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Introduction Inundative releases of natural enemies
to augment activity in the field and to improve pest control have been
practiced for many years (King et al. 1985a, Kogan et al. 1999). Annual short
term crops are particularly suited for the inundative release approach in
biological control, because they often fail to provide a stable environment
for the continuous abundance of natural enemies. By manipulating the kinds of
natural enemies, the stage of their development for release, the numbers
released and the time and modes of release, a much more active role is
required of field managers than in classical biological control. Kogan et al. (1999) showed that successes of inundative
releases may depend upon (1) the nature of the crop plant, (2) the
developmental stage of the crop, (3) the developmental stage of the pest, (4)
the absolute density of pest organisms, (5) the quality of the natural
enemies, including host specificity, searching capacity and proper identity,
(6) the density of natural enemies, (7) the climate, (8) complementary or
antagonistic effects of other natural enemies, (9) ability to integrate
releases with other control methods, particularly insecticides, and (10) the
cost of the release program. Five examples of inundative or inoculative
release programs will represent the range of crop/pest systems and spectrum
of natural enemies for which augmentation has adopted. Trichogramma spp. For Annual Crops. Trichogramma species are
presently the most widely used insects in inundative and augmentive control
(Ridgway & Morrison 1985). The area of crops covered by Trichogramma releases has
increased annually and amounts to ca. 11 million ha. in the former Soviet
Union (Voronin 1982), 5,500 ha. in Western Europe (Hassan et al. 1986),
355,000 ha. in the United States and about 2 million ha. in the People's
Republic of China (van Lenteren 1987), and extensive areas in Mexico (Jimenez
1980). Annual crops on which Trichogramma
are used include rice, crucifers, sorghum, millet, sugar beets, cotton, corn
and cassava. However, documentation of the release data and the outcomes of
release programs are mostly lacking, which makes it difficult to evaluate
results and to draw conclusions about the applicability of particular
practices to the overall use of Trichogramma
for biological control. Consequently, a scientific discussion is limited to a
few prominent examples. Trichogramma in Cotton.--Cotton, Gossypium hirsutum, is a perennial plant that is usually grown as a
warm season annual. In the Northern Hemisphere it is typically planted in March
or April and, according to the variety, harvested between August and
November. The plant is attacked by many insects, particularly Homoptera,
Hemiptera and Lepidoptera, from germination to picking time (Reynolds et al.
1982). However, the early season, which is typified by vigorous plant growth,
is often characterized by a relatively smaller risk of insect damage than is
the later part of the season. This is because of the relatively high
abundance of natural enemies in the early part of the season (Bar et al.
1979), to the lower susceptibility of plants to damage because no mature
fruits are present and because of their capacity for compensatory growth
(Wilson 1986a), and the lower numbers and less damaging characteristics of
many early season pests. Therefore, the use of early season insecticide
treatments in cotton is particularly unnecessary and its avoidance may enable
growers to extend substantially the insecticide-free period of the crop. Trichogramma pretiosum
(Riley) and T.
australicum Girault are used
in Colombia to control early season lepidopterous cotton pests such as Alabama argillacea (Hübner), Trichoplusia
ni (Hübner), Pseudoplusia
includens Walker, Sacadodes
pyralis Dyar and Heliothis
spp. (Amaya 1982). Releases are intended to prevent damage by these pests
and to facilitate an insecticide-free period of about 100 days, after which
treating against boll weevils, Anthonomus
grandis Boheman is often
necessary (Kogan et al. 1999). The parasitoids are mass reared on Sitotroga cerealella Zeller, mostly in local insectaries and
released in the field as pupae within host eggs that are glued to small (6 cm2)
cardboard strips, each bearing 3,000 eggs, 85% of which are
parasitized. Releases start 20-25 days after germination and continue
throughout the season. The first three releases are made at five day
intervals to establish an overlapping parasitoid population. Subsequent
releases are made every eight days. Each release consists of 20 cards (ca.
51,000 parasitoids) up to square formation, and 30 cards (ca. 76,000
parasitoids) thereafter (Amaya 1982). Sometimes parasitoid releases may begin
at planting time when parasitoids are liberated along field margins to kill
Lepidoptera that develop on the surrounding vegetation, and continue within
the field at about weekly intervals for some three months. Parasitoids are
released as freshly emerged adults, first at the rate of 40-50,000 per ha.
and later 30-36,000 per ha. The exact timing of releases is determined by
field scouting performed twice weekly by the growers (Kogan et al. 1999).
When unavoidable insecticide applications occur, Trichogramma releases are made as soon as two days after
applications to maintain continuous control (Amaya 1982). Inundative
programs using Trichogramma
resulted in a marked reduction of insecticide treatments. In a 15,000 ha.
area of the Valle del Cauca, Colombia, the number
of treatments changed from 20 per year in 1975 to four in 1981; on 6,000 ha.
in the norther part of that region, the number was reduced to 1.2 treatments
per year. The success in Colombia and in Mexico is due partly to the
relatively inexpensive and efficient Trichogramma
production methods available. However, emphasis on applied research for the
improvement and maintenance of parasitoid quality through continuous
selection, the development of parasitoid storage techniques, the accurate
determination of the quantities of parasitoids to be released and the correct
timing of the releases have been crucial to the success of the program (Amaya
1982, Kogan et al. 1999). In
the former Soviet Union, special races of T.
euproctidis Girault are used
for the control of Heliothis armigera
Hübner and of cutworms, Agrotis
sp. in Central Asian cotton (Voronin 1982). In the case of H. armigera, three releases of the parasitoids at the rate of
1:1 or 1:2 pest:parasitoid are used with a resulting parasitism of 66-90%.
Release rates against cutworms are 200,000 per ha., three times, once every
5-7 days when cotton is in the seedling stage. This procedure provides
complete protection of the crop. Release thresholds in Tadjikistan against H. armigera are such that only 50% parasitization efficiency
is sufficient for economic control (Voronin 1982). In
the People's Republic of China about 680,000 ha. of cotton are treated with Trichogramma (Huffaker 1977).
In over 100,000 ha. of the Shaanxi Province, control of Heliothis is achieved with the release of T. chilonis Ishii ( = T.
confusum Viggiani). The
parasitoids are applied at the rate of 120,000 per ha. in a total of three
releases at 3-4 day intervals during the F2 host generation; 75%
parasitization is achieved (King et al. 1985b). In the Jiangang farm, H. armigera has been controlled on 3,546 h. yearly between
1975-1984 by releasing 414,000 parasitoids/ha. Parasitization reaches 45%
with a residual worm density of 4/100 plants. The large amount of data
obtained permitted the construction of a reliable model for predicting the
efficiency of T. confusum in cotton fields
during the third and fourth host generations (Zhou 1988). Studies
on the practicality of using Trichogramma
species, especially T. sp.
nr. pretiosum for the
control of the bollworms H. zea and H. virescens
have been conducted in the United States (King et al. 1985c). A three-year
pilot test was conducted in southeastern Arkansas in 1981 and North Carolina
in 1983 to evaluate Trichogramma
for controlling Heliothis
species in cotton King et al. (1985b) summarized the project and its
achievements and concluded that mean parasitism rate of 47.4% of Heliothis spp. by T. sp. nr. pretiosum
augmented in cotton was insufficient to provide adequate control (King et al.
1985c). Explanations for the failure of Trichogramma
in the United States were presented in contrast to its successes in China,
South America and Mexico. A key reason was the higher production cost of the
parasitoids in the United States, especially compared to the lower cost of
insecticides. Low insecticide costs in the United States also create lower
economic thresholds for Heliothis,
which in turn promote numerous insecticide treatments. An additional factor
that plays an important role in many other areas of the world where cotton is
grown, is the frequent need to use insecticides against other pests, with the
result that such treatments further disrupt parasitoid performance. Trichogramma in Corn.--Corn, Zea mays, is an annual crop that, like cotton, is grown during
the warm season of the year. The growth cycle from planting to harvest varies
from two to five or six months according to the variety and growing
conditions. Corn originated in the Western Hemisphere and has spread
worldwide (Aldrich et al. 1975). It has become a cosmopolitan staple. In
addition to the indigenous pest complex on corn, many local insect species
have adapted to the crop, and presently each geographic region has both
cosmopolitan and local corn pests (Chiang 1978). The European corn borer, Ostrinia
nubilalis Hübner, originally
fed on unknown hosts, but readily moved onto corn, spreading from Europe to
reach the status of a severe pest of worldwide importance in temperature and
cold climate countries (Balachowski 1951, Kogan et al. 1999). Along the
northern boundaries of its distribution in Germany, Switzerland and the former
Soviet Union, China and Canada, the corn borer has only one generation per
year. Here it may be the main or only serious corn pest (Hassan 1982). The
number of generations per year increases at lower latitudes just as the
complex of pests associated with corn expands. Therefore, insecticide
treatments against the corn borer in its univoltine range are not only
expensive and environmentally disruptive but may cause the outbreak of
secondary pests such as aphids, which would otherwise be controlled by natural
enemies (Hassan 1982). Efforts to control the corn borer by releasing Trichogramma were first
reported from the former Soviet Union (Zimin 1935) and such efforts have
continued ever since (Voegele 1988). However, commercial efforts to use Trichogramma were initiated
only during the last decade after successful field trials were carried out in
Europe (Bigler 1986, Voegele 1988). The number of countries using commercial Trichogramma rose within a few
years from two (former Soviet Union and People's Republic of China) to 10
(Austria, Bulgaria, Colombia, France, Italy, Germany, Switzerland and the
United States). Several
reasons propelling the commercial use of Trichogramma
as a principal means of corn borer control are (1) concern over the
disadvantages of chemical pesticides, (2) increase in the efficiency of Trichogramma production, (3)
awareness of the importance of the specific biological characteristics of the
parasitoid to be used, leading to the acquisition of more efficient
parasitoid species (Beglyarov & Smetnik 1977, Huffaker 1977, Bigler et
al. 1982, Voronin 1982, Hawlitzky 1986, Voegele 1988), and (4) identity of
the requirements for optimal field releases (Stengel 1982, Voronin 1982,
Hassan et al. 1986, Hawlitzky 1986). Most
researchers maintain that parasitoids must be in the field before the first
oviposition wave of corn borer, and various methods have been devised to
accomplish this. Hassan et al. (1986) used
light traps to detect the first appearance of adult moths. In France, Stengel
(1982) and Hawlitzky (1986) discuss a day-degree calculation based on records
of the development and flight of the moths since 1963. These data, together
with the emergence of moths from caged pupae, are used to determine the onset
of oviposition. Economic threshold is reached when 10-12% of the eggs have
been laid about three weeks after first flight. This threshold varies
according to climatic conditions and corn variety, ranging from 6% for early
and 15% for late varieties. Inclement
weather and predators may cause mortality of the parasitized eggs that are
placed in the field. Egg predation becomes more severe with longer exposures.
Methods are available to minimize such mortality factors. In France,
Hawlitzky et al. (1987) placed parasitized eggs in specially designed
perforated capsules 1-3 days before emergence. In Germany Hassan (1982)
placed egg cartons within a 3 x 6 cm screen saran bag as protection against
predators and a plastic cover as protection against rain. In the People's
Republic of China plastic bags are employed (Coulson et al. 1982). It
is especially important to guarantee the quality of parasitoids, as was
demonstrated by a reduction of parasitism from 75.2% in 1978 to 18.8% in 1979
in Switzerland when mass produced wasps deteriorated (Bigler et al. 1982).
Stock quality is usually assured by rearing at least one generation annually
on O. nubilalis eggs (Bigler et al. 1982, Hassan 1982, Voronin
1982). Stock can also be strengthened by introducing field collected
material, a practice that is very common in the People's Republic of China
(Coulson et al. 1982). Voegele (1988) discussed the preservation of stock
quality through the retention of original traits and improvement of parasitoids.
He recommended that in addition to the cyclic return to natural hosts, to use
isogenic females, manipulate the nutrition of the parasitoids in artificial
rearing media, optimize the host/parasitoid ratio in culture, manipulate
parasitoid diapause, use semiochemicals from the plant or from the host
insect, and select for insecticide resistance. Kogan et al. (1999) suggested that genes for insecticide
resistance as well as genes for response to certain environmental stimuli may
also be introduced into the parasitoid cultures. Other Crops.--In the former
Soviet Union Trichogramma
was used to control lepidopterous pests of peas and cabbage. Parasitization
of 89-96% of the eggs of Laspereysia
dorsana F. and 67% of the
eggs of Autographa gamma L. attacking peas was
achieved following the enrichment of the environment with nectariferous
plants (e.g., Phacelia tanecetifolia). The use of
nectar sources marked an improvement over the 29 and 31% control that had
been obtained without those sources, largely because of increased parasitoid
longevity. Similar results were obtained in the control of A. gamma on cabbage, where improvement was from 50-60% to
80-90% parasitization (Voronin 1982). Noctuid larvae that infest sugar beets
and potatoes were controlled by releasing 20-60,000 parasitoids per ha.,
which resulted in a 60-90% reduction in infestation levels (Beglyarov &
Smetnik 1977). The rice leaf roller, Cnaphalocoris
medinalis Guenee, and other
rice pests are controlled in the People's Republic of China by five seasonal
releases of from 150,000 to 600,000 T.
australicum per ha.,
depending on the host density (Kogan et al. 1999). The resulting parasitism
amounts to 80% and the total cost is half that for insecticidal control
(Huffaker 1977). Shen et al. (1988) reported successful results with
inoculative releases of only 15,000 T.
dendrolimi per ha. on seven
experimental hectares of rice. In Colombia Trichogramma is used for the biological control of various
crop pests in addition to those on cotton and corn. These include beans and
soybeans, where the pests are Anticarsia
gemmatalis and Heliothis sp. and cassava,
where the principal pest is the sphingid moth Enrinnyis ello
(L.). Parasitoids are released on egg cards at the rate of 51,000 per ha from
10 days after germination for beans, and 76,500 per ha. starting 30 days
after plant emergence for cassava. Initial releases were spaced five days
apart; later releases eight days apart, and satisfactory control was reported
(Amaya 1982). Predatory
Mites in Short Term Crops (also please
see <bc-40.htm>) Spider
mites have been controlled biologically for over two decades with
considerable success (Huffaker et al. 1970), with most work involving
glasshouses (see section on glasshouses). Outdoor crops are either treated
with acaricides or efforts are made to conserve naturally occurring predatory
mites (Jeppson et al. 1975). The active suppression of spider mites in fields
was studied by Oatman et al. (1976, 1977a, 1981), who used three species of
phytoseids, Amblyseius
californicus (McGregor), Phytoseiulus
persimilis Athias-Henriot,
and Typhlodromus
occidentalis Nesbit to
suppress Tetranychus
urticae Koch in California strawberry
fields. Phytoseiulus
persimilis was the most efficient
of the three predatory species. This predator was successfully established in
southern California where it survived in strawberry and lima bean fields as
well as on weed species in the genera malva,
Solanum, and Convolvulus. The weeds served
as reservoirs for the predators, from which they dispersed to strawberry and
lima beans during the season. However, in most cases the economic thresholds
in these crops were too low to enable a complete reliance on these predators
for control (Oatman et al. 1981). In
Israel, the Netherlands and France, commercial use of inundative predatory
mite releases in open fields has been practiced effectively. In Israel,
spring melons, cantaloupes and watermelons grown in the Jordan and Arava
Valleys, have been subjected each season to attacks by T. cinnabarinus
(Boisduval) and T. urticae. The normal practice of
using acaricides against these mites was expensive and in many cases
insufficient due to an increase in resistance. This enabled commercial
companies to culture A. persimilis for inundative
releases. Fields are surveyed every week for germination, and predatory mites
are released when spider mites are found. The release rate is 20,000
predators per ha. or about one predator to 10 spider mites when the plants
are at the four leaf stage and double that amount when plants are larger and
have formed runners. This method has the disadvantage of dispersing predators
evenly throughout the field, whereas spider mites are usually found in
aggregates. The result is that local epizootics may occur, and the
introduction of additional natural enemies may be required. The problem can
be circumvented with preemptive releases of a mixture of 5/1 spider
mites/predators in fields not yet infested (Kogan et al. 1999). Biological
control in Israel has resulted in an average net savings of ca. $300 per ha,
and growers experienced better yields due to the absence of phytotoxic
pesticides and a reduction in soil compaction that had been caused earlier by
ground spraying equipment. Aphid attacks were also substantially reduced. The
commercial control of T. urticae in vegetable crops
through the release of P. persimilis has been gaining
acceptance in France and The Netherlands. The system is based on the
integration of pesticide treatments against diseases and thrips and on two
widespread releases of 4-5 predaceous mites per m2. Treatments
against thrips with mevinphos are made two days before the first mite release
about 2-3 weeks after planting. Treatments are accompanied by inspection and
monitoring of infestation levels. Infestations usually decline below the
economic injury level following the second release. This system is integrated
with treatments against Botrytis,
mildew and Pseudoperenospora
and it has been applied successfully to strawberries in France and to
strawberries and pickling cucumbers in Holland. The major advantages are
healthier and stronger plants that last longer and extend the growing season
(Kogan et al. 1999). In
cassava there have been reported about 50 species of phytophagous mites, in
the genera Tetranychus and Mononychellus, which are
particularly destructive both in South America and in Africa, mainly when
they reach high infestation levels during dry seasons (Bellotti et al. 1982,
Mesa & bellotti 1987). The South American species Mononychellus tanaioa
(Bondar), or cassava green mite, was first detected in east Africa in 1971
(Bellotti & Schoonhoven 1978, IITA 1987a). It spread rapidly throughout
most cassava-growing areas of Africa. The green mite seems to be specific to
species of Manihot and a few
other Euphorbiaceae. Yield losses range from 13 to 80 percent, mainly as a
result of defoliation (IITA 1987a). Bellotti & Schoonhoven (1978) report
several predators feeding on cassava mites, including coccinellids of the
genera Stethorus, Chilomenes and Verania; the staphylinid Oligota minuta; the anthocorid Orius
insidiosus; several species
of cecidomyiids and thrips; and the phytoseid mites, Typhlodromalus limonicus,
and T. rapax. The phytoseid mites and Oligota minuta
seem to be the predominant predators. Later studies showed that some 19
species of predaceous mites were present in cassava fields infested by the
green mite in Colombia (Bellotti et al. 1982, Mesa & Bellotti 1987). A
comprehensive biological control program of the cassava green mite complex in
Africa involves cooperation among national and international research
centers. According to this plan, five species of predaceous mites, Typhlodromalus
limonicus, Neoseiulus anonymus, N. idaeus,
Galendromus
annecteres, and Euseius concordis are mass produced at
CIAT, Colombia, on Mononychelus
progressivus with a method
that was developed by Mesa & bellotti (1987). Predaceous mite shipments
are routed through CIBC quarantine in London and then forwarded to Africa for
field release. This biological control effort, coupled with the propagation
of resistant cassava varieties and cultural control methods are expected to
alleviate the impact of the green mites on cassava in Africa (IITA 1987a,
Kogan et al. 1999). Misc.
Natural Enemies in Short Term Crops The
Mexican bean beetle, Epilachna
varivestis Mulsant, has been
under a control program that involves inoculation releases of an imported
parasitoid. Importation of the tachinid Aploymyiopsis
epilachnae (Aldrich) from
Mexico during 1922-1923 was the first attempt to control this beetle on
common bean, Phaseolus vulgaris L. (Smyth 1923, Jones
et al. 1983). The parasitoid failed to become established despite extensive
releases of the flies in 19 states. Although as much as 90% parasitization
was attained the fly could not survive the winter (Landis & Howard 1940).
Importations specifically aimed at controlling this beetle on soybeans were
made in 1966 when two parasitoids of Oriental species of Epilachna were brought from India (Angalet et al. 1968).
The egg parasitoid Tetrastichus
ovulorum Ferriere did not
adapt to the new host, but the eulophid Pediobius
foveolatus (Crawford), a
larval-pupal parasitoid, selectively attacked E. varivestis
but not the larvae of beneficial coccinellids. Although the parasitoid
produced various generations with a season, thereby attaining high levels of
parasitization, it could not overwinter in the central United States. Inoculative
releases were begun on an areawide basis in 1974, based on the establishment
of nurse crops of common bean (Stevens et al. 1975b). Patches of common bean
were strategically established early in the growing season in areas adjacent
to soybean fields. The Mexican bean beetle was attracted to the bean patches
and established healthy colonies that served as breeding hosts for P. foveolatus kept over the winter in laboratory colonies
(Stevens et al. 1975a). From these patches the parasitoids readily spread to
soybean fields where levels of parasitization remained between 60-90%. The
program is presently conducted in Maryland, Delaware and Virginia (Schultz
& Allen 1976) and has been tested in South Carolina (Shepard &
Robinson 1976). Pediobius foveolatus releases in central
Florida in 1975 and 1976 reduced Mexican bean beetle populations to barely
detectable levels in commercial fields, although in home gardens, common
beans continued to be damaged. The success of the parasitoid in Florida has
been attributed to the long growing season that allows up to 10 generations of
the parasitoid. Additionally there is an abundance of beggar weed, Desmodium tortuosum, a preferred wild host of the beetle that serves
the natural inoculum of the parasitoid (Jones et al. 1983, Kogan et al.
1999). This program is an example of the use of a nurse crop in connection
with inoculative releases of a parasitoid originally obtained from a host
species different from that of the species targeted for biological control.
The economic feasibility of the program has been demonstrated (Reichelderfer
& Bender 1979). Current research focuses on strains of P. foveolatus imported from Japan (Honchu) at latitudes
comparable to those in regions of the United Sttes affected by the Mexican
bean beetle, but no new strains have thus far overwintered (Jones et al.
1983). Microbial
Pesticides
For Short Term Crops Microbial
agents that have been investigated for controlling pests in short term crops
include entomopathogenic viruses, bacteria, fungi and protozoa. Although many
pathogens have shown promise in field trials, very few microbial insecticides
are commercially available for use on short term crops. Bacillus thuringiensis, the
spore-forming bacterium, is the most widely used microbial insecticide. It
produces a toxic crystal at the time of sporulation that is very active
against Lepidoptera, but also safe to humans and natural enemies. The insect
for mortality to occur must ingest the crystal. Burges & Daoust (1986)
estimated that total annual sales in the United States were $40 million, most
of which were used to control forest Lepidoptera. As about 50 percent of all
insecticides used in the United States are applied to cotton, it might be
expected that B. thuringiensis would be used
extensively on that crop, which it is not. Control has been too unreliable
and variable, probably because Heliothis
spp. and Pectinophora gossypiella, major cotton
pests, bore into squares and bolls before ingesting enough of the leaf
surface to cause mortality. Vegetables sustain the greatest use of B. thuringiensis. In 1985 between $5-10 million was spent on
this bacterium for the control of Plutella
xylostella (L.), Artogeia rapae (L.) and Trichoplusia
ni (Hübner). A portion of
the genome, which produces the toxic crystal of B. thuringiensis, has been
incorporated into other bacteria and in higher plants. Very recent
information (1999) on the effectiveness of the toxin applied in the manner in
plants is that it is not as toxic as when applied directly to plant surfaces.
There is even some evidence that when incorporated into the genome of
potatoes, it causes illness in humans who consume the tubers (P. Maddon,
pers. commun.). No
commercial fungal products are available for insect pests of annual crops in
the United States, but government-sponsored mass production of Beauveria bassiana is prevalent in the former Soviet Union,
primarily for control of the Colorado potato beetle. Species of Metarhizium have been
extensively tested for the control of planthoppers in sugarcane and pasture
grasses in Brazil (Kogan et al. 1999). Although many additional fungi have
been field-tested, there is no commercial availability expected in the near
future. Kogan et al. (1999) report
that with the emphasis on lepidopteran defoliators of soybeans, three
strategies have been considered in the experimental development of Nosema rileyi as a biological
control agent: (1) inundative releases (Getzen 1961, Mohamed 1978), (2)
induced epizootics (inoculative releases) (Sprenkel & Brooks 1975,
Ignoffo et al. 1976), and (3) manipulation of the ecosystem (Sprenkel et al.
1973). However, it is doubtful that N.
rileyi will ever be used
extensively as a microbial insecticide. Ignoffo (1981) listed characteristics
of this microsporidian that limit its success as a microbial insecticide: (1)
it kills slowly, allowing older caterpillars to cause considerable damage
before dying, 92) it requires free water for germination, growth and
sporulation, (3) it has a temperature requirement of 15-30°C, and extreme
field temperatures may limit its effectiveness, and (4) to be effective large
spore dosages must be directed at young insects. Carner
& Turnipseed (1977) isolated a nuclear polyhedrosis virus from larvae of Anticarsia
gemmatalis collected in
southern Brazil. the virus was imported into the United States and examined
for pathogenicity. Small plot field tests gave significant reductions of A. gemmatalis, which were confirmed in Florida by Moscardi
(1977). Since the early 1980's extensive field and laboratory studies were
continued in Brazil (Moscardi & Correa Ferreira 1985). The virus (AgNPV)
is highly specific to A. gemmatalis (Moscardi &
Corso 1981) and is effective at field dosages above 10 LE/ha. Populations are
reduced below the economic injury level with a single application of the
virus suspension, and mortality soars to 80% at 40 LE/ha. (Moscardi 1983).
Leaf consumption by diseased larvae was reduced by about 75% and although the
half-life of crude preparations or a purified preparation with a clay
adjuvant was either six or seven days, respectively, a single application was
sufficient to control the caterpillars (Moscardi 1983). Kogan et al. (1999) report
that large-scale field testing with this virus started in Parana during 1980.
The virus was applied as a crude preparation at 50 LE/ha. when A. gemmatalis larvae were less than 1.5 cm long. Applications
were made with ground equipment at rates of 100-200 L of water per ha. Virus
used in the field experiments were extracted from batches of 50 cadavers of
large caterpillars (>2.5 cm). The dead larvae were macerated in water and
filtered through several layers of cheesecloth. The suspension was then
transferred to a sprayer tank containing the amount of water needed to cover
one ha. Infectivity after four days was 80%. Experiments were conducted in
areas of high incidence of A.
gemmatalis and check plots
were either treated with standard insecticides or left untreated. In all
cases, yields were as high with the virus treatment as they were with
insecticides (Moscardi 1983). An estimated 11,000 ha. of soybeans were
treated with the virus in 1983 and the area was expected to increase to
300,000 ha. in 1984 (Moscardi & Correa Ferreira 1985). At
the present time there is only one virus available as a commercial produce in
the United States, which is the nuclear polyhedrosis virus of Heliothis zea. Field trials have been conducted on control of insect
pests of short term crops with many different baculoviruses. Many of these
trials have produced encouraging results, but the costs of production make
large-scale commercialization difficult. Many of these viruses can best be
produced in a cottage industry environment and in areas where hand labor is
inexpensive (Kogan et al. 1999). Biological
Control & Plant Resistance The
IPM approach most compatible with biological control is the development of
plant resistance (Kogan 1982). Nevertheless, incompatibilities arise when
mechanisms of resistance indiscriminately affect both pests and natural
enemies, or when natural enemies are indirectly affected through their hosts
or prey. Experimental evidence of incompatibilities is shown in tomato
(Duffee & Isman 1981, Duffey & Bloem 1986, Duffey et al. 1986). This
may be illustrated with Heliothis
zea, Spodoptera exigua
and the endoparasitic wasp, Hyposoter
exiguae (Vier.). When host
larvae ingest a diet with the glycoalkaloid tomatine, the development of the
parasitoid is detrimentally affected (Duffey & Bloem 1986). Kogan et al. (1999) warn that such studies demonstrate
that depending on the mechanism of resistance, natural enemies may be
detrimentally affected; and that when exploiting such mechanisms one should
weigh the risk of reducing the natural enemy load versus the benefit of the
particular resistance trait. Obrycki (1986)
studying the impact of potato glandular trichomes on Edovum puttleri
(Grissell, an egg parasitoid of the Colorado potato beetle, drew similar
conclusions. He showed that E.
puttleri readily parasitizes
L. decemlineata eggs on Solanum
tuberosum but that the
parasitoid is entrapped in glandular trichomes of Solanum berthaultii.
On S. tuberosum, egg mortality is increased not only due to
parasitism but probably also to host feeding and superparasitism. But aphid
parasitoids that are equally affected by S.
berthaultii trichomes in the
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levels of trichomes and the biological control of potato aphids are not
incompatible. Therefore, it is apparent that both biochemical and physical
plant defenses are potentially detrimental to natural enemies. As behavioral
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