Large Cabbage White Butterfly Research Articles

Antifeedant effects of crofton weed, Ageratina adenophora extracts and their phytochemical compounds against three cabbage insect pest species

AbstractGiven that synthetic pesticides used in agricultural fields have negative side effects, safer alternatives such as botanical insecticides are required for insect pest management. Here, solvent extracts (hexane, ethyl acetate, chloroform, acetone, and methanol) of crofton weed, Ageratina adenophora (Spreng.) King & H.Rob. (Asteraceae), were studied for antifeedant activity against third instars of large cabbage white butterfly, Pieris brassicae L. (Pieridae), diamondback moth, Plutella xylostella L. (Plutellidae), and cabbage cluster caterpillar, Crocidolomia pavonana Fabricius (Crambidae, all Lepidoptera). Among the solvent extracts tested, A. adenophora hexane extract at 5% concentration exhibited maximum antifeedant activity against P. xylostella (92.1%), C. pavonana (87.4%), and P. brassicae (81.7%). The effective concentration (EC50) values were 1.4, 2.1, and 2.8%, respectively. The hexane extract was characterised using gas chromatography–mass spectrometry (GC–MS) analysis and it was determined that its main compounds were 9,11‐octadecadiynoic acid, 8‐oxo‐, methyl ester (27.5%), 2(3H)‐naphthalenone, 4,4a,5,6,7,8‐hexahydro‐4a, (R)‐7,7‐trimethyl‐ (20.4%), germacrene D (6.1%), 2(1H)‐naphthalenone (5.8%), 5,6‐decadien‐3‐yne, 5,7‐diethyl (5%), and α‐bisabolene (2.1%). The results suggest that hexane extract of A. adenophora could be used for the development of botanical insecticides.

Response of Parasitic Wasp Cotesia glomerata L. (Hymenoptera: Braconidae) to Cabbage Plants of Two Varieties: Olfactory Spectra of Males and Females.

The parasitoid Cotesia glomerata L. (Hymenoptera: Braconidae) oviposits in larvae of the large cabbage white butterfly (Pieris brassicae L.). Many parasitoids are attracted by volatile organic compounds (VOCs) emitted by the plant that the host insects feed on. The objectives of the study were to identify the VOCs in leaves of two varieties of cabbage (white cabbage, Brassica oleracea var. capitata f. alba and cauliflower, B. oleracea var. botrytis) damaged by P. brassicae caterpillars which elicit antennal responses of C. glomerata, and characterize the olfactory spectra of females and males. Leaf extracts were analyzed by using gas chromatography equipped with an electroantennographic detector (GC-EAD) and GC-mass spectrometry. In total, 32 olfactory-active compounds for C. glomerata in cauliflower and 24 in white cabbage were revealed. The females perceived more compounds than males. Hexan-1-ol, (E)-3-hexen-1-ol, (E)-2-octenal, benzylcianide, tetradecanal, and two unidentified compounds elicited EAG responses in females but not in males. Females were more sensitive to (E)-3-hexenal, (Z)-3-hexenal, and pentadecenal, whereas males showed higher sensitivity to (Z)-3-hexenyl butanoate, heptanal, (Δ)-2-pentenal, (E)-2-hexenol, and octanal. The olfactory spectrum of C. glomerata was expanded from 18 to 41 VOCs emitted by different varieties of cabbage damaged by P. brassicae caterpillars. Eight EAG-active VOCs were common for all cabbage varieties. In tritrophic interactions, benzylcyanide can serve as an important signal for C. glomerata females indicating damage of cabbage caused by P. brassicae caterpillars. The data are useful for development of a push-pull strategy for P. brassicae control, based on parasitoid behavior regulation by VOCs.

Studies on Pest Complex of Broad Leaf Mustard, Brassica juncea var. rugosa Roxb. Tsen and Lee in the Valley of Manipur

A experiment was conducted at Entomological Research Farm, Department of Entomology, College of Agriculture, Central Agricultural University, Imphal. Total eleven insect pest species were found to infest the crop viz. Large cabbage white butterfly, Pieris brassicae; Asian cabbage white butterfly, Pieris canidia; mustard aphid, Lipaphis erysymi and striped flea beetle, Phyllotreta striolata were observed frequently while, Cabbage aphid, Brevicoryne brassicae; green peach aphid, Myzus persicae; crucifer flea beetle, Phyllotreta cruciferae; mustard sawfly, Athalia lugens proxima; cabbage semilooper, Trichoplusia ni; fungus beetle, Monolepta signata and tobacco grasshopper, Atractomorpha crenulata were observed rarely.

The phytotoxic air-pollutant O3 enhances the emission of herbivore-induced volatile organic compounds (VOCs) and affects the susceptibility of black mustard plants to pest attack

Stress-induced changes to plant biochemistry and physiology can influence plant nutritional quality and subsequent interactions with herbivorous pests. However, the effects of stress combinations are unpredictable and differ to the effects of individual stressors. Here we studied the effects of exposure to the phytotoxic air-pollutant ozone (O3), feeding by larvae of the large cabbage white butterfly (Pieris brassicae), and a combination of the two stresses, on the emission of volatile organic compounds (VOCs) by black mustard plants (Brassica nigra) under field and laboratory conditions. Field-grown B. nigra plants were also measured for carbon-nitrogen (C–N) content, net photosynthetic activity (Pn), stomatal conductance (gs) and biomass. The effects of O3 on interactions between plants and a herbivorous pest were addressed by monitoring the abundance of wild diamondback moth larvae (Plutella xylostella) and feeding-damage to B. nigra plants in an O3-free air concentration enrichment (O3-FACE) field site. Herbivore-feeding induced the emission of VOCs that were not emitted by undamaged plants, both under field and laboratory conditions. The combination of O3 and herbivore-feeding stresses resulted in enhanced emission rates of several VOCs from field-grown plants. Short-term O3 exposure (of 10 days) and P. brassicae-feeding did not affect C–N content, but chronic O3 exposure (of 34 and 47 days) and P. brassicae-feeding exacerbated suppression of Pn. Ozone exposure also caused visible injury and decreased the plant biomass. Field-grown B. nigra under elevated O3 were infested with fewer P. xylostella larvae and received significantly less feeding damage. Our results suggest that plants growing in a moderately polluted environment may be of reduced quality and less attractive to foraging herbivores.

Oviposition Preference for Young Plants by the Large Cabbage Butterfly (Pieris brassicae) Does not Strongly Correlate with Caterpillar Performance

The effects of temporal variation in the quality of short-lived annual plants on oviposition preference and larval performance of insect herbivores has thus far received little attention. This study examines the effects of plant age on female oviposition preference and offspring performance in the large cabbage white butterfly Pieris brassicae. Adult female butterflies lay variable clusters of eggs on the underside of short-lived annual species in the family Brassicaceae, including the short-lived annuals Brassica nigra and Sinapis arvensis, which are important food plants for P. brassicae in The Netherlands. Here, we compared oviposition preference and larval performance of P. brassicae on three age classes (young, mature, and pre-senescing) of B. nigra and S. arvensis plants. Oviposition preference of P. brassicae declined with plant age in both plant species. Whereas larvae performed similarly on all three age classes in B. nigra, preference and performance were weakly correlated in S. arvensis. Analysis of primary (sugars and amino acids) and secondary (glucosinolates) chemistry in the plant shoots revealed that differences in their quality and quantity were more pronounced with respect to tissue type (leaves vs. flowers) than among different developmental stages of both plant species. Butterflies of P. brassicae may prefer younger and smaller plants for oviposition anticipating that future plant growth and size is optimally synchronized with the final larval instar, which contributes >80% of larval growth before pupation.

Role of Large Cabbage White butterfly male-derived compounds in elicitation of direct and indirect egg-killing defenses in the black mustard.

To successfully exert defenses against herbivores and pathogens plants need to recognize reliable cues produced by their attackers. Up to now, few elicitors associated with herbivorous insects have been identified. We have previously shown that accessory reproductive gland secretions associated with eggs of Cabbage White butterflies (Pieris spp.) induce chemical changes in Brussels sprouts plants recruiting egg-killing parasitoids. Only secretions of mated female butterflies contain minute amounts of male-derived anti-aphrodisiac compounds that elicit this indirect plant defense. Here, we used the black mustard (Brassica nigra) to investigate how eggs of the Large Cabbage White butterfly (Pieris brassicae) induce, either an egg-killing direct [i.e., hypersensitive response (HR)-like necrosis] or indirect defense (i.e., oviposition-induced plant volatiles attracting Trichogramma egg parasitoids). Plants induced by P. brassicae egg-associated secretions expressed both traits and previous mating enhanced elicitation. Treatment with the anti-aphrodisiac compound of P. brassicae, benzyl cyanide (BC), induced stronger HR when compared to controls. Expression of the salicylic (SA) pathway- and HR-marker PATHOGENESIS-RELATED GENE1 was induced only in plants showing an HR-like necrosis. Trichogramma wasps were attracted to volatiles induced by secretion of mated P. brassicae females but application of BC did not elicit the parasitoid-attracting volatiles. We conclude that egg-associated secretions of Pieris butterflies contain specific elicitors of the different plant defense traits against eggs in Brassica plants. While in Brussels sprouts plants anti-aphrodisiac compounds in Pieris egg-associated secretions were clearly shown to elicit indirect defense, the wild relative B. nigra, recognizes different herbivore cues that mediate the defensive responses. These results add another level of specificity to the mechanisms by which plants recognize their attackers.

Plant‐mediated effects of butterfly egg deposition on subsequent caterpillar and pupal development, across different species of wild Brassicaceae

1. Herbivory can change plant quality, which may have consequences for interactions between the inducing herbivore and other insect community members.2. Studies investigating the effects of plant quality on herbivore performance often have neglected the egg stage, and instead introduced larvae onto the plant. Recently, we reported that herbivore oviposition by Pieris brassicae (Linnaeus) (Large Cabbage White Butterfly) reduced the plant quality of Brassica nigra L. (black mustard) for subsequent herbivores.3. It remains unclear how persistent and common these plant‐mediated effects of oviposition are. Here, five species of wild Brassicaceae were used (B. nigra L., Brassica oleracea L., Sinapis arvensis L., Moricandia arvensis L., and Moricandia moricandioides Boiss). The response to oviposition by the specialist P. brassicae was determined by following the natural sequence of events: oviposition, egg, larval, and pupal development. All tested plant species are known to interact with P. brassicae in nature. Caterpillar, pupal mass, and development time on plants exposed to butterfly eggs were assessed compared with egg‐free plants.4. It was shown that the plant‐mediated effects of oviposition are not specific for B. nigra but occur in most of the tested plant species except for M. arvensis. However, the strength of the plant‐mediated effect on caterpillar growth depended on plant species. Thus, across different members of the Brassicaceae family, oviposition can influence plant quality and has negative consequences on P. brassicae growth. Further studies are needed to assess to what extent this trait might be phylogenetically conserved.

To be in time: egg deposition enhances plant-mediated detection of young caterpillars by parasitoids.

Animals use information from their environment while foraging for food or prey. When parasitic wasps forage for hosts, they use plant volatiles induced by herbivore activities such as feeding and oviposition. Little information is available on how wasps exploit specific plant volatiles over time, and which compounds indicate changes in host quality. In experiments investigating the role of herbivore-induced plant volatiles in wasp foraging, induction of plant response is usually achieved by placing larvae on clean plants instead of allowing the natural sequence of events: to let eggs deposited by the herbivore develop into larvae. We compared the attraction of the parasitoid Cotesia glomerata to volatiles emitted by black mustard (Brassica nigra) plants induced by eggs and successive larval stages of the Large Cabbage White butterfly (Pieris brassicae) to the attraction of this parasitoid to black mustard plant volatiles induced only by larval feeding in a wind tunnel setup. We show that wasps are attracted to plants infested with eggs just before and shortly after larval hatching. However, wasp preference changed at later time points towards plants induced only by larval feeding. These temporal changes in parasitoid attraction matched with changes in the chemical compositions of the blends of plant volatiles. Previous studies have shown that host quality/suitability decreases with caterpillar age and that P. brassicae oviposition induces plant defences that negatively affect subsequently feeding caterpillars. We investigated parasitoid performance in hosts of different ages. Wasp performance was positively correlated with preference. Moreover, parasitism success decreased with time and host stage. In conclusion, the behaviour of Cotesia glomerata is fine-tuned to exploit volatiles induced by eggs and early host stages that benefit parasitoid fitness.

Seasonal phenology of interactions involving short‐lived annual plants, a multivoltine herbivore and its endoparasitoid wasp

Spatial-temporal realism is often missing in many studies of multitrophic interactions, which are conducted at a single time frame and/or involving interactions between insects with a single species of plant. In this scenario, an underlying assumption is that the host-plant species is ubiquitous throughout the season and that the insects always interact with it. We studied interactions involving three naturally occurring wild species of cruciferous plants, Brassica rapa, Sinapis arvensis and Brassica nigra, that exhibit different seasonal phenologies, and a multivoltine herbivore, the large cabbage white butterfly, Pieris brassicae, and its gregarious endoparasitoid wasp, Cotesia glomerata. The three plants have very short life cycles. In central Europe, B. rapa grows in early spring, S. arvensis in late spring and early summer, and B. nigra in mid to late summer. P. brassicae generally has three generations per year, and C. glomerata at least two. This means that different generations of the insects must find and exploit different plant species that may differ in quality and which may be found some distance from one another. Insects were either reared on each of the three plant species for three successive generations or shifted between generations from B. rapa to S. arvensis to B. nigra. Development time from neonate to pupation and pupal fresh mass were determined in P. brassicae and egg-to-adult development time and body mass in C. glomerata. Overall, herbivores performed marginally better on S. arvensis and B. nigra plants than on B. rapa plants. Parasitoids performance was closely tailored with that of the host. Irrespective as to whether the insects were shifted to a new plant in successive generations or not, development time of P. brassicae and C. glomerata decreased dramatically over time. Our results show that there were some differences in insect development on different plant species and when transferred from one species to another. However, all three plants were of generally high quality in terms of insect performance. We discuss ecological and evolutionary constraints on insects that must search in new habitats for different plant species over successive generations.

Insect Egg Deposition Induces Indirect Defense and Epicuticular Wax Changes in Arabidopsis thaliana

Egg deposition by the Large Cabbage White butterfly Pieris brassicae on Brussels sprouts plants induces indirect defense by changing the leaf surface, which arrests the egg parasitoid Trichogramma brassicae. Previous studies revealed that this indirect defense response is elicited by benzyl cyanide (BC), which is present in the female accessory reproductive gland (ARG) secretion and is released to the leaf during egg deposition. Here, we aimed (1) to elucidate whether P. brassicae eggs induce parasitoid-arresting leaf surface changes in another Brassicacean plant, i.e., Arabidopsis thaliana, and, if so, (2) to chemically characterize the egg-induced leaf surface changes. Egg deposition by P. brassicae on A. thaliana leaves had similar effects to egg deposition on Brussels sprouts with respect to the following: (a) Egg deposition induced leaf surface changes that arrested T. brassicae egg parasitoids. (b) Application of ARG secretion of mated female butterflies or of BC to leaves had the same inductive effects as egg deposition. Based on these results, we conducted GC-MS analysis of leaf surface compounds from egg- or ARG-induced A. thaliana leaves. We found significant quantitative differences in epicuticular waxes compared to control leaves. A discriminant analysis separated surface extracts of egg-laden, ARG-treated, untreated control and Ringer solution-treated control leaves according to their quantitative chemical composition. Quantities of the fatty acid tetratriacontanoic acid (C34) were significantly higher in extracts of leaf surfaces arresting the parasitoids (egg-laden or ARG-treated) than in respective controls. In contrast, the level of tetracosanoic acid (C24) was lower in extracts of egg-laden leaves compared to controls. Our study shows that insect egg deposition on a plant can significantly affect the quantitative leaf epicuticular wax composition. The ecological relevance of this finding is discussed with respect to its impact on the behavior of egg parasitoids.

Digestive enzymes of large cabbage white butterfly, Pieris brassicae L. (Lepidoptera: Pieridae) from developmental and site of activity perspectives

Investigation of digestive enzymes in the alimentary tract of insects is very important for knowledge on physiology of nutrition. The overall results of this study showed the presence of the majority of digestive enzymes including α-amylase, α- and β-glucosidase and galactosidase, lipases and proteases in the midgut of Pieris brassicae. It was found that the fourth and fifth instar larvae had the highest enzymatic activity, but the obvious differences were found in the cases of sites of enzyme activity known as soluble and membrane-bound fractions. The highest enzymatic activities in the soluble fractions were found in cases of α-amylase, lipases and proteinases such as trypsin-like, chymotrypsin-like and elastase. In contrast, glycosidases and exopeptidases (amino and carboxypeptidases) demonstrated the highest enzymatic activity in the membrane-bound fractions. However, no enzymatic activity was observed on cysteine protease as a type of proteinases. By using line regression, parameter β (slope) showed larval dependency on enzymes along with developmental and feeding processes. It was found that α-glucosidase had the highest slope and the membrane-bound fraction of general proteases had the lowest, showing high and low dependencies of larvae to α-glucosidase and exopeptidases, respectively, in addition to second priority on β-glucosidase and α-amylase. Optimal pHs of soluble and membrane-bound α-amylase were found at 9 while all assayed glycosidases had the highest activity at pH 8 for both fractions. Soluble fraction of lipase had the highest enzymatic activity at pH 8 in comparison with membrane-bound fraction at pH 10. The soluble fraction showed the highest proteolytic activity at pH 9, while the highest proteolytic activity was observed at pH 8 in the case of the membrane-bound one.

Root and shoot jasmonic acid induction differently affects the foraging behavior of Cotesia glomerata under semi-field conditions

Plants can accumulate and release defensive chemicals by activating various signaling pathways when they are damaged by herbivores or pathogens. The jasmonic acid pathway is activated after damage by chewing herbivores. Here we used jasmonic acid (JA) as an exogenous elicitor to induce feral cabbage plants. In this study, the effects of root JA (RJA) and shoot JA (SJA) induction on the foraging behavior of Cotesia glomerata, a parasitoid of the large cabbage white butterfly Pieris brassicae, was investigated under semi-field conditions. In all combinations of differently induced plants (RJA, SJA and control plants), the percentages of shoot induced plants that were visited by at least one wasp were significantly higher than those of controls or root induced plants during 3 h of foraging. Consequently, parasitism rates of P. brassicae on shoot-JA induced plants were significantly higher than on plants induced with JA to the roots or control plants in all tests. However, this behavioral preference was not reflected in the allocation of offspring. The clutch sizes of C. glomerata eggs on control, root induced and shoot induced plants were not significantly different from each other in two-choice or three-choice experiments, but did differ with clutch size in the two-choice experiment of uninduced control plants versus SJA. This semi-field study helps to further understand the choice behavior and preferences of parasitoids in natural multitrophic communities in which plants induced with root or shoot herbivores occur together.

The Biosynthesis of Hexahydrofarnesylacetone in the Butterfly Pieris brassicae

Hexahydrofarnesylacetone (6,10,14-trimethylpentadecan-2-one, 1) is a widespread ketone occurring in plants and insects. Several species use this compound or the respective alcohol as part of their pheromone bouquet. Here, we showed by using deuterium labeled phytol (3) and GC-MS experiments that the Large Cabbage White butterfly Pieris brassicae can take up phytol in the larval stage and transforms it into 1 by oxidative degradation.

Chemical espionage on species-specific butterfly anti-aphrodisiacs by hitchhiking Trichogramma wasps

Parasitic wasps employ a wide range of chemical cues to find their hosts. Very recently, we discovered how 2 closely related egg parasitoids, Trichogramma brassicae and Trichogramma evanescens, exploit the anti-aphrodisiac pheromone benzyl cyanide of one of their hosts, the gregarious large cabbage white butterfly Pieris brassicae that deposits a clutch of more than 20 eggs per oviposition bout. The pheromone is transferred by male butterflies to females during mating to enforce female monogamy. On detecting the anti-aphrodisiac, the tiny parasitic wasps ride on a mated female butterfly to a host plant and then parasitize her freshly laid eggs. The present study demonstrates that both wasp species similarly exploit the anti-aphrodisiac mixture of methyl salicylate and indole of another host, the more common solitary small cabbage white butterfly Pieris rapae that deposits only one egg at a time. Interestingly, this behavior is innate in T. brassicae, whereas T. evanescens learns it after one successful ride on a mated female butterfly. Moreover, we show that the wasps only respond to the anti-aphrodisiacs of the 2 cabbage white butterflies when the ubiquitous compounds are part of a complete mated female odor blend. Obviously, parasitic wasps use the sophisticated espionage-and-ride strategy to find eggs of different gregarious and solitary host species. From the wasps’ perspective there seems to be a trade-off between the abundance and egg-laying behavior of the butterflies. Our findings suggest that Pieris butterflies are under strong selective pressure to minimize the use of an anti-aphrodisiac.

Do parasitized caterpillars protect their parasitoids from hyperparasitoids? A test of the ‘usurpation hypothesis’

Caterpillars that are attacked by some species of parasitoid wasps are known to survive for several days after the parasitoid larvae emerge and pupate. It has been argued that the behaviour of the parasitized larva is ‘usurped’ by the parasitoid and that it ‘guards’ the parasitoid cocoons against their own natural enemies such as hyperparasitoids (the ‘usurpation hypothesis'). We tested this hypothesis in the association involving a gregarious endoparasitoid, the wasp Cotesia glomerata; caterpillars of its host, the large cabbage white butterfly Pieris brassicae; and a pupal hyperparasitoid, the wasp Lysibia nana. In laboratory experiments, we presented cocoon broods of C. glomerata to single females of L. nana in arenas for 6 h. We tested several treatments for rates of primary parasitoid survival, including variation in the position of the caterpillar and the presence or absence of an additional silk web spun by parasitized caterpillars. Parasitized P. brassicae larvae survived longer than the period necessary for C. glomerata adults to emerge. Rates of parasitoid survival were, however, unaffected by the presence of a P. brassicae larva on the cocoon brood, although significantly more parasitoids emerged when the silk web was present. Analyses of the foraging behaviour of individual L. nana females in arenas, performed using Observer software, revealed that the wasps showed a greater tendency to leave cocoons when caterpillars and silk were present. The laboratory experiments only partially support the usurpation hypothesis. In nature, usurpation of the host of the primary parasitoid may be a more effective strategy against generalist predators than against more specialized and better-adapted hyperparasitoids.

Associative learning of visual and gustatory cues in the large cabbage white butterfly, Pieris brassicae

AbstractThe landing response of the large cabbage white butterfly Pieris brassicae was studied under controlled optical and gustatory stimulus conditions. Experience-based changes in landing behaviour were examined by offering cardboard circles of two different shades of green, treated with either an oviposition stimulant or a deterrent. We employed two training situations. In one situation the two shades of green, carrying either the stimulant or the deterrent, were offered simultaneously, in the other sequentially. During the 1 hour training periods, butterflies were either landing and drumming spontaneously or they were caught at the end of the period and placed on the artificial leaves until tarsal drumming ensued. Our experiments demonstrated that P. brassicae females can learn to associate visually detected substrate characteristics with contact-chemosensory information available only after landing. Furthermore, a learned preference for a substrate could be turned into a preference for the alternative substrate by exposing the insect to a deterrent on the previously preferred substrate. These results provide indications of aversion learning, thus far undocumented in oviposition behaviour of Lepidoptera. Bringing the butterflies into forced contact with the oviposition stimulant resulted in similar effects on landing preference compared to those of spontaneous landing, but spontaneous landing had a stronger effect on preference for associations involving the deterrent. The simultaneous training regime, which supposedly requires a less important role for short-term memory, was more effective in modifying landing preferences.

The aerodynamics of hovering insect flight. III. Kinematics

The aerodynamics of hovering insect flight. III. Kinematics

New chlorinated juvenile hormone analogues and their biological activity

AbstractIn the search for new juvenile hormone analogues (JHAs), a general approach was chosen based on the substitution of the isoprene unit, that carries the epoxy function in the natural hormone JHIII, by 3,3,3‐trichloropropyl, 2,2‐dichlorocyclopropyl‐ methyl, 3,3‐dichloroallyl, or 3‐chloroprop‐2‐ynyl groups. Accordingly, several terpenoid aromatic JHAs were prepared with, or without, oxygen atoms in the aliphatic chain. Structure‐biological activity relationships of the most active compounds toward the yellow mealworm (Tenebrio molitor) and the large cabbage white butterfly (Pieris brassicae) are given. With a view to preparing synthetically accessible analogues, some alkyl phenyl ethers were also made, which were not branched in the aliphatic chain and were related terminally to the most active compounds. They were only moderately active against insects when tested for morphogenetic activity, but exhibited a good field persistence. The occurrence of a moderate acaricidal activity among these analogues prompted the examination of several 1,4‐disubstituted‐phenyl derivatives. The ovicidal effectiveness of these products against a susceptible strain of the two‐spotted spider mite, Tetranychus urticae, was ten or 100 times higher than that of the well known and highly potent JHAs.

Effects of Lovo Spray Additives on the Performance of Low-Volume Sprays of Bacillus thuringiensis Berliner

Laboratory results showed that Lovo® (amine stearate) spray additives produced a significant improvement in the technical characteristics required for the application of Thuricide® (Bacillus thuringiensis Berliner) in aqueous suspension at ultra low volume (0.5 to 2 Imperial gal./acre). The effects of the additives were three-fold leading to improved suspensibility, reduced evaporation of spray, and improved adhesion of deposits. Cage tests using large cabbage white butterfly (Pieris brassicae) larvae demonstrated that improved rain-fastness of Thuricide by Lovo produced a four-fold increase in dosage efficiency.

Birds attacking Butterflies and Moths

As this subject is again interesting your readers, I would repeat that both my gardener and myself have independently observed robins capture and swallow the large cabbage white butterfly.