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INTRODUCTIONFor most insects, adult nutrition affects not only survival of individuals butalso their reproductive output. In particular, fecundity of females commonlydepends on ingestion of protein necessary for egg development, whereasmale fertility is not highly protein dependent (Chapman, 1982; House, 1974;Engelmann, 1999). Furthermore, because insects generally are unable toconvert lipids to monosaccharides (Bignell, 1981), carbohydrates and pro-teins serve as primary energy sources in both sexes (House, 1974).Recentstudiesrevealthatanumberofinsectsexhibitsex-specificdiffer-ences in feeding behavior that are correlated with adult nutritional require-ments. For example, young female tephritid fruit flies (Diptera: Tephritidae)that are protein-hungry are more attracted to odors of protein food baitsthan to odors of host fruit, whereas males and protein-fed females areattracted more toward fruit odors (Cornelius

Insects, including cockroaches, have the ability to select a proper diet from different nutrient choices. We have showed previously that various neurotransmitters and neuromodulators appear to regulate certain aspects of feeding in the cockroach, Rhyparobia madera. In the current study, we examined the role of octopamine in feeding behavior of cockroach nymphs. By either injection or direct incorporation into the diet blocks, an octopamine agonist (octopamine or synephrine) or antagonist (phentolamine) was effective in altering feeding in R. madera nymphs. Both octopamine and synephrine increased feeding slightly, while phentolamine decreased feeding dramatically. Phentolamine was able to decrease feeding, but not motor activity, when injected directly into the nymphs. Octopamine appears to cause increased feeding in the cockroach.

The immature stages of some parasites live in prey animals (intermediate hosts) and only reach reproductive maturity when they are eaten by final host predators. Some of these parasites alter intermediate host behavior in ways that increase the likelihood of predation (parasite transmission). Using the acanthocephalan (Moniliformis moniliformis) in the cockroach (Periplaneta americana), we show that infected cockroaches experienced a decrease in wind-evoked escape responses, a predator avoidance behavior, that consisted of fewer escape responses, longer latency and higher threshold for escape behavior. We found no correlates of the impairment of the escape behavior in the abdominal portion of the escape neuronal circuitry. This study demonstrates a parasite-induced alteration of a behavior dedicated to predator avoidance.

Experiments were performed to test for nutrient-specific olfactory learning in the American cockroach, Periplaneta americana L. In a conditioning period, cockroaches were presented with two complementary foods, one of which contained protein and the other carbohydrate, this combination allowing them to select a nutritionally balanced diet. The foods were separated in space, and each was paired with one of two odors, citral or carvone. The cockroaches were then selectively deprived of one of the nutrients for 24 or 48 h. In the final (test) phase of the experiment the movement of the cockroaches toward the nutrient-associated odors was monitored. Associative learning was demonstrated with respect to protein, with protein-deprived cockroaches moving more frequently toward the protein-associated odor. No learned associations between carbohydrate and odor were demonstrated. These data are contrasted with similar experiments on an herbivorous insect, the locust Locusta migratoria.

In the courtship behavior of the German cockroach, the male presents tergal glands to the female and feeds her with glandular secretions to place her in the appropriate precopulatory position. The phagostimulant activity of the secretions was quantitatively examined using the polyethylene glycol film method. The methanol extract of the glands on the eighth tergite induced a potent feeding response in 6-day-old virgin females (EC 50 = 0.0037 male equivalent/40 μg PEG spot). However, there was no temporal relation between the feeding response and the sexual receptivity of the females. Moreover, besides virgin females, the extract induced a feeding response in gravid or mated females, males, and the last-instar nymphs. These results strongly suggest that the secretions function as a dietary feeding stimulant in principle but as a courtship pheromone in the context of courtship behavior where the stimulants are offered as a nuptial gift.

The sexual behavior of males and females ofEurycotis floridana was investigated and the various associated behavioral sequences are described. Olfactometer data proved that the male produces a volatile sex pheromone attractive at a distance to conspecific females. The male initiates courtship behavior by exposing the glandular areas on the anterior parts of abdominal tergites 2, 7, and 8. This male calling behavior was observed throughout the day. The males can mate when 8 days old, whereas virgin females are sexually receptive 18 days after becoming adults. Once attracted near the male, the female opens her genital atrium and climbs on the back of the male, where she feeds on the glandular secretions that oozed around a little tuft of setae on the first tergite. These setae are mechanoreceptors and they are stimulated when licked by the females, which informs the male that she is in a proper position for copulation.

We studied agonistic interactions among male Madagascar hissing cockroaches,Gromphadorhina portentosa, in groups of five (low-density) or 10 (high-density) males. Consistent with previous studies of male pairs, we observed aggression (Abdomen Flick, Abdomen Push, Butt, Lunge), submission (Crouch, Retreat), and noncontact behavior (Abdominal Extension, Abdomen Thrash, Agonistic Hiss, Stilt). Males at both densities performed all acts. However, males in the high-density group performed Abdomen Push significantly more often than males at a low density. The rate of each remaining act was unaffected by density. Regardless of density, males within social groups varied in aggression. More aggressive males utilized frontal assaults (Butt and Lunge) during interactions, while males displaying lower levels of aggression preferentially used the abdomen during interactions. More aggressive males performed Abdomen Flick more frequently, while males displaying lower levels of aggression performed Abdomen Push. We also investigated the relationship between male aggression and the four noncontact behaviors. We found that Abdominal Extension, Abdomen Thrash, and Agonistic Hiss were positively correlated with our aggregate measure of male aggression suggesting these are aggressive displays. Stilt was positively correlated with measures of both aggression and submission, leaving its function unclear. None of the behavioral acts examined in this study were highly correlated with male weight. Our results are discussed in light of possible hypotheses addressing the function of specific behavior during male-male competition.

Two chemical signals are essential in all cockroach sexual behavioral sequences: the sex pheromone released by one partner, generally the female (for long distance attraction), and an aphrodisiac sex pheromone produced exclusively by male tergal glands (for female mounting and tergal contact or “feeding” behavior). Unlike the other cockroach groups, the males of the Oxyhaloinae species produce both chemical signals: the pheromone and the aphrodisiac. The occurrence of three patterns of mating behavior (A, B, and C), the production of male sex pheromones, and the existence in the male of developed sternal and tergal glands in seven related Oxyhaloinae species, make these cockroaches a useful model for studying the evolution of mating behavior patterns. The various types of mating behavior were not classified in the previous studies by Roth and Barth. In this report, they have been named type A (female in upper position), B (male in upper position), and C (male and female end to end). In type A mating, the male tergal glands, which are licked by the females, are well developed, whereas in types B and C, there is no licking of the male's tergal secretion by the females and the tergal glands are much less developed; the aphrodisiacs secreted by the tergal glands may no longer act in this case through contact chemoreception, but through an olfactory process involving volatile components. One common sex pheromone component seems to be acetoin. I suggest that the mating behavior tends from A toward B and C during the evolutionary process with a concomitant regression of the tergal glands and changes in the aphrodisiac emission levels. The mating behavioral sequences of cockroaches (Dictyoptera) and crickets (Orthoptera) show a striking degree of similarity and are probably examples of convergent evolution.

Virgin German cockroach females, Blattella germanica(L.), were observed, for the first time, to exhibit a characteristic calling behavior during which females emit a volatile sex pheromone. Under a photoperiod of 12L∶12D, the percentage of 7-day-old virgin females that exhibited this behavior peaked before the end of the scotophase in a similar pattern to the diel periodicity of mating. A clear relationship was evident between calling and stages of sexual receptivity during successive gonotrophic cycles. Females initiated calling 5–6 days after the imaginal molt, when their basal oocytes were 1.6 mm long. If not mated, females continued to exhibit bouts of calling during the next 3–4 days until 24 h before ovulation. Calling was completely suppressed by mating as well as the presence of an egg case in the genital atrium in both virgin and mated gravid females. We suggest that calling and the emission of a volatile sex pheromone serve to attract males from a distance as well as to potentiate responses to contact sex pheromone in aggregations.

Parasites are known to alter the behavior of their hosts, but little is known about their effects on responses to sexual stimuli. Periplaneta americanainfected with the acanthocephalan Moniliformis moniliformiswere compared to uninfected animals in their behavioral and electroantennogram responses to a synthetic P. americanapheromone component, periplanone-B, and a pheromone mimic, bornyl acetate. In a t-maze there was no significant difference between infected animals' responses to periplanone-B and a random binomial distribution; the responses of uninfected animals were significantly nonrandom. The electroan-tennogram responses of infected and uninfected animals to bornyl acetate or periplanone-B did not differ significantly, however, indicating that the alteration probably does not occur at the peripheral level but at a central nervous system level.

Effects of dehydration on the behavioral response to humidity have been examined in the cockroach, Periplaneta americanaL. Under normal conditions cockroaches prefer lower humidity, but the preference is reversed by dehydration. Dehydrated cockroaches can distinguish smaller differences between two air streams compared with normal individuals. Cockroach behavioral responses to humidification include swing of antennae and locomotion. The period from onset of humidification to behavioral response decreases with increasing dehydration, indicating that the humidity threshold required for behavioral response decreases. Observed modulations of the behavioral response are discussed in relation to the contribution of the antennal hygroreceptors.

When juvenile praying mantids (Tenodera sinensis)were exposed to unpalatable prey (the milkweed bug Oncopeltus fasciatus),they attacked, sampled, and then usually rejected the prey. About 70% of the handling time was spent feeding. When offered a second milkweed bug, the mantids usually attacked the prey. However, the overall time required for the mantids to sample, recognize, and then reject the unpalatable prey decreased by half. The proportion of handling time that was spent feeding remained the same as in the first encounter. In contrast, when the second prey individuals encountered by mantids were Drosophila melanogaster,the flies were completely consumed and the proportion of handling time that was spent feeding significantly increased. When praying mantids were exposed to the milkweed bugs for the first time, up to 33% of the bugs survived attack by the mantids. Survival of milkweed bugs increased to 55% when mantids had been previously exposed to the bugs. In contrast, flies that were caught never survived.

The diel pattern of pheromone-releasing behavior (calling) of the adult female brown-banded cockroach, Supella longipalpa (F.),was examined. Calling occurs discontinuously throughout most of the scotophase in a 12L: 12D photoperiodic regime. Females exhibited a free-running calling rhythm after transfer to continuous light or dark conditions. Shifts in the temporal pattern of calling following changes in the timing of the photoperiodic cues indicated that “lights-off” is the Zeitgeber. Studies of insects under long- and short-day conditions suggest that, although insects within a population may call synchronously or asynchronously, respectively, the duration of an individual's calling bout is innately limited.

The social behavior of five species of Zetoborinae cockroaches is compared with respect to inter-individual interactions of nymphs in the laboratory. These species belong to the same Neotropical subfamily and were characterized as gregarious (Lanxoblatta emarginata, Parasphaeria boleiriana, Phortioeca nimbata, Schultesia lampyridiformis) and solitary (Thanatophyllum akinetum) by previous field studies. Our results show that gregarious species accept closer contacts than does the solitary one. The solitary species did not display especially short, infrequent or less diverse behavioral sequences when forced to remain aggregated, but its interactions are characterized by fewer acts promoting contact and more dominance-like acts. The solitary species symmetrically interacts with conspecifics and does not show specific dispersal-promoting behaviors. This suggests that the solitary behavior observed in the field for species of Zetoborinae mainly results from a passive spacing tendency and a lack of attraction for conspecifics. One of the gregarious species, P. boleiriana, was previously described as subsocial with nymphs remaining with the female in a wood chamber after brood birth. This species does not show a peculiar behavioral repertoire but its interactions are characterized by more dominance-like behaviors than are those of the non-subsocial gregarious species.

The effects of female sex pheromone concentration and time of day on behavioral responses of male brown-banded cockroaches were monitored with a two-choice olfactometer. The duration of each behavior or behavioral event in a deterministic response sequence and the probabilities of behavioral transitions were analyzed. Pheromone concentration had a significant effect on all behaviors in the response sequence. The time of testing, relative to the entrainment photoperiodic regime, significantly affected behavioral events early in the sequence but not later-occuring behaviors. Although the probability of males choosing the pheromone rather than the solvent control was a function of the pheromone concentration, it was independent of the time of testing. By affecting the probability of behavioral transitions, the pheromone concentration and time of testing determined the number of insects exhibiting specific behaviors in the response sequence. The behavioral response sequence can be characterized by probabilities of behavioral transitions along with latencies of early behavioral acts.