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NUTRITION
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Overview The nutrition of entomophagous arthropods was originally discussed
in detail by Doutt (1964) and Hagen (1964). Slansky (1982, 1986) and Thompson
& Hagen (1999) illustrate the complex interactions of behavioral,
physiological and nutritional factors in arthropod nutrition. Nutrition is
thus the action or processes of transforming substances found in foods into
body materials and energy to do all the things attributed to life.
Nutritional requirements are dependent on the synthetic abilities of the
organism, which is controlled genetically. House (1977) stated that "...
through nutrition we have a direct and essential connection between an
environmental factor, foodstuff and the vital processes of the insect
organism." Most nutrition research with insects has been aimed at
improving rearing and not developing a basic understanding of their
nutrition. Research has emphasized feeding and the development of artificial
diets, which are concerned with dietetics (Beck 1972). Although critical to
insect rearing, such research has given only a little understanding of insect
nutrition per se. Qualitative nutritional requirements of all insects are very
similar in spite of a great diversity of feeding habits (Beck 1972, Dadd
1973, Hagen 1986b). Although knowledge of dietetics and nutrition has
advanced, practical application of principles to insect rearing to support
biological control is lacking. Rearing each insect species is a unique
challenge as there is meager knowledge of nutrition principals that might
provide a broad and sound basis for approaching insect husbandry (house
1977). With entomophaga, foodstuff is in a continuous state of qualitative
and quantitative change, and very little is known of the quantitative
nutritional requirements for various life stages and physiological functions
of these insects. The requirements for many nutrients are often dependent on
the presence and concentration of others and correct nutrient balance may be
critical for successful nutrition. Those parasitoids and predators for which
artificial diets have been developed may serve as models for in vitro investigation on quantitative requirements for
specific nutrients. Thompson (1976a, 1982) used a defined artificial medium to
examine the quantitative requirements for supporting larval growth of Exeristes roborator. Parasitoid development is intimately associated
with host physiology. Changes in the host's physiology following parasitism
are adaptive for the parasitoid, which insures successful development (Vinson
& Iwantsch 1980, Thompson 1986). Parasitoids overcome potential nutrient
constraints by altering their host's behavior and physiology (Slansky 1986).
Changes in composition of the host's internal milieu may have significant
nutritional consequences for a parasitoid (Grenier 1986, Thompson 1989).
Endocrine interactions seem critical to successful parasitoid development.
Synchrony in development between many larval endoparasitoids and their hosts
occurs (Beckage 1985), and this suggests that the host's hormones and
endocrine physiology influence parasitoid development (Lawrence 1986a.). The
physiological basis of developmental synchrony is not well understood and
knowledge is restricted to investigation of the relationship of Biosteres longicaudatus with its host Anastrepha suspensa
(Lawrence 1982, 1986b). Some studies have tested the effects of hormones on
the development of parasitoids in
vitro with little success.
The potential of using insect hormone supplements in artificial media to
achieve successful growth and development of parasitoids in vitro
deserves research emphasis. The importance of ecological considerations in the nutrition
of insects was discussed by Slansky (1982). It was emphasized that behavior
and regulatory physiology of insects are in a state of continuous flux in
response to food supply, and that nutrition can be fully understood only by
considering the insects "nutritional ecology." With entomophaga
both the ecology of the entomophage as well as that of the host or prey needs
to be known. Dietary and nutritional requirements are genetically based and
genetic manipulation holds promise as a way to modify the nutrition of
entomophages. Chabora (1970) suggested that nutritional content varies
between strains of insects when he demonstrated that the yields of two
parasitoids, Nasonia vitripennis (Walker) and M. raptor Girault & Sanders were significantly increased
when they were reared on a hybrid of two strains of the host, Musca domestica L. The selection of desired traits for insect
rearing was discussed by Collins (1984). The potential for genetic
improvement of entomophages was outlined by Rousch (1979) and Hoy (1979,
1986). Most genetic selection has been directed to increase field
effectiveness of entomophages, such as improving sex ratio, host finding
ability, host preference, pesticide resistance and improved climatic
tolerance. However, genetic improvement must also guarantee the preservation
of vigor and vitality of the entomophage. Because these are intimately
associated with nutrition, genetic programs may involve selection for nutritionally
related traits. Advances in recombinant DNA technology indicate a possibility
for genetic manipulation of the nutrition of entomophages (Thompson 1989).
The incorporation of foreign or in
vitro altered genes for the
expression of desirable traits by an organism, is rapidly advancing
(Beckendorf & Hoy 1985), but is still not suited for practical
application as of 1991. History of
Parasitoid Nutrition.--Salt (1941) probably was the first to emphasize the complexity
of parasitoid nutrition in studies that demonstrated that the host influences
growth and survival of the developing parasitoid as well as sex ratio,
fecundity, longevity and vigor of the adult wasp (Clausen 1939, Salt 1941).
Such complexities were demonstrated in work by Arthur & Wylie (1959),
Wylie (1967), Nozato (1969) Sandlan (1979a) and others (Vinson & Iwantsch
1980). It has long been known that there is a relationship between host
biomass and size of solitary parasitoids, larger parasitoids developing from
larger hosts. This relationship exists for parasitoids which attack every
host developmental stage, but applies more generally to parasitoids of host
eggs and pupae where host size is fixed (Sandlan 1982). The relationship
applies when a parasitoid is reared on different host species of variable
size as well as when reared on different sized individuals of a single host
species (Salt 1940, Jowyk & Smilowitz 1978, Mellini & Campadelli
1981, Sandlan 1982, Mellini & Beccari 1984). It does not seem to hold
with ectophagous parasitoids, however (Legner 1969 ). The size of
adult Trichogramma pretiosum Riley reared on the
eggs of five hosts showed a direct correlation between parasitoid size and
the volume of the host egg from which it emerged (Bai et al. 1989). A
correlation also exists between total parasitoid biomass and/or numbers with
host size in the case of gregarious larval parasitoids (Wylie 1965,
Bouletreau 1971, Thurston & Fox 1972). The means by which gregarious
organisms moderate their development relative to host size has been shown
(Beckage & Riddiford 1983). The relationship between size of host and parasitoid is
closely associated with food quality and quantity (Arthur & Wylie 1959,
Sandlan 1982). Salt (1940) found that adult Trichogramma evanescens
Westwood display behavioral dimorphism related to host size. Large females
obtained from large hosts failed to oviposit on small hosts, whereas small
females accepted hosts of all sizes. Male wing development was influence by
host size, and this was also found in Gelis
corruptor (Foerster) by Salt
(1952). Adult female Coccygomimus
(= Pimpla) turionellae (L.) did not show
morphological and behavioral polymorphism, but larger females found it
difficult to oviposit in small hosts. On the other hand small females were
more efficient in attacking small hosts. Fecundity was influenced by
longevity with the greatest longevity reported for larger individuals reared
from large hosts (Sandlan 1982). The success of parasitoids in parasitization activity is
directly related to nutritional factors. Smith (1957) found differences in
larval mortality and adult size, sex ratio and reproductive rate of several
species when reared on Aonidiella
aurantii (Maskell) and Comperiella bifasciata Howard maintained on
different food plants. Habrolepis
rouxi Compere displayed
limited mortality on A. aurantii when feeding on
citrus, but 100% mortality when feeding on sago palm. Pimentel (1966) and
Altahtawy et al. (1976) showed differences in parasitoid fecundity and
longevity depending on host food source. Thurston & Fox (1972) reported
that nicotine influenced the emergence of Cotesia
(= Apanteles) congregata (Say) when reared on
Manduca sexta (L.) feeding on tobacco. Hyposoter exiguae
(Viereck) was harmed by tomatine in Heliothis
zea (Boddie) feeding on
tomato (Campbell & Duffey 1979). Aphelinus asychis
Walker required a longer larval developmental time and showed a decreased
adult longevity when reared on Myzus
persicae (Sulzer) fed on
defined diets deficient in sucrose or iron (Zohdy 1976). The effects seemed
related to decreased host size rather than a difference in nutritional
quality of the host, however. The survival of Aphaereta pallipes
(Say) was affected by the balance of amino acids and glucose in the
artificial diet used for rearing its host, Agria housei
(= affinis) Shewell (House
& Barlow 1961). Differences in larval development and adult size,
fecundity and sex ratio were observed in Tetrastichus
israeli (Mani & Kurian)
when reared on several host species, which was correlated to the total level
of essential amino acids in host tissues (Nadarajan & Jayaraj 1975). Even
though parasitoids reared from some host species with high levels of
essential amino acids were larger and longer-lived, the results were
variable, as were the specific amino acid compositions of the different
hosts. In general it may be assumed that parasitoid fecundity, reproductive
size, sex ratio and longevity are correlated with host size and nutritional
factors (Charnov et al. 1981, Charnov 1982, Luck et al. 1982, Mackauer 1986,
Strand 1986). The importance of rearing Chelonus
sp. nr. curvimaculatus on
the natural host for vigor retention was demonstrated by Legner &
Thompson (1977), as discussed in previous sections. In contrast to parasitoids, few studies have been done on the
effects of various natural foods on the biological character of predators. Smith
(1965) reported that 10 coccinellid species fed dried, powdered aphids, grew
larger and faster when feeding on Acyrthosiphon
pisum (Harris) and Rhopalosiphum maidis (Fitch) than on Aphis fabae Scopoli. Coccinella
septempunctata L. gained
more weight when feeding on Lipaphis
erysimi (Kaltenbach) than on
two other aphid species, and it was demonstrated that L. erysimi
had higher protein levels (Atwal & Sethi 1963). Parasitoids have been thought to show high efficiencies in
food utilization. Larvae consume food of high nutritional content and are
mostly inactive within the host which offers a limited food supply, which
points to selection for high food efficiency (Fisher 1971, 1981; Slansky
& Scriber 1985, Wiegert & Petersen 1983). Parasitoids examined for
food utilization include Coccygomimum
(= Pimpla) instigator (F.), Pteromalus puparum (L.) (Chlodny 1968), Gelis macrurus
(Thompson), Hidryta frater (Cresson) (= sordidus) (Edgar 1971), Brachymeria intermedia (Nees) and C. turionellae (Greenblatt et al. 1982), Diadromus pulchellus
Wesmael (Rojas-Rousse & Kalmes 1978) and Trypatgilum (= Trypoxylon)
politum (Say) (Cross et al.
1978), Phanerotoma flavitestacea Fischer
(Hawlitzky & Mainguet 1976), Venturia
(= Nemeritis) canescens (Gravenhorst) (Fisher
1968), Cidaphus alarius Gravenhorst and Phygadeuon dumetorum Gravenhorst (Varley 1961), and Cotesia glomerata (L.) (Slansky 1978). In these species, the mean
net conversion efficiency (= proportion of assimilated food converted to body
mass (Petrusewicz 1967, Calow 1977, Hagen et al. 1984) varied broadly
(11-62%), with a mean of 37% that was <
than for many groups of insect herbivores and detritivores. Cameron &
Redfern (1974) of two studied parasitoids, Eurytoma tibialis
Boheman and Habrocytus elevatus (Walker), were at the
high end of this range. Net conversion efficiencies may not be very high
because selection might have been for rapid rather than efficient growth
(Slansky 1986). Possibly the well known inverse relationship between growth
efficiency and assimilation (Welch 1968) may also be important. In contrast
to net conversion efficiency, the above parasitoids had relatively high
percentages of assimilation (= percentage of ingested food that is
assimilated) ranging from 55-94%, with mean of 67%, compared with means of
40-50% for most herbivores and detritivores. Howell & Fisher (1977) reported the highest nutritional
efficiencies for a parasitoid in the ichneumonid V. canescens.
Larvae had a 65% net conversion efficiency and 95% assimilation when
maintained on the host Anagasta
(= Ephestia) kuehniella (Zeller); net
conversion efficiency to the adult was 20%. The proportion of food/host available that is consumed by the
parasitoid and converted to parasitoid biomass was calculated by Slansky
(1986) and Howell & Fisher (1977). Calculated exploitation indices varied
among species from 3-80%, and V.
canescens larvae consumed
90% of its host's biomass and converted 55%, but there was no clear
correlation between host size and parasitoid size nor biomass conversion. Food utilization by predators has also been thought to be
highly efficient, for reasons similar to that for parasitoids. This is
especially true when predators spend much time waiting for their food (Lawton
1971), thus avoiding metabolic expenditure. Studies on food utilization of 11
predators was reviewed by Slansky & Scriber (1985). All had similar net
conversion efficiencies (4-64%, mean = 34%), but higher assimilation
efficiencies (37-98%, mean = 86%) than those of parasitoids. Cohen (1984,
1989) in studies of food utilization by Geocoris
punctipes (Say) when reared
from 1st instar nymphs to adults on eggs of Heliothis virescens
(F.), found an assimilation efficiency of ca. 95%, gross conversion
efficiency of 53% and net conversion efficiency of 55%. Nutritional
Requirements in Development Qualitative nutritional requirements of insects, determined by
use of defined and deficient artificial diets, were presented by several
authors (Dadd 1973, 1977, 1985; Friend & Dadd 1982, Hagen et al. 1984).
All insects have similar requirements for ca. 30 chemicals that include
protein and/or 10 essential amino acids (arginine, histidine, isoleucine,
leucine, lysine, methionine, phenylalanine, threonine, tryptophan and valine),
the B-vitamin complex (biotin, folic acid, nicotinic acid, panthothenic acid,
pyridoxine, riboflavin and thiamin), as well as other water soluble growth
factors, including choline and inositol, some fat soluble vitamins,
cholesterol or a structurally similar phytosterol, a polyunsaturated fatty
acid, minerals and an energy source usually provided by simple or complex
carbohydrates and/or lipids. Nutritional requirements of entomophagous insects are similar,
and similar to those of nonentomophagous species. House (1977) referred to
this common characteristic of insect nutrition as the "rule of
sameness" (House 1966a, 1974). The rule has been confirmed by recent
studies with parasitic and predaceous insects. In assessing the need for
nutrients, it is important to consider that rearing a single generation on a
synthetic or semi-synthetic diet did most studies. Some investigations
overlooked the potential contribution of nutrients stored within the egg.
Stored nutrients may support limited development and, in the case of trace
nutrients, supply a sufficient quantity to ensure development of at least one
generation. Studies with Itoplectis
conquisitor (Say) (Yazgan
1972) and Exeristes roborator (F.) (Thompson 1981a)
demonstrated partial larval development on diets lacking various essential
amino acids and B-complex vitamins. Other studies have demonstrated that
entomophagous insects have no unusual qualitative nutritional requirements. A
requirement for asparagine by Eucelatoria
bryani Sabrosky (Nettles
1986a) and the absence of a requirement for a polyunsaturated fatty acid by A. housei (House & Barlow 1960) were consistent with
findings for nonparasitic Diptera (Dadd 1977). The quantitative balance of different nutrients is a critical
and dominating factor determining dietary acceptability and suitability
(House 1969, 1974). The predominant foods of both parasitic and predaceous
insects are of animal origin and, thus, are generally high in protein content
and low in carbohydrate and fat (House 1977). Thompson (1986a) found a high
requirement for protein and/or amino acids in parasitoids. Exeristes roborator at the 6% amino acid level completed larval
development without glucose and/or fatty acids (Thompson 1976a). However,
glucose markedly improved survival when the amino acid level was reduced to
3% and at 1% amino acid, no development occurred with the carbohydrate.
Similar effects of amino acid level on larval development were reported by
Yazgan (1972) for I. conquisitor. Adult eclosion was
reduced by dietary amino acid levels of <6% and by deletion of glucose.
Fatty acids were only marginally beneficial in enhancing growth and
development rates of both species. A polyunsaturated fatty acid, however, was
required in small amounts. Adult I.
conquisitor (Yazgan 1972)
and E. roborator (Thompson 1981a) displayed crumpled wings and/or
bent ovipositors without a polyunsaturated fatty acid in the larval diet.
Linolenic acid alleviated these deformities in I. conquisitor,
and linoleic and linolenic acids were provided together in the case of E. roborator . Thompson (1983a) described the effect of nutritional balance
on larval growth of Brachymeria
lasus (Walker). Media
containing 0-10% glucose with 2% amino acids, and 1-8% amino acids with or
without 2% glucose were tested. All media contained 15% albumin and 2.5%
lipids. Weight gain increased on diets containing 2% glucose when the amino
acid level was increased from 1-4%, but was reduced at the higher amino acid
levels. Similar effects of varying the amino acid level were obtained with
diets lacking glucose, but the overall weight gain was less than observed
with the diets containing glucose. On diets containing 2% amino acids, weight
gain increased dramatically when glucose was increased from 0.5-4%, but
decreased at higher glucose levels. Growth rates on the above diets were
generally in the range of 15-200 mg/g/day. The maximal rate, 260 mg/g/day,
was obtained on a medium containing 2% glucose and 2% amino acids. The effects
of nutrient balance were closely related to the osmolality of the artificial
medium (Thompson 1983b). House (1966b) demonstrated similar quantitative requirements
to those of hymenopterous parasitoids in the dipteran Agria housei
Shewell. Maximal growth and survival were achieved when all nutrients were
increased proportionately over the levels in a basal diet that contained
2.25% amino acids, 0.05% salts, 1.16% lipids and 2.25% other ingredients,
including glucose, ribonucleic acid, vitamins and agar. When amino acid level
alone was increased, survival was reduced. On a diet containing nutrient
levels equivalent to pork liver (= 20% amino acids, 4% glucose, 3.5% lipids,
2% salts and 0.75% ribonucleic acid), survival was >80%. House (1967,
1970) showed that the relative balance of amino acids and glucose was
critical in determining growth or development and that A. housei
larvae selected diets for feeding on the basis of nutrient balance. Quantitative nutritional studies with parasitoids have
generally evaluated the effects of nutritional balance by univariate or
monofactorial analysis. Grenier et al. (1986) thought that such an approach had severe limitations
because it ignored potential interactions between nutrients, including
"...additivity, competitivity, antagonism or synergy." Thus,
interpretation of effects of nutrient variation aimed at medium optimization
was difficult, and it was suggested that nutritional studies be designed and
analyzed in a multidimensional manner that accounted for interactions between
all nutrients and biological criteria. Canonical correlation analysis, which constructs maximum
correlations between all linear combinations of variables within sets, such
as between growth and development, and dietary parameters, were reported with
Lixophaga diatraeae (Townsend) by Bonnot
(1986, 1988). Because biologically meaningless correlations may be generated,
accurate interpretation requires knowledge of biological correspondence
between variables. Bonnot varied the concentrations of 30 medium components
and determined the effects on 9 developmental criteria. Nine linear
correlations were obtained and three had correlation coefficients of
>0.95. There is little information about the effects of developmental
nutrition on the behavior of parasitoid larvae apart from measurements
related to growth and development rate. However, Veerman et al (1985)
reported that a photoperiodic response by C.
glomerata was influenced by
the carotenoid content of its host's diet. Vitamin A was essential for
photoperiodic induction of diapause and it was suggested that this vitamin or
a derivative may function as a photoreceptor pigment. Optimal nutritional balance can be influenced by environmental
factors, as was shown by House (1966b) with A. housei
that the effects of dietary glucose level on larval survival and development
could be modulated by temperature. The nutritive value of a basal medium
(House 1966a) was increased by increasing the temperature from 20 to 25 and
30°C at glucose levels between 0-1.5%. At higher glucose levels larval
survival and development were reduced with increasing temperature. Two media
of different composition were formulated whose superiority for larval growth
and development of A. housei was reversed at two
different temperatures (15 & 30°C) (House (1972). Such nutritional
effects might have ecological significance in affecting insect host range
(House 1966b). It was thought that in establishing host range, an insect
might be affected differently if the nutrient composition of its food were
uniform but the temperature varied within the range rather than if the
temperature were uniform but the composition of food was variable. On the
other hand, the insect might not be affected if variation in food composition
was accompanied by compensatory changes in temperature. Therefore, an insect
species that attacks a particular foodstuff in a region with a specific
temperature might, if introduced into another area with a different
temperature, adapt to a different food source whose nutrient composition is
favored at the new temperature . Non-nutritional factors are intimately and intrinsically
involved in food acceptance and ingestion. These include physical properties
such as form, texture, etc., but also non-nutritive chemicals that elicit
specific behavioral and/or physiological responses essential for finding and
accepting foodstuff and in some cases for initiating behaviors associated
with the feeding process itself (Bernays & Simpson 1982, Bernays 1985).
Although such factors have been best shown in phytophagous insects, they also
play a role in the biology of entomophaga and will likely be of importance in
the development of continuous in
vitro culture. Predator Culture
In Vitro The artificial rearing of predators has stressed maintenance
of the adult stage for maximizing egg production rather than complete in vitro culture. Predator larvae are the preferred
biological control agent, and eggs and larvae produced by adults are placed
directly in the field. However, some effort has been aimed at complete
artificial culture of predators.. Among the first reared artificially from
egg to adult was the coccinellid Coleomegilla
maculata maculata (DeGeer) by Szumkowski
(1952). Adults fed on raw liver or meat being kept for months on these food
in the absence of prey. However, survival of larvae was poor on meat products
alone and only 38% reached the adult stage. Supplementing vitamins resulted
in ca. 86% of the larvae reaching adults. Oviposition and egg viability were
increased by addition of vitamin E to the adult diet. The culture methods
were refined and a diet of fresh yeast and glucose supported larval
development (Szumkowski 1961a,b). Smith (1965, 1966) reared several
coccinellid species including C
maculata lengi on dried aphids
supplemented with pollen. Success also was achieved on a diet of 40% brewer's
yeast, 55% sucrose, inorganic salts, cholesterol, RNA, wheat germ oil and
vitamins. Adults were fed the same diet supplemented with powdered liver.
Attallah & Newsom (1966) reared 8 generations of this coccinellid on a
defined diet of casein, sucrose, wheat germ, soybean hydrolysate, glycogen,
butter fat, corn oil, a liver factor, dextrose, cotton leaf extract (with
carotenoids and steroids), brewer's yeast, ascorbate, inorganic salts,
vitamins and agar. Adults reared in
vitro were fecund and mating
was stimulated by addition of vitamin E to the diet. The medium failed to
support growth of Coccinella
novemnotata Herbst, Cycloneda spp., Hippodamia convergens Guérin and Olla
v-nigrum (= abdominalis) (Mulsant). The
last species was successfully cultured in
vitro by Bashir (1973).
Optimum egg production required inclusion of vitamin E in the larval diet,
which was in contrast to the results of Szumkowski (1952) where
supplementation of the adult diet alone was insufficient for maximum egg
production. Several coccinellid species were reported to be successfully
cultured in vitro by Smirnoff (1958). These
included Psyllobora (= Thea) virgintiduopunctata (L.), C. septempunctata,
Oenopia (= Harmonia) doublieri (Mulsant), O.
(= Harmonia) conglobata (L.), Rhizobius lophantae (Blaisdell), R.
litura (F.), Rodolia cardinalis (Mulsant), Exochomus
anchorifer Allard, E. quadripustulatus (L.), E.
nigromaculatus Erhorn, Scymnus suturalis Thunberg, S.
pallidivestis Mulsant, S. kiesenwetteri Mulsant, Stethorus
punctillum Weise, Chilocorus bipustulatus (L.), Clitostethus
arcuatus Rossi, Pharoscymnus numidicus Pie, P. ovoideus Sicard and Mycetaea
tafilaletica Smirnoff
(Endomychidae). The diet contained sucrose, honey, alfalfa flour, yeast,
royal jelly and agar supplemented with dried pulverized prey. Larval rearing
in a few species was improved by adding beef jelly. All species developed
more rapidly and lived longer on the artificial diet compared with insects
reared under natural conditions, and the adults were very healthy. Harmonia axyridis (Pallas), C.
septempunctata and Chilocorus kuwanae Silvestri were reared on Smirnoff's (1958) diet
and other artificial media by Tanaka & Maeta (1965). Successful culture of
all species was obtained but adults failed to lay eggs. Chumakova (1962)
reared Crytolaemus montrouzieri Mulsant on similar
crude diets supplemented with dried prey. Okada et al (1971a,
1972) and Matsuka et al. (1972)
successfully reared H. axyridis on diets containing
powdered larvae and pupae of drone honeybees (Apis mellifera
L.). Sixteen generations of H.
axyridis and three
generations of Menochilus sexmaculatus (F.) were cultured
in vitro. Okada & Matsuka (1973) and Matsuka et al.
(1982) later improved the rearing method for maintaining adult Rodolia cardinalis. Chilocorus
rubidus Hope, Scymnus hilaris Motschulsky, S.
otohime Kamiya, Vibidia duodecimguttata Poda and S. hilaris
adults were maintained on the diet but did not lay eggs (Matsuka et al. 1972).
Niijima
et al. (1986) described the use of drone honeybee powder for rearing several
coccinellids including A. bipunctata, Anatis halonis Lewis, Coccinella
explanata Miyatake, C. septempunctata, Coccinula
crotchi (Lewis), Eocaria muiri, H.
axyridis, Harmonia yedoensis Takizawa, H.
convergens, Hippodamia tredecimpunctata L., Lemnia
beplagiata (Swartz), M. sexmaculatus, Propylea
japonica, S. hilaris and S.
otohime. Variable results were obtained, but 11, 16 and 25 successive
generations of E. muiri, H. axyridis
and M. sexmaculatus respectively were cultured from the egg to
adult stage. Larval development, adult longevity and fecundity were
satisfactory. The fractionation of honeybee powder was described by Matsuka
& Okada (1975) who found that the active factor stimulating predator
growth was unstable but nonproteinaceous. Expanded attempts to analyze bee
powder was described by Niijima et al. (1977). Niijimi et
al. (1986) then
formulated several chemically defined diets for rearing H. axyridis.
Larvae developed from the 1-3rd instar on a diet containing 18 amino acids,
sucrose, cholesterol, 10 vitamins and 6 minerals. Kariluoto et al. (1976) described rearing of A. bipunctata.
About 60 variations of seven artificial diets were tested. These contained
varying amounts of wheat germ, brewer's yeast, casein, cotton-leaf extract,
egg yolk, sucrose, liver fractions, honey, glycogen, soybean hydrolysate,
butter fat, corn oil, amino acids, dextrose, ascorbate, choline, inorganic
salts, vitamin E and antibiotics. The best diets yielded 60-80% of larvae
that became adults, but development time was slowed and adult weight lowered.
Kariluoto (1978) modified the medium, and Kariluoto (1980) obtained fecund
adults of A. bipunctata, C. septempunctata and others
reared in vitro. In vitro
culture attempts with Chrysopa
species did not succeed until Hagen & Tassan (1965) got a complete
culture of Chrysoperla carnea (Stephens) on an
encapsulated liquid medium (in paraffin droplets). The diet consisted of
enzymatic yeast, protein hydrolysate, ascorbate, fructose, choline and casein
hydrolysate. Adults were fecund but development time from the egg stage was
ca. 2X that of insects reared on aphids. Vanderzant (1969) then successful
cultured C. carnea for 7 generations on
pieces of cellulose sponge soaked in enzymatic casein and soy hydrolysates,
fructose, inorganic salts, lecithin, cholesterol, choline, ascorbate,
vitamins and inositol. Development on this diet was slow, but 50-65% of
larvae reached the adult stage compared with 85% when reared on natural
insect eggs. Hassan & Hagen (1978) reported obtaining three generations
of C. carnea on an artificial diet of honey, yeast flakes,
sucrose, casein, yeast enzymatic hydrolysates and egg yolk. Developmental
time and pupal weights were similar to those of insects on eggs of Sitotroga cerealella (Olivier). Chrysoperla
sinica (Tjeder) was cultured
for 10 generations on a diet of egg, brewer's yeast, sucrose, honey and
ascorbate (Ye et al. 1979). Adults were fed powdered liver, honey and
brewer's yeast. Cai et al. (1983)
reared this species on an encapsulated medium of soybean and beef
hydrolysates, egg yolk, sucrose, honey, brewer's yeast, ascorbate and
linoleic acid, with similar success reported by Zhou & Zhang (1983). The hemipteran predator, Geocoris
punctipes may be reared on
several diets (Dunbar & Bacon 1972). Media were nevertheless supplemented
with insects. Cohen (1981) reported in
vitro culture of G. punctipes from 1st stage nymph to adult on encapsulated
semidefined diets. Six media containing casein hydrolysates, yeast, sucrose,
cholesterol, corn oil, lecithin, agar, inorganic salts, phenylalanine and a
vitamin mixture were formulated and encapsulated in different forms. The
latter included mixtures of polybutene 32, dental impression wax, Vaseline,
epoline C-16, candelilla wax, Sunoco, and Paraplast. Best results were with
vitamin-enriched medium encapsulated in a mixture of 5% polybutene 32 and 95%
dental impression wax. Development of G.
punctipes in vitro was better than when reared on Spodoptera exigua
(Hübner). The percent of nymphs that reached adults and survival of the in vitro reared predators were significantly greater on the
artificial diet. Cohen (1983) then described modifications of media content,
preparation and encapsulation and could rear two generations of G. punctipes. Geocoris
pallens Stal, H. convergens, H.
axyridis and Nabis spp. also successfully
fed on the encapsulated medium. In all cases superior results were obtained
on medium encapsulated with 30% polybutene 32 and 70% dental wax. A diet
composed of equal parts of fresh ground beef and beef liver supplemented with
sucrose for continuous rearing of G.
punctipes was produced
(Cohen 1985). The ingredients were blended into a paste and small aliquots wrapped
in stretched Parafilm presented to developing nymphs for feeding. Twelve
generations were successfully cultured, and artificially reared predators
displayed greater fecundity and adult weight than individuals reared on
insect eggs and coddled larvae (Cohen & Urias 1986). Nevertheless,
development was slower on the artificial diet. Parasitoid Cultures
In Vitro In vitro
culture offers a simple alternative for mass culture (Mellini 1978, Greany et
al. 1984), and also enables dietary and nutritional manipulations for
fundamental studies of nutrition and biochemistry. Some benefits of in vitro culture were given by Greany et al. (1984). However,
the physiological and metabolic adaptations exhibited by insect parasitoids
in relation to their parasitic way of life are of critical importance for
successful in vitro culture (Mellini 1975a,
Thompson 1981a, Grenier et al. 1986, Campadelli & Dindo 1987).
Parasitoid/host relationships are often incorrectly thought to lack the
complex physiological interactions typical of the host associations of other
Metazoa (Thompson 1985, 1986a, Dindo 1987). The immature stages of many
parasitoids are truly parasitic and such parasitoid/host relationships are
characterized by extensive physiological and biochemical interaction (Beckage
1985, Thompson 1985, 1986a; Lawrence 1986a). Such interactions are often
intimately associated with nutrition and successful development of the
parasitoid in the host (Beckage & Riddiford 1983, Thompson 1983a, 1986a).
The potential importance of the host endocrine system and of hormonal
interaction in in vitro culture was discussed by
Mellini (1975b, 1978, 1983) and Grenier et al. 1986). Greany (1986) discussed
physiological interaction with reference to the culture of hymenopterous
larval endoparasitoids. The extent that parasitoid/host physiological
interactions need to be considered in the successful development of in vitro culture must still be determined but will
undoubtedly vary with the parasitoid species. Diptera.--A variety of natural foodstuffs, including fish and liver
products, were utilized in early rearing attempts with parasitoids. House
& Traer (1948) reared the sarcophagid A.
housei for many generations
on a diet of salmon and liver. Contrasted to 38% pupation among larvae reared
on the host, Choritoneura fumiferana (Clemens), 88%
pupated when reared on the artificial medium. A related species, Sarcophaga aldrici Parker was reared on the same medium and on liver
alone (Arthur & Coppel 1953) and subsequently Coppel et al. (1959) maintained
A. housei in the laboratory on fresh pork liver. About 1,000 A. housei larvae were reared on 1/2 lb. of sliced liver and
were not affected by putrefaction of the tissue. Smith (1958) maintained Kellymyia kellyi (Aldrich) for 40 generations on pork liver and was
also able to rear larvae on a mixture of powdered milk, powdered egg and
brewer's yeast moistened with water to form a thick paste. House (1954) developed the first chemically defined medium for
rearing a parasitoid, using A.
housei. The diet contained
19 amino acids, ribonucleic acid, dextrose, inorganic salts (U.S.P. XII), B
vitamins, choline and inositol. It was prepared aseptically and gelled with
agar. About 84% of the larvae reached the 3rd instar, 60% of those pupated
and 32% of the pupae emerged as adults. The medium was later refined and many
of the developmental nutritional requirements of A. housei
were determined (House 1977). Vitamin E was necessary for reproduction (House
1966c). Other dipterous parasitoids have been more difficult to
culture outside the host. Many of these species have specialized
physiological adaptations associated with parasitism that are lacking in
sarcophagids. Tachinids, for example, have relatively high respiratory rates
(Ziser & Nettles 1979, Bonnot et al. 1984) and during or immediately
following the first stadium form a direct connection to the host's tracheal
system (Kellen 1944, Fisher 1971). First instar larvae of the parasitoid E. bryani attach to the host's tracheal system 12 hrs after
hatching, and respiratory considerations were critical for the development of
in vitro cultures (Nettles et al. 1980). During initial
studies, first instar larvae dissected from the host were placed directly in
a liquid artificial diet. They were then transferred to diets gelled with
agar, thereby exposing larvae directly to atmospheric oxygen. Improvements in
the methods allowed development without transfer. Powdered artificial diet
containing 1.5% agar was preconditioned by maintaining it at 5% RH for 24
hrs. The diet was then poured into petri dishes and held at 90% RH. Young
larvae dissected from the host 18-24 hrs after larviposition fed on the
liquid diet covering the surface of the gelled medium, and this was
consistent with the normal feeding habit of first instar larvae that feed on
and develop in the host's hemolymph. As the liquid was slowly absorbed by the
agar gel, the surface of the gelled medium dried and larvae were exposed to
the atmosphere. The artificial medium for rearing E. bryani
was composed of mixtures of organic acids, amino acids, nucleic acid bases, B
and fat soluble vitamins, phospholipids and derivatives as well as ATP,
lactalbumin hydrolysate, bactopeptone, yeastolate, albumin, cholesterol,
triolein, glucose and trehalose. When thus reared, larvae developed at an
equivalent rate as when reared on the host, H. virescens,
and 13% developed into adults with a sex ratio of ca. 66% females. Adults
were fecund but produced fewer progeny than host reared insects. The medium
was later refined and simplified and some of the basic developmental
nutritional requirements of E.
bryani were determined
(Nettles 1986a). The nutritive values of adding albumin or soy flower to the
medium was tested, which greatly increased adult yields and fecundity
(Nettles 1986b). Other tachinid parasitoids have been successfully reared on
artificial media. Larval development of Phryxe
caudata Rondani to the 3rd
instar was obtained with a liquid artificial diet (Grenier et al. 1974).
However, in contrast to the results of Nettles et al. (1980) with E. bryani, development of P.
caudata was not improved by
rearing larvae on gelled diets (Grenier et al. 1975). It was suggested that
this may have resulted from the slower development rate and respiratory
requirements of the latter when reared in
vitro (Nettles et al. 1980).
Bonnot (1986) discussed the importance of respiratory requirements in the in vitro culture of P.
caudata. The first tachinid
that was successfully cultured in
vitro on artificial media
from the first instar larvae to the adult was Lixophaga diatraeae
(Townsend) (Grenier et al. 1978). This medium contained organic acids, amino
acids, B and fat soluble vitamins, gelatin, enzymatic hydrolysates of casein,
soy protein, lactalbumin, ovalbumin, ATP, cholesterol, lecithin and gelled
with agarose. Adults were fecund and their progeny developed normally on Galleria mellonella. One critical factor for successful development
of both P. caudata and L. diatraeae was osmolality, which could not exceed 450
mOs/Kg (Grenier et al. 1986). Grenier (1979) investigated the embryonic development of P. caudata and L.
diatraeae on artificial
media. Newly fertilized eggs were removed from adult females and placed on an
agarose-gelled medium similar to that for the larvae. Larval yield was equal
to that observed in vivo and was much greater than
when reared on a liquid diet. Again, respiratory requirements seemed critical
for success. Hymenoptera.--Simmonds (1944) made the first attempt to rear hymenopterous
parasitoids in vitro. Three species of
ichneumonid ectoparasitoids were maintained as larvae for extended periods on
raw beef and gelatin. Although some growth was observed, none could complete
their development. Bronskill & House (1957) did succeed in rearing C. turionellae on a slurry of pork liver in 0.8% saline. An
autoclaved homogenate of the liver was dispensed into sterile test tubes and
surface sterilized eggs were dissected from host pupae and transferred to
this medium. Mature larvae were placed in gelatin capsules for pupation and
7% of the eggs developed to adults. When reared naturally on G. mellonella, 50% parasitoid adults were obtained. Culture
of the ichneumonid I. conquisitor on a diet similar
to that developed by House (1977) for A.
housei was obtained by
Yazgan & House (1970). The first holidic diet for rearing a hymenopterous
parasitoid in vitro was reported by Yazgan
(1972) for I. conquisitor. The diet was a
mixture of amino acids, fatty acids, fat soluble vitamins, B vitamins and
lipogenic growth factors, and glucose, RNA and gelled with agar. It was
ground into a viscous slurry. Parasitoid eggs dissected from the host were
placed directly on this medium, and development from egg to fecund adult was
obtained with a development time twice that observed on the natural host, G. mellonella. Exeristes
roborator was reared on a
diet with a similar nutrient composition (Thompson 1975), but unlike I. conquisitor, larvae of this parasitoid would not tolerate
direct contact with gelled media. Direct exposure to atmospheric oxygen was
important for successful in vitro culture of E. roborator and success was achieved by retaining
suspensions of the liquid diet in lipipholic Sephadex LH-20 gel filtration
medium. Mortality, size and development time of the parasitoid reared in vitro were similar to those of individuals reared on Pectinophora gossypiella (Saunders). Many of
the developmental nutritional requirements of I. conquisitor
and E. roborator were determined by Yazgan (1972) and Thompson
(1976a,b). Thompson (1980, 1981d) described the various chemically
defined diets for rearing various chalcids of the genus Brachymeria. Complete development of B. lasus
from egg to adult at rates approximating those observed in G. mellonella were obtained on diets containing heat-denatured
albumin, amino acids, glucose, B vitamins, inorganic salts, lipogenic growth
factors and Intralipid. The latter, a phospholipid emulsion of soybean oil,
was necessary for complete development. Larvae were reared from eggs
dissected from host pupae immediately following oviposition and parasitoids
were cultured individually in the wells of micro tissue culture plates.
Development of larvae was ca. 2X as long on the synthetic medium compared to
the insect host, and ca. 80% reached the active adult stage. Interestingly,
the yellow coloration of the femur did not develop if vitamin A was lacking. A critical factor in formulating the artificial media for B. lasus was osmotic pressure (Thompson 1983b). The effect of
both carbohydrate and amino acid levels was similar and appeared closely
related to osmolality. Optimum osmotic pressure in the artificial diets
ranged from 550-700 mOs/Kg which was much greater than the 350-450 mOs/Kg of
host hemolymph and tissues. Complete development of the pteromalid Pachycrepoideus vindemiae
(Rondani) was not obtained on an artificial medium similar to that used
successfully for in vitro culture of Brachymeria (Thompson 1981c).
When the amino acid component was replaced with a mixture of the corresponding
polymerized amino acids and the osmolality was reduced to ca. 390 mOs/Kg,
development from egg to adult was obtained (Thompson et al. 1983c). These studies demonstrate that the importance of osmotic
pressure varies with the parasitoid species. Parasitoids such as I. conquisitor and E.
roborator are very tolerant
of osmotic pressures. Artificial diets that supported in vitro
culture of these species had osmolalities of ca. 2,000 mOs/Kg. On the other
hand, the tachinids, P. caudata and L. diatraeae (Grenier et al. 1986), and the pteromalid P. vindemiae did not develop at osmolalities of >450
mOs/Kg. Pteromalus puparum was
cultured in vitro by Bouletreau (1968,
1972). Complete development on host hemolymph in hanging drop slide mounts
was obtained. Similar results were reported by Hoffman et al. (1973). Hoffman
& Ignoffo (1974) had limited success with an artificial medium containing
yeast hydrolysate, fetal bovine serum and Grace's tissue culture medium.. Tetrastichus schoenobii Ferriere was reared
on modified Gardiner's tissue culture medium supplemented with egg yolk, milk
and hemolymph from Anteraea pernyi Guérin (Ding et al.
1980a). About 60% of the parasitoids completed development to the adult stage
with no deformities nor abnormal fecundities. Greany (1980, 1981) described
studies on the in vitro embryonic development of
the braconid Cotesia (= Apanteles) marginiventris (Cresson), maintained in Grace's tissue
culture medium supplemented with fetal bovine serum, bovine serum albumin and
whole egg ultrafiltrate. Insects were reared from the embryonic germ band
stage to mature first instar larvae on this diet cocultured with host fat
body tissue. Greany (1986) obtained similar results with Microplitis croceipes.
Emphasis was placed on the importance of protein nutrition for success and
protein secretion by the fat body was considered a factor to explain the need
for this tissue for successful embryonic development. Vinson & Iwantsch (1980) found that teratocytes (cells
derived from the embryonic membrane of the parasitoid egg) are released into
the host hemocoel at the time of egg hatching. It was suggested that the
teratocytes may play a role in parasitoid nutrition. Sluss (1968)
demonstrated that the teratocytes of Perilitus
coccinellae (Shrank) increased
in volume several times in the coccinellid host and where then subsequently
eaten by the developing parasitoid larvae. Greany (1980) found that
teratocytes present in artificial culture medium for C. marginiventris
caused dissociation of cocultured fat body and suggested that the teratocytes
might facilitate larval growth. Rotundo et al (1988) obtained complete larval
development of the braconid Lysiphelebus
fabarum (Marshall) on a
similar artificial diet that was lacking in fat body and teratocytes. Strand et al. (1988) demonstrated a role for teratocytes in
the successful in vitro culture of the egg
parasitoid Telenomus heliothidis Ashmead. Embryonic
development of T. heliothidis was obtained in
Hinks TNH-FH medium containing 30% w/v M.
sexta hemolymph. Mature
embryos were transferred to a medium containing 40% M. sexta
hemolymph, chicken egg yolk, trehalose and milk. Development to the adult
stage required one day more than on the host H. virescens
and 42% of the larvae became adults. The sex ratio was ca. 50% females. The
presence of teratocytes had no effect on larval development to the third
instar. However, when teratocytes were removed from the medium during larval
development, pupation was greatly reduced and the development time of parasitoids
that completed development increased. The authors concluded that the
teratocytes aided larval feeding by dispersing the particulate material in
the medium and solubilizing nutrients. It was suggested by Strand et al.
(1986) that teratocytes of T.
heliothidis aided in
decomposition and necrosis of host tissue partially due to release of lytic
enzymes. Therefore their function in
vitro might be the same that
occurs during the normal development of the parasitoid in the host, Heliothis virescens. Culture of Trichogramma
pretiosum in vitro was first attained by Hoffman et al. (1975)
following unsuccessful attempts by Rajendram (1978) with T. californicum
Nagaraja & Nagarkatti. Trichogramma
pretiosum completed
development on filter paper discs soaked in sterile H. zea
hemolymph. In vitro culture to the adult
stage required ca. 25% more time than observed on the host, Trichoplusia ni (Hübner). Even though most
adults did not fully expand their wings, they mated and laid eggs without
difficulty. Progeny from eggs of in
vitro cultured parasitoids
had a sex ratio of 1.2:1 males/females when reared on host eggs. Hoffman et al. (1975) reported development to the prepupal stage on a
semisynthetic artificial diet similar to that described by Hoffman &
Ignoffo (1974) for P puparum, but supplemented with
wheat germ oil. Strand & Vinson (1985) obtained complete in vitro culture of T.
pretiosum on an artificial
medium similar to that used by Thompson (1981d) for B. lasus
but supplemented with ca. 40% M.
sexta hemolymph, the latter
being required to induce pupation. Survival to the adult stage was 70s% and
the sex ratio ca. 1:2 males/females. Xie et al. (1986a) also reported that host hemolymphs was required for
pupation of T. pretiosum and that factors in
the host egg influenced adult emergence. Irie et al. (1987) reported that the
requirement of host hemolymph for the complete in vitro
development was due to the presence of specific factors that could be
extracted in 76% ethanol. Purification of the pupation factor by
chromatographic methods showed the presence of two active carbohydrate
containing factors. Trichogramma dendrolimi
Matsumura was cultured in vitro in hanging drop mounts of
hemolymph from A. pernyi Guan et al (1978). Liu
et al (1979) reported success in hanging drop mounts containing media with A. pernyi or Attacus
cynthia (Drury) hemolymph
and chicken egg yolk, bovine milk, organic acids and procine serum. The
extent of development of Trichogramma
japonicum Ashmead, T. australicum Girault and T. evanescens
was not reported, however. Wu et al. (1980, 1982) and Wu & Qin (1982a)
obtained successful culture of T.
dendrolimi to the adult
stage on media without host hemolymph but containing chicken egg yolk,
chicken embryo fluid, bovine milk, and amino acid mixture and peptone.
However, only 16% of the eggs completed development, and most adults were
females of poor vitality. The results did suggest that in contrast to T. pretiosum, the in
vitro culture of T. dendrolimi does not require host factors (Xie et al.
1986a, Irie et al. 1987). Liu & Wu (1982) reported on in vitro culture of T.
dendrolimi, Trichogramma confusum Viggiani and T. pretiosum on a medium of yeast hydrolysate, fetal calf
serum, Grace's tissue culture medium, chicken embryo extract, bovine milk and
chicken egg yolk. However, adults were less viable than normal and displayed
abnormal wing development. The cooperative Research Group of Hubei Province,
China (CRGHP 1979) has carried out extensive studies on the complete in vitro culture of T.
dendrolimi in artificial
media encapsulated in artificial eggs into which the adult females
oviposited. Gao et al (1982) reported rearing 35 continuous generations of
this species in hanging drop mounts of the artificial medium. Some studies have tried to determine the effects of hormone
supplementation on parasitoid development in
vitro, with generally
negative results. The tachinid Gonia
cinerascens Rondani depends
on its host's endocrine system for growth and development, but was not
induced to mold from the 1-2nd instar by addition of 20-hydroxy (b) ecdysone
to an artificial medium of host tissue homogenate and Grace's tissue culture
medium. Development from the 2nd instar to adult was reported on artificial
medium in the absence of hormones, indicating that some hormones may be
necessary for the 1-2nd instar molt in
vitro. The 20-Hydroxy
ecdysone failed to stimulate development of B. intermedia
in vitro (Thompson 1980); however, Greany (1980, 1981)
reported that this hormone inhibited egg hatching in C. marginiventris
and ecdysone, 20-hydroxy ecdysone and the juvenile hormone analog hydroprene
had no effect on larval growth or development. The deleterious effect of this
hormone could be overcome by simultaneous application of hydroprene. Nenon (1972a,b) demonstrated that hormones greatly increased in vitro survival of developing embryos and larvae of the
encyrtid Ageniaspis fuscicollis (Dalman). The
parasitoid was maintained on a diet of chicken embryo extract, beef peptone
and equine serum. Ecdysteroid or juvenile hormone added in the medium had
little effect, but when included together, resulted in nearly 100% survival
to the 2nd instar. Further study of the effects of host hormones in vitro culture systems must require careful and detailed
experimental design. Hormones act in a complex and often synergistic way, and
the timing of their application as well as the method of exposure may prove
critical to assessing their potential. There is no doubt as to the importance
of hormonal interaction to the successful development of parasitoids in vivo, particularly with regard to synchronizing parasitoid
development to the host's life cycle. Adults of many entomophaga must feed, and although adult
parasitoids and predators are usually fed in the laboratory, early workers
had largely ignored the significance of such feeding in nature. Bierne (1962)
considered that many biological control attempts failed as a result. Leius
(1967a) gave one of the first field demonstrations of the importance of adult
feeding when he reported a relationship between the natural abundance and
variety of wild flowers in apple orchards and the incidence of parasitism of Malacosoma americanum (F.) and Laspeyresia
(= Carpocapsa) pomonella (L.) by the
parasitoids, I. conquisitor, Apophua (= Glypta) simplicipes
(Cresson), Scambus hispae Harris, Telonomus sp., Ooencyrtus clisiocampae (Ashmead), and Eupelmus spongipartus
Foerster. Eighteen times as many M.
americanum pupae, four times
as many M. americanum eggs and five times
as many L. pomonella eggs were parasitized
in orchards with an undergrowth of wild flowers when compared with other
orchards lacking such flora. The early literature describing how adult parasitoids feed
from flowers and other plant parts was reviewed by Leius (1960). Generally
insects fed on floral and extrafloral nectars as well as pollens. Although
knowledge of the specific nutritional requirements of adult entomophagous
insects is limited, much data are available on the chemical and nutritional
requirements of adult entomophaga is limited, much is available on the
chemical and nutritional composition of these plant products. Floral nectars
contain up to 75% by weight of simple sugars, mainly sucrose, fructose and
glucose (Baker & Baker 1983), but considerable qualitative and
quantitative differences exist between plant species. Free amino acids are
also abundant in nectars although most nectars do not contain all 10
essential amino acids. Small amounts of proteins, lipids, dextrins and
vitamins that are nutritionally beneficial are also found. The composition of
extrafloral nectars is also complex (Baker et al. 1978). Pollens have a
complex composition of small molecular nutrients and many pollens have high
levels of free amino acids (Barbier 1970, Stanley & Linskens 1974). By
comparison, pollens generally have higher levels of protein, lipid and
polysaccharides. Pollens and nectars together can provide a complete diet for
successful growth, development and reproduction. The predator Coleomegilla maculata lengi Timberlake can complete larval development on pollen
alone (Smith 1961); therefore, when prey are scarce, plant products may play
a critical role in maintaining predators (Hodek 1973). Hagen (1986a)
discussed the complex ecological and evolutionary interactions between plant
flowers, nectars and pollens and several insect groups. Leius (1960) examined the plant feeding habits of I. conquisitor, Scambus
buolianae (Hartig) and Orgilus obscurator (Nees). The attractiveness of the flowers of
wild mustard, white sweetclover, wild parsnip, silky milkweed and annual
sowthistle were tested. Except for annual sowthistle, I. conquisitor
was attracted to and fed from all flowers tested, but was most attracted to
wild parsnip. Similar results were shown with S. buolianae.
Orgilus obscurator was attracted to and fed on wild parsnip only,
but further tests revealed that this parasitoid also fed on other
umbelliferous plant flowers, including those of wild carrot and water
hemlock. The nutritive value of various pollens for fecundity and longevity
of S. buolianae was reported by Leius (1963). Itoplectis conquisitor and S.
buolianae accepted various
semi-natural foods also, including honey, sucrose solution with or without
plant pollens and raisins. Plant feeding behavior of O. obscurator
examined by Syme (1975) showed a broad range of food plants, including
species from five families. Adult parasitoids may emerge prior to the
availability of the insect host, and Syme (1977) suggested that a variety of
plant species be provided as food to ensure sufficient longevity of the adult
female. Lingren & Lukefar (1977) demonstrated that adult Campoletis sonorensis (Cameron), a parasitoid feeding on the
extrafloral nectar of cotton, lives longer when exposed to extrafloral
nectaried cotton than nectariless cotton. Parasitism of hosts was higher on
the nectaried form. Adejei-Maafo & Wilson (1983) showed that 15
categories of entomophaga, including the predators Deraeocoris signatus
(Distant), Geocoris lubra (Kirkaldy), Nabis capsiformis Germar, Chrysopa
spp., Laius bellalus Guérin, Coccinella repanda (Thunberg) and Verania
frenata Erichson, were
present at densities of 2-3 times higher on nectaried versus nonnectaried
cotton. Although contribute to
attraction for plants in these insects, the nutrition provided by nectars and
pollens seems to be important. Hemptinne & Desprets (1986) reported that
following hibernation Adalia
bipunctata (L.) fed on
pollens as an alternate food which allows the predators to lay eggs as soon
as prey become available. As was discussed in an earlier section, in addition to feeding
plants and plant products many parasitoids are host-feeders. Adult female
Hymenoptera puncture or damage host larvae or pupae and feed on the hemolymph
and/or internal tissues. Kidd & Jervis (1989) estimated that as much as
1/3rd of the world's parasitoid fauna (>100,000 species) host feed. Some
parasitoids may kill more host individuals by host feeding including
ovipositor probing followed by host rejection, than by parasitization
(Johnston 1915, DeBach 1943, 1954). Legner (1979) emphasized
that consideration of a parasitoid's host destructive capacity was important
to correctly evaluate the impact of periodic inundative field releases on
pest populations, and Greathead (1986) and Yamamura & Yano (1988)
suggested that host-feeding behavior was important for assessing the
potential of a biological control agent. Kidd & Jervis (1989) recently
discussed the significance of host-feeding on parasitoid-host population
dynamics. Bartlett (1964) in examining host-feeding in the encyrtid, Microterys flavus Howard, was among the first to correlated
host-feeding behavior with nutrition. He hypothesized that host feeding
developed coincidentally with depletion of eggs and suggested that host
mutilation was a reflection of "frustrated" host feeding when the
host failed to bleed readily. Host feeding by M. flavus
was usually displayed following egg-laying, and oviposition resumed after
host feeding. Reviewing this predatory habit for adults from 20 families of
Hymenoptera, Bartlett concluded that the behavior was indicative of the
necessity for dietary supplementation of some ubiquitous substances required
by many diverse species. He reported that a food supplement of enzymatic
yeast and soy hydrolysate with honey satisfied the nutrient requirements for
sustaining reproductive activity in M.
flavus, and suggested that a
protein nutrient source may be necessary. The difference between proovigenic and synovigenic Hymenoptera
was discussed earlier, categories proposed by S. E. Flanders (1950). Females
of proovigenic parasitoids complete oogenesis prior to or shortly after
emergence and lay eggs over a relatively short period of time principally on
larval stages of their host. Host feeding is important for ensuring that the
female lives long enough to deposit all eggs. In contrast, females of
synovigenic species eclose with a minor fraction of their total egg complement
as mature eggs. Synovigenic parasitoids attack primarily host eggs and pupae,
are longer lived than proovigenic species and produce eggs throughout their
adult lives. To sustain oogenesis the females of many synovigenic species
require additional nutrients. Based on the egg type, Dowell (1978) described
two types of synovigenic parasitoids: (1) those producing large anhydropic or
yolk-rich eggs that contain sufficient nutrient for completion of embryonic
development prior to oviposition. Parasitoids that produce anhydropic eggs
obtain nutrition for sustaining egg production by host-feeding; (2) those
producing hydropic or yolk-deficient eggs. Embryonic development in hydropic
eggs occurs in the host following oviposition, in which case the adult does
not require additional nutrient to support egg development and has no
requirement to host feed. Legner & Gerling (1967) showed the
importance of early host feeding and oviposition to pteromalids of the first
type, as was previously discussed. Leius (1962, 1967b) demonstrated the
importance of feeding habits to fecundity of S. buolianae.
Egg production was reduced to 1/3rd and longevity to 2/3rds, when females
were permitted to host-feed intermittently or were deprived after 15 days of
age. No eggs were laid if females were deprived for 20 days. The effects of
feeding host body fluids, in conjunction with honey, pollen and raisins on
fecundity and longevity of S.
buolianae and I. conquisitor were examined by Leius (1961a,b). Maximum
fecundity and longevity of both species were obtained when host fluids and
seminatural foods were provided together. Host feeding was nevertheless
essential, and S. buolianae did not lay eggs when
deprived of host hemolymph or tissues. The feeding behavior of 140 hymenopterous parasitoids was also
reviewed by Jervis & Kidd (1986), who concluded that host feeding was
important for egg fecundity or egg production, while non-host foods were
important for maintenance and longevity. Four types of host feeding
distinguished were (1) concurrent feeding where the female used the same host
individual for feeding and oviposition, (2) nonconcurrent if the female used
different host individuals for feeding and oviposition, (3) the feeding habit
may be nondestructive or destructive (the host may survive or may die), and
(4) destructive feeding which generally resulted in a host that was
unsuitable for oviposition. Parasitoids were found to differ in their
lifetime and diurnal patterns of feeding, and it was concluded by Jervis
& Kidd (1986) that concurrent/nondestructive feeding was most likely when
hosts were readily available and that destructive feeding was advantageous
when host density was low. Jervis & Kidd (1986) also gave several models to assess
how the energetic demands and constraints on a parasitoid affect its
host-feeding strategy. One model predicted the feeding strategy for
maximizing egg production of a single synovigenic female (see Thompson &
Hagen 1999, for formulae). Host feeding also occurs among dipterous parasitoids but is
not as common as in Hymenoptera (Clausen 1940). Host feeding by tachinid
parasitoids may affect longevity and fecundity (Shahjahan 1968)(. Nettles
(1987b) demonstrated that fecundity was prolonged by feeding E. bryani host hemolymph compared with feeding a sucrose
solution. The effect of host feeding on fecundity could not be simulated by
substituting a solution of free amino acids or bovine serum albumin. The excretion of various Homoptera, such as honeydew, may
serve as a food for many adult entomophaga. Neuropterans of the genus Chrysoperla and other genera
with nonpredaceous adults feed actively on honeydew as well as on nectar and
pollen (Principi & Canard 1984). Although honeydew does not contain all
the essential amino acids, yeast symbiotes residing in the gut can provide
the missing amino acids in some nonpredaceous species (Hagen & Tassan
1972). Neuropteran predatory adults also feed on honeydew, but reproductive
activity ensues only after prey are eaten (Hagen 1986a). Hagen (1962) found
that honeydew alone will not stimulate egg production in coccinellid
predators. Dipterous and hymenopterous parasitoids also have been found to
feed on honeydew (Clausen 1940, Zoebelein 1956). The importance of honeydew
as a supplementary food was suggested by Clausen et al. (1933) in work with Tiphia matura Allen & Jaynes. Female adults traveled long
distances from the location of their host to feed on honeydew, which
migration occurred annually. Ichneumonids of the genus Rhyssa appeared dependent on honeydew for maintaining the
longevity necessary to parasitize and regulate populations of Sirex (Hocking 1967). The
nutritional value of honeydew for parasitoids varies with the homopteran
source, as Wilbert (1977) showed considerable differences in longevity of
several Hymenoptera when fed aphid or coccid honeydew. Nutritional requirements of adult entomophagous insects are
obscure. Bracken (1965, 1966, 1969) examined some requirements of the
parasitoid Exeristes comstockii (Cresson), finding
that adult females fed an artificial medium containing amino acids, sucrose,
fatty acids, cholesterol, vitamins and inorganic salts produced eggs at an
equivalent rate as individuals fed Galleria
mellonella (L.) larvae and
sucrose. Egg production was reduced or eliminated when amino acids, sucrose,
vitamins or salts were deleted. Sucrose, pantothenic acid, folic acid and
thiamine were all essential for egg-laying. Nutritional requirements of adult
predators similarly are not well known. Numerous semi-natural diets have been
successfully developed for maintaining chrysopid predators and various adult
coccinellids. It seems that predators require a complete and well balanced
diet to ensure maximum longevity and reproductive potential. The effects of
various diets on fecundity of some chrysopids was summarized by Hagen
(1986b), and nutritional data for adults of several other species by Roussett
(1984). Continuous
Culture on Artificial Media The ultimate goal of studies on in vitro
culture of entomophagous insects is continuous artificial culture without the
host insect. In order to achieve this goal, careful scrutiny of factors that
otherwise would not be considered of direct important to nutrition must be
made. Commercial parasitoid culturing requires the direct deposition of eggs
or larvae onto an artificial substrate. Artificial food must be acceptable
for feeding by all stages of a predator. Behavioral considerations may be
critical for the successful continuous culture of many entomophaga. Successes
with in vitro culture thus far reflect the level of complexity of
behavioral interactions between parasitoid and host or predator and prey. The
first success with parasitoids was achieved with Sarcophagidae, many of which
readily oviposit and develop on carrion. Sarcophaga
aldrici and K. kellyi were reared for many generations on fish and liver,
respectively (Arthur & Coppel 1953, Smith 1958). Agria housei
was reared continuously for 756 generations on pork liver. However, the
behavioral interaction between many parasitoids and their hosts are complex,
involving numerous physical and chemical cues that initiate specific behavior
which leads to oviposition. Host selection and successful parasitism is a
multistep process which involves host habitat location, host location, host
acceptance, host suitability and host regulation, as was discussed in
previous sections (Doutt 1959, Vinson 1976, 1984). Factors that influence
host acceptance in particular are critical for continuous culture. The different
events which lead to successful oviposition, including examination of the
host, probing with the ovipositor, insertion and oviposition (Schmidt 1974)
may each be stimulated by different chemical as well as physical cues (Arthur
1981, Vinson 1984). These cues may be associated with the host species, the
plant or other food source of the host, or may result from interactions
involving both the host and its food (Vinson 1975). Physical factors
associated with the host's food plant are essential for successful
oviposition and parasitism by G.
cinerascens (Mellini et al.
1980). This tachinid deposits microtype eggs on the leaves of certain plants,
and host larvae become infected by ingesting the eggs. Leaf color, shape,
thickness, size and reflectivity are among the several factors which
influence oviposition in this species. Mellini et al (1980) constructed
polished, thin, yellow oval-pointed artificial bee's wax leaves, 2-7 cm2,
on which large numbers of parasitized eggs were laid. This parasitoid readily
developed in G. mellonella after host feeding
on the artificial leaves. Complex combinations of physical cues, including
size, shape, color, texture and movement have been demonstrated to have
influence on oviposition behavior in parasitoids (Arthur 1981, Jones 1981,
Nordlund et al. 1981). Important roles are played by chemicals in both
parasitoid-host and predator-prey interactions (Arthur 1981, Greany and Hagen
1981, Vinson 1984, Hagen 1986a). The involvement of chemicals in host
acceptance and oviposition by parasitoids is well documented. During
predator-prey relationships, produced
by the prey may serve as attractants, arrestants and/or phagostimulants. Chrysopa carnea adults, e.g., are attracted to a variety of
chemicals such as tryptophan byproducts (Hagen et al. 1976). Although studies
with numerous predaceous insects have demonstrated the role of in prey
finding and recognition, their role in feeding is not well established. Deployment of behavior modifying chemicals in continuous
artificial culture has involved only a few species. Itoplectis conquisitor
accepts a host and oviposits following detection of specific components of
host hemolymph during ovipositor probing (Arthur et al. 1969). This
parasitoid even oviposited into host hemolymph that was placed on paraffin
tubes. The active fraction was colorless, water soluble and gave a strong
reaction to ninhydrin and folinphenol reagents. It had a molecular weight of
ca. 7,000, was heat stable and nondializable. Arthur et al (1973) concluded
that the stimulant was proteinaceous and they were successful in stimulating
similar oviposition activity with a variety of amino acid mixtures containing
trehalose and/or MgCl2. The best results were with a mixture of
serine (0.5M), leucine (0.065 M), arginine (0.05 M) and MgCl2
(0.025 M). The ovipositional activity observed greatly exceeded that
stimulated by the host hemolymph. House (1978) then developed a synthetic
artificial host comprised of an artificial diet encapsulated in paraffin. The
diet was based on that described by Yazgan (1972) and contained gelatin,
casein, inorganic salts, amino acids, glycogen, lipids, trehalose, glucose,
water and fat soluble vitamins and agar. Female parasitoids readily accepted
and oviposited into the artificial host, and the first successful complete
artificial culture of a hymenopterous parasitoid was realized. However, only
one single adult male was obtained. There have been considerable studies to determine how chemical
and other factors influence adult reproductive capacity in in vitro cultures. Larviposition by E. bryani
is stimulated by kairomones which emanate from the host's cuticle, and female
adults examine artificial hosts coated with cuticular extracts with great
care (Burks & Nettles 1978). Tucker & Leonard (1977) extracted a
kairomone from the pupae of Lymantria
dispar that appeared
responsible for ovipositional behavior by Brachymeria
intermedia. Tetrastichus schoenobii was stimulated to
oviposit in artificial eggs coated with host scales (Ding et al. 1980b). The parasitoid group, which has received the most attention,
is the Trichogrammatidae. There have been more extensive efforts to develop
continuous artificial culture with Trichogramma
spp. than with other parasitoids. Many aspects of the ovipositional behavior
of this genus were described by Salt (1934, 1940) in studies on T. evanescens (Fisher 1986). Recent studies demonstrate the
importance of physical (Rajendram & Hagen 1974) and chemical factors,
including kairomones (Nordlund et al. 1985) for eliciting oviposition.
Rajendram (1978a,b) obtained artificial oviposition by T. californicum
into physiological saline or Neisheimer's salt solution encapsulated in
paraffin. Nettles et al (1982, 1983) reported that a dilute solution of KCl
and MgSO4 induced oviposition by T. pretiosum
into artificial wax eggs (Nettles et al. 1984). Leucine, Phenylalanine and/or
isoleucine stimulated oviposition by T.
dentrolimi in artificial
eggs (Wu & Quin 1982b). Adult females laid more eggs than when insect
hemolymph was used then employing a complete mixture of all three amino
acids, 600, 400 and 320 mg/100ml. A synthetic membrane was developed as an
alternative for paraffin through which T.
pretiosum would oviposit
(Morrison et al. 1983). The silicone-polycarbonate copolymer was clear,
highly elastic and adult females oviposited through the surface into an
ovipositional stimulant at rates that were comparable to host eggs. The use
of polyethylene as an alternative to wax for producing artificial eggs for
oviposition by T. dendrolimi was described by the
Chinese CRGHT (1985). Xie et al. (1986b) demonstrated the potential for large scale continuous
artificial culture of T. pretiosum. Three in vitro culture methods were developed as a follow up to
earlier work by Xie et al. (1986a) and Nettles et al. (1985). These utilized
microtiter tissue culture plates, multiple drop rearing in petri plates and
flooded petri plat rearing. The basic diet was 50% heat treated insect
hemolymph, 25% egg yolk, 15 g/100 ml dried milk suspension and 0.15%
gentamycin. Each method supported large populations of parasitoid larvae.
Microbial contamination and subsequent loss of entire petri plats was a major
obstacle but several antibiotics were available for reducing losses. Field trials
with in vitro reared Trichogramma
have been made. Continuous artificial mass culture of T. dendrolimi
was described by Li (1982) and Gao et al. (1982), who reported that field
release of in vitro reared parasitoids
resulted in 93% parasitism of Heliothis
armigera (Hübner) eggs in
cotton. Artificial mass culture of Chrysopa
carnea was described by
Yazlovetskij & Nepomnyashchaya (1981) after the development of a suitable
artificial medium for supporting larval development (Nepomnyashchaya et al.
1979). The medium was microencapsulated and composed of casein hydrolysate,
brewer's yeast extract, soybean oil. wheat germ extract, sucrose, lecithin,
choline, cholesterol and ascorbate. The effectiveness of the artificially
reared larvae against Myzus persicae was equal to that of
insects reared on eggs of S.
cerealella. A
microencapsulation technique for mass producing artificial eggs for C. carnea was also described by Morrison et al. (1975). Considerations of how nutrition currently applies in
biological control programs, focuses on its purpose as being restricted to
use of food and food supplements to enhance the activity and effectiveness of
entomophagous insects in the field as suggested earlier (Hagen & Hale
1974, Hagen & Bishop 1979. Greenblatt & Lewis 1983, Hagen 1986a,
Gross 1987). Such use is dictated by a lack of synchrony between natural
enemies and their hosts and/or isolation of entomophagous insects from the
natural environment that normally supplies alternate food sources such as
nectars and honeydews (Hagen 1986a). These factors occurring in crop
monoculture may intensify following pesticide application. The importance of
nutritional supplements for adult parasitoids and predators is well known,
and recent studies with Trichogramma
demonstrated that fecundity and longevity could be increased by feeding adult
insects (Anunciada & Voegele 1982, Bai et al. 1988). The future use of
feeding prior to or following release in the field may have a significant
effect on biological control successes. Few studies on the effects of feeding
parasitoids on field performance are available, however. Temerak (1976)
reported spraying honey solution on sorghum stalks during winter to provide
supplementary food to Bracon
brevicornis Wesmael in the
absence of pollen, honeydew and nectars. Parasitoid cocoons significantly
increased after spraying and the prevalence of hosts decreased. Despite field
trials employing kairomones for attracting and stimulating host searching by Trichogramma sp. (Lewis et al.
1979, 1982), no attempt has been made to use kairomones in combination with
supplemental foods to maintain parasitoid populations when host numbers are
low. Supplementary food sprays have been successfully deployed with
predaceous insects. Ewert & Chiang (1966) sprayed sucrose solutions on
corn to aggregate coccinellid and chrysopid adults. Increased predator
density and reproductive activity significantly lowered aphid populations.
The numbers of Chrysopa sp.
and Glischiochilus quadrisignatus (Say) were
increased in corn sprayed with sugar or molasses solutions (Carlson &
Chiang 1973), with resultant increased predation resulting in significant
reductions of Ostrinia nubilalis (Hübner). Hagen et al. (1976) working in sugar sprayed alfalfa plots were able to
retain larger numbers of C. carnea and Hippodamia sp. in the field during periods of low host
density. Within 24 hrs the population of coccinellid adults increased 20X and
that of C. carnea 200X. Populations of Lygus spp. also increased after
application of sugar sprays (Lindquist & Sorenson 1970), and Hagen et al.
(1971) concluded that sucrose was an arrestant for adult Lygus and coccinellids. Chrysopa carnea,
coccinellids and Lygus were
attracted to potato plants sprayed with honey, which suggested a critical
role for volatile components (Ben Saad & Bishop 1976a,b). Adding semiochemicals to supplemental foods for C. carnea is useful. The complex interactions of
semiochemicals and food in influencing the behavior of C. carnea
was described by Hagen & Bishop (1979). The adult responds to a volatile
signal, a , from plant habitats in which prey are located and is then
attracted to the prey by tryptophan breakdown products from the honeydew (Van
Emden & Hagen 1976). Specific behavioral and flight patterns shown by C. carnea in response to these interactions were discussed by
Duelli (1980). The habitat synomone affecting the behavior of C. carnea in cotton was shown by Flint et al. (1979) to be
caryophyllene, but several chemicals from other plants also displayed
synomone activity for this species (Hagen 1986b). Chrysopa carnea
adults were successfully attracted to alfalfa fields by applying artificial
honeydews composed of various yeast (Wheast) products and sugar (Hagen et al.
1971). Although the specific synomone of alfalfa is unknown, application of
caryophyllene with the kairomone from tryptophan greatly improved attraction
of C. carnea during the beginning of flowering (Hagen 1986a).
Application of artificial honeydew was also successful for aggregating Hippodamia spp. as well as
coccinellids and other predators. Subsequent trials employing the yeast
mixture in combination with sucrose and honey or molasses applied to various
crops were successful in manipulating C.
carnea populations (Hagen
& Hale 1974). The sugar was essential for retaining C. carnea
adults in the field after attraction. Butler & Ritchie (1971) reported
that C. carnea adults were attracted to the yeast/sugar mixtures
sprayed on cotton, but no increase in egg deposition was noted. Similar
studies demonstrated inconsistent oviposition in grape culture (White &
Jubb 1980). There was no attraction of chrysopid adults in treated apple
orchards (Hagley & Simpson 1981) nor in potato fields when only the yeast
was applied. Dean & Satasak (1983) gave reasons why food sprays might not
be practical in control programs for cereal aphids in England, which included
the variable abundance of univoltine C.
carnea populations, low
plant growth form and the development of sooty mold on plants where food
sprays containing sugar were applied. Duelli (1987) did not find an increase
in oviposition by chrysopids when artificial honeydews were applied to
alfalfa, corn, sunflowers and in prune orchards. It was suggested that the
different responses of sibling species of C.
carnea in Europe and North
America may be related to behavioral differences. REFERENCES: references may be found atMELVYL
Library ] Adejei-Maafo, I. K.
& L. T. Wilson. 1983. Factors affecting the relative abundance of arthropods
on nectaried and nectariless cotton. Environ. Ent. 12: 349-52. Altahtawy, M. M., S. M. Hammad & E. M.
Hegazi. 1976. Studies on the dependence of Microplitis rufiventris
Kok (Hym.: Braconidae) parasitizing Spodoptera
littoralis (Boisd.) on its
own food as well as its host. Z. ang. Ent. 81: 3-13. Anunciada, L.
& J. Voegele. 1982. L'importance de la nourriture dans le potentiel
biotique de Trichogramma maidis Pintureau et Voegele et T. nagarkattii Voegele et Pintureau (Hym.: Trichogrammatidae)
et l'oosorption dans les femelles en contention ovarienne. Les Trichogrammes.
Les Colloques de l'INRA 9: 79-84. Arthur, A. P. 1981. Host acceptance by
parasitoids, p. 97-120. In:
D. A. Nordlund, R. L. Jones & W. J. Lewis (eds), Semiochemicals: Their
Role in Pest Control. John Wiley, New York. Arthur, A. P. & H. C. Coppel. 1953.
Studies on dipterous parasites of the spruce budworm, Choristoneura fumiferana
(Chem.) (Lepidoptera: Tortricidae). I. Sarcophaga aldrichi Park. (Diptera: Sarcophagidae). Canad. J. Zool. 31: 374-91. Arthur, A. P. & H. G. Wylie. 1959.
Effects of host size on sex ratio, development time and size of Pimpla turionellae (L.) (Hymenoptera:
Ichneumonidae). Entomophaga 4: 297-301. Arthur, A. P., W. W. Batsch & L.
Rollins. 1969. Component of the host haemolymph that induces oviposition in a
parasitic insect. Nature 223: 966-67. Arthur, A. P., B. M. Hegdekar & W. W.
Batsch. 1973. A chemically defined, synthetic medium that induces oviposition
in the parasite Itoplectis conquisitor (Hymenoptera:
Ichneumonidae). Canad. Ent. 105: 787-93. Ashby, M. D. & P. Singh. 1987. A
glossary of insect rearing terms. Dep. Sci. Ind. Res., New Zealand Bull. 239.
p. 46. Attallah, Y. H. & L. D. Newsom. 1966.
Ecological and nutritional studies on Coleomegilla
maculata DeGeer (Coleoptera:
Coccinellidae). I. The development of an artificial diet and a laboratory
rearing technique. J. Econ. Ent. 59: 1173-79. Atwal, A. S. & S. L. Sethi. 1963. Biochemical
basis for the food preference of a predator beetle. Curr. Sci. 32: 511-12. Bai, B., R. F. Luck, J. Jansen & B.
Stevens. 1989. Trichogramma
quality. The effects of honey, host egg size and number of wasp progeny per
host egg on the size, fecundity and longevity of Trichogramma pretiosum.
Ent. Exp. Appl. (in press). Baker, H. G. & I. Baker. 1983. A brief
historical review of the chemistry of floral nectar, p. 126-52. In: B. Bentley & T. Elias (eds.),
The Biology of Nectaries. Columbia Univ. Press, New York. Baker, H. G., P. A. Opler & I. Baker.
1978. A comparison of amino acid complements of floral and extrafloral
nectars. Bot. Gaz. 139: 322-32. Barbier, M. 1970. Chemistry and biochemistry of pollens.
Prog. Phytochem. 2: 1-34. Baronio, P. & F.
Sehnal. 1980. Dependence of the parasitoid Gonia cinerascens
on the hormones of its lepidopterous hosts. J. Insect Physiol. 26: 619-26. Bartlett, B. R. 1964. Patterns in the host-feeding habit of
adult parasitic Hymenoptera. Ann. Ent. Soc. Amer.
57: 344-50. Bashir, M. O. 1973. Effect of nutrition on development
and reproduction of aphidophagous coccinellids with species reference to Olla abdominalis (Say). Ph.D. dissertation, University of
California, Berkeley. Beck, S. D. 1972. Nutrition, adaptation and
environment. p. 1-6. In: J.
G. Rodriguez (ed.), Insect and Mite Nutrition. North-Holland, Amsterdam. Beckage, N. E. 1985. Endocrine interactions
between endoparasitic insects and their hosts. Ann. Rev. Ent. 30: 371-413. Beckage, N. E. & L. M. Riddiford. 1983.
Growth and development of the endoparasitic wasp Apanteles congregatus
dependence on host nutritional status and parasite load. Physiol. Ent. 8:
231-41. Beckendorf, S. K. & M. A. Hoy. 1985.
Genetic improvement of arthropod natural enemies through selection,
hybridization or genetic engineering techniques. p. 167-87. In: Biological Control in
Agricultural IPM Systems. Academic Press, New York. Bellows, T. S., Jr. & T. W. Fisher,
(eds) 1999. Handbook of Biological Control: Principles and Applications.
Academic Press, San Diego, CA. 1046 p. Ben Saad, A. A. &
G. W. Bishop. 1976a. Attraction of insects to potato plants through use of
artificial honeydews and aphid juice. Entomophaga 21: 49-57. Ben Saad, A. A. & G. W. Bishop. 1976b.
Effect of artificial honeydews on insect communities in potato fields.
Environ. Ent. 5: 453-57. Bernays, E. A. 1985. Regulation of feeding
behavior. p. 1-32. In: G. A.
Kerkut & L. I. Gilbert (eds.), Comprehensive Insect Physiology, Biochemistry
and Pharmacology, Vol. 4. Pergamon Press, New York. Bernays, E. A. & S. J. Simpson. 1982.
Control of food intake. Adv. Ins. Physiol. 16: 59-118. Bierne, B. 1962. Trends in applied
biological control of insects. Ann. Rev.
Ent. 7: 387-400. Bonnot, G.
1986. Les particularités de la nutrition des insectes parasites. p. 227-40. In: C. L. Leger (ed.), La
Nutrition de Crustacés et des Insectes. Compte rendu du Colloque du CNERNA,
Paris, France. Bonnot, G. 1986. Multivariate analysis of
nutritional experiments with parasitoids. Colloques de l"INRA. (in press). Bonnot, G.,
B. Delobel & S. Grenier. 1984. Elevage, croissance et développement de Phryxe caudata (Diptera, Tachinidae) sur son hote de substitution
Galleria mellonella (Lepidoptera, Pyralidae)
et sur milieu artificiel. Bull. Soc. Linéenne, Lyon 53: 313-20. Bouletreau, M. 1968. Premiers résultats de
l'élevage des larves d'un Hyménoptere chalcidien (Pteromalus puparum
sur hémolymphe de lépidoptere. Entomophaga
13: 217-22 Bouletreau,
M. 1971. Croissance larvaire et utilisation de l'hote chez Pteromalus puparum (Hym.: Chalc.): influence de la densité de
population. Ann. Zool. Ecol. Anim. 3: 305-18. Bouletreau, M. 1972. Développement et
croissance larvaires en conditions semi-artificielles et artificielles chez
un Hyménoptere entomophage: Pteromalus
puparum L. (Chalc.).
Entomophaga 17: 265-73. Bracken, G. K. 1965. Effects of dietary
components on fecundity of the parasitoid Exeristes
comstockii (Cress.)
(Hymenoptera: Ichneumonidae). Canad. Ent. 97: 1037-41. Bracken, G. K. 1966. Role of ten dietary
vitamins on fecundity of the parasitoid Exeristes
comstockii (Cresson)
(Hymenoptera: Ichneumonidae). Canad. Ent. 98: 918-22. Bracken, G. K. 1969. Effects of dietary
amino acids, salts and protein starvation on fecundity of the parasitoid Exeristes comstockii (Hymenoptera: Ichneumonidae). Canad. Ent. 101:
92-6. Bronskill, J. & H. L. House. 1957.
Notes on rearing a pupal endoparasite, Pimpla
turionellae (L.) (Hymenoptera:
Ichneumonidae), on unnatural food. Canad. Ent. 89: 483. Burks, M. L. & W. C. Nettles, Jr. 1978.
Eucelatoria sp.: effects of
cuticular extracts from Heliothis
virescens and other factors
on oviposition. Environ. Ent. 7: 897-900. Butler, G. D. & P. L. Ritchie. 1971.
WheastR and the abundance and fecundity of Chrysopa carnea.
J. Econ. Ent. 64: 933-34. Cai, C., X. Zhang
& J. Zhao. 1983. Studies on the artificial diet for rearing larvae of Chrysopa sinica Tjeder. Nat. Enemies of Insects 5(2): 82-85. Calow, P. 1977. Conversion efficiencies in
heterotrophic organisms. Biol. Rev. 52: 385-409. Cameron, R.
A. D. & M. Redfern. 1974. A simple study in ecological energetics using a gall-fly
and its insect parasites. J. Biol. Educ. 8: 75-82. Campadelli,
G. & M. L. Dindo. 1987. Recenti progressi nello studio delle diete
artificiali per l'allevamento di insetti entomologi parassiti. Boll. Ent.
Bologna 42: 101-18. Carlson, R.
E. & H. C. Chiang. 1973. Reduction of Ostrinia
nubilalis population by
predatory insects attracted by sucrose sprays. Entomophaga 18: 205-11. Charnov, E.
L. 1982. The Theory of Sex Allocation. Princeton Univ. Press,
Princeton. New Jersey. Charnov, E. L., R. L. Hartogh Los-den, W.
T. Jones & J. van den Assem. 1981. Sex ratio evolution in a variable
environment. Nature 289: 27-33. Chlodny, J. 1968. Evaluation of some
parameters of the individual energy budget of the larvae of Pteromalus puparum (L.) (Pteromalidae)
and Pimpla instigator (Fabr.) (Ichneumonidae).
Ekol. Polska (Ser. A) 16: 505-12. Chumakova, B. M. 1962. Significance of
individual food components for the vital activity of mature predatory and
parasitic insects. Vop. Ekol. Kievsk. 8: 133-34. Clausen, C. P. 1939. The effect of host size upon the sex
ratio of hymenopterous parasites and its relation to methods of rearing and
colonization. J. New York Ent. Soc. 47: 1-9. Clausen, C. P. 1940. Entomophagous Insects. McGraw-Hill,
New York. Clausen, C. P., H. A.
Jaynes & T. R. Gardner. 1933. Further investigations of the parasites of Popillia japonica in the far east. USDA Tech. Bull. 366: 68. Cohen, A. C. 1981. An artificial diet for Geocoris punctipes. Southwest Ent. 6: 109-13. Cohen, A. C. 1983. Improved method of
encapsulating artificial diet for rearing predators of harmful insects. J. Econ. Ent. 76: 957-59. Cohen, A. C. 1984. Food consumption, food utilization,
and metabolic rates of Geocoris
punctipes (Het.: Lygaeidae)
fed Heliothis virescens (Lep.: Noctuidae)
eggs. Entomophaga 29: 361-67. Cohen, A. C.
1985. Simple method for rearing the insect predator Geocoris punctipes (Heteroptera: Lygaeidae) on a meat diet. J. Econ. Ent. 78: 1173-75. Cohen, A. C. 1989. Ingestion efficiency and protein
consumption by a heteropteran predator. Ann. Ent. Soc. Amer. 82: 495-99. Cohen, A. C.
& N. M. Urias. 1986. Meat-based artificial diets for Geocoris punctipes
(Say). Southwest Ent. 11: 171-76. Cooperative Research Group of Hubei
Province, China. 1979. Studies on the artificial host egg of the
endoparasitoid wasp Trichogramma.
Acta Ent. Sin. 22: 301-09. Cooperative Research Group of Hubei
Province, China. 1985. Study on artificial host egg-shell for Trichogramma. J. Wuhan Univ. 4: 1-10. Coppel, H. C., H. L. House & M. G. Maw.
1959. Studies on dipterous parasites of the spruce budworm, Choristoneura fumiferana (Clem.)
(Lepidoptera: Tortricidae). VII. Agria
affinis (Fall.) (Diptera:
Sarcophagidae). Canad. J. Zool. 37: 817-30. Cross, E. A., A. E. S. Mostafa, T. R.
Bauman & I. J. Lancaster. 1978. Some aspects of energy transfer between
the organ-pipe mud dauber Trypoxylon
politum and its araneid
spider prey. Environ. Ent. 7: 647-752. Dadd, R. H.
1973. Insect nutrition: current developments and metabolic
implications. Ann. Rev. Ent. 18: 381-420. Dadd, R. H. 1977. Qualitative requirements
and utilization of nutrients: insects. p. 305-46. In: M. Rechcigl (ed.), Handbook Series in Nutrition and
Food, Vol. 1, Sec. D: Nutritional Requirements. CRC Press, Cleveland. Dadd, R. H. 1985. Nutrition: organisms. p. 313-90. In: G. A. Kerkut & L. I. Gilbert (eds.), Comprehensive
Insect Physiology, Biochemistry and Pharmacology, Vol. 4. Pergamon Press, New
York. Dean, G. J. & C. Satasook. 1983.
Response of Chrysopa carnea (Stephens) (Neuroptera: Chrysopidae)
to some potential attractants. Bull. Ent. Res. 73: 619-24. DeBach, P. 1943. The importance of host
feeding by adult parasites in the reduction of host populations. J. Econ. Ent. 36: 647-58. DeBach, P. 1954. Relative efficacy of the
red scale parasites Aphytis chrysomphali Mercet and Aphytis "A" on citrus
trees in southern California. Bull. Lab. Zool. Gen. Agric. (Filippo
Silvestri), Portici 33: 135-51. Dindo, M. L. 1987. Effetti indotti da
parassitoidi lmenotteri nei loro ospiti. Boll. Ent. Bologna 42: 1-46. Ding, D.-C., H.-G Qiu & C.-B. Hwang.
1980a. In vitro rearing of an
egg-parasitoid Tetrastichus schoenobii (Hymenoptera:
Tetrastichidae). Contr. Shanghai Inst. Ent. p. 55-8. Ding, D.-C., T.-P.
Zhang & Y.-K. Zhong. 1980b. Studies on Tetrastichus
schoenobii Ferriere
(Hymenoptera: Tetrastichidae): oviposition into artificial media. Contr.
Shanghai Inst. Ent. p. 59-61. Doutt, R. L. 1959. The biology of parasitic
Hymenoptera. Ann. Rev. Ent. 4: 161-82. Doutt, R. L. 1964. Biological
characteristics of entomophagous adults. p. 145-67. In: P. DeBach (ed.), Biological Control of Insect Pests
and Weeds. Chapman & Hall, London. Dowell, R. 1978. Ovary structure and
reproductive biologies of larval parasitoids of the alfalfa weevil
(Coleoptera: Curculionidae). Canad. Ent. 110: 507-12. Duelli, P. 1980. Adaptive dispersal and
appetitive flight in the green lacewing, Chrysopa
carnea. Ecol. Ent. 5: 213-20. Duelli, P.
1987. The influence of food on the oviposition site selection in a
predatory and a honeydew-feeding lacewing species (Planipennia: Chrysopidae).
Neurop. Int. 4: 205-10. Dunbar, D. M. & O.
G. Bacon. 1972. Feeding, development and reproduction of Geocoris punctipes (Heteroptera: Lygaeidae) on eight diets. Ann. Ent. Soc. Amer. 65: 892-95. Edgar, W. D. 1971. Aspects of the ecology
and energetics of the egg sac parasites of the wolf spider Pardosa lugubris (Walckenaer). Oecologia 7: 155-63. Ewert, M. A. & H. C. Chiang. 1966.
Dispersal of three species of coccinellids in corn fields. Canad. Ent. 98:
999-1003. Fisher, R. C. 1968. Conversion studies in
parasitic Hymenoptera. Proc. 13th Intern. Cong. Entomology 1: 376-77. Fisher, R. C. 1971. Aspects of the
physiology of endoparasitic Hymenoptera. Biol. Rev. 46: 243-78. Fisher, R. C. 1981. Efficiency of parasites
in assimilating host tissues into their own. Parasitology 82: 33-4. Fisher, R. C. 1986. George Salt and the
development of experimental insect parasitology. J. Ins. Physiol. 32: 249-53. Flint, H. M., S. S.
Salter & S. Walters. 1979. Caryophyllene: an attractant for the green lacewing.
Environ. Ent. 8: 1123-25. Friend, W. G. & R. H. Dadd. 1982.
Insect nutrition - a comparative perspective, p. 205-47. In: H. H. Draper (ed.), Advances in Nutritional Research,
Vol 4. Plenum Press, New York. Gao, Y. G., K. J. Dai, L. C. Shong et al.
1982. Studies on the artificial host egg for Trichogramma. p. 181. In: Les Trichogrammes
Antibes (France), 20-23 Avril 1982. Les Coloques de l'INRA, No. 9. Greany, P. 1980. Growth and development of
an insect parasitoid in vitro. Amer. Zool. 20: 946. Greany, P. 1981. Culture of hymenopteran
endoparasites in vitro. In Vitro 17: 230. Greany, P. 1986. In vitro
culture of hymenopterous larval endoparasitoids. J. Ins. Physiol. 32: 409-19. Greany, P. & K. S. Hagen. 1981. Prey
selection. p. 121-35. In: E.
A. Nordlund, R. L. Jones & W. J. Lewis (eds.), Semiochemicals: Their Role
in Pest Control. John Wiley & Sons, New York. Greany, P., S. B. Vinson & W. J. Lewis.
1984. Insect parasitoids: finding new opportunities for biological control.
Biosci. 34: 690-96. Greathead, D. J. 1986. Parasitoids in
classical biological control. p. 289-318. In:
J. K. Waage & D. J. Greathead (eds.), Insect Parasitoids. 13th Symp. of
the Royal Ent. Soc. of London. Academic Press, London. Greenblatt, J. A. & W. J. Lewis. 1983.
Chemical environmental manipulation for pest insect control. Environ. Manag.
7: 35-41. Greenblatt, J. A., P. Barbosa & M. E.
Montgomery. 1982. Host's diet effects on nitrogen utilization efficiency for
two parasitoid species: Brachymeria
intermedia and Coccygomimus turionellae. Physiol. Ent. 7: 263-67. Grenier, S. 1979. Developpement embryonnaire in
vitro, en milieu artificiel
defini de deux parasitoides ovolarvipares, Phryxe caudata
et Lixophaga diatraeae (Diptera, Tachinidae).
Ent. Exp. Appl. 26: 13-23. Grenier, S., G. Bonnot
& B. Delobel. 1974. Définition et mise au point de milieux artificiels
pour lélevage in vitro de Phryxe caudata
Rond. (Diptera, Tachinidae). I. Survie du parasitoide sur milieux dont la
composition est basée sur celle de l'hémolymphe de l'hote. Ann. Zool. Ecol. Anim. 6: 511-20. Grenier, S., G. Bonnot
& B. Delobel. 1975. Définition et mise au point de milieux artificiels
pour l'élevage in vitro de Phryxe caudata
Rond. (Diptera, Tachinidae). II. Croissance et mues larvaires du parasitoide
en milieux définis. Ann. Zool. Ecol. Anim. 7: 13-25. Grenier, S., G. Bonnot
& B. Delobel. 1986. Physiological considerations of importance to the
success of in vitro culture: an overview. J. Ins. Physiol. 32: 403-08. Grenier, S., G.
Bonnot, B. Delobel & P. Laviolette. 1978. Développement en milieu
artificiel du parasitoide Lixophaga
diatraeae (Towns.) (Diptera, Tachinidae). Obtention de l'imago a partir de l'ouef. C. R. Acad. Sci. Paris, Ser. D 287: 535-38. Gross, H. R. 1987. Conservation and
enhancement of entomophagous insects - a perspective. J. Ent. Sci. 22:
97-105. Guan, X., Z. Wu, T. Wu
& H. Feng. 1978. Studies on rearing Trichogramma
dendrolimi in vitro. Acta. Ent. Sin. 21: 221-26. Hagen, K. S. 1962. Biology and ecology of
predaceous Coccinellidae. Ann. Rev. Ent. 7: 289-326. Hagen, K. S. 1964. Nutrition of entomophagous insects
and their hosts. p. 356-80. In:
P. DeBach (ed.), Biological Control of Insect Pests and Weeds. Chapman &
Hall, London. Hagen, K. S. 1986a. Ecosystem analysis: plant cultivars
(HPR), entomophagous species and food supplements. p. 151-97. In: D. J. Boethel & R. D.
Eikenbary (eds.), Interactions of Plant Resistance and Parasitoids and
Predators of Insects. Halsted Press: John Wiley & Sons, New York. Hagen, K. S. 1986b. Dietary requirements for mass rearing
of natural enemies for use in biological control. p. 35-85. In: Y. Hussein & A. G.
Ibrahim (eds.), Biological Control in the Tropics: Proc. of the First
Regional Symp. on Biological Control. Penerbit Universiti Pertanian,
Malaysia. Hagen, K. S. & G. W. Bishop. 1979. Use
of supplemental foods and behavioral chemicals to increase the effectiveness
of natural enemies. p. 49-60. In:
D. W. Davis, S. C. Hoyt, J. A. McMurtry & M. T. Ali Niazee (eds.),
Biological Control and Insect Pest Management. Univ. Calif. Agric. Sci. Publ. 4096. Hagen, K. S. & R.
Hale. 1974. Increasing natural enemies through use of supplementary
feeding and non-target prey. p. 170-81. In:
F. G. Maxwell & F. A. Harris (eds.), Proc. Summer Institute on Biological
Control of Plant Insects and Diseases. Univ. Press, Mississippi. Hagen, K. S. & R. L. Tassan. 1965. A
method of providing artificial diets to Chrysopa
larvae. J. Econ. Ent. 58: 999-1000. Hagen, K. S. & R. L. Tassan. 1972.
Exploring nutritional roles of extracellular symbiontes on the reproduction
of honeydew feeding adult chrysopids and tephritids. p. 323-51. In: J. G. Rodriguez (ed.),
Insect and Mite Nutrition. Nort-Holland, Amsterdam. Hagen, K. S., R. H. Dadd & J. Reese.
1984. The food of insects. p. 79-112. In:
C. B. Huffaker & R. L. Rabb (eds.), Ecological Entomology. John Wiley
& Sons, New York. Hagen, K. S., E. F.
Sawall, Jr. & R. L. Tassan. 1971. The use of food sprays to increase
effectiveness of entomophagous insects. Proc. Tall Timbers Conf. on Ecol.
Animal Control by Habitat Management. 2: 59-81. Hagen, K. S., P. Greany, E. F. Sawall, Jr.
& R. L. Tassan. 1976. Tryptophan in artificial honeydews as a source of
an attractant for adult Chrysopa
carnea. Environ. Ent. 5: 458-68. Hagley, E. A.
C. & C. M. Simpson. 1981. Effect of food sprays on numbers of predators in an
apple orchard. Canad. Ent. 113: 75-77. Hassan, S. A. & K. S. Hagen. 1978. A
new artificial diet for rearing Chrysopa
carnea larvae (Neuroptera,
Chrysopidae). Z. ang. Ent. 86: 315-20. Hawkes, O. A. M. 1920. Observations on the
life-history, biology, and genetics of the lady-bird beetle, Adalia bipunctata (Mulsant). Proc. Zool. Soc. London. p. 475-90. Hawlitzky, W. & A. M. Mainguet. 1976. Analyse quantitative des lipides, des substances azotées et du
glycogene chez la larve d'un insecte parasite ovo-larvaire Phanerotoma flavitestacea (Hymenoptera:
Braconidae). Ent. Exp. Appl. 20: 43-55. Hemptinne, J. L. & A. Desprets. 1986.
Pollen as a spring food for Adalia
bipunctata. p. 29-35. In: I. Hodele (ed.), Ecology of
Apidophaga. Academia, Prague. Hodek, I.
1973. Biology of Coccinellidae. W. Junk, Academia, Prague. Hoffman, J. D. &
C. M. Ignoffo. 1974. Growth of Pteromalus
puparum in a semisynthetic
medium. Ann. Ent. Soc. Amer. 67: 524-25. Hoffman, J. D., C. M. Ignoffo & W. A.
Dickerson. 1975. In vitro rearing of the
endoparasitic wasp, Trichogramma
pretiosum. Ann. Ent. Soc. Amer. 68: 335-36. Hoffman, J. D., C. M. Ignoffo & S. H.
Long. 1973. In vitro cultivation of an
endoparasitic wasp, Pteromalus
puparum. Ann. Ent. Soc. Amer. 66: 633-34. House, H. L. 1954. Nutritional studies with
Pseudosarcophaga affinis (Fall.), a dipterous
parasite of the spruce budworm, Choristoneura
fumiferana (Clem.). 1. A
chemically defined medium and aseptic-culture technique. Canad. J. Zool. 32:
331-41. House, H. L. 1966a. The role of nutritional
principles in biological control. Canad. Ent. 98: 1121-34. House, H. L. 1966b. Effects of varying the
ratio between the amino acids and other nutrients in conjunction with a salt
mixture on the fly Agria affinis (Fall.). J. Ins.
Physiol. 12: 299-310. House, H. L. 1966c. Effects of vitamins E
and A on growth and development, and the necessity of vitamin E for reproduction
in the parasitoid Agria affinis (Fallen) (Diptera:
Sarcophagidae). J. Ins. Physiol. 12: 409-17. House, H. L. 1976. The role of nutritional
factors in food selection and preference as related to larval nutrition of an
insect, Pseudosarcophaga affinis (Diptera:
Sarcophagidae), on synthetic diets. Canad. Ent. 99: 1310-21. House, H. L. 1969. Effects of different
proportions of nutrients on insects. Ent. Exp. Appl. 12: 651-69. House, H. L. 1970. Choice of food by larvae
of the fly, Agria affinis, related to dietary
proportions of nutrients. J. Ins. Physiol. 16: 2041-50. House, H. L. 1972. Inversion in the order
of food superiority between temperatures affected by nutrient balance in the
fly larva Agria housei (Diptera:
Sarcophagidae). Canad. Ent. 104: 1559-64. House, H. L. 1974. Nutrition. p. 1-62. In: M. Rockstein (ed.), The
Physiology of Insecta, Vol. 5. Academic Press, New York. House, H. L. 1977. Nutrition of natural
enemies. p. 151-82. In: R.
L. Ridgway & S. B. Vinson (eds.), Biological Control by Augmentation of
Natural Enemies. Plenum Press, New York. House, H. L. 1978. An artificial host:
encapsulated synthetic medium for in
vitro oviposition and
rearing the endoparasitoid Itoplectis
conquisitor (Hymenoptera:
Ichneumonidae). Canad. Ent. 110: 331-33. House, H. L. & J. S. Barlow. 1960.
Effects of oleic and other fatty acids on the growth rate of Agria affinis (Fall.) (Diptera:
Sarcophagidae). J. Nutr. 72: 409-14. House, H. L. & J. S. Barlow. 1961.
Effects of different diets of a host, Agria
affinis (Fall.) (Diptera:
Sarcophagidae), on the development of a parasitoid, Aphaereta pallipes
(Say) (Hymenoptera: Braconidae). Canad. Ent. 93: 1041-44. House, H. L. & M. G. Traer. 1948. An
artificial food for rearing Pseudosarcophaga
affinis (Fall.), a parasite
of the spruce budworm Choristoneura
fumiferana (Clem.). 79th
Ann. Rep. Ent. Soc. Ontario. p. 1-4. Howell, J. & R. C. Fisher. 1977. Food
conversion efficiency of a parasitic wasp, Nemeritis canescens.
Ecol. Ent. 2: 143-51. Hoy, M. A.
1976. Genetic improvement of insects: fact or fantasy. Environ. Ent. 5: 833-39. Hoy, M. A.
1979. The potential for genetic improvement of predators for pest
management programs. p. 106-15. In:
M. A. Hoy & J. J. McKelvey (eds.), Genetics in Relation to Insect
Management. Working Papers, The Rockefeller Found., New York. Hoy, M. A.
1986. Use of genetic improvement in biological control. Agric.
Ecosystems Environ. 15: 109-19. Irie, K., Z.-N Xie, W. C. Nettles, Jr., R.
K. Morrison, A. C. Chen, G. M. Holman & S. B. Vinson. 1987. The partial
purification of a Trichogramma
pretiosum pupation factor
from hemolymph of Manduca sexta. Ins. Biochem. 17: 269-75. Jervis, M. A. & A.
C. Kidd. 1986. Host-feeding strategies in hymenopteran parasitoids. Biol. Rev. 61: 395-434. Jones, R. L.
1981. Chemistry of semiochemicals involved in parasitoid-host and
predator-prey relationships. p. 239-50. In:
D. A. Nordlund, R. L. Jones & W. J. Lewis (eds.), Semiochemicals: Their
Role in Pest Control. John Wiley & Sons, New York. Jowyk, E. A. & Z. Smilowitz. 1978. A
comparison of growth and developmental rates of the parasite Hyposoter exigua reared from two instars of its host, Trichoplusia ni. Ann. Ent. Soc. Amer. 71: 467-72. Kariluoto, K. T. 1978.
Optimum levels of sorbic acid and methyl-p-hydroxy-benzoate in
an artificial diet for Adalia
bipunctata (Coleoptera,
Coccinellidae) larvae. Ann. Ent. Fenn. 44: 94-97. Kariluoto, K. T. 1980. Survival and
fecundity of Adalia bipunctata (Coleoptera, Coccinellidae)
and some other predatory insect species on an artificial diet and a natural
prey. Ann. Ent. Fenn. 46: 101-06. Kariluoto, K. T., E. Junnikkala & M.
Markkula. 1976. Attempts at rearing Adalia
bipunctata L. (Col.,
Coccinellidae) on different artificial diets. Ann. Ent. Fenn. 42: 91-7. Deilin, D. 1944. Respiratory systems and
respiratory adaptations in larvae and pupae of Diptera. Parasitology 36:
1-66. Kidd, N. A. C. & M. A. Jervis. 1989.
The effects of host-feeding behaviour on the dynamics of parasitoid-host
interactions, and the implications for biological control. Res. Popul. Ecol. (in press). Lawrence, P. O. 1982. Biosteres longicaudatus:
Developmental dependence on host (Anastrepha
suspensa) physiology. Exp.
Parasitol. 53: 396-405. Lawrence, P. O. 1986a. Host-parasite
hormonal interactions: an overview. J. Ins. Physiol. 32: 295-98. Lawrence, P. O. 1986b. The role of
20-hydroxyecdysone in the moulting of an endoparasitoid, Biosteres longicaudatus.
J. Ins. Physiol. 32: 329-37. Lawton, J. H.
1971. Ecological energetics studies on larvae of the damselfly Pyrrhosoma nymphula (Sulzer) (Odonata: Zygoptera). J. Anim. Ecol. 40: 385-423. 1969 Legner, E. F. 1969.
Adult emergence interval and reproduction in parasitic Hymenoptera
influenced by host size and density.
Ann. Entomol. Soc. Amer.
62(1): 220-226. 1979 Legner, E. F. 1979.
The relationship between host destruction and parasite reproductive potential
in Muscidifurax raptor, M. zaraptor, and Spalangia endius [Chalcidoidea: Pteromalidae]. Entomophaga 24(2): 145-152. 1967 Legner, E. F. & D.
Gerling. 1967. Host-feeding and oviposition on Musca domestica by Spalangia cameroni, Nasonia vitripennis, and Muscidifurax raptor
(Hymenoptera: Pteromalidae) influences their longevity and fecundity. Ann. Entomol. Soc. Amer. 60(3): 678-691. 1977 Legner, E. F. &
S. N. Thompson. 1977. Effects of the parental host on host
selection, reproductive potential, survival and fecundity of the egg-larval
parasitoid Chelonus sp. near curvimaculatus, reared on Pectinophora gossypiella and Phthorimaea
operculella. Entomophaga
22(1): 75-84. Leius, K. 1960. Attractiveness of different foods and
flowers to the adults of some hymenopterous parasites. Canad. Ent. 92:
369-76. Leius, K. 1961a. Influence of various foods
on fecundity and longevity of adults of Scambus
buolianae (Htg.) (Hymenoptera:
Ichneumonidae). Canad. Ent. 93: 1079-84. Leius, K. 1961b. Influence of food on
fecundity and longevity of adults of Itoplectis
conquisitor (Say)
(Hymenoptera: Ichneumonidae). Canad. Ent. 93: 771-80. Leius, K. 1962. Effects of body fluids of
various host larvae on fecundity of females of Scambus buolianae
(Htg.) (Hymenoptera: Ichneumonidae). Canad. Ent. 94: 1078-82. Leius, K. 1963. Effects of pollen on
fecundity and longevity of adult Scambus
buolianae (Htg.)
(Hymenoptera: Ichneumonidae). Canad. Ent. 95: 202-07. Leius, K. 1967a. Influence of wild flowers
on parasitism of tent caterpillar and codling moth. Canad. Ent. 99: 444-46. Leius, K. 1967b. Food sources and
preferences of adults of a parasite, Scambus
buolianae (Hym.: Ich.), and
their consequences. Canad. Ent. 99: 865-71. Lewis, W. J., M. Beevers, D. A. Nordlund,
H. R. Cross & K. S. Hagen. 1979. Kairomones and their use for management
of entomophagous insects. IX. Investigations of various kairomone-treatment
patterns for Trichogramma
spp. J. Chem. Ecol. 5: 673-80. Lewis, W. J., D. A. Nordlund, R. C.
Gueldner, P. E. A. Teel & J. H. Tumlinson. 1982. Kairomones and their use
for management of entomophagous insects. XII. Kairomone activity for Trichogramma spp. of abdominal
tips, feces, and a synthetic pheromone blend of Heliothis zea
(Boddie) moths. J. Chem. Ecol. 8: 1323-32. Li, L. Y. 1982. Trichogramma sp. and their utilization in People's
Republic of China. Les Trichogrammes, Antibes (France,
20-23 Avril 1982). Les Colloques de l'INRA, No. 9 Lindquist, R. K. & E. L. Sorenson. 170.
Interrelationships among aphids, tarnished plant bugs and alfalfas. J. Econ. Ent. 63: 192-95. Lingren, P. D. &
M. J. Lukefahr. 1977. Effects of nectariless cotton on cages populations of Camplletis sonorensis. Environ. Ent. 6: 586-88. Liu, W. & Z. Wu. 1982. Recent results
in rearing Trichogramma in vitro with the artificial media devoid of insect
additives. Acta Ent. Sin. 25: 160-63. Liu, W., Z.
N. Xie, G. F. Xiao, Z. F. Zhou, D. H. Ou Yang et al. 1979. Rearing of Trichogramma dendrolimi
in artificial diets. Acta Phyto. Sin. 6: 17-24. Luck, R. F.,
H. Podoler & R. Kfir. 1982. Host selection and egg allocation behavior
by Aphytis melinus and A. lingnanensis: comparison of two facultatively gregarious
parasitoids. Ecol. Ent. 7: 397-408. Mackauer, M. 1986. Growth and developmental interactions
in some aphids and their hymenopterous parasites. J. Ins. Physiol. 32: 275-80. Matsuka, M. & I.
Okada. 1975. Nutritional studies on an aphidophagous coccinellid, Harmonia axyridis. (1) Examination of artificial diets for the
larval growth with special reference to drone honeybee powder. Bull. Fac.
Agric. Tamagawa Univ. 15: 1-9. Matsuka, M. & S. Takahashi. 1977.
Nutritional studies of an aphidophagous coccinellid Harmonia axyridis.
2. Significance of minerals for larval growth. Apl. Ent. Zool. 12: 325-29. Matsuka, M., M. Watanabe & K. Niijima.
1982. Longevity and oviposition of vedalia beetles on artificial diets.
Environ. Ent. 11: 816-19. Matsuka, M., D. Shimotori, T. Senzaki &
I. Okada. 1972. Rearing some coccinellids on pulverized drone honeybee brood.
Bull. Fac. Agric. Tamagawa Univ. 12: 28-38. Mellini, E. 1975a. Possibilita di allevamento di insetti entomofagi parasiti su diete
artificiali. Boll. Ent. Bologna 32: 257-90. Mellini, E.
1975b. Studi sui ditteri larvevoridi XXV. Sul determinismo ormonale delle
influenze esercitate dagli ospiti sui loro parassiti. Boll. Ent. Bologna 31:
165-203. Mellini, E.
1978. Moderni problemi di entomoparassitologia. Estratto dagli Atti XI Congresso
Nazionale Italiano di Entomologia Portici-Sorrento. p. 263-92. Mellini, E.
1983. L'ipotesi della dominazione ormonale, esercitata dagli ospiti sui
parasitoidi, alla luce delle recenti scopterte nella endocrinologia degli
insetti. Boll. Ent. Bologna 38: 135-66. Mellini, E.
& G. Beccari. 1984. Relazioni tra dimensioni degli ospiti e percentuali
di parassitizzazione nella coppia ospite-parassita Galleria mellonella
L.-Gonia cinerascens Rond. Boll. Ent.
Bologna 38: 71-88. Mellini, E.
& G. Campadelli. 1982. Potenziale megetico del parasitoide Gonia cinerascens Rond. misurato sull'ospite di sostituzione Galleria mellonella L. Mem. Soc. Ent. Ital., Genova 60: 239-52. Mellini, E.,
M. Malagoli & L. Ruggeri. 1980. Substrati artificiali per
l'ovideposizione dell'entomoparassita Gonia
cinerescens Rond. (Diptera:
Larvaevoridae) in cattivita. Boll. Ent. Bologna 35: 127-56. Morrison, R. K., V. S. House & R. L.
Ridgway. 1975. Improved rearing unit for larvae of a common lacewing. J. Econ. Ent. 68: 821-22. Morrison, R. K., W. C. Nettles, D. Ball
& S. B. Vinson. 1983. Successful oviposition by Trichogramma pretiosum
through a synthetic membrane. Southwest Ent. 8: 248-51. Nadarajan, L. & S. Jayaraj. 1975.
Influence of various hosts on the development and reproduction of pupal
parasite, Tetrastichus israeli M. & K.
(Hymenoptera). Curr. Sci. 44: 458-60. Nenon, J. P. 1972a. Culture in vitro des embryons d'un Hyménoptere endoparasité
polyembryonnaire: Ageniaspis
fuscicollis (=Encyrtus fuscicollis). Role des hormones de synthese. C. R. Acad.
Sci. Paris 274D: 3299-3302. Nenon, J. P. 1972b. Culture in vitro des larves d'un Hyménoptere endoparasite
polyembryonnaire: Ageniaspis
fasicollis. Role des
hormones de synthese. C. R. Acad. Sci. Paris 274D: 3409-12. Nepomnyashchaya, A., E. Maintcher & I.
Yazlovetskij. 1979. New approach to elaboration of artificial nutritive diets
for mass rearing of entomophagous insects. Optimization of nutritive diets by
the simplex lattice method. Biochem. Physiol. Ins., Kishinev,
Shtiintsa. p. 29-35. Nettles, W. C., Jr. 1986a. Asparagine: a
host chemical essential for the growth and development of Eucelatoria bryani, a tachinid parasitoid
of Heliothis spp. Comp.
Biochem. Physiol. 85A: 697-701. Nettles, W. C., Jr. 1986b. Effects of soy
flour, bovine serum albumin and three amino acid mixtures on growth and
development of Eucelatoria bryani (Diptera: Tachinidae)
rearing on artificial diets. Environ. Ent. 15: 1111-15. Nettles, W. C., Jr. 1987a. Amino acid
requirements for growth and development of the tachinid, Eucelatoria bryani.
Comp. Biochem. Physiol. 86A: 349-54. Nettles, W. C., Jr. 1987b. Eucelatoria bryani (Diptera: Tachinidae):
effect on fecundity of feeding on hosts. Environ. Ent. 16: 437-40. Nettles, W. C., Jr., C. M. Wilson & S.
W. Ziser. 1980. A diet and methods for the in vitro
rearing of the tachinid, Eucelatoria
sp. Ann. Ent. Soc. Amer. 73: 180-84. Nettles, W. C., Jr., R. K. Morrison, Z.-N.
Xie, D. Ball, C. A. Shenkir & S. B. Vinson. 1982. Synergistic action of
potassium chloride and magnesium sulfate on parasitoid wasp oviposition.
Science 218: 164-66. Nettles, W. C., Jr., R. K. Morrison, Z.-N.
Xie, D. Ball, C. A. Shenkir & S. B. Vinson. 1983. Effect of cation,
anions and salt concentrations on oviposition by Trichogramma pretiosum
in wax eggs. Ent. Exp. Appl. 33: 283-89. Nettles, W. C., Jr., R. K. Morrison, Z.-N.
Xie, D. Ball, C. A. Shenkir & S. B. Vinson. 1984. Ovipositional stimulant
for Trichogramma spp. U. S.
Patent 4,484,539. Nettles, W. C., Jr., R. K. Morrison, Z.-N.
Xie, D. Ball, C. A. Shenkir & S. B. Vinson. 1985. Effect of artificial
diet media, glucose, protein hydrolyzates, and other factors on oviposition
in wax eggs by Trichogramma pretiosum. Ent. Exp. Appl. 38:
121-29. Niijima, K., M. Matsuka & I. Okada.
1986. Artificial diets for an aphidophagous coccinellid, Harmonia axyridis,
and its nutrition (Minreview). p. 37-50. In:
I. Hodek (ed.), Ecology of Aphidophaga. Academia Publ. House, Czechoslovak
Acad. Sci., Prague. Niijima, K., R. Nishimura & M. Matsuka.
1977. Nutritional studies of an aphidophagous coccinellid, Harmonia axyridis. III. Rearing of larvae using a chemically
defined diet and fraction of drone honeybee powder. Bull. Fac. Agric.,
Tamagawa Univ. 17: 45-51. Nordlund, D. A., R. B. Chalfant & W. J.
Lewis. 1985. Response of Trichogramma
pretiosum females to
extracts of two plants attacked by Heliothis
zea. Agric. Ecosyst.
Environ. 12: 127-33. Nordlund, D. A., R. L. Jones & W. J.
Lewis (eds.). 1981. Semiochemicals: Their Role in Pest Control. John Wiley
& Sons, New York. Nozato, K. 1969. The effect of host size on
the sex ratio of Itoplectis cristatae Momoi (Hymenoptera:
Ichneumonidae), a pupal parasite of the Japanese pine shoot moth, Petrova (= Evetria) cristata
(Walsingham) (Lepidoptera: Olethreutidae). Kontyu 37: 134-46. Okada, I. & M.
Matsuka. 1973. Artificial rearing of Harmonia axyridis
on pulverized drone honeybee brood. Environ. Ent.
2: 301-02. Okada, I., H.
Hoshiba & T. Maruoka. 1971a. An artificial rearing of a coccinellid beetle, Harmonia axyridis Pallas, on drone honeybee brood. Bull. Fac.
Agric., Tamagawa Univ. 11: 91-7. Okada, I., M. Matsuka & H. Hoshiba.
1971b. Utilization of drone larvae and pupae as semiartificial diet for
aphidophagous insects. Apiacta 6: 119-20. Okada, I., H.
Hoshiba & T. Maehava. 1972. An artificial rearing of a coccinellid beetle, Harmonia axyridis Pallas, on pulverized drone honeybee brood. Bull.
Fac. Agric. Tamagawa Univ. 12: 39-47. Petrusewicz, K. 1967. Concepts in studies
on the secondary productivity of terrestrial ecosystems. p. 17-49. In: K. Petrusewicz (ed.),
Secondary Productivity of Terrestrial Ecosystems, Vol. 1. Panstwowe
Wydawnictwo Naukowe, Warsaw. Pimentel, D. 1966. Wasp parasite (Nasonia vitripennis) survival on its fly host (Musca domestica) reared on various foods. Ann. Ent. Soc. Amer. 59: 1031-38. Prell, H. 1915. Zur
Biologie der Tachinen Parasitiogena
segregata Rdi und Panzeria rudis Fall. Z. angew. Ent. 2: 57-148. Principi, M. M. & M. Canard. 1984.
Feeding habits, p. 76-92. In:
M. Canard, Y Séméria & T. R. New (eds.), Biology of Chrysopidae. W. Junk,
The Hague, Netherlands. Rajendram, G.
F. 1978a. Oviposition behavior of Trichogramma
californicum on artificial
substrates. Ann. Ent. Soc. Amer. 71: 92-4. Rajendram, G.
F. 1978b. Some factors affecting oviposition of Trichogramma californicum
(Hymenoptera: Trichogrammatidae) in artificial media. Canad. Ent. 110:
345-52. Rajendram, G. F. & K. S. Hagen. 1974. Trichogramma oviposition into
artificial substrates. Environ. Ent. 3: 399-401. Rojas-Rousse, D. & R. Kalmes. 1978. The
development of male Diadromus
pulchellus (Hymenoptera:
Ichneumonidae) in the pupae of Acrolepiopsis
assectella (Lepidoptera:
Plutellidae): comparison of assimilation and energy losses under two
temperature regimes. Environ. Ent. 7: 469-81. Rotundo, G.,
R. Cavalloro & E. Tremblay. 1988. In
vitro rearing of Lysiphlebus fabarum (Hym.: Braconidae).
Entomophaga 33: 261-67. Rousch, R. T. 1979. Genetic improvement of
parasites. p. 97-105. In: M.
A. Hoy & J. J. McKelvey (eds.), Genetics in Relation to Insect
Management. Working Papers, Rockefeller Found., New York. Roussett, A. 1984. Reproductive physiology
and fecundity. p. 116-29. In:
M. Canard, Y Séméria & T. R. New (eds.), Biology of Chrysopidae. W. Junk,
The Hague, Netherlands. Salt, G. 1934. Experimental studies in
insect parasitism. II. Superparasitism. Proc. Roy. Ent. Soc. London 114B:
455-76. Salt, G. 1940. Experimental studies in
insect parasitism. VII. The effects of different hosts on the parasite Trichogramma evanescens Westw. (Hym.:
Chalcidoidea). Proc. Roy. Ent. Soc. London 15A: 81-124. Salt, G. 1941. The effects of hosts upon
their insect parasites. Biol. Rev. 16: 239-64. Salt, G. 1952. Trimorphism in the
ichneumonid Gelis corruptor. Q. J. Microsc. Sci.
93: 453-75. Sandlan, K. P. 1979a. Sex ratio in Coccygomimus turionellae Linnaeus
(Hymenoptera: Ichneumonidae) and its ecological implications. Ecol. Ent. 4:
365-78. Sandlan, K. P. 1979b. Host-feeding and its effects
on the physiology and behavior of the ichneumonid parasitoid, Coccygomimus turionellae. Physiol. Ent. 4:
383-92. Sandlan, K. P. 1982. Host suitability and
its effects on parasitoid biology in Coccygomimus
turionellae (Hymenoptera:
Ichneumonidae). Ann. Ent. Soc. Amer. 75: 217-21. Schmidt, G. T. 1974. Host acceptance behavior of Compoletis sonorensis toward Heliothis
zea. Ann. Ent. Soc. Amer. 67: 835-44. Shahjahn, M. 1968. Effect of diet on the
longevity and fecundity of the adults of the tachinid parasite Trichopoda pennipes pilipes.
J. Econ. Ent. 61: 1102-03. Shannon, R. C. 1923. Rearing dipterous
larvae on nutrient agar. Proc. Ent. Soc. Wash. 25: 103-04. Simmonds, F. J. 1944. The propagation of
insect parasites on unnatural hosts. Bull. Ent. Res. 35: 219-26. Slansky, F., Jr. 1978. Utilization of
energy and nitrogen by larvae of the imported cabbageworm, Pieris rapae, as affected by parasitism by Apanteles glomeratus.
Environ. Ent. 7: 179-85. Slansky, F., Jr. 1982. Toward a nutritional
ecology of insects. p. 253-59. In:
J. H. Viser & A. K. Minks (eds.), Proceedings of the 5th Intern. Symp. on
Insect-plant Relationships. Pudoc, Wageningen, Netherlands. Slansky, F., Jr. 1986. Nutritional ecology
of endoparasitic insects and their hosts: an overview. J. Ins. Physiol. 32: 255-61. Slansky, F., Jr. & J. M. Scriber. 1985.
Food consumption and utilization. p. 87-163. In: G. A. Kerkut & L. I. Gilbert (eds.), Comprehensive
Insect Physiology, Biochemistry and Pharmacology. Pergamon Press, New York. Sluss, R. 1968. Behavioral and anatomical
responses of the convergent lady beetle to parasitism by Perilitus coccinellae
(Shrank) (Hymenoptera: Braconidae). J. Invert. Path. 10: 9-27. Smirnoff, W. A. 1958. An artificial diet
for rearing coccinellid beetles. Canad. Ent. 90: 563-65. Smith, B. C. 1961. Results of rearing some
coccinellid (Coleoptera: Coccinellidae) larvae on various pollens. Proc. Ent.
Soc. Ontario 91: 270-71. Smith, B. C.
1965. Growth and development of coccinellid larvae on dry foods
(Coleoptera: Coccinellidae). Canad. Ent. 97: 760-68. Smith, B. C. 1966. Effect of food on some
aphidophagous Coccinellidae. p. 75-81. In:
I. Hodek (ed.), Ecology of Aphidophagous Insects. Academia Publ. House,
Czechoslovak Acad. Sci., Prague. Smith, J. M. 1957. Effects of the food
plant of California red scale, Aonidiella
aurantii (Mask.) on
reproduction of its hymenopterous parasites. Canad. Ent. 89: 219-30. Smith, R. W. 1958. Parasites of nymphal and
adult grasshoppers (Orthoptera: Acrididae) in Western Canada. Canad. J. Zool.
36: 217-62. Stanley, R. G. & H. F. Linskens. 1974.
Pollen - Biology, Biochemistry, Management. Springer-Verlag, Berlin. Strand, M. R. 1986. The physiological
interactions of parasitoids with their hosts and their influence on
reproductive strategies. p. 97-136. In:
J. Waage & D. J. Greathead (eds.), Insect Parasitoids. Academic Press,
London. Strand, M. R. & S. B. Vinson. 1985. In vitro culture of Trichogramma
pretiosum on an artificial
medium. Ent. Exp. Appl. 39: 203-09. Strand, M. R., S. M. Meola & S. B.
Vinson. 1986. Correlating pathological symptoms in Heliothis virescens
eggs with development of the parasitoid Telenomus
heliothidis. J. Ins. Physiol. 32: 389-402. Strand, M. R., S. M. Meola, W. C. Nettles
& Z. N. Xie. 1988. In vitro culture of the egg
parasitoid Telenomus heliothidis: the role of
teratocytes and medium consumption in development. Ent. Expt. Appl. 46: 71-8. Syme, P. D. 1975. The effects of flowers on
the longevity and fecundity of two native parasites of the European pine
shoot moth in Ontario. Environ. Ent. 4: 337-46. Syme, P. D. 1977. Observations on the
longevity and fecundity of Orgilus
obscurator (Hymenoptera:
Braconidae). and the effects of certain foods on longevity. Canad. Ent. 109:
995-1000. Szumkowski, W. 1952. Observations on
Coccinellidae. II. Experimental rearing of Coleomegilla on a non-insect diet. Trans. IX Intern. Cong. Ent. 1: 781-85. Szumkowski, W. 1961a. Dietas sin insectos vivos para la cria de Coleomegilla maculata
Deg. (Coccinellidae, Coleoptera). Agronomía Trop. Venezolana 10: 149-54. Szumkowski,
W. 1961b. Aparición de un coccinelido predator nuevo para Venezuela.
Agronomía Trop. Venezolana 11: 33-37. Tanaka, M.
& Y. Maeta. 1965. Rearing of some predacious coccinellid beetles by the
artificial diets. Bull. Hort. Res. Sta. Japan D(3): 17-35. Temerak, S. A. 1976. Studies on certain
mortality factors affecting distribution and abundance of sugarcane borers in
upper Egypt. Ph.D. dissertation, College of Agriculture, Assiut University,
Egypt cf. Temerak (1983). Thompson, S. N. 1975. Defined meridic and
holidic diets and aseptic feeding procedures for artificially rearing the
ectoparasitoid Exeristes roborator (Fabricius). Ann. Ent. Soc. Amer. 68: 220-26. Thompson, S. N. 1976a. Effects of dietary
amino acid level and nutritional balance on larval survival and development
of the parasite Exeristes roborator. Ann. Ent. Soc. Amer. 69: 835-38. Thompson, S. N. 1976b. The amino acid
requirements for larval development of the hymenopterous parasitoid Exeristes roborator Fabricius (Hymenoptera: Ichneumonidae). Comp.
Biochem. Physiol. 72B: 211-13. Thompson, S. N. 1977. Lipid nutrition
during larval development of the parasitic wasp, Exeristes. J. Ins. Physiol. 23: 579-83. Thompson, S. N. 1980. Artificial culture
techniques for rearing larvae of the chalcidoid parasite, Brachymeria intermedia. Ent. Expt. Appl.
27: 133-43. Thompson, S. N. 1981a. The nutrition of
parasitic Hymenoptera. Proc. 9th Intern. Cong. Plant Prot. 1: 93-6. Thompson, S. N. 1981b. Essential amino acid
requirements of four species of parasitic Hymenoptera. Comp. Biochem.
Physiol. 69A: 173-74. Thompson, S. N. 1981c. Brachymeria lasus
and Pachycrepoideus vindemiae: sterol requirements
during larval growth of two hymenopterous insect parasites reared in vitro on chemically defined diets. Expt. Parasitol. 51:
220-35. Thompson, S. N. 1981d. Brachymeria lasus:
culture in vitro of a chalcid insect
parasite. Expt. Parasitol. 32: 414-18. Thompson, S. N. 1982. Exeristes roborator:
quantitative determination of in
vitro larval growth rates in
synthetic media with different glucose concentrations. Expt. Parasitol. 54:
229-34. Thompson, S. N. 1983a. Larval growth of the
insect parasite Brachymeria lasus reared in vitro. J. Parasitol. 69: 425-27. Thompson, S. N. 1983b. Brachymeria lasus:
effects of nutrient level on In
vitro larval growth of a
chalcid insect parasite. Expt. Parasitol. 55: 312-19. Thompson, S. N. 1983c. Metabolic and
physiological effects of metazoan endoparasites on their host species. Comp.
Biochem. Physiol. 74B: 183-211. Thompson, S. N. 1985. Metabolic integration
during the host associations of multicellular animal endoparasites. Comp.
Biochem. Physiol. 81B: 21-42. Thompson, S. N. 1986a. Nutrition and in vitro culture of insect parasitoids. Ann. Rev. Ent. 31:
197-219. Thompson, S. N. 1986b. The metabolism of
insect parasites (parasitoids): an overview. J. Ins. Physiol. 32: 421-23. Thompson, S. N. 1989. Nutritional
considerations in propagation of entomophagous species. p. 389-404. In: R. Baker & P. Dunn (eds.),
New Directions in Biological Control. UCLA Symposia on Molecular &
Cellular Biology. Alan R. Liss Publ., New York. Thompson, S. N. &
K. S. Hagen. 1999. Nutrition of entomophagous insects. In: Bellows, T. S., Jr. & T. W. Fisher, (eds) 1999. Handbook
of Biological Control: Principles and Applications. Academic Press, San
Diego, CA. 1046 p. Thompson, S.
N., L. Bednar & H. Nadel. 1983. Artificial culture of the insect
parasite Pachycrepoideus vindemiae. Ent. Expt. Appl. 33:
121-22. Thurston, R. & P. M. Fox. 1972.
Inhibition by nicotine of emergence of Apanteles
congregatus from its host,
the tobacco hornworm. Ann. Ent. Soc. Amer. 65: 547-50. Tucker, J. E. & D.
E. Leonard. 1977. The role of kairomones in host recognition and host
acceptance behavior of the parasite, Brachymeria
intermedia. Environ. Ent. 6: 527-31. Vanderzant,
E. S. 1969. An artificial diet for larvae and adults of Chrysopa carnea, an insect predator of crop pests. J. Econ. Ent. 62: 256-57. Vanderzant,
E. S. 1973. Improvements in the rearing diet for Chrysopa carnea
and the amino acid requirements for growth. J. Econ. Ent. 66: 336-38. Van Emden, H. F. & K. S. Hagen. 1976.
Olfactory reactions of the green lacewing, Chrysopa carnea
to tryptophan and certain breakdown products. Environ. Ent. 5: 469-73. Varley, G. C.
1961. Conversion rates in hyperparasitic insects. Proc. Roy. Ent.
Soc. London 26C: 11. Veerman, A., M. E.
Slagt, M. F. J. Alderlieste & R. L. Veenendaal. 1985. Photoperiodic induction of
diapause in an insect is vitamin A dependent. Experimentia 41: 1194-95. Vinson, S. B.
1975. Biochemical coevolution between parasitoids and their hosts.
p. 14-48. In: P. Price
(ed.), Evolutionary Strategies of Parasitic Insects and Mites. Plenum Press,
New York. Vinson, S. B.
1976. Host selection by insect parasitoids. Ann. Rev. Ent. 21:
109-33. Vinson, S. B. 1984. Parasitoid-host
relationship. p. 205-33. In:
W. J. Bell & R. T. Cardé (eds.), Chemical Ecology of Insects. Chapman
& Hall, London. Vinson, S. B. & G. F. Iwantsch. 1980.
Host suitability for insect parasitoids. Ann. Rev. Ent. 25: 397-419. Waage, J. K., K. P. Carl, N. J. Mills &
D. J. Greathead. 1984. Rearing entomophagous insects. p. 45-66. In: P. Singh & R. F. Moore
(eds.), Handbook of Insect Rearing. Elsevier, Amsterdam, Netherlands. Welch, H. E. 1968. Relationships between assimilation
efficiencies and growth efficiencies for aquatic consumers. Ecology 49:
755-59. White, T. D. & G. L. Jubb. 1980.
Potential of food sprays for augmenting green lacewing populations in vineyards.
p. 35-42. Melsheimer Ent. Ser. #29. Wiegert, R. G. &
C. E. Petersen. 1983. Energy transfer in insects. Ann. Rev. Ent. 28: 455-86. Wilbert, H. 1977. Der
Honigtau als Reiz und Energiequelle für Entomophage Insekten. Apidologie 8: 393-400. Wu, Z.-X. & J. Quin. 1982a. Culturing Trichogramma dendrolimi in vitro
with media devoid of insect materials. Proc. Chinese Acad. Sci. - United
States Nat. Acad. of Sci. Joint Symp. on the Biological Control of Insects.
p. 431-35. Wu, Z.-X.
& J. Quin. 1982b. Ovipositional response of Trichogramma
dendrolimi to the chemical
contents of artificial eggs. Acta Ent.
Sinica 25: 363-72. Wu, Z., Z. Zhang, T.
Li & D. Liu. 1980. Artificial media devoid of insect additives for rearing larvae
of the endoparasitoid wasp Trichogramma.
Acta Ent. Sinica 23: 232. Wu, Z., J. Quin, T.-X.
Li, Z.-P. Chang & T.-M. Liu. 1982. Culturing Trichogramma in
vitro with artificial media
devoid of insect materials. Acta Ent. Sinica 25: 128-35. Wylie, H. G. 1965. Effects of superparasitism on Nasonia vitripennis (Walk.) (Hymenoptera: Pteromalidae). Canad. Ent. 97: 326-31. Wylie, H. G. 1967. Some effects of host size on Nasonia vitripennis and Muscidifurax
raptor (Hymenoptera:
Pteromalidae). Canad. Ent. 99: 742-48. Xie, Z.-N., W. C. Nettles, Jr., R. K. Morrison,
K. Irie & S. B. Vinson. 1986a. Effect of ovipositional stimulants and
diets on the growth and development of Trichogramma
pretiosum in vitro. Ent. Expt. Appl. 42: 119-24. Xie, Z.-N., W. C. Nettles, Jr., R. K.
Morrison, K. Irie & S. B. Vinson. 1986b. Three methods for the in vitro culture of Trichogramma
pretiosum Riley. J. Ent.
Sci. 21: 133-38. Yamamura, N.
& E. Yano. 1988. A simple model of host-parasitoid interaction with
host-feeding. Res. Popul. Ecol. 30: 353-69. Yazgan, S. 1972. A chemically defined
synthetic diet and larval nutritional requirements of the endoparasitoid Itoplectis conquisitor (Hymenoptera). J. Ins. Physiol. 18: 2123-41. Yazgan, S. & H. L. House. 1970. An
hymenopterous insect, the parasitoid Itoplectis
conquisitor, reared axenically
on a chemically defined diet. Canad. Ent.
102: 1304-06. Yazlovetskij,
I. G. & A. M. Nepomnyashchaya. 1981. Essai d'elevage massif de Chrysopa carnea sur milieux artificiels micro-encapsules et de son application
dans la lutte contre les pucerons en serre. Lutte Biologique et Intégrée
Coutre les Pucerons. Colloque Franco-Soviétique, Rennes, 26-27 Sept. 1979.
Paris, France. Institue National de la Recherche Agronomique. p. 51-8. Ye, Z., Y. Han, D.
Wang, S. Liang & S. Li. 1979. Studies on the artificial diets of larvae
and adults of Chrysopa sinica Tjeder. Acta Phyto. Sinica 6(2): 11-16. Zhou, W. & X.
Zhang. 1983. Preliminary study on the use of encapsulated diet for
rearing Chrysopa sinica. Acta Phyto. Sinica 10(3): 161-65. Ziser, S. & W. C.
Nettles. 1979. The rate of oxygen consumption by Eucelatoria sp. in relation to larval development and
temperature. Ann. Ent. Soc. Amer. 72: 540-43. Zoebelein, G. 1956.
Der Honigtau als Nahrung der Insekten. Z. angew. Ent. Teil I 38: 369-416;
Teil II 39: 129-67. Zohdy, N. M. 1976. On the effect of the food of Myzus persicae Sulz. on the hymenopterous parasite Aphelinus asychis Walker. Oecologia, Berlin 26: 1985-191. |