Taken from Slayers, Saviors, Servants, and Sex: An Exposé of Kingdom Fungi by David Moore; published by Springer-Verlag, New

York: 2001, ISBN 0387950982.

Chapter 6 Turning the tables. Using fungi to control other pests

Just about everything we do is affected by pests of one form or another and over the years we have
invented an armoury of chemical pesticides which have permitted enormous improvements in
agricultural and horticultural yields. But most of these chemicals are powerful and indiscriminate
poisons and worries over the adverse environmental impact of heavy usage of chemicals like these
are increasing. As we have seen so far in this book, fungi are very effective pests of other creatures
in their own right so it’s not surprising that attention is turning to the use of fungi as control agents
by harnessing their natural antagonisms to pests of our crops - and there is potential for us to use
fungi in controlling other fungi that cause diseases of crops, as well as insect pests, nematode
worms and even weeds.

Over two thousand years ago, the Chinese were writing about the fungus diseases of silkworms and
cicadas. So the importance of fungi that infect insects (they’re called entomogenous fungi) in
natural populations of insects has been recognized for a long time. It’s interesting that early support
for the germ theory of disease came from a fungus disease of silkworms. The disease was called
muscadine disease and it killed silkworms in Europe so effectively that the future of the silk
industry there was in peril. A lawyer-turned farmer named Agostino Bassi found that the silkworm
corpses produced masses of fungus spores. The fungus, now called Beauveria bassiana in Bassi’s
honour, is one of today’s prime candidates as an alternative to chemical pesticides. Bassi is worthy
of honour. The people who are remembered by histories of biology and medicine as establishing the
germ theory of disease are Pasteur, Lister, and Koch. Great men indeed, but they did their work in
the 1870s, and Bassi had already settled the issue in 1835/36 (ten years before Berkeley’s work on
the Late Blight of Potato discussed in chapter 2). Unfortunately, choosing to work on insects and
fungi is a sure way of being forgotten, then as now, and Bassi never got the credit he deserved. Still,
they did name the fungus after him! The idea that these fungi might be useful as biological control
agents was first explored by two pioneering scientists in the Ukraine, Metchnikoff and
Krassilstschik, in the 1880s. These two visionaries mass-produced the spores of an insect-attacking
fungus (called Metarhizium) and tested it against insect pests of wheat and sugar beet. Throughout
the twentieth century insect-infecting fungi have been assessed as possible control agents for a
variety of insect pests. There’s a repetitive pattern. Field experiments are carried out, products are
developed, and may even be successfully marketed for several years. But then the fungal products
are replaced by more effective chemical insecticides. The sequence of development,
commercialization and later withdrawal has been repeated with a number of other fungal products.
Currently, there are no fungal products which are widely used for insect control, although in
particular parts of the world (including Russia, China, Brazil, and the UK) and, more importantly,
for particular insect pests in very particular circumstances, there are fungal products which are in
use.

During recent years, of course, there has been a renewed interest in these insect-attacking fungi
because of increasing insecticide resistance and environmental concerns over pesticide use. New
strains of fungi have been isolated from a wide range of hosts, and these have even further
emphasized the potential that exists for the use of fungi in insect control. But the continuously
repeated cycle of development of a promising fungal product followed by its eventual failure has
emphasized the importance of ecological features in determining their success as biological control
agents. The problems encountered relate to the biology and life style demands of the fungi which
we try to use as control agents. Over one hundred different sorts of fungi (representing all the main
fungal subdivisions) have been shown to parasitize living insects. They are more common in
tropical areas where temperature and humidity favor their growth. But they occur in most
ecosystems, ranging from natural water habitats to the high technology horticultural production
systems we create around the world. The host range of individual fungi is variable, with some

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© David Moore 2001
Taken from Slayers, Saviors, Servants, and Sex: An Exposé of Kingdom Fungi by David Moore; published by Springer-Verlag, New
York: 2001, ISBN 0387950982.

attacking only one insect whilst others, and Metarhizium is an example, have a broad host range.

The key controlling feature that determines the usefulness of these natural control agents is their
dependence on humidity for germination and growth. For them to attack their insect prey their
spores must germinate on the insect and the processes of spore germination and growth of the
spores on the surface of the insect are highly dependent on both available moisture and temperature.
Even ninety percent relative humidity might result in only half the spores surviving. In the real
world this means that the microclimate in the tiny area that the host insect prefers might determine
the success or otherwise of the control agent. For example, in attempts to control aphids in
glasshouses there was a major difference in efficiency against aphids which fed on the undersides of
leaves (very effective control) where the humidity is high, compared to the more exposed stem
(virtually no control). In similar experiments the spread of infection through aphid populations was
maximal when free water was present but completely prevented at ninety-three percent relative
humidity. Still rather damp to you and me, but dry as a desert to the fungal spores. Temperature also
affects germination and growth. Both are markedly reduced below fifteen degrees Celsius and
above thirty-five degrees, so the window of effectiveness is fairly narrow.

Once the fungus spore has germinated and grown through the insect’s skin it produces cells which
circulate within the insect and proliferate by budding. Insect death usually occurs three to fourteen
days after the fungus spore alighted on it. Death results from a combination of mechanical damage
resulting from tissue invasion by the fungus, loss of nutrients to the fungus and reaction to toxins
generated by the fungus. After the death of the insect, the fungus grows on its corpse, eventually
producing a new crop of spores. So the initial application of fungus spores results in the production
of even more spores after the first successful infections. It is this amplification of the control agent
that makes the biological control mechanism so attractive. A chemical agent will be diluted, washed
away in the rain or used up. But as long as there are hosts to infect the biological control agent will
grow stronger and stronger, producing an epidemic that can destroy the target insect completely.
Unfortunately, this promise is rarely realized, though there are sufficient successes on the record for
us to keep trying.

Rice pests are potentially good targets for these insect-attacking fungi; the high humidity in rice
paddies and warm temperatures in rice growing areas combine to produce close to ideal conditions
for epidemic infection of the insects. Attempts to control rice pests in the field by applications of
fungal spores have indeed been relatively successful. Pests of glasshouse crops can also be
controlled well by biocontrol agents. This is probably the most ideal circumstance because, being
within the glasshouse, the environment can be controlled to optimize the infestation. Even here,
though, the environmental demands of the parasites can cause difficulties. For example, control of
whitefly on cucumber crops is effective only after the crop has reached four weeks old. This is
because the smaller leaves of younger crops greatly reduce the humidity around the plants and the
fungal spores cannot germinate. The age of the insect pest can influence results, too. Ninety-four
percent kill was obtained when eggs of the white fly were treated (because the first larvae are then
infected as soon as they emerge from the egg), but only twenty-eight percent of older larvae were
killed when the treatment was delayed. The fungus spore preparation has to be applied several times
to maximize and the spray must cover the entire plant to get good spread through the whitefly
populations because the larvae themselves are not sufficiently mobile to spread the fungus disease.
If all these requirements can be met, the development of fungal infections in the whitefly when
humidity is high is very rapid and it is possible that a single night of high humidity in a glasshouse
would be enough to destroy a whitefly infestation. There are no chemicals and no chemical
residues, so biocontrol of whitefly, although difficult and demanding, is very promising.

Another successful application of fungi is in the biocontrol of the Lucerne aphid in Australia. This

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© David Moore 2001
Taken from Slayers, Saviors, Servants, and Sex: An Exposé of Kingdom Fungi by David Moore; published by Springer-Verlag, New
York: 2001, ISBN 0387950982.

pest was introduced to Australia with none of its natural enemies and so it rapidly became a major
problem. A search for natural Lucerne aphid disease agents found several likely pathogens in Israel.
One of these was introduced into Australia; it spread rapidly and now helps to maintain the pest
population at a low level.

Beauveria bassiana, that belated monument to Agostino Bassi, has been tried as a microbial control
agent of several insect pests in different parts of the world, although most development work on this
organism was done in the then Soviet Union where it was mainly used for control of Colorado
potato beetle. Field trial showed that it was more active against weakened insects and the standard
treatment developed was to use the fungus in conjunction with a quarter of the usual rate of
insecticide application. The reduced dose of insecticide weakens the insects to such an extent that
the fungus can easily infect and kill virtually all of them. In the 1970s through to the 1990s
preparations of Beauveria bassiana spores were used on several crops over an area of at least half a
million hectares in China. The fungus was used to help control pests like the corn borer, pine
caterpillars and leafhoppers. Biocontrol agents of this sort are particularly useful in peasant farming
and commune-based agricultural systems because the peasants can produce their own supplies of
fungal spores. In this case, spores were produced by growing the fungus on boiled rice. This puts
production of the control agent into the hands of the community which needs it in a way which
simply cannot be done with chemical insecticides, which usually have to be bought with hard-
earned cash from multinational chemical companies. This ‘empowerment’ of the community is such
an attractive proposition that it may outweigh any lesser effectiveness of biocontrol agent when
compared against the chemical treatment. There’s a similarly successful example from Brazil. The
fungus known as Metarhizium is used commercially in Brazil to control spittlebug of cane sugar
plants. The fungus spores are produced by growing the fungus on sterilized rice and once a good
crop of spores have appeared the rice grains are dried and grains and spores are ground up and the
powder sold under a variety of trade names. Again, this process is very simple and is well suited to
production on a local basis by grower co-operatives. The yield of spores is very good, about a
million-million spores per kilogram of rice, which is enough to treat a whole hectare of cane sugar.
Spittlebug control is one of the few major success stories in biocontrol. Conversely, trials of the
organism Beauveria bassiana in the USA have been less successful. During a three-year test,
control of the potato beetle was highly variable. In only eight out of twenty-four trials was potato
yield from fungus-treated plots significantly greater than yield from control (untreated) fields, but
only two of these gave yields on a par with insecticide treatment. It seems likely that the more
industrial farming practices result in high pest numbers at the time of fungal application and that
this reduces the effectiveness of the fungus.

As the stories above show, there have been several major successes in particular places, under
particular circumstances and with particular pests. These successes are a spur to continued research,
but there has been no really dramatic breakthrough. Nothing has been found that has wide
effectiveness and can be used under a range of conditions. What is most urgently needed to set this
topic on fire is a fungal formulation that is not only safe to use (the common characteristic of these
agents) but also cheap to produce, easy to apply, not too dependent on specific environmental
conditions and that can consistently control the target pest.

Unfortunately, we are not very close to being able to achieve that. There are no commercially
available ‘mycoinsecticides’ in the USA or Europe which get anywhere near this ideal. The history
of the topic is littered with products being launched in great hope and then being withdrawn because
they failed to provide reliable pest control. Astonishingly, despite this disappointing experience,
commercial interest continues and it is remarkably widely accepted that fungi have the potential to
control insect, aphid and other pests. It seems to be a topic in which blind optimism wins out over
bitter experience. I suppose that to a great extent what drives this blind optimism are the possible

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© David Moore 2001
Taken from Slayers, Saviors, Servants, and Sex: An Exposé of Kingdom Fungi by David Moore; published by Springer-Verlag, New
York: 2001, ISBN 0387950982.

advantages of mycoinsecticides compared to conventional chemical insecticides. Fungi infect most

sorts of insect, aphid and mite, but individual strains of fungus can be very specific and may only
infect one type of host. So, you can imagine fungi being used to control important pests without
affecting innocuous insects in the same environment - that’s part of their inherent safety. Of course,
insect-attacking fungi cannot infect any other type of animal (or plant, for that matter), so unlike
some of the more toxic chemical treatments the environmental impact of a mycoinsecticide is
strictly limited to the host species it was developed to control. Another advantage over chemicals is
that there is no evidence so far of fungal resistance occurring in insect populations. Now, this might
be due to the fact that they’ve been used to only a limited extent so far, but it is a sharp contrast
with chemicals which seem to select out resistance strains of the pest very quickly. The holy grail,
therefore, is that widespread mycoinsecticide use could provide great benefits to us and our
environment. Like the real holy grail, the problem is to find it.

The generally disappointing experiences of the past hundred years or so, and to be fair the few
successes, do give some indications as to what directions we need to take in the future to realize the
high hopes which so many people have for mycoinsecticides. The first point is the crucial
importance of selecting the right strain of fungus. There are an enormous number of fungi out there
and just about all features of the pathogen-host relationship can vary greatly between strains of any
one species. By searching through a large collection of different strains, one with the desired
combination of characters will be found. Eventually! Molecular biology might help. Genetically-
modified pathogens of pest insects might be designed which bring together toxins, virulence or
other pathogen functions from organisms which cannot be mated together conventionally. Progress
is being made in understanding pathogenicity mechanisms, particularly things like how the fungus
first penetrates the outer defences of the host, so it might become possible to increase the speed of
insect kill by genetic manipulation of the fungus. This is important because most mycoinsecticides
can only be applied after the crop has become infested by the pest. That being the case it is essential
that the pest is killed rapidly, otherwise you’ll end up with the unsatisfactory situation of killing the
pest after it’s done its damage.

Once you’ve selected, engineered and genetically modified a really mean insect pathogen, you need
to be able to produce it on a large scale if it is to become a commercial proposition. The most
attractive production method is liquid fermentation using stirred tank reactors. The engineering of
this technology is well understood because it is used for other useful products like citric acid and
antibiotics. However, as I’ve explained above, two of the most successful examples used a method
using cereal grains as a semi-solid medium to produce the fungus spores. This is a traditional way
of producing several human food products so it is a type of production that offers possibilities, but
each new product requires new research so it can be a slow development program. Another
commercial requirement is that the product should be easy to store. One expert has suggested that a
storage life of more than18 months is necessary for a commercial product. Few farms around the
world can afford refrigerated and/or humidity-controlled storage, so we’re talking about storage in a
barn or shed with no special precautions. These aspects of the ‘recipe’ can be controlled by the
nature of the formulation with which the fungus is mixed to make the commercial product. The
formulation can also assist initial infection if it includes materials which help maintain favourable
humidities while the spore germinates on the host. There is a limit to this, of course, and then the
product is at the mercy of the real world, depending on atmospheric factors and the density of the
host population to achieve the desired catastrophic infection of the target organism, prolonged pest
control, reduced risk of resistance and a high degree of safety to non-target organisms with all the
associated environmental benefits. It’s a long job.

It’s not only insects that are pests. Weeds pose a large problem in both agricultural and natural
ecosystems, not just in gardens and amenity sites in urban areas. Most people would agree that an

4
© David Moore 2001
Taken from Slayers, Saviors, Servants, and Sex: An Exposé of Kingdom Fungi by David Moore; published by Springer-Verlag, New
York: 2001, ISBN 0387950982.

insect which eats its way through the leaves of a plant is something of a pest, but weeds are not that
easy to spot. What is a weed to one person is a wild flower to another. A plant might be designated
as a weed because it has a detrimental affect on agriculture or amenity plantings, but the same plant
may actually benefit beekeepers, wild flower and fruit gatherers, herbalists and others. Not
surprisingly, attempts to control the ‘weed’ may generate heated arguments! There’s an interesting
story about a plant called Chromolaena odorata from Brazil which became what is known as a
pantropical weed causing a biologically devastating invasion of Ghana and the Côte d’Ivoire in
1993. In Brazil, where the plant was native, it was not considered a weed, and though classed as
common in Brazil, it never formed the dense, continuous stands which characterized its growth
pattern in West Africa (and Asia). The plant caused significant agricultural and environmental
damage over a large area of West Africa and Asia, and was suggested as a good candidate for a
biocontrol programme. This led to an outcry, however. Some people suggested using chemical
herbicides, despite the enormous area that would have to be sprayed. But others argued that the
weed should not be controlled at all because the plant had medicinal properties and because it
formed a rapid plant cover in the ‘slash and burn’ agriculture of the region. The counter arguments
were put so strongly that funding was withdrawn from the proposed biocontrol project.

Agricultural losses due to weeds are difficult to estimate but there is broad agreement that at least
ten percent of the world’s agricultural production is lost each year. For the US this means that
annual crop losses due to weeds range up to thirty billion US-dollars. Estimates like this do not
include the negative effects of environmental weeds in natural ecosystems. Another measure of the
importance of weeds is to estimate how much people are willing to spend to get rid of them by
using herbicides. Herbicide sales account for about seventy percent of all pesticides sold (the
proportion was only twenty percent in the early 1950s) and have a retail value of around twenty
billion US-dollars. Invasions by weeds which are ‘foreign’ to a region have become a particular
problem as international travel and commerce have increased. Such introductions (usually unwitting
and unintentional) can invade native vegetation and out-compete the resident plants so disrupting
the balance of the whole ecosystem.

The recognition of fungi as important natural enemies of weeds which might be safely exploited to
our benefit is not new, but it wasn’t until the 1970s that they began to be used in serious attempts at
weed biological control. The approaches used for control of insect pests are just as valid for
biocontrol of weeds with fungi. Where the target is a plant which has become a pest outside its
natural range by being introduced into some ‘exotic’ area, then the best strategy is to introduce a
fungal natural enemy of a target weed from its native range into the exotic area. An alternative
strategy is to use fungal pathogen from the exotic area but to mass produce them and apply at such a
high concentration that the weed population is inundated with fungal disease. Just as with attempts
to control insect pests, it is a long job and is dependent for success on enormous amounts of
scientific research. It is also essential to make proper economic evaluations of the project. No
matter how attractive the idea might be from scientific and conservation points of view, someone
will have to foot the bill and economic feasibility and cost/benefit analyses must be done.

At first glance the use of fungi in weed biocontrol seems like enlightened management which is
likely to solve important problems and contribute towards sustainable agricultural development. But
the truth is not quite so clear cut. Yes, you can find numerous examples of weed infestations which
have caused major problems like disrupting fishing activities, harbouring disease vectors, and major
crop losses. In some instances the effects have overtaken whole cultures and led to agriculture being
abandoned or major population movements to escape the problem. No one can deny that weed
control (with fungi or with chemicals) in such dramatic cases is desirable in social and cultural
terms. But these are the dramatic extremes which are, thankfully, rare. In most places, most weeds
are just a few plants growing out of place in a field or other cultivated plot. Careful husbandry

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© David Moore 2001
Taken from Slayers, Saviors, Servants, and Sex: An Exposé of Kingdom Fungi by David Moore; published by Springer-Verlag, New
York: 2001, ISBN 0387950982.

should be able to cope with them. In fact, weeding is still one of the most common activities of a
large part of the human population. So think of it this way: in a sense, weeds could be seen as being
beneficial because they generate honest employment. You might counter that by pointing out that
physically removing weeds is such a demeaning human activity that it is a waste of human
resources. Freeing the human population from this burden then becomes a noble goal for scientists
and technologists alike. The attractions of the ‘honest toil’ argument are deceptive, indeed illusory.
There’s a direct analogy with the development of industrial machines which change the nature of
the industrial shop floor. Supplementation, and eventual replacement, of human toil is the desirable
aim and herbicides have the capacity to do that. When mycoherbicides are common in the
marketplace they will expand the options available to major users of herbicides.

Control of weeds with mycoherbicides is an ecologically ‘clean’ method. It uses a natural agent and
leaves no chemical residues. There are concerns that fungal plant diseases which are used as
mycoherbicides might also pose a threat to crops or native plant species. Knowledge is the key,
here; particularly knowledge about host specificity, disease severity and possible tolerance to the
disease. Paradoxically, a fungus that is capable of attacking a native or even a crop plant may have
potential as a biocontrol agent under particular circumstances. It’s then a matter of weighing the
cost of causing disease in native plants against the benefit of using that same disease to control the
weed. If the weed population is less tolerant of the disease then the balance shifts in favour of using
the fungus to control the weed. This type of consideration applies to chemicals, too. Most pest
control chemicals are toxic to a wide range of organisms - the useful ones are more toxic to the pest
than to the crop. It’s a matter of scientific research again. The risks of plants other than the target
weed being affected by an introduced mycoherbicide fungus can only be judged if appropriate
scientific knowledge about the mechanisms of fungal host-specificity is available.

The potential economic value of mycoherbicide is becoming easier to judge as more trials are
completed. Introduction of a rust fungus into Australia to a weed which originated from the
Mediterranean region was certainly biologically effective. It resulted in better than ninety-nine
percent reduction in infestations by the weed. But the estimated annual saving due to increased crop
yields and reduced chemical herbicide use was sixteen million Australian dollars. Since the cost of
the whole project was only three million Australian dollars, the return on this initial scientific
investment has been enormous. Unfortunately, benefits on this scale are not often seen. The
mycoherbicide ‘business’ has a history, similar to the mycoinsecticide business, of products being
researched, brought to the market and sold apparently successfully for several years, but then being
withdrawn. Often the market is too limited to repay the high initial development costs and then
support the ongoing costs of further development to cope with competition. Several products which
were successfully sold in the 1980s were withdrawn in the 1990s because of the costs of registration
procedures imposed by environmental protection agencies.

This last point brings home the fact that a major obstacle to the introduction of biological control is
(unjustified) fear of disease spread. This ‘pathophobia’ has led to overzealous rules and seemingly
endless tests and trials. Oddly enough, this bias against introduction of fungal plant diseases as
biocontrol agents is not also applied to introductions of insect pests and most countries pay little
attention to precautions against new weed introductions. About half of the weeds in United States
and thirteen of the top fifteen weeds are introduced species. Similarly, seventy-eight of the one
hundred and seven most noxious weeds in Canada were introduced to the country. The best way to
control such ‘aliens’ is to introduce diseases from their home territories. But the regulatory
authorities make that process unreasonably difficult (and unrealistically costly). They do so despite
the fact that the overall rate of effectiveness that has been achieved with the fungal diseases which
have been introduced in the past, is sixty-seven percent. We’re often quite good at shooting
ourselves in the foot!

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© David Moore 2001
Taken from Slayers, Saviors, Servants, and Sex: An Exposé of Kingdom Fungi by David Moore; published by Springer-Verlag, New
York: 2001, ISBN 0387950982.

Plant parasitic nematode worms are also candidates as targets for using fungi as biological control
agents. These nematode worms spend part of their life cycle in soil or on the root surface where
they are exposed to nematode-attacking (called nematophagous) fungi that could provide useful
alternatives to chemical nematicides. Current evidence is that trying to enhance the activities of
resident nematophagous fungi in soils requires too much material to be generally acceptable. On the
other hand, adding biological control fungi to soils where they are scarce or absent could lead to
commercial products. But many problems remain to be solved before predictable control can be
achieved at practical rates of application. If nothing else, greater knowledge of the pest and its
interactions with soil, host and fungal disease agent might lead to identification of more acceptable
chemical nematicides.

One thorny problem that all biocontrol projects face is that of what to do about patenting the
biocontrol fungus. Commercial companies would expect to be able to get some patent protection for
any agent in which they have invested. A chemical is relatively easy - you can patent the chemical
itself, the way you produce it and whole families of chemical derivatives. You can then put the
material on the market, confident that even if a competitor buys a supply of the material they cannot
make more of it very easily. When the agent is a live organism, though, any competitor with a half-
way competent biologist would be able to grow more of that organism from even the smallest
sample of the commercial material. If the biocontrol agent is a genetically modified organism their
identity is not likely to be difficult to establish and the company which first created them will,
therefore, be able to identify and claim what rightly belongs to them. But suppose the biocontrol
agent is a fungus which was isolated from nature. Say, your staff collected fungi from a particular
site and found one of them to be especially virulent in controlling a specific pest. What’s to stop a
competitor going out to the same place and finding a related fungus that’s equally virulent? What’s
to stop an unscrupulous competitor growing up your fungus and simply claiming that it’s something
they’ve newly found? There’s no difficulty in writing patents, registrations or other legal devices
which assign ownership. The real difficulty is in having a catalogue of identifying features of the
commercialized organism which is sufficiently good to ensure that you could recognize it in any
competitive product. Genetic markers help a lot with organisms, like fungi, that have few
distinguishing features of their own. That’s why there’s so much interest in ‘genetic fingerprinting’.
These techniques really would allow you to identify that a competitor had stolen your organism. Of
course, you can never stop competitors going out to find a biocontrol organism of their own. The
thing to do in that case is to license your patented product for your competitor to sell at a price less
than the cost of developing a separate competing organism. Badge-engineering for fungi!

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