A Symbiotic Relationship

Ancient fossils in a Wisconsin cutbank shed light on a modern erosion

control project.
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By Michael Amaranthus, David Steinfeld

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When Alexander Fleming discovered penicillin, he was trying to perfect an antiseptic formula based on
nasal mucus. The nasal mucus formulation never did materialize (we all can breathe a sigh of relief!), but
his unforeseen discovery of antibiotics changed the world. Great discoveries often have been
unanticipated or appreciated only years after the initial findings. Walter Alvarez was attempting to figure out
the geologic origins of Italy's Apenine Mountains; years later, he realized that he had stumbled upon
evidence that dinosaurs had been rendered extinct by a large comet or asteroid. When scientists were
trying to find an alternative form of Freon, they invented Teflon. Rayon and nylon were botched
experiments. Clearly serendipity has played a role in many great discoveries.

This brings us to a recent discovery in a Wisconsin

roadcut (Redecker et al., 2000). The roadcut
contained newly discovered fossilized roots
containing the threads and microscopic spores of
fungi some 460 million years old. At first, scientists
believed these ancient fungi had a negative effect on
the plant. After all, fungi are parasites and
saprophytes and are likely robbing the plant roots of
their energy or rotting them like a moldy piece of
bread. Upon closer examination, however, this group
of fossilized fungi clearly was different. They were
mycorrhizal fungiÑthe same mycorrhizal fungi that
form a beneficial association with the fine roots of
terrestrial plants growing throughout the world today.

Photo 1: Mycorrhizal fungal filaments

radiating from a mycorrhizal colonized
root
These specialized fungal filaments form a symbiotic organ called the mycorrhiza (photo1). Through this
structure, the plant provides energy for the fungus, and in return the fungus provides the nitrogen,
phosphorous, and other minerals and water necessary for plant establishment and growth. The
mycorrhizal fungus also promotes fine root development and soil structure and protects plant roots from
diseases and soil toxicities (Trappe and Fogel, 1977; Harley and Smith, 1983).

The discovery in the Wisconsin cutbank is further evidence that fungi and green plants moved from water
onto land together, bolstering the theory that these specialized fungi were the key factor that allowed
plants to successfully invade the land. The newly discovered fossils push the origin of land-based fungi
back some 55 million to 60 million years to about the same era that green plants first grew on land. The
mycorrhizal fungi attached to the roots of aquatic plants allowed plants to access water and nutrients on
the relatively harsh earth surface. Before the mycorrhizal arrangement, these aquatic plants could not
survive on land.

The plant and fungus mycorrhizal association is widespread today, with 85-90% of all green plants forming
mycorrhizae. By far, the most common mycorrhizal type is the arbuscular mycorrhizae (AM), which
penetrate the plant root cells but do not modify the external appearance of the root. This type of
mycorrhiza forms with more than 100 fungal species in the family Endogonaceae; predominates in
grasses, forbs, bryophytes, pteridophytes, and most tropical tree genera; and is associated with most
nursery and agricultural crops.

Most undisturbed natural areas today contain an abundance of mycorrhizal fungi. However, areas
disturbed by human activity often lack sufficient mycorrhizal populations that are important for plant
establishment and growth (Amaranthus and Perry, 1994; Amaranthus et al., 1996; Dumroese et al., 1998).
Disturbance, compaction, invasion by weeds, erosion, removal of topsoil, mixing, land clearing, and
frequent pesticide use can adversely affect beneficial mycorrhizae. The ancient Wisconsin fossils provide
insight into establishing plants in the disturbed and hostile environments of today. A good example is a
recent study of the McDonald Basin, a chronic erosion-prone site in the Siskiyou Mountains of southwest
Oregon.

McDonald Basin

McDonald Basin is a denuded, high-elevation site

that contributes large quantities of granitic sand-size
sediments to the Applegate watershed. The need for
soil cover by native plants has been identified as a
resource objective for the area for nearly four
decades. Debbie Whiteall, hydrologist for the United
States Department of Agriculture Forest Service, has
worked in the region for more than 15 years and
describes the area's condition: "McDonald Basin is
certainly one the more disturbed areas along the
Siskiyou crest. The site's elevation, aspect, and
prevailing weather patterns contribute to the naturally
occurring, highly erosive, and nutrient deficient
condition. For over a hundred years, human impacts,
including grazing and road building, have
Photo 2: Planting Agrostis plug seedlings exacerbated the condition. The basin is situated in
at McDonald basin the headwaters of the Little Applegate watershed and
provides water for domestic uses and endangered
salmon downstream and is a priority for restoration activities. Many years of operational plantings and
studies of native grasses and trees in McDonald Basin have yielded little improvement in soil cover."

Objective
From a management standpoint, the restoration
objective at McDonald Basin is to establish an
adequate soil cover over time. This can occur in
several ways: directly through the presence of live
or dead vegetation or indirectly through more seed
yield or a better environment for seed germination
and establishment. Any measures that improve soil
cover should improve watershed conditions. Poor Above: Photo 3a: A McDonald basin area
fertility at the McDonald Basin site often has been before planting
cited as one of the primary reasons for the mortality
Below: Photo 3b: after planting
of planted vegetation. The use of time-release
fertilizer in the nursery soil of containerized
plantings could be a mechanism for maintaining
plant vigor in the harsh outplanting environment.
New information from the Wisconsin cutbank also
has sparked interest in using the primary
relationship between plants and mycorrhizal fungi in
restoration planting. This study investigated the role
that AM and slow-release fertilizers in the nursery
play in establishing grass seedlings outplanted at
McDonald Basin.

Methods

We sowed native grass seed, Agrostis pallens

(dune bent grass) in 4-in.3 containers in late May
1999. Prior to sowing, we mixed mycorrhizae and slow-release fertilizers into the media in the following
combinations:

• Mycorrhiza only
• Mycorrhiza and slow-release fertilizer
• Slow-release fertilizer only
• No mycorrhiza or slow-release fertilizer (standard greenhouse practices)

The mycorrhiza inoculum was composed of the species Glomus intraradices and had a concentration of
130 propagules per gram. We applied it at a rate of 1.2 grams inoculum per seedling. We used Osmocote
18-6-12 with an eight- to nine-month longevity for the slow-release fertilizer. After the seeds were sown, we
placed the containers in a greenhouse and grew the seedlings for two months. By that time, the snows had
melted on the slopes of McDonald Basin and we could assess the site.

We planted the seedlings in mid-July in a random block outplanting design, where each block included
each treatment randomly placed (photo 2). Although the growing season is short at this elevation, the
planted seedlings became established and grew before the snows covered the slopes in late October and
remained covered until the following summer (photos 3a and 3b).

Results

We revisited the study site in mid-September to take measurements and collect seedling samples. Within
a two-month period we found dramatic differences in the survival rate between mycorrhizal inoculated
grass seedlings and noninoculated seedlings (Figure 1). Grass seedlings that had been inoculated with
mycorrhiza in the nursery had 100% survival. Seedlings that had been inoculated with mycorrhiza in
combination with slow-release fertilizer application survived at an 81.4% rate. Seedlings fertilized with
slow-release fertilizer, but not inoculated with mycorrhiza, resulted in 26.4% survival rate. Seedlings grown
under standard nursery practices of soluble fertilizer, but without mycorrhizal inoculation and slow-release
fertilizer, had only a 17.2% survival rate.
From each treatment, we excavated five seedlings and brought them back to the lab to measure the dry
weights of roots and leaves and also to evaluate the amount of mycorrhizal colonization on the root
systems of each seedling. We found that mycorrhizal-inoculated seedlings had significantly greater root
biomass and mycorrhizal colonization after one year in the field compared to seedlings receiving fertilizers
only (Figures 2 and 3). Very little mycorrhizal colonization occurred on seedling roots with fertilizer-only
treatments. Seedlings inoculated with mycorrhizae in the nursery also had high nutrient concentrations
compared to other treatments. Foliar levels of phosphorous, potassium, calcium, and sulfur levels were all
significantly higher (Figure 4).
Discussion

The results of this study highlight basic differences between the function of mycorrhizal fungi and
fertilizers. "Can't I just fertilize?" is a common question from growers. Inoculated Agrostis seedlings grown
without additions of time-release and soluble fertilizers had higher foliar concentrations of important
macronutrients. It is well established that mycorrhizal fungi facilitate the capture and uptake of nutrients
from the soil and produce enzymes important for the extraction of minerals from soil particles. Studies
have shown that grass species in the family Poaceae benefit greatly from mycorrhizal colonization in terms
of growth and nutrient acquisition (Gemma and Koske, 1989; Hall et al., 1984). Mycorrhiza also perform
many important activities beyond the capture and uptake of nutrients that might improve plant
establishment. For example, mycorrhizal fungi are important for disease suppression, drought protection,
improved soil structure, enhanced leaf chlorophyll levels, and tolerance of nutrient imbalances (Linderman,
1994). Research studies have shown that mycorrhizae can enhance the ability of grasses to avoid water
stress (Koske et al., 1995; Auge et al., 1995; Allen, 1991). Recent studies indicate that creeping bent grass
inoculated with the mycorrhizal fungus Glomus intraradices tolerated drought conditions significantly
longer than nonmycorrhizal turf (Gemma et al., 1997). It is also well documented that inoculating grasses
with mycorrhizal fungi in soil with low phosphorous concentrations can produce greater shoot and root
biomass (Hall et al., 1984; Petrovic, 1984; Hetrick et al., 1986). Further research on mycorrhizae has
shown that mycorrhizae promotes the development of early successional tallgrass prairie communities. A
significant increase in the percent cover of native species Andropogon gerardii, Panicum virgatum, and
Sorghastrum nutans was observed (Smith et al., 1998). Mycorrhizal fungi can tap into the soil resource by
extending their hyphae far from the roots' surface. Mycorrhizal filaments are two to five times smaller than
the roots themselves and have a greater surface area per unit volume. This allows the filaments to explore
more soil volumes and smaller spaces in the soil not accessible by roots. Once in these small spaces, the
mycorrhizal fungi excrete enzymes and chelates that access immobile nutrients.

"In nature, most plants require mycorrhizae to compete effectively; without mycorrhizal fungi they are
competitively inferior," says Jim Trappe, Ph.D., professor of botany at Oregon State University. "In stressful
environments this means the difference between life and death." "Some of the biggest success stories
have been the use of mycorrhizal inoculants on mine reclamation sites," remarks Efren Cazares, assistant
professor of forest science at Oregon State University. Over the years he has been able to study the
effects of human activities on mycorrhizal activity. "While mycorrhizal inoculants should not be considered
a silver bullet, it is clear that linking plants to the soil resource via the appropriate mycorrhizal fungi greatly
increases the chances for a successful revegetation project," says Cazares.

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A common question from erosion control practitioners is "When should I apply mycorrhizal fungi?"
Mycorrhiza can be incorporated during propagation, nursery production, or transplanting, or to existing
trees and shrubs. Common methods include incorporating granules into soil-less bark/sand media at the
nursery or banding or rototilling into soils at field sites. Other methods include using mycorrhizal seed
coats, sprinkling granular materials around root balls, and injecting soluble powder blends into soils
surrounding existing plants. Still other methods include inserting mycorrhizal tablets into the rooting zone
and using hydromulchers to spray mycorrhizal inoculum onto sites. Whatever the method, mycorrhizal
fungi need access to roots, and as with any living organisms, mycorrhizal fungi prefer some living
situations to others, so diverse species mixes are best.

Mycorrhizal fungi have been critical to the establishment of plants for more than 460 million years, but it
has been only in the last 40 years that scientists truly have appreciated the importance of the relationship.
Today there are more than 50,000 scientific papers on mycorrhiza and their function. The significant
difference in plant survival and root growth in the McDonald Basin is part of a growing body of scientific
studies that demonstrate that mycorrhizal inoculation in disturbed environments can help plants become
established more effectively and with less chemical inputs. Microscopic root fossils in a Wisconsin cutbank
remind us of a simple and obvious restoration tool that lies right under our feet.

Author's Bio: Michael Amaranthus, Ph.D., is an associate professor (adjunct) in the Department of Forest
Science at Oregon State University and president of Mycorrhizal Applications Inc.

Author's Bio: David Steinfeld is a USDA soil scientist with J. Herbert Stone Nursery in Central Point, OR.

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