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Host-Plant Selection by the Mexican Bean Beetle, Epilachna Varivestis1

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Host-Plant Selection by the Mexican Bean Beetle, Epilachna Varivestis1 Host-Plant Selection by the Mexican Bean Beetle, Epilachna varivestis1 MITHRA G. AUGUSTINE,2' * F. W. FISK,2 R. H. DAVIDSON,2 J. B. LAPIDUS,3 AND R. W. CLEARY8 ABSTRACT The Mexican bean beetle, Epilachna varivestis Mul- fraction from foliage of host-plants yielded positive ol- sant, feeds selectively on certain species of Phaseolus to factory responses from Mexican bean beetle larvae, and the exclusion of the remaining representatives of this only indirect evidence of similar responses from the and other genera of beans. In a study of the chemical adults. These findings indicate the presence of a short- factors underlying this selective feeding, the phagostimu- range attractant in the volatile constituents of host- lant fraction in the seeds of P. vulgaris was isolated and plants. The influence of both attractants and arrestants identified as sucrose. Bioassay of 13 sugars and 15 in host-plant selection by insect pests is discussed, and amino acids showed that only sucrose and, to a lesser the significance of quantitative differences in essential degree, its hexose components induced feeding by the nutrients in host-plants is stressed. Phytogeographical beetle. The seeds as well as leaves of the nonresistant factors relating to the 10 common species of Phaseolus species of Phaseolus were found to have a higher con- also are discussed, emphasizing that in host-plant selec- centration of sucrose, glucose, and fructose, thereby re- tion their role, if any, is only subordinate to stimulant vealing the role these sugars play as arrestants in host- chemicals in plants. plant selection by the beetle. Bioassay of the volatile Host preference in many phytophagous species of selection or rejection by phytophagous insects; and insects is well known, and the factors, in the host and (iii) physiological changes in the plant and their im- of the pest, that make possible this preference are pact on the food-finding behavioral pattern of the currently receiving an extensive probe from several pest. Despite these 3 different approaches—nutrients, investigators. Factors in the plant concern its physi- secondary chemicals, and physiological changes within cal and chemical makeup, while those of the pest con- the plant—it is obvious that the emphasis in all these cern its perceptive powers. Recognition and subse- investigations of host-plant selection in phytophagous quent selection of the food-plant by the insect call insects remains essentially one of finding specific into play a harmonious coordination of all these fac- chemotactic stimuli resting in or emanating from the tors. Several of the easily discernible physical at- plants. tributes of plants, size, shape, and color, are too The Mexican bean beetle, Epilachna varivestis variable and possess no uniqueness that could possibly Mulsant (Coleoptera: Coccinellidae), exhibits host- aid their recognition by the pest. Vision, phototaxis, specificity. Of the 5 genera of beans, Vigna, Phaseo- geotaxis, and hygrotaxis play a part in directing in- lus, Dolichos, Glycine, Vicia, the beetle shows marked sects to the proper environment for feeding and ovi- preference to only certain species of Phaseolus. All position; but the ultimate close-range forces, operat- others (barring a negligible number of interspecific ing in the food-plant selection by the host-specific crosses showing "slight to intermediate" susceptibility, pests, are presumed to be largely chemical—regulated but never preferred) are categorized as resistant to for the most part by smell and taste. the pest (Wolfenbarger 1961). The insect may, under Ever since the first accounts by Verschaffelt experimental and occasionally even field conditions, (1910) and Mclndoo (1919) invited attention to the live on certain plants totally unrelated to legumes possible relationships between the chemical constit- (Auclair 1959). Elmore (1949) provides a list of uents of plants and host-specific phytophagous in- plants the pest did not accept under experimental sects, considerable progress has been made in un- conditions, and a few it fed upon but without sub- ravelling some of the causal factors underlying this sequent reproduction. The investigation reported in phenomenal host-specificity of certain insects. Deth- this paper was made to identify the plant constituents ier (1953), Kennedy (1953), Lipke and Fraenkel eliciting those chemotactic responses in the beetle (1956), Beck (1956), and Thorsteinson (1960) pro- that aid its unerring recognition and discreet accept- vide some of the very helpful reports in this field of ance of its preferred hosts. study. These reports give an adequate insight into the intricacies of the problem and list an extensive MATERIALS bibliography to the work of other investigators since Throughout the period of this study, a large cul- Verschaffelt. A survey of this literature reveals ture of Mexican bean beetles was maintained in the emphasis along 3 basic lines of investigation: (i) con- greenhouse. The beetles were reared mostly on red sideration of the nutritional and other factors resident kidney and lima varieties of bean plants. Separate in plants and leading to the insect's "passive" selec- cages for adults, larvae, pupae, and newly emerged tion of host-plants; (ii) search for nonnutritious, at- adults facilitated the use of samples of uniform age tractive or repellent factors responsible for "active" in all bioassays. Where feeding activity was the cri- terion sought, young adults (not older than 8-10 1 This investigation was supported by Research Grant EF days) and early second- and third-instar larvae were 00185-03 from the Public Health Service. Accepted for publica- tion March 18, 1963. found to yield more consistent results than other 2 Department of Zoology and Entomology, Ohio State Uni- stages of the beetle. versity, Columbus. 3 College of Pharmacy, Ohio State University, Columbus. Seeds for analysis were secured from commercial 4 Fulbright, Smith-Mundt. and Danforth Grantee from the Madras Christian College, Tambaram, India. as well as various other sources (Augustine 1962). 127 128 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA [Vol. 57 Plantings for foliage analysis were at the rate of 40 Extraction of seeds.—Storage of seeds is easier to 45 seeds per each 9-inch pot. Agrico water-soluble than that of foliage, and seeds also assure a ready plant food 17-17-17 was mixed to half strength, and supply of uniform material for chemical analysis. All each pot received 16 oz of the solution. The water varieties of P. vulgaris are known to be susceptible to supply in the saucers holding the pots was replenished severe attack by the beetle and, therefore, seeds of every other day, and the foliage for analysis was one of its common varieties (red kidney bean) were picked always from plants about the same stage of chosen for thorough fractionation to isolate the prin- growth—when the trifoliate leaves were just unfold- cipal feeding stimulant for the Mexican bean beetle. ing. A mixture of ground seeds of P. vulgaris and a small quantity of sea sand was placed in a sintered METHODS AND RESULTS glass-bottom vessel and extracted in a soxhlet appara- Most of the damage done by the Mexican bean tus : first with ether absolute and then with 95% ethyl beetle, E. varivestis, to the susceptible bean plants alcohol. Each sample was extracted for 20 to 22 is defoliation, although both larvae and adults are hours. The extract fractions, after the removal of the known to feed on pods of some varieties after foliage solvent in vacuo, were dried in a vacuum desiccator is destroyed. Adults or larvae, feeding from below, and bioassayed. The fractionation scheme and the re- eat ragged areas in the lower surface of the leaf, sults of the bioassay of the fractions are found in often also cutting through the upper surface. The Fig. 2. By techniques listed in Table 1, the active damaged leaf is a peculiar network which charac- ingredient in the ethanolic fraction was identified as terizes the attack of this pest (Howard 1941). Be- sucrose. cause of the pest's preferential feeding on leaf juices, Bioassay.—The seed extracts were dissolved in it was postulated that the insect may attack filter known volumes of the solvent, and Whatman No. 1 paper treated with any attractant fraction from its (4.25 cm) filter paper discs were treated with meas- host-plant tissues. Initial investigations (Byers 1961) using the leaf juices obtained by mincing leaves of Seeds of Phaseolus vu Iqo ris Phaseolus vulgaris L. in a Waring blendor proved (ground and sieved) the procedure satisfactory. A damaged leaf and the pattern of feeding on filter paper with leaf juices are Extract with anhydrous shown in Fig. 1. ethyl ether Ether distillate Ether insoluble residue Evaporate Extract with 957. ethanol Deep yellow, Alcohol soluble Alcohol insoluble greasy precipitate fraction residue Evaporate Add MgO Reddish-brown, Yellow-green syrupy precipitate solution Decant Distillate Residue I Treat with activated charcoal and filter Filtrate Residue Reduce volume under pressure and add anhydrous ether Yellow-white, flocculent Filtrate precipitate (-) (•••+) Dissolve in absolute ethanol and repeat process of purification I White crystalline substance (identified as sucrose) (++•+) FIG. 2.—Fractionation scheme of seeds of Phaseolus vulgaris, and bioassay of the fractions. ( —), inactive as feeding stimulant; (-)—1--|—|-), very active feeding stim- FIG. 1.—Pattern of feeding on leaves and filter paper. ulant. 1964] Arc.usxiNE ET AL. : HOST-PLANT SELECTION BY MEXICAN BEAN BEETLE 129 Table 1.—Identification of the phagostimulant crystal- from 12 to 18 hours. The sugar solutions were ap- line material from the seeds of Phascolns vulgaris. plied in 0.2 ml portions to each disc. Only 3 of the 13 sugars tested stimulated feeding by the beetle. Results (LaPidus et al., in press). The degree of the in- Test Unknown crystals Sucrose sect's preference for these three sugars may be in- dicated as: Melting point 185° C.
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