Chapter
3
One
Thousandth Of A Metre
We’re
coming back to familiar things now. The great blobs on the lens that were just
misty patches are finally slipping into some sort of focus, and a whole raft of
life forms are dropping by the wayside as we pull out and change our scale to
something far larger: the tiny fragments of plankton that fill the oceans; the
dusty fungal clouds in the evening air; the singular amoebas that live wherever
there is moisture – all wonderful subjects, but for another time. I just
can’t seem to get the focus right though: the nematodes are everywhere. Where
should I start?
What
about soybeans?
I have
just found out that The Society of Nematologists is advertising the 4th National Soybean Cyst Nematode
Conference. A whole conference about the egg filled bodies of a specific
nematode worm that affects a specific crop. This being the fourth one you might
be forgiven for accusing the organisers of being a little overenthusiastic –
maybe they are scientists ensuring they have their research grants for another
year; maybe they are companies trying to sell a product; maybe soybean cyst
nematodes are actually very important indeed. Actually it’s all three.
Scientists need to justify their work so they can keep on working:
unfortunately, whereas justification used to be on mostly scientific grounds,
justification in many modern universities requires evidence of commercial
potential. Pest control companies need to raise the profile of the “pests”[i]
they sell control products for, so they can sell their products to worried
consumers. Finally, according to the US Department of Agriculture[ii],
the cultivation of soybeans is not economically possible unless soybean nematode
cysts are sufficiently controlled.
A
glance at the literature on nematodes reveals two things: they are apparently
almost all damaging pests, and there are an awful lot of them. To answer the
latter point, many writers turn to the words of N.A.Cobb, legendary
nematologist, and stalwart of the US Department of Agriculture in the first half
of the 20th
century:
In short,
if all the matter in the universe except the nematodes were swept away, our
world would still be dimly recognizable, and if, as disembodied spirits, we
could then investigate it, we should find its mountains, hills, vales, rivers,
lakes, and oceans represented by a film of nematodes. The location of towns
would be decipherable, since for every massing of human beings there would be a
corresponding massing of certain nematodes. Trees would still stand in ghostly
rows representing our streets and highways. The location of the various plants
and animals would still be decipherable, and, had we sufficient knowledge, in
many cases even their species could be determined by an examination of their
erstwhile nematode parasites.[iii]
Stirring
stuff, indeed. Nathan Cobb could certainly move the soul when writing about his
foremost passion; and he needed to, because if ever a biological subject needed
a higher profile, it was the much-maligned, but utterly fascinating world of
nematodes. Cobb himself recognised this problem, writing: “[nematodes] offer
an exceptional field of study, and probably constitute almost the last great
organic group worthy of a separate branch of biological science comparable with
entomology.”[iv]
But was Cobb right? Do nematodes really create this film of organic matter
around every object in contact with the Earth?
There
is a certain difficulty in gaining realistic statistics about the variety and
quantity of nematodes; after all nematodes were not formally discovered until
1808, principally because they are too small to observe properly with the naked
eye. Victor Dropkin made a more sober assessment than Cobb of the nematode
population in 1980, stating: “Take a handful of soil from almost anywhere in
the world…and you will find elongate, threadlike, active animals. These are
nematodes. Or catch a fish, a bird or a mammal almost anywhere in the
world…and in most cases you will find some nematodes inside.”[v]
Although nematodes are aquatic animals, in that they need water to survive, the
best place to find them is in soil. Simon Gowen of the University of Reading,
tells his students that in temperate grasslands there are around nine million
nematodes for every square metre of soil – then the same
students are expected to count them for themselves (not all nine million of
them, I hasten to add), just to get an idea of what this means. That is an
astounding figure for something that is not a virus or a bacterium, but an
animal. This means that the lush grasslands of New Zealand that produce rich
butter, high quality lamb and 150 thousand tonnes of wool[vi]
each year, but only constitute 5.5 percent of New Zealand’s land area, also
hold something like 132,660,000,000,000,000 nematodes. That’s 132 quadrillion,
for those of you who ever wanted to know how large a quadrillion is. Compare
this to the apocryphal (but believable, and slightly disturbing!) figure of one
million spiders per acre of grassland, and you find that nematodes outnumber
spiders by 36,000 to 1.
Globally
– and I’m going to have to take an outrageous stab in the dark here – you
are probably looking at between 100 quintillion (that’s 20 zeros) and 1000
quintillion (21 zeros) nematodes on and in the land. To put this into
perspective somewhat: for each human on Earth, there are something like a
trillion nematodes. Nematodes in the oceans are far less abundant, but there are
still lots and lots of them – they may, in fact, account for ninety percent of
all life at the bottom of the sea[vii].
Sorry for boggling you with figures, but that’s what often happens in
nematode-land.
The
pest control industry ensure that the dangers that would be unleashed in a world
where nematodes are not controlled are writ large in the minds of farmers, so it
is the “pest” nematodes that are given the biggest exposure, at the expense
of other types. Despite the commercial world’s propensity to invent problems
in order to sell products they may, in this case, be right – but for all the
wrong reasons. The pressure we place on already exhausted soils and the effort
we go to in order to extract every last gram of nutrition from industrially
farmed crops to feed a growing human population (both in number and, in rich
countries, appetite), means that the slightest drop in the production of a
staple crop is treated as a potential catastrophe.
In
the majority of European countries, the impact of the potato cyst nematode (PCN)
is such that the movement of untested seed potatoes and the planting of potatoes
on untested land is banned, and the quarantine of land on which PCN is found is
mandatory[viii].
PCN is a global problem for potato growers, being found across Europe (since
1913, and possibly the 1880s), in Australia (since 1986), in the USA (since
1941) – in fact just about everywhere that potatoes are grown on a large
scale. There are quite a few varieties of potato that are naturally resistant to
the effects of PCN which, essentially, means not being in danger of having
entire crops wiped out within two seasons of growing on the same spot; and there
are lots of sensible, non-chemical methods of avoiding the problem, such as the
aforementioned quarantine, crop rotation and the use of natural predators. But
the chemical companies persist in pushing their wares, both in the form of
pesticides and genetically modified organisms (GMOs):
The
chemical group BASF has expressed optimism that within a few weeks the European
Commission may approve the genetically modified “Amflora” potato to be grown
in Europe. In early December, Hans Kast, Managing Director of BASF Plant
Science, spoke with journalists in Brussels and stated the expectation the
decision be made in any case early enough for the growing season of 2008.[ix]
It
won’t be too long before natural resistance to PCN is engineered into
non-resistant potato varieties. Now, I am no scaremonger when it comes to
genetic modification, but when economic gain comes before concerns for
environmental welfare – the GMO producing companies still obstinately refuse
to accept liability for any negative effects of their products – and the
number of discovered nematode species is less than ten percent of the number
that potentially exist in the wild, then I start to get a little worried.
Then
there is the question of pest versus friend:
Figure
4 : Nematode Adverts (Source: Author’s image)
Yes,
those are adverts from Google. It’s remarkable what is advertised on the
Internet: not so much the availability of salacious activities and products to
enhance your performance in all sorts of ways, but the wide range of friendly
nematodes that you can use in your garden, and can buy on line. Nematodes in a
box. I think it’s about time we stopped for a little and went back to first
principles.
What
Are Nematodes?
Remember
me saying that I had problems getting the focus right? There are teeny-tiny
nematodes and there are, relatively, very large ones indeed. I admit the title
of this chapter takes a few liberties, but there are many species of nematode
that are around a millimetre in length. There are many that are less than a
millimetre, and some parasitic types that are a few centimetres long. One type
(which no one alive seems to have seen) was measured at eight metres long, in
the placenta of a sperm whale. This species is, unironically, known as Placentonema
gigantissima.
Nematode
is the name given to any one of at least 20,000 species of unsegmented worm,
which have a single end-to-end digestive tract, no limbs or other appendages,
and a surprisingly well-developed nervous system, considering their antiquity.
They are commonly known as “roundworms”, which describes their cross
section, not their overall shape, and which distinguishes them from many other
types of worm, including flatworms and bristle worms.
Such
is their age and diversity (although this does not always follow) that they
occupy their own Phylum, separate from the arthropods (insects, spiders etc.)
and molluscs. There are as many as twenty different Orders of nematode, ranging
from those that attack plants and fungi, to those that feed on other animals, to
those that drift around feeding on whatever bacteria or single-celled animal
might be available. Despite nematodes being aquatic in origin, the vast majority
of Orders describe land-based varieties. This strange inconsistency is most
likely simply because the world’s oceans have been so poorly researched
compared to the land masses we are so familiar with. Our natural, possibly
ancestral attraction to the sea, a place that has always (until recently)
provided us with a rich source of food, only goes so far. Like gasping fish on
the water’s edge, humans immersed in water will only survive for a few minutes
– less if it is particularly cold. Maybe it is for the good that much of the
vast oceanic world has been left unexplored – the level of exploitation by the
oil, gas and industrial fishing industries is a dire warning of what can happen
– but it does leave us with a large empty space in our knowledge, and the
consequent skewing of information that suggests that the oceans are a vast,
barren place. Sadly, that lack of knowledge also hides the inexorable, and
possibly irreversible changes that we may be causing to the oceans.
Because
nematodes can look very similar, regardless of their size, they are most
commonly distinguished by their mouth-parts, which define what they are able to
eat. Nematologists do have what some might consider to be an unhealthy obsession
with mouth parts, but when you have a great mass of seething, wormy matter to
identify, then it’s usually best to take the easy route. Such efforts are not
without their rewards though: success in the field of nematology can be quite
lucrative if you don’t have any qualms about taking the corporate shilling.
The
multi-segmented tapeworm that can live for years inside humans and most other
mammals is not a nematode; human parasitic threadworms and hookworms, on the
other hand, are nematodes. I’m sorry to enter the bowel, as it were, at this
stage of the chapter, but if you have had anything to do with children’s
health or education then you will probably have come across threadworms.
Unfortunately for human health, children have a propensity to pick, scratch and
probe anything and everything with their fingers: noses, scabs on knees, eyes,
bottoms. Under the nails of a good proportion of kindergarten children just
about anywhere in the world lie a little cluster of threadworm eggs waiting to
be passed into the digestive system (you can guess how) of that child, or any
other person they may meet: “How do you do?” and with a shake of the hand
the eggs are passed on. Fortunately for us, threadworms are relatively harmless.
There
are a number of other parasitic nematodes that infect humans, and other mammals:
in dogs, hookworms can cause severe anaemia, and in both dogs and cats the
roundworm Toxocara is endemic. The latter is of particular concern to
humans because of the potentially severe symptoms that the resulting
Toxocariasis can lead to, including blindness and pneumonia. Studies carried out
between 1985 and 2000 found that children’s sandpits in public parks contained
Toxocara eggs a minimum of 25 percent of the time, with one study in Greece
finding 97.5 percent of play parks infected.[x]
A child who touches dog or cat faeces will almost certainly have nematodes on
his or her fingers. This level of infection may be shocking, but it is the
hygiene failures of humans that have turned something pretty benign into
something approaching epidemic proportions in certain parts of the world. This
lack of hygiene makes the little press nematodes get predominantly negative –
which is a shame because, like humans, not all of them are bad.
The
Good Guys
Good
organic gardeners know how to deal with pests – the kinds that damage the
crops they are trying to grow. A piece of fruit or a vegetable is at its most
appealing to birds, insects, slugs and snails at just the time when it is at its
most appealing to us; the sugar-rich strawberry, the fat-to-bursting pea pod,
the succulent red tomato, the crisp crimson and white radish – all perfect for
eating, regardless who the final consumer may be. Good organic gardeners don’t
need to spray chemicals across their gardens, to be caught by the wind and
misted across the neighbouring crops, flowers and ponds, and into the lungs of
playing children: they just need to understand the natural interactions between
plants, soil, weather and the organisms that may protect or attack what they are
trying to grow.
The
history of pesticides (that kill animals), herbicides (plants) and fungicides is
littered with toxins that no sensible person would let anywhere near their
mouths. We see arsenic being widely used for pest control on plants and even as
sheep dip; mercury, formaldehyde and hydrogen cyanide used to fumigate buildings
and glasshouses; and the surprisingly lethal copper sulphate applied as a common
weed killer[xi].
Paris Green derived its name from its colour and its use as a rat control agent
in the sewers of Paris in the 19th century. Alternatively known as Parrot Green and Emerald
Green, amongst other names, it is a compound of copper and arsenic, and is still
widely used as a barnacle prevention measure on the hulls of ships, and a wood
preservative, as well as an insecticide. Seven drops of this, surprisingly
unregulated, substance is enough to kill a normal sized human[xii],
and its death toll almost certainly includes many artists who keenly made use of
its vivid tones; not to mention the poor souls who made the stuff. It seems that
if a substance is useful enough then being lethal in tiny doses is not enough
reason to regulate it.
The
use of cyanide, mercury and formaldehyde may have been dramatically reduced
during the 20th century, but given the boom in the use of
organophosphates and organochlorines (“organo” simply means something that
is carbon based), such substances were not required much anyway. The world now
had cheap, highly effective and controllable – so it seemed – agents that
could be applied at will. Of course, as we know now (and, no doubt, the
manufacturers already knew early on) the legacy of these chemicals was passed
into the water, and from mother to child in countless animal species, including
humans. There is no way of knowing how many cancer deaths have been caused by
chemical pesticides, nor any way of predicting how many more deaths will come as
their legacy lives on both in the bodies of fish and marine mammals, and also
those parts of the world where such pesticides are still commonly used with
relish.
Organic
gardening and farming have been practiced for far longer than chemical based
growing, and nematodes can play an important part in this. Remember those I
mentioned that feed on other animals? Well, there is a whole range of different
species that not only leave the plants you are growing well alone, but also
actively destroy the very creatures that would otherwise cause damage.
Technically, these are known as Entomopathogenic[xiii]
nematodes, and there are two main species that are used: Steinernema and Heterorhabditis
(don’t worry; I won’t be testing you later). They are similar in form
and effect, both killing a wide range of insects and related organisms, like
caterpillars, by entering their bodies as juveniles then releasing bacteria that
are toxic to the host. This bacterium kills the insect, after which the nematode
is free to mate or, in the case of Heterorhabditis, reproduce alone.
The
downside? Well, there really isn’t one, unless you count having to make sure
they are not fried by ultraviolet light, or overheated. I include this quotation
from Cornell University, just to show I am not overstating the advantages of
these wonders:
Entomopathogenic
nematodes are extraordinarily lethal to many important soil insect pests, yet
are safe for plants and animals. This high degree of safety means that unlike
chemicals nematode applications do not require masks or other safety equipment;
residues, groundwater contamination and pollinators are not issues. Most [other]
biologicals require days or weeks to kill, yet nematodes, working with their
symbiotic bacteria, kill insects in 24-48 hr.
Dozens of
different insect pests are susceptible to infection, yet no adverse effects have
been shown against nontargets in field studies. Nematode production is easily
accomplished for some species using standard fermentation in tanks up to 150,000
liters. Nematodes do not require specialized application equipment as they are
compatible with standard agrochemical equipment.[xiv]
“But
wait!” you may say, “if these creatures are such efficient killers then
surely they can multiply, spread and kill off everything they touch, even
human-beneficial insects.” A fair point, but one that isn’t backed up in
practice. The aim of applying commercially available biological control
nematodes is in order to overload the natural system and kill many more insects
than would be killed by nematodes naturally[xv].
After they are applied, they do indeed destroy their targets very quickly, but
once the target is destroyed then there is little for the juvenile worm to
mature within and nematode numbers rapidly decline.
So why
aren’t nematodes used all over the world, making most types of pesticides
redundant? There are three reasons. First, not a lot of widely read research has
been carried out on the usefulness of such nematodes; in fact many nematologists
still believe that every nematode is a pest[xvi].
Second, although nematode insect parasites were identified as effective controls
in the 1930’s, the availability of cheap, effective chemical pesticides in the
1940s caused this research to be largely ignored, and it was not until some
chemicals were banned that research started up again[xvii].
Finally, and linking these two together, it is clear from the continued lobbying
of powerful companies like BASF, Monsanto and Syngenta, that the chemical
industry will not give up without a fight. It is no coincidence that DDT was not
widely banned until 20 years after clear evidence of its terrible impacts on
wildlife was made public, and that the 2007 European Union REACH legislation –
which enforces the control of hundreds of previously uncontrolled chemicals –
took ten difficult years to come into force. Industry still calls the shots,
even in an age when it is so obvious that natural ecosystems cannot cope with
the torrent of chemicals being washed into them day after day.
I may
come back to this later.
Moving
With The Climate
How fast
can a nematode move? One study suggests that 3cm in five hours is a fair guess[xviii];
although there are so many variables that all we can truly say is they move
pretty well considering their size. Despite their elegant, sinuous propulsion
method, the problems nematodes have in movement are manifold, largely related to
their diminutive length. Even in water, something about a millimetre long will
experience considerable pressures from all sides, and have to swim through the
thick soup of tightly interconnected molecules to make any headway – if you
have ever tried to run in the sea then you will understand how it feels. In the
soil the problems are multiplied: air gaps have to be traversed, boulder-like
grains circumvented and anything like a solid object simply accepted as
impassable. Viruses and bacteria can be carried in water flows, or in droplets
through the air, but animal vectors are the smart way to travel, whether this be
within parasitised insects or the gut of a human. A flying insect, bird or
aircraft will lap up distance with ease, meaning that anything able to take
advantage of a mobile host is definitely one more rung up the evolutionary
ladder. Nematodes are also easily transmitted from one place to another on
plants, as they are moved from nursery to farm and on agricultural equipment.
The latter is particularly significant. It only requires a farmer to plough a
field infected with, for instance, Root Knot Nematode, and then plough an
uninfected field with the same plough for the nematode to become ensconced in
the next field.[xix]
In
terms of climate change, though, speed is not of the essence. The rate of global
heating, although significant in its impact on the forces that drive weather and
other processes that rely on heat, is slowly creeping across the Earth. Slowly,
but inexorably, altering environments as the swath of change moves across the
land and the sea. Gradual movement is what nematodes can best take advantage of,
and that gradual movement is what is starting to concern farmers. There is a
concept used by phenologists (people who study the timescales and cycles of
natural events) called Degree Days. A degree day is simply a measure of the
amount of time available for an event to occur depending on temperature: one day
at one degree above the lowest temperature an organism will breed at is one
degree day. Using this system it is possible to predict the lengths of the
lifecycles of many organisms, including nematodes, according to the measured air
temperature. For example, if a certain nematode requires the temperature to be
above 5°C and below 30°C to carry out its lifecycle, four days at a constant
10°C makes twenty degree days.
Using
degree days, not only is it possible to work out how long the lifecycle of a
nematode will take at different temperatures, but you can also determine if an
area of soil is warm enough for the lifecycle to take place at all. The Root
Knot Nematode is widely regarded as one of the world’s most destructive
pathogens[xx].
If a particular species of root knot nematode needs 1000 degree days to produce
an entirely new generation of worms[xxi]
in a new crop of potatoes or carrots in a new field, then a one degree average
temperature increase could certainly make the difference between a new field
being a favourable breeding ground or not, and the difference between a crop
being successful or not. One ploughing is enough to distribute a few plants’
worth of nematodes across an entire field; just because nematodes move slowly,
doesn’t mean that they can’t spread extraordinarily quickly. If temperatures
in a field never drop below the lower threshold for root knot nematode, then the
nematode will happily keep multiplying there all the time food is available:
impervious, because of sheer numbers, to all but the most toxic applications of
pesticide. I’ll leave you to imagine what the impact of increasing temperature
on our food supply could be.
A
Singularity Of Bananas
Here are
some facts about bananas:
1.
They grow on the stems of ground-loving plants, not trees.
2.
The fruit of the banana plant can be yellow, green, purple or even red.
3.
In their natural form, bananas have large seeds.
4.
A single variety, “Cavendish”, accounts for the vast majority of the
world’s banana trade. It is virtually seedless.
5.
Bananas are an analogy for the whole of the industrial economy.
Ok,
that last one you won’t find in any text books or scientific journals, but
I’m not just making this up on the spot; you may consider society to have gone
“bananas” in more ways than one, but even that isn’t what I’m getting
at. The simple fact is that the bananas that most of us eat are in deadly peril,
and it is likely that the global supply will be largely wiped out within a few
years. It was only in the 1950s that the previous reigning variety “Gros
Michel” was almost totally destroyed by a fungus called Panama Disease. Gros
Michel had many of the characteristics of Cavendish, except it wasn’t
resistant to the particular type of fungus that Cavendish is; but that is set to
change dramatically. The problem is that every Cavendish plant is genetically
identical to the original variety that was brought to the Caribbean from South
East Asia in the early 19th century[xxii],
regardless of the small differences in texture, size and colour that derive from
different growing methods and climates. In order for genetic variety to occur in
plants that reproduce sexually, two sets of chromosomes, one male, one female,
have to be combined. Making cuttings doesn’t create genetic variety, and this
is basically the reason why family interbreeding amongst humans has been
outlawed in most human cultures for centuries, possibly even thousands of years.
It is not possible for different sexes to be genetically identical, but if
brothers and sisters, or other close relatives breed over a number of
generations, then any damaging genetic mutations will remain within the family
line, eventually leading to a much higher rate of abnormalities, including poor
resistance to disease.
Evolution
occurs in order to ensure that a particular species remains hardy enough to
continue its line. The Cavendish, and the Gros Michel before it, are perfect
examples of what happens when evolution is not allowed to occur. In the 40 years
since Gros Michel was almost wiped off the map, the fungus that caused Panama
Disease in that plant has evolved so that it can now do the same to Cavendish.
In the words of one writer: “the banana is too perfect, lacking the genetic
diversity that is key to species health. What can ail one banana can ail all. A
fungus or bacterial disease that infects one plantation could march around the
globe and destroy millions of bunches, leaving supermarket shelves empty.”[xxiii]
Lack of bananas may not cause huge numbers of deaths, but lack of genetic
diversity most certainly can:
From 1845
to 1846 Ireland's potato crop consisted of one or two closely related varieties.
Both were wiped out by blight. In the ensuing famine, nearly a million people
died and more than a million others were forced to emigrate. By 1851 Ireland’s
population had diminished by 23 percent. If Irish farmers had been growing many
varieties of potatoes with different genetic backgrounds the disaster would
never have happened.[xxiv]
You
may ask why this is an analogy of the industrial economy. The reason is that
throughout the 20th
century and into the 21st century, money and the possession of
material goods have come to dominate the way societies are run, especially in
the industrial West. The market economy, which governs the way most commerce and
a great deal of politics in the world operates, does not favour diversity –
positively discourages it, in fact. The spoils almost always end up going to the
individual, company or country that can provide the most of some thing or
another – whether that be a raw material, a consumer product, a service or a
variety of banana or potato – at the cheapest price, in the shortest time and,
often as a result, at the lowest quality. There is no way out of this; it is
just the way this type of economic system works: if you want variety (and
quality) then you have to operate outside of the market economy.
Of
course there are notable exceptions, for instance products that fail strict
safety guidelines in one country will not be sold there, but that does not mean
they cannot be successful in countries where those guidelines don’t exist. The
much-touted sub-$2000 car will, no doubt, be a roaring success in India, its
country of manufacture, but can never be sold in Europe, Canada or the USA due
to its poor construction. But in the main, big, fast and cheap wins out; so
while the Cavendish banana is the type chosen by the largest producers, who
effectively have the banana market cornered, then that will be the banana that
sits on supermarket shelves, market stalls and in fruit bowls around the world.
You
may also ask what all this talk about bananas is doing in a chapter about
nematodes. Like almost all types of food crop, bananas are vulnerable to attack
by nematodes; and in many countries they can causes losses[xxv]
of 30-60 percent – that is the difference between making a living from banana
sales, and not being able to afford to grow the crop. Two particular species of
nematode, Pratylenchus coffeae and Radopholus similis, exist right
across the world, from the Caribbean, to Ecuador, to Central Africa, to the
Philippines – in fact everywhere bananas are grown on a commercial basis.
There can be little doubt that this vast distribution is the result of a single,
genetically identical variety of banana having a virtual monopoly. Even if the
new strain of Panama Disease doesn’t finish off the world’s banana crop,
then a tiny writhing worm may well do so; a tiny little worm whose relations we
hardly notice, but which exist in uncountable numbers in almost every animal,
every piece of soil, every plant and all the way down at the bottom of the
ocean.
References
[i] In the world of commercial “pest” control almost everything that moves is a potential pest. A few commentators have observed that maybe the only true pests are humans, e.g. http://query.nytimes.com/gst/fullpage.html?res=9801E2DB123FF931A25757C0A9619C8B63 (accessed 13 June, 2008).
[ii] “Plant Parasitic Nematodes”, USDA ARS, http://www.ars.usda.gov/Services/docs.htm?docid=9628 (accessed 3 January, 2008)
[iii] N. A. Cobb, “Nematodes and their relationships,” Dept. Agric. Yearbook, 1914 (quoted in R. N. Huettel and A. M. Golden, “Nathan Augustus Cobb”, Ann. Rev. Phytopathology (29), 1991).
[iv] ibid.
[v] Victor H. Dropkin, “Introduction to Plant Nematology”, John Wiley and Sons, 1980.
[vi] Meat & Wool New Zealand, “Wool Exports”, http://www.meatandwoolnz.com/main.cfm?id=259#331 (accessed 3 January, 2008)
[vii] R. Danovaro et al, “Exponential Decline of Deep-Sea Ecosystem Functioning Linked to Benthic Biodiversity Loss”, Curr. Biology (17), 2007.
[viii] The Scottish Government, “Potato cyst nematodes - a technical overview for Scotland”, http://www.scotland.gov.uk/consultations/agriculture/PCN_Technical_Paper_Scotland_SEERAD.pdf (accessed 30 December, 2007)
[ix] SeedQuest, “BASF expects European Union approval of Amflora potato within weeks”, http://www.seedquest.com/News/releases/2007/december/21250.htm (accessed 30 December, 2007)
[x] M. Toparlak et al, “Contamination of Childrens Playground Sandpits with Toxocara
eggs in Istanbul, Turkey [MÜFİT TOPARLAK, AYŞEN GARGILI, ERKUT TÜZER, VEDAT KELEŞ, MELTEM ULUTAŞ ESATGİL, HANDAN ÇETİNKAYA]”, Turk J Vet Anim Sci (26), 2002.
[xi] “A History of Crop Protection and Pest Control in our Society”, Croplife Canada, http://www.croplife.ca/english/pdf/Analyzing2003/T1History.pdf (accessed 7 January, 2007)
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[xiii] Sorry about the terminology, it just means “insect killer”.
[xiv] Randy Gaugler, “Nematodes”, Cornell University, http://www.nysaes.cornell.edu/ent/biocontrol/pathogens/nematodes.html (accessed 2 February, 2008)
[xv] William T. Crow, “Using Nematodes to Control Insects: Overview and Frequently Asked Questions”, http://edis.ifas.ufl.edu/IN468 (accessed 8 January, 2008).
[xvi] Simon Gowan, University of Reading, personal communication.
[xvii] G. C. Smart Jr. “Entomopathogenic Nematodes for the Biological Control of Insects”, Supp. J. Nematology (27), 1995.
[xviii] A.H. Jay Burr and A Forest Robinson, “Locomotion Behavior” in Eds. Randy Gaugler, Anwar L. Bilgrami, “Nematode Behavior”, CABI Publishing, 2004.
[xix] Southern Illinois University Carbondale, “Root-knot nematode moving into Illinois fields”, 2001, http://news.siu.edu/news/June02/060402k2114.html (accessed 9 January, 2008)
[xx] Iowa State University, “Researchers Bioengineer Plants Resistant to Devastating Pathogen”, 2006, http://www.ag.iastate.edu/aginfo/news/2006releases/baum.html (accessed 10 January. 2008)
[xxi] University of California IPM, “Phenology Model Database: Columbia Root Knot Nematode”, http://ucipm.ucdavis.edu/PHENOLOGY/mn-columbia_root_knot.html (accessed 10 January, 2008)
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[xxv] “Burrowing and Lesion Nematodes of Banana”, Secretariat of the Pacific Community, http://www.spc.int/PPS/PDF%20PALs/PAL%2005%20Banana%20Burrowing%20Nematode.pdf (accessed 10 January, 2008)