JANUARY/FEBRUARY 2003 1

WINEGROWING

Encouraging
WHY AND HOW AM fungal hyphae are finer than
plant root hairs, which allows them to

beneficial
penetrate smaller soil pores. This
allows a plant with AM fungi to absorb
water and nutrients from areas of the
soil profile that roots can’t reach, pro-
viding plants with a higher tolerance

AM fungi
for drought and increased uptake of
certain nutrients, such as phosphorus.
Extraradical hyphae can colonize
multiple roots of either the same plant
or different plants, forming a mycor-
rhizal network. This network can

in vineyard soil
mediate the transfer of water and/or
nutrients between different plant
species. In one study, soybean plants
received nutrients via the AM fungal
network from adjacent weeds follow-
ing an herbicide application.6
AM fungi produce thick-walled
spores on extraradical hyphae. The
spores are released into the soil sur-
rounding roots colonized by AM fungi
and are spread by anything that moves

A
soil, including wind, water, and
BY Kendra Baumgartner, they cannot live in the absence of a machinery. Spores germinate near
plant host except as dormant spores. roots in response to root exudates,
USDA Research Plant Pathologist including various phenolic com-
Arbuscular mycorrhizal fungi are
Davis, CA pounds (flavonoids in particular).
so named because they form a fungal
structure known as an arbuscule. After germination, the AM fungus col-
rbuscular mycorrhizal (AM)
Arbuscules are tree-shaped structures onizes fine roots (germinating spores
fungi are beneficial organisms
that the fungus forms inside plant do not colonize woody roots).
that have a close association
cells. These structures are responsible Soil fumigants kill AM fungal
with the roots of a variety of
for nutrient exchange between the spores. Deep, uniform soil fumigation
plants, ranging from corn to redwoods.
plant and the fungus. may result in slow recolonization
They have been shown to increase crop
Some AM fungi also produce a (greater than two years after fumiga-
yield and nutrient uptake, enhance
structure known as a vesicle, which is tion) of grapevines by indigenous AM
resistance to plant pathogens, and to
thought to have a function in nutrient fungal species and can lead to stunting
stabilize soil structure.
storage. Vesicle production was once in some nutrient-poor soils.5
AM fungi live in symbiosis with
believed to be common among all AM Research on AM fungi and grape-
plants. This means they live in close
fungi, which were then called vesicular- vines in both the greenhouse 1,4,7,8 and
association with each other, enjoying a
arbuscular mycorrhizal (VAM) fungi. the field 5 has shown that mycorrhizal
mutually beneficial relationship. AM
Later research identified a family of vines with certain AM fungal species
fungi live partially within the roots of
supposedly VAM fungi that didn’t pro- have greater shoot and root growth
their plant hosts. The plant provides
duce vesicles. Since all types produce when compared to non-mycorrhizal
the fungus with carbon (a product of
arbuscules, the name was amended to vines. Rootstocks and ungrafted
photosynthesis), while the fungus
become AM fungi. cultivars kept free of AM fungi were
provides the plant with phosphorus
Arbuscules are connected to approximately two to three times
and other nutrients absorbed from the
strands of AM fungal tissue that smaller than mycorrhizal vines.4
soil.
extend into the soil (called extraradi- Mycorrhizal vines had denser, more
Other direct benefits provided by
cal hyphae). These hyphae absorb compact root systems, presumably due
mycorrhizal fungi to some plant hosts
water and nutrients for the fungus to the presence of numerous extra-
include improved drought tolerance 2
and the plant. They extend the root radical hyphae that branched out in the
and resistance to specific pests and
system and, in effect, increase the soil.7
diseases.3 AM fungi are biotrophs;
plant’s root surface area.
2 JANUARY/FEBRUARY 2003

WINEGROWING

In greenhouse experiments in which

various rootstocks were colonized by
different AM fungal species, some
were found to have higher concentra-
tions of phosphorus and zinc, but these
benefits were dependent on the AM
fungal species and the soil in which the
rootstock was planted.1
These studies, along with other
research conducted on many other crops,
demonstrate that the specific combina-
tion of plant species, fungal species,
and soil has a significant effect on how
a plant benefits from AM fungi. Some
combinations of plant, fungus, and soil
may increase zinc uptake, while others
may improve drought tolerance.
In some combinations, the plant
may not benefit at all. If the AM fungus
provides the plant with nothing, the
fungus starts to act like a parasite.
Plants do have some control over this
situation, although the mechanism of
this control is not known. For example,
under high rates of phosphorus fertil-
ization, plants are found to have fewer
Microscopic spores of arbuscular mycorrhizal fungi are produced on the roots of mycorrhizal
AM fungal structures in their roots. plants and are spread to new roots with the help of wind, equipment, rodents, and anything
With adequate levels of phosphorus, else that moves soil. Three arbuscular mycorrhiza fungal spores are connected to each other
the cost of “feeding” AM fungi (pro- by vegetative fungal tissue.
viding them with carbon) outweighs Propagation of inoculum and subse- can do to encourage AM fungal popu-
the benefit (more phosphorus). quent evaluation of the effects of spe- lations if you are concerned about a
Since the combination of plant, fun- cific AM fungal species on grapevine lack of them in your vineyard.
gus, and soil affects how a plant bene- growth in the greenhouse takes • Plant a cover crop. Most vineyard
fits from being an AM fungal host, it approximately one year to accomplish. cover crops are hosts of AM fungi (with
would seem important to find the com- Further greenhouse testing would be the notable exception of mustard).
bination that works best for specific required for evaluations in different Maintenance of cover crops during the
agricultural crops. This is, however, soil types. Finding the right combina- winter will likely result in a higher con-
easier said than done. Finding the ideal tion of AM fungal species for grape- centration of AM fungal spores in vine-
AM fungal species for grapevines vines may not be necessary, since AM yard soil than will be found in vineyards
would require identification of AM fungi occur naturally in vineyards where no cover crop is grown. Cover crop
fungal species from healthy vines in worldwide. mixes are likely to encourage a greater
the field, propagation of inoculum, and When land is cleared for vineyard diversity of AM fungal species, which
subsequent inoculation of vines grown establishment, AM fungal spores and may increase the chances of encouraging
in the greenhouse to determine which roots from mycorrhizal hosts that pre- those that are best for vines in your soil.
species are the most beneficial. viously inhabited the site serve as • Mow the cover crop in spring instead
Identification of AM fungi is very inoculum for grapevines. Most AM of incorporating it into the soil (if your
difficult. Since they cannot be cultured fungal species are thought to have cover crop is not too competitive). Soil
on artificial nutrient agar in the lab, broad host ranges. Nearby grapevines disruption breaks up mycorrhizal net-
they have to be grown on roots of liv- can serve as a source of AM fungal works, which are important for provid-
ing plants. AM fungal taxonomy is spores. AM fungi also occur on the ing mycorrhizal plants with increased
based on microscopic, morphological roots of field-propagated nursery nutrient and/or water uptake.
characteristics of the spores, and since stock. Therefore, even if soil is fumi- • Plant with field-grown nursery
many species are undescribed, keys are gated before planting, AM fungi will stock. Vines raised in the field have
useful only for identifying some of the eventually colonize young grapevines. AM fungi in their roots.
species. Vineyards become colonized by AM Commercial inocula of AM fungi
fungi naturally. There are things you are available. These products are more
 JANUARY/FEBRUARY 2003 3

WINEGROWING

practical for use in newly planted
rather than established grapevines.
Adding inoculum may be more
important when planting into fumi-
gated soils. Be sure to obtain the
freshest inoculum possible (within
days or weeks of it coming off the
production line) and apply it imme-
diately. The longer the spores stay in
the container, the lower their chances
of germinating once they are placed
on plant roots.
AM fungi likely have an unappreci-
ated role in helping grapevines tolerate
drought induced by deficit irrigation
and in the maintenance of sufficient
root growth and nutrient uptake in
marginal vineyard soils.
Adopting practices that encourage
AM fungal populations is easy, since
these beneficial fungi naturally colo-
nize grapevine roots. However, these
practices should not be substituted for
nutrient amendment, irrigation, or pest
management, especially when deficien-
cies or diseases/pests are already wide-
spread in the vineyard. n Hemstreet, C. L. 1983. “Interactions be-
References
1. Biricolti, S., Ferrini, F., Rinaldelli, E.,
Tamantini, I., Vignozzi, N. 1997. “VAM
fungi and soil lime content influence root-
stock growth and nutrient content.” Amer.
Journal of Enology & Viticulture 48: 93–99.
2. Fitter, A. H. 1988. “Water relations
of red clover, Trifolium pratense L., as
affected by VA mycorrhizal infection and
phosphorus supply before and during
drought.” Journal of Experimental Botany
39: 595–604.
3. Hooker, J. E., Jaizme-Vega, M., Atkin-
son, D. 1994. “Biocontrol of plant pathogens
using arbuscular mycorrhizal fungi.” In:
Gianinazzi, S., Schüepp, H., eds. Impact of
Arbuscular Mycorrhizas on Sustainable Agri-
culture and Natural Ecosystems. Basel,
Switzerland: Birkhäuser Verlag. p 191–200.
4. Linderman, R. G., Davis, A. A. 2001.
“Comparative response of selected grape-
vine rootstocks and cultivars to inoculation
with different mycorrhizal fungi.” Amer.
Journal of Enology & Viticulture 52: 8–11.
5. Menge, J. A., Raski, D. J., Lider, L. A.,
Johnson, E. L.V., Jones, N. O., Kissler, J. J.,
tween mycorrhizal fungi, soil fumigation,
and growth of grapes in California.” Amer.
Journal of Enology & Viticulture 34:117–121.
6. Mujica, M., Fracchia, S., Ocampo, J.,
Godeas, A. 1999. “Influence of the herbi-
cides chlorsulfuron and glyphosate on myc-
orrhizal soybean intercropped with the
weeds Brassica campestris or Sorghum
halepensis.” Symbiosis 27: 73–81.
7. Schellenbaum, L., Berta, G., Ravo-
lanirina, F., Tisserant, B., Gianinazzi, S.,
Fitter, A. H. 1991. “Influence of endomyc-
orrhizal infection on root morphology in
a micropropagated woody plant species
(Vitis vinifera L.).” Annals of Botany
68:135–141.
8. Schubert, A., Cammarata, S., Eynard,
I. 1988. “Growth and root colonization of
grapevines inoculated with different
mycorrhizal endophytes.” HortScience 23:
302–303.

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