ALBERT

Notes: Abstract The nutritional preferences of storage insects are evaluated from the viewpoint of feeding habits, composition and utilization of foodstuffs as well as the involvement of attractants and feeding stimulants. On the basis of their feeding habits, coleopterous species infesting stored produce may be divided into four major groups: (a) Species which in the larval and adult stage consume dried foodstuffs from plant sources (e.g., Sitophilus granarius), (b) Species which in the larval and/or adult stage consume dried foodstuffs of plant and animal origin (e.g., Oryzaephilus surinamensis, Cryptolestes ferrugineus, Tenebroides mauretanicus), (c) Species which feed on material of animal origin in the larval stage and feed on nectar and pollen as adults (e.g., Attagenus pellio), (d) Species feeding on material of animal origin as larvae and adults (e.g.,Dermestes maculatus). Most lepidopterous species (except forHofmannophila pseudospretella) feed only in the larval stage on stored seeds and other plant tissues. Adequacy of stored foodstuffs for insect propagation mainly depends on availability of the proper nutrients and conditions which vary with different groups of insects. Olfactory attraction of storage insects can be induced by a blend of food volatiles such as the aroma of wheat or other cereals, while aggregation and feeding may be stimulated by less volatile food components including salts, sugars and lipids. Several saturated or unsaturated fatty acids with a chain length of 12 to 18 carbon atoms induce aggregation and/or feeding in adults and/or larvae of some curculionid, clerid, dermestid, gelechiid and pyralid species. Certain triglycerides elicit aggregation in adultSitophilus granarius, Tribolium castaneum andTribolium confusum as well as in larvae ofSitotroga cerealella. Mono- and disaccharides are phagostimulatory forTribolium andPloida breeding in food with considerable sugar content, while the cations K and Na stimulate feeding of carpet beetle larvae which usually grow on contaminated wool. Conditioning to strange chemical stimuli is discussed as a possible factor involved in the diversification of the feeding habits of storage insects.

Notes: Abstract Interspecific responses among severalTrogoderma species have been correlated with their pheromone components. The most important component emitted by four of the species is (Z)- or (E)-14-methyl-8-hexadecenal, which is not detectable in extracts of macerated beetles. The response to macerated beetles is probably due to the corresponding alcohol and ester. The recency of common origin of seven species is discussed.

Notes: Abstract Several polyphagous coleopteran and lepidopterous species, presently known as “storage insects”, have presumably evolved from free-living ancestral species, being capable of growth and reproduction on stored, desiccated and often nutritionally deficient foodstuffs. These potentially harmful insect species have probably adapted themselves to the newly acquired storage biotope by means of a well-developed sensory equipment serving food acquisition, aggregation and mate finding. Information by molecules may be communicated among the individuals of an insect species by means of relatively volatile pheromones (Greek, pherō=convey) being emitted by exocrine glands and mainly carried by moving air to the sensilla of responsive individuals, or among the internal organs of an insect by means of relativelynonvolatile hormones (Greek, hormaō=impel), secreted fromendocrine glands and transported by the haemolymph to the receptors of target organs. It was postulated that pheromones were among the first chemical messengers utilized during evolution of animal behaviour, and that the pheromones of primitive protozoans could have been precursors of the hormones of metazoans. Hormones of the neurosecretory cells and corpora allata were found to induce sex pheromone biosynthesis in femaleTenebrio molitor, while dietary intake of a juvenile hormone analogue was shown to significantly enhance the production of aggregation pheromones in the males of certain silvanid and cucujid species. Aggregation pheromones are usually produced by the longlived and feeding males of several coleopteran species (Table 2) which deposit those chemical messengers to the substrate, where they induce the formation of bisexual assemblies supporting feeding, mating and reproduction. Sex pheromones are mostly produced by the short-lived and non-feeding females of several coleopteran and lepidopterous species (Table 2); females of those species usually release their sex pheromones to the air space during calling, and thus attract conspecific males for mating (Fig. 5 a–c). In some dermestid species, pheromone emission differs from the above scheme. Females of the short-lived and non-feedingTrogoderma granarium andT. inclusum release a phromone acting as a sex attractant for conspecific males and—in synergistic combination with tactile stimuli—as an assembling scent for conspecific females (Figs. 1 a, b, 2 and Table 1), females of the short-lived and feedingAntbrenus verbasci, Attagenus megatoma andAtt. elongatulus produce a sex pheromone for conspecific males, while females of the long-lived and feedingAn. scrophulariae emit a sex pheromone which lures conspecific males. Males of the long-lived and non-feeding bruchid speciesAcanthoscelides obtectus release a sex pheromone which attracts conspecific females. Androconial pheromones are discharged during courtship from the alar scales and abdominal tufts found in males of several microlepidopteran species (Phycitidae) includingAnagasta kuebniella, Cadra cautella, Ephestia elutella andPlodia interpunctella (Fig. 6 b–c); those aphrodisiac pheromones are known to enhance the specific responsiveness of the females to their mates. Electrophysiological recordings revealed that aggregation pheromones elicit considerable receptor potentials in the antennal olfactory sensilla of both sexes, whereas sex pheromones induce high receptor potentials in the antennal olfactory sensilla of one sex only. It was assumed that aggregation pheromones may be the evolutionary precursors of sex pheromones. Pheromone-producingexocrine glands are essentially groups of modified epidermals cells which are found in different body regions of male and/or female storage insect species. A simple pheromone gland, consisting of a single layer of adjacent secretory cells beneath the endocuticle of the 5th visible abdominal sternite, occurs in femaleTrogoderma granarium (Fig. 3 a). A more complex design, comprising an intra-abdominal semiglobular pheromone gland with numerous secretory cells being connected to tubuli which lead to an invaginated cuticular cribellum, is available in maleDermestes maculatus (Figs. 3 c, d and 4 c). The cribellum, provided with a caudally curved brush of fluted brisles, occurs in the centre of the 4th visible abdominal sternite (Figs. 4 a, b and 7 b). An apodemous exocrine gland is found in the lumen of the second abdominal segment of femaleLasioderma serricorne (Fig. 3 b). This lobate gland comprises many secretory cells, being connected by numerous tubuli to a sheath-like conical duct enveloping a V-shaped skeletal apodeme, which terminates in the abdominal tip. In maleTribolium castaneum, the secretory cells of both pheromone glands are connected by tubuli to two cribella, being densely covered by fluted bristles, and found in the femora of both forelegs (Fig. 7 a). Females of the phycitid speciesAnagasta kuebniella, Cadra cautella, Ephestia elutella andPlodia interpunctella are equipped with an intersegmental pheromone gland, situated between the 8th and 9th abdominal segment near the genital opening. The exocrine gland of the four moth species consists of a single layer of columnar secretory cells, lined by a spongy cuticle which seems to be permeable to the sex pheromone (Fig. 6 a). The latter is disseminated by calling females (Fig. 5 a, b) while their exocrine glands are widely exposed. Males of the above phycitid species are furnished with alar and abdominal androconia which become exposed during courtship and discharge aphrodisiac pheromones. The base of each of the androconial bristles and scales is immersed to an underlying unicellular, pheromone-producing gland (Fig. 6 d, e). The aphrodisiac pheromones, being secreted by the above glandular cells, are passing the lumen and walls of the bristles and scales, and evaporate from the surface of the latter. For example, malePlodia interpunctella possess 2 pairs of scent tufts (a small and a large one) on both sides of the 8th abdominal tergiet as well as 2 pairs of scent tufts (a small and a large one) near the base of the costal margin of the forewings (Fig. 6 b, c). Females of several phycitid species respond to the aphrodisiac pheromone of conspecific males by a pronounced readiness to mate. In the course of time, about 3 dozens of insect species (⊃3/4 coleopteran and ⊃ 1/4 lepidopterous species) have undergone sympatric speciation by sharing desiccated food in stores as a common habitat. Fertile matings between such heterogeneous species are often prevented by morphological and anatomical incompatibilities as well as physiological and behavioural barriers. Most of the species living in the storage habitat are reproductively isolated due to the molecular structure and blend composition of their pheromones (Table 2). Interestingly, some species (listed below) deviate from the majority by sharing the structure of their main pheromone components (mentioned in parenthesis), and are thus poorly separated: the curculionidsSitophilus oryzae andS. zeamais ((4S,5R)-5-hydroxy-4-methyl-3-heptanone), the tenebrionidsTribolium castaneum andT. confusum ((4R,8R)-dimethyldecanal) as well as the dermestidsTrogoderma inclusum andT. variabile ((R,Z)-14-methyl-8-hexadecenal). Theinsufficient reproductive isolation of the above species is compensated, i.a., by additional availability of a sex pheromone in femaleTribolium confusum, by different calling periods and emission rates of (R,Z)-14-methyl-8-hexadecenal in females of the forementionedTrogodema species.Trogoderma glabrum andT. granarium areincompletely isolated by sharing (R,E)-14-methyl-8-hexadecenal as a pheromone component; they are indeed capable of cross-mating, but produce sterile hybrids. Moreover, maleOryzaepbilus mercator andO. surinamensis incorporate (Z,Z)-3,6-dodecadien-11R-olide as a common chiral component to their aggregation pheromones. The females of 5 phycitid species share (Z,E)-9,12-tetradecadien-1-yl acetate as their main pheromone

Notes: Abstract A major sex pheromone component of each of fourTrogoderma species was isolated by aeration of the female beetles and absorption of the volatiles on Porapak-Q. (Z)-14-Methyl-8-hexadecenal was identified as the major component inT. inclusum andT. variabile, and (E)-14-methyl-8-hexadecenal was identified inT. glabrum. Both (Z)- and (E)-14-methyl-8-hexadecenal were found inT. granarium (Khapra beetle), in the ratio 92Z∶8E. In laboratory bioassays, male beetles exhibited arousal and mating responses to the aldehydes, and could discriminate between the geometric isomers. The daily production of the aldehyde was calculated for each species, and other active components were detected. These aeration-absorption studies contrast with earlier studies on macerated beetles, in which the aldehyde was not detected. The efficacy of the aeration-absorption system for collection of the sex pheromones is also described. The absorbent (Porapak-Q) efficiently collected the active pheromone; only minor amounts of activity were left in the other parts of the system.

Notes: Abstract On the basis of the antennal receptor potentials and the extent of attraction and copulation induced in unmated male khapra beetles, (Z)- and (E)-14-methyl-8-hexadecenal were recognized as the most important components of the pheromone system of femaleTrogoderma granarium (Everts), and were named (Z)- and (E)-trogodermal. Air blown over 10−5 to 10−4 μg of (Z)-trogodermal produced receptor potentials equivalent to that elicited by one virgin femaleT.granarium, while ∼10−2 μg of (Z)-trogodermal was required to cause complete attraction and copulation of unmated males. (Z)-Trogodermal was about 10 times more active than (E)-trogodermal. (Z)-8-Hexadecenal was ∼10−2 times less effective than (Z)-trogodermal in causing attraction and 104 time less active in stimulating copulation. (Z)- and (E)-14-methyl-8-hexadecen-1-ol and methyl (Z)- and (E)-14-methyl-8-hexadecenoate displayed a relatively low activity for unmated male khapra beetles. Methyl and ethyl oleate, ethyl linoleate, ethyl palmitate, and ethyl stearate were less effective than (Z)-trogodermal by 6–8 orders of magnitude and are nonspecific attractants. The intensity of response to a particular compound was consistent when assessed by the essential components of mating behavior: receptor potentials, attraction, and copulation.