Mycorrhizae
Nature's minute miracle-workers C
O N T E N T S
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IN RECENT YEARS we've
read and heard a lot about the benefits of mycorrhiza in association with
pines - how that tacky-looking white fluff can actually benefit the tree.
But this is only part of the story and there is a lot more to this miracle
of nature.
Before we go further, lets just examine exactly what we mean by mycorrhiza.
The term mycorrhiza (plural mycorrhizae) is rather like the term "marriage".
It describes an association, a relationship. If both parties to the relationship
are compatible, and conditions suitable, they will both benefit. Otherwise
the relationship is of no benefit to either and it ceases.
One party to this relationship is your tree, the other
is a microscopic beneficial fungus. One type of fungus forms a sheath-like
structure at the root tips through which it passes to the tree various
nutrients it has gathered from the soil in exchange for food the tree has
produced through photosynthesis (remember, fungi are not able to produce
their own food in this way).
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In fact almost all vascular plants (which excludes mosses, other fungi
etc) benefit in nature from a mycorrhizal association of one kind or another.
Although mycorrhizae are by no means essential to the well-being of any
plant, their associations are of tremendous benefit in less than ideal circumstances.
For example, a tree planted in fertile, moist yet well-drained soil with
a good supply of readily available nutrients will already be growing at
its maximum rate with maximum health, and so has little need of mycorrhiza.
Indeed, as we will see later, mycorrhizal fungi would probably not survive
for long in such conditions anyway. On the other hand, trees planted in
marginal conditions would probably not survive without a mycorrhiza and
it is in these conditions that mycorrhyzae will thrive.
This begs the question; "Is bonsai soil in a bonsai pot
ideal or less than ideal?" The answer has to be less than ideal. Bonsai
containers provide "marginal conditions" for any tree and it's only
the dedication and knowledge of the grower that enables the tree to thrive. In a
bonsai pot the roots are subjected to the extremes of temperature - becoming as
cold as the ambient temperature in winter and as hot as a tin roof in summer.
They are also subjected to daily drenching and drying during the growing period.
The soil is largely inert or lifeless (Akadama, baked clay, grit, pumice) and
any nutrients are rapidly leached out with daily watering. These are exactly the
conditions in which mycorrhizae can be of profound benefit to plants.
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Whatever happened
to mycelium?
When mycorrhiza first came to the attention of bonsai growers,
there was much misguided talk of a fungus called "mycelium".
There's no such thing.
Fungi comprise fine tubular structurescalled hyphae. These
become modified to perform different tasks under different names. They fuse
together in large numbers to form fruiting bodies called mushrooms, they
group in smaller numbers and extend through the soil under the title of
rhizomorphs, they also extend singly increase colony size and to propagate
under the name of mycelium. |
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Ectomycorrhiza
on juniper roots
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Are Mycorrhizae
species-specific?
Not as a rule. There are a few mycorrhizal fungi species that will only
associate with one host species such as Orchids, but the vast majority have a broad range
of potential hosts. Likewise, virtually all plants - and almost certainly
all trees - are perfectly happy to form mycorrhizal associations with a
number of different fungi, the eventual choice being dictated largely by
which fungi are available in those particular soil conditions.
A closer look
Types of Mycorrhiza
There are six kinds of mycorrhiza: Vesicular Arbuscular
(Endomycorrhiza), Ectomycorrhiza, Ectendomycorrhiza, Arbutoid, Monotropoid, Ericoid
and Orchid. Of these six, there are two major types of mycorrhizal fungi based on the anatomy of their
association with the host roots: ectomycorrhizae and endomycorrhizae.
An important fact about mycorrhizal fungi is that they are
not all equal in the benefits supplied to plants. Some species deliver
more benefits to plants than others.
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Ectomycorrhizae typically grow in the intercellular spaces of the root
cortex (outer layer or "skin") and for a thick mantle of tissue
around the exterior of the root tip. Some hyphae (fine, thread-like filaments)
extend out from the roots and into the surrounding soil to gather water
and nutrients. The network of intercellular filaments, the Hartig net, forms
the exchange sites where the host swaps carbohydrates for nutrients from
the fungus. Ectomycorrhizae occur primarily on members of the Pinaceae,
Betulaceae and Fagaceae families.
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| Left:Transverse section of an ectomycorrhiza on roots of Pseudotsuga
menziesii showing the mantle (deep yellow). Right: Cross section
of a root cell completely surrounded by hyphae of the Hartig net. |
Endomycorrhizae grow mainly inside the cortical cells (INTRAcellular
spaces. These don't form any external mantle so they are impossible to detect
with the naked eye, but they do also send out extensive hyphae into the
surrounding soil.
Some endomycorrhizae form structures called vesicles and
arbuscules within the root's cortical cells. These are known, naturally enough,
as vesicular-arbuscular mycorrhizae, or VAM for short. This is the type of
mycorrhiza we find on 90% of the world's higher plant groups. The arbuscules are
tightly bunched hyphae which take carbohydrates from the cells, growing as they
do so. Once they have completely filled the cells, they break down, releasing
their nutrients to the host and the fungus proceeds to colonize another cell. Vesicles
are thick-walled structures of varying
shapes, from ovoid, irregularly lobed
to box-like, depending on the species
of fungus and where the vesicle is
formed. They contain abundant
lipids and numerous nuclei and it is
likely that they are important storage
organs and may play a significant role
as propagules within root
fragments. Nevertheless, little
is known of their biology, in
particular with respect to either
germination or mobilization of the
reserves.
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Talking
trees?
There are instances where trees appear somehow to communicate
with each other.
One involves a tree which is preyed upon by an insect which
strips the tree bare faster than it can regenerate foliage, usually killing
it. As soon as one tree in a cluster becomes infested, all the others immediately
begin to secrete a substance toxic to the insects, thus keeping them away.
Scientists now believe that the chemical message is passed
through the mycorrhiza which commonly become bonded with mycorrhiza from neighboring
trees.
Another involves a plant phenomenon called aggressive
competition, where one species won't allow any other to grow near it.
Researchers are now looking into the possibility that mycorrhizae are also
responsible for this.
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The group called Ectendomycorrhiza, as you might have
guessed, combines some of the features of the Ecto and Endomycorrhizae
groups.
How do Mycorrhizae get there?
First, the potential fungal symbiont must produce viable mycelium (see panel on
Anatomy of Fungus, below) in the vicinity of the roots of the potential host.
Usually this involves the germination of either spores or "resting"
hyphae or host plant root fragments. This mycelium must then find its way to the roots of the host, which it
does not entirely by chance. The area of soil around a plant's roots - the
rhizosphere - contains millions of minute organisms (micro flora) which are
influenced by the presence of the roots. By detecting this, the mycelium can
navigate its way to the roots remarkably efficiently. Having reached the roots,
the mycelium must penetrate them.
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Reproduction
VAM reproduce from chlamydospores, which are long-lived,
thick-walled spores, produced by the fungus near the surface of the host root,
and are able to withstand the rigors of underground life until the roots of a
potential host grow close by.
Ectomycorrhizae can reproduce from spores or vegitatively from various
types of clusters of hyphae or from resting hyphae.
In all cases, germination is stimulated by near proximity of
roots of a potential host plant, via their effect on the micro flora in the
rhizosphere.
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Some you can eat
Although many mycorrhizae are microscopic, producing single
spores (chlamydospores), others produce quite large fruiting bodies. Miniature
fungi that appear in the pots of birch, hornbeam, larch and spruce, may well
belong to ectomycorrhizal fungi.
The most famous ectomycorrhizal mushroom is the truffle. In
nature animals feed on these and the spores pass through the body to be
distributed around the forest as and when...!
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Benefits of Mycorrhizae
Research is continuing into the many benefits to plants of mycorrhizal
associations and there are probably many yet to be discovered. However,
in the light of current knowledge, benefits can be divided into six categories.
Water and Nutrient Uptake
Mycorrhiza greatly increases the roots efficiency at nutrient
and water uptake largely because of the vastly increased absorptive surface
area. The combined surface area of the millions of hyphae is far greater than
that of non-mycorrhizal roots. Increased water uptake by mycorrhizal plants is
due to physiological improvements which are made as a result of increased
nutrient uptake into the plant. The water uptake is through the roots, not
the mycorrhizal fungi. In addition, the extending hyphae are able to draw
on more distant or inaccessible supplies of nutrients from than the roots can
reach. (Bear in mind that the rhizosphere is always in effect nutrient deficient
by virtue of the presence of the functioning root which has taken the available
nutrients!)
Using radioactively-labeled nutrients, scientists have
shown that ectomycorrhizae are especially clever at absorbing phosphate
and potassium as well as alkali metals. VAM were shown to be efficient at
absorbing phosphorus, copper, iron, zinc and calcium. Potassium
uptake by VAM is also indicated, but is governed by levels of nitrogen,
potassium and calcium that are present.
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Alleviation of Stress and Disease
Environmental and cultural stresses influence the plants susceptibility to and
ability to combat bacterial diseases and are known to actually cause some
non-bacterial diseases. VAM greatly reduce the environmental stresses -
nutritional (too much or too little), drought, root pathogens, soil toxicity etc
- which predispose a plant to disease. The increased uptake of nutrients,
particularly micro-nutrients which are "locked" to soil particles and
unavailable to the roots, make the plant less susceptible to the ingress of
plant pathogens, and more resistant to other environmental stresses such as cold
and heat.
Protection Against Root Pathogens
Ectomycorrhiza, in particular, have recently been shown to resist attack by
soil-borne pathogens. For example, there are several mycorrhizal fungi known to
protect pines from pathogens such as phytophthora, Fusarium and Rhizoctomia.
There are several mechanisms by which this occurs, many of which are thought to
operate simultaneously.
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production of antibiotics by the fungus itself, which
inhibit root pathogens
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the physical barrier created by the mantle of
ectomycorrhizal hyphae
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production of chemical inhibitors by the host, induced by
their reaction to invasion by the mycorrhizal fungus
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the establishment of populations of protective microbes in
the rhizosphere.
Altered Root Physiology
Researchers have demonstrated that ectomycorrhizae produce growth hormones
and regulators which are responsible for the altered metabolism and growth
of the roots themselves. These substances enhance the ramification of root
tips, the proliferation of roots, enlargement of cells, and enhanced
rooting of cuttings.
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Fossilised structures
from the
roots of early land plants thought
to be vesicles of an early
endomycorrhizal fungus
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Jurassic Mycorrhiza
Phyto-archaeologists have discovered correlations between
the geologically sudden disappearance of tree species from large areas with
the disappearance of evidence of associated mycorrhizae. The interesting
aspect is that the mycorrhizae seems to have disappeared first, indicating
the potential long-term effect of mycorrhizal deficiency. |
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Detoxification of Soils
This is still a very sketchy area, as research is still in the early stages.
However, scientists are now investigating what appears to be the capacity of
mycorrhizae to assist plants to colonize soils which would otherwise be
chemically toxic to the plants.
Maintenance of Soil Structure
Mycorrhizae accelerate the decomposition of primary minerals and secrete
organic 'glue' (extra cellular polysaccharides) which bond the finer soil
particles into larger, water-stable aggregates.
Significance of
Mycorrhizae in bonsai culture
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If your bonsai is in a pot that is large enough, with a suitable soil
and an adequate regular supply of water, nutrients and micro-nutrients,
it's probably in reasonable health and growing well. But that doesn't necessarily
mean it's performing to its full potential. One can get so accustomed to
experiencing early autumns, weak second growth flushes, mid-summer shut-down,
poor leaf condition in late summer and so on, that it becomes accepted as
the norm.
We're familiar with the benefits to pines from mycorrhiza, but lets see
how they can benefit bonsai in general by looking at the five points again.
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A portion of endomycorrhizal
mycelium bearing some of the
very large spores |
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Water and nutrient uptake
Newly repotted or collected trees don't have access to the entire growing medium
simply because their roots don't fill the container. Mycorrhizal hyphae will
extend from the existing roots throughout the container in a fraction of the
time it would take non-mycorrhizal roots, thus utilizing all available moisture
and nutrients. They also regulate the rate of nutrient uptake, thereby reducing
the danger of 'root burn'. Later in the season, when the tree's water demand is
higher, mycorrhiza can still help, even though the pot appears to be full of
roots. Many soil ingredients such as the harder grade of Akadama, calcined
(baked) clay, pumice and even some bark chips, are impenetrable to roots. The
hyphae, however, are able to penetrate the micro-pores in these particles and
and retrieve nutrients and micronutrients stored therein. In addition, they
supply these to the tree in a form which the tree can use immediately.
Alleviation of stress
Bonsai, by definition, are always under some form of stress (albeit controlled,
hopefully) and this is made all the more significant with the increased usage of
non-organic, inert growing media and synthetic fertilizers. Therefore, bonsai
are more susceptible to serious damage by disease and stress-related disorders
than field-grown trees. Symptoms such as mid-summer shut-down and early autumn,
or discolored and tired foliage are all indications of stress or stress-related
disease. In fact, if was only one category of plant crying out for the
additional protection offered by mycorrhizae, it would be bonsai.
There is evidence to suggest that VAM increase
resistance to some viral diseases, but no evidence they can decrease occurrences of viral
diseases. In certain cases they can limit
the severity of attack.
Protection against root pathogens
Traditional bonsai wisdom states that if there's a problem with the tree's vigor,
the cause is in the roots. Not all root pathogens are fatal - but more become
fatal in a bonsai container than in the field, simply because of the slow rate
of root growth and absence of the roots of other plants. Good tool, pot, soil
and water hygiene, plus the choice of reputable organic fertilizers, should
prevent most soil pathogens from entering the container. However, some are
air-borne and can arrive at any time, but many of these are unlikely to become a
danger in a good bonsai soil. That still leaves the the few that could become a
danger. The added protection afforded by mycorrhiza could give the bonsai grower
the confidence to say that if there's a problem with the tree, it's probably NOT
caused by the roots.
Altered root physiology
Increased ramification, increased root proliferation, enlargement of cells
(greater efficiency) and enhanced rooting of cuttings. Need more be said?
Detoxification of soils
Once again, good soil and water hygiene should eliminate the possibility
of accidental toxicity of bonsai soil. But there is some concern that calcined
clays and other mineral soil ingredients can accumulate a toxic level of
salts which could eventually harm the plant. If mycorrhizae can assist here,
and it's not yet certain that they can, then better with than without!
Maintenance of soil structure
As bonsai soil's organic matter - as well as it's akadama, loam or clay
content - naturally breaks down into fine particles, they are re-bonded by
the mycorrhiza's polysaccharide secretion, thus maintaining an open,
free-draining and well aerated soil. You may have noticed how the soil in
the pot of a pine with mycorrhiza is more 'friable' and granular than that
in the pot of a non-mycorrhizal pine.
Mycorrhiza
inoculation
We all save some old mycorrhiza from our pines, and re-introduce it into
the new soil when we repot. Does this work? Well, yes it does.
In fact there could well be enough spores, chlamydospores,
sclerotia, rhizomorphs and resting hyphae left on the remaining roots to
colonize the pot ten times over. But because you have pruned away the root tips,
where the mycorrhiza forms, and your loose, granular soil has left you with an
almost bare-rooted tree, you can never be sure, so re-introducing it is a very
good idea.
The same goes for other species with endomycorrhiza, which you
can't see. Re-introducing chopped-up pieces of the pruned-away root tips will
help to ensure re-colonization of the pot.
However, there is one other important point. Remember we
discovered that when the fragments or spores germinate, they are
stimulated to do so by the microbial changes in the rhizosphere - which
you don't have in your new soil and clean roots. The roots that the
inoculated mycorrhiza is adhered to are now dead. One answer is to make
sure that, when you introduce the chopped-up mycorrhizal roots, they are
in good close contact with living FEEDER roots. Another is to include a
proportion of the previous soil in your new mix. Since the entire pot was
probably completely filled with roots, practically all the soil would
qualify as rhizosphere. Research has demonstrated that
endomycorrhizal fungi occurs naturally at low levels. For plants benefited
by endomycorrhizae, it can be more effective to apply a good commercial inoculum
on an annual basis rather than trying to rely upon chopped-up mycorrhizal root
fragments.
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