost of AMF inocu-
lant production, which is linked to the biotrophic nature of
AMF, inoculation technologies are currently largely restrict-
ed to a few special crop production systems. With progress
in AMF inoculant production techniques and the develop-
ment of inoculants for field use, AMF inoculation could
become profitable for agronomic crops. The identification
of molecules that stimulate AMF could lead to the develop-
ment of inoculant formulation including compounds stimu-
lating the AMF contained in the product. The value of the
AMF inoculants could be enhanced by the inclusion of
endophitic or hyphosphere-inhabiting PGPRs. These organ-
isms could enhance or complement the growth-stimulating
effect of AMF, or their ability to reduce plant pathogenic
populations in soils. In turn, the management of AMF in
crop production would insure good extraradical mycelium
development leading to soil quality improvement and
reduced activity of soil-borne pathogens.
In addition to environmental conditions and AMF geno-
type, plant genotype is a factor determining the AMF effect
(Liu et al. 2000; Linderman and Davis 2004b). If there is
variation in plant response to AMF within a plant species, it
should be possible to breed plants for their ability to form
efficient symbioses with AMF. Breeding techniques involv-
ing the selection of genotypes in soil containing selected
AMF strains may lead to the development of cultivar/inocu-
lant packages for low-input crop production. In light of the
large body of literature reporting the biocontrol effect of
AMF (Dehne 1982; St-Arnaud et al. 1995), it appears that
this approach might be particularly interesting for the pro-
duction of pesticide-free food crops, at a time where food
safety has become a priority in Canada and elsewhere. Plant
breeding, however, is a business driven by the requests of
the industry. In Canada and in other countries that have
favoured high-input crop production, breeding for mycor-
rhizal effectiveness is awaiting the implementation of crop-
ping systems considering AMF.
Arbuscular mycorrhizal biotechnologies are already used
in some countries where farmers could not afford the cost of
high input agriculture. The governments of India (Sharma et
al. 2006) and Cuba (Rivera and Fernández 2003; Rivera et
al. 2007), for example, have developed successful cropping
practices based on AMF inoculation to increase crop pro-
duction efficiency and reduce production costs. Effective
mycorrhizae formation in crop increases fertilizer use effi-
ciency by enhancing crop nutrient uptake efficacy. High soil
fertility levels, in particular high P fertility, stimulate plant
growth but repress AMF development. Maximizing AMF
contributions through precise fertilization allows the pro-
duction of good yields in soils maintained at lower fertility
levels, reducing nutrient loss to the environment while
improving soil physical and biological quality. The produc-
tion of crops with fertilizer levels favouring AMF develop-
ment also brings AMF-related benefits in the form of
reduced disease incidence, improved crop tolerance to
drought and other stresses, and improved soil physical qual-
ity, which translate into reduced soil erosion and improved
aeration and water infiltration, providing a better environ-
ment for plant growth.
In wealthier countries, crop production relies heavily on
agrochemicals because these products are reliable and easy
to use on large highly mechanized farms although they may
have negative impacts on the environment. Concerns relat-
ed to the environmental impacts of crop production effluents
are now an incentive for the development of AMF tech-
nologies also in these countries. At the same time, scientific
and technological progresses are coming to a point where it
may be possible to integrate AMF management into inten-
sive cropping systems. The adoption of crop production
practices considering the management of AMF would lead
the improvement of the quality of agricultural soil and crop
resistance to stresses. Experts predict that global climate
change will increase the frequency of extreme climatic
events (Patz et al. 2005), thus increasing risks of crop fail-
ure. The temperate latitudes, which are projected to warm
disproportionately, are among the potentially vulnerable
regions. Stress-resistant crop plants growing in high-quality
soils will certainly be an asset in the future. Arbuscular myc-
orrhizal fungi have a place amongst the biotechnologies of
tomorrow’s agriculture.
CONCLUSION
The real nature of AMF and the important contribution of
these fungi to soil quality maintenance and proper function
are just being realized. Research on AMF has progressed
slowly due largely to the biotrophic nature of AMF, which
limited our ability to study and understand these fungi.
However, technologies are now in place for the develop-
ment of the tools required for the management of AMF in
crop production as well as to increase knowledge on the
ecology of these fungi. With better understanding of AMF
in agricultural soil and with tools to monitor and manage
AMF, we will be able to optimize the contribution of AMF
to agricultural production and finally move toward the man-
agement of more sustainable cropping systems, for the ben-
efit of societies and agro-industries.
ACKNOWLEDGEMENT
Thanks to Keith Hanson for comprehensively reviewing and
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