Publications

1999
Bloch G. Regulation of queen-worker conflict in bumble-bee (Bombus terrestris) colonies. PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES. 1999;266 :2465-2469.Abstract
In annual colonies of bumble-bees overt queen-worker conflict is limited to a distinct `competition phase' (CPh). In unmanipulated Bombus terrestris colonies, the queen's switch to male production (the `switch point', SP) accounted for only 22% of the variation in the onset of the CPh. In some colonies, the CPh even began before the SP. The CPh was more strongly correlated with the transition in queen production (r=0.79). Replacing the queen eggs with male eggs or doubling the number of workers in young colonies resulted in a significantly earlier onset of the CPh and a significantly earlier transition to queen production. Replacing queen eggs with female eggs did not have this effect. These manipulations did not affect the timing of the queen's switch from female to male production. These findings show that the mechanism underlying the queen-worker conflict in insect societies is more complex than previously appreciated. The onset of queen-worker conflict cannot be attributed simply to a single factor such as the queen's switch to male production or a decrease in queen inhibition. Rather, multiple cues are important.
Bloch G, Hefetz A. Regulation of reproduction by dominant workers in bumblebee (Bombus terrestris) queenright colonies. BEHAVIORAL ECOLOGY AND SOCIOBIOLOGY. 1999;45 :125-135.Abstract
The mechanisms of regulating worker reproduction in bumblebees were studied by direct behavioral observations and by measuring ovarian development and juvenile hormone (JH) biosynthesis rates in workers under different social conditions. Workers in the last stage of Bombus terrestris is colony development (the competition phase) had the lowest ovarian development and JH biosynthesis rates. Gallows introduced into colonies immediately after queen removal (dequeened colonies) demonstrated a significant increase in ovarian development before, bur not during, the competition phase. These findings differ from the higher ovarian development in colonies during the competition phase predicted by the prevailing hypothesis that worker reproduction starts in response to a decrease in queen inhibition. Reproduction of callows housed with dominant workers in small queenless groups was inhibited as in queenright colonies. This suggests that the reduced ovarian development and JH biosynthesis rates observed in dequeened and normally developing colonies during the competition phase also reflect inhibition by dominant workers. Thus, two distinct stages of inhibition of reproduction seem to exist: (1) before the competition phase, when the queen slows down worker ovarian development and prevents oviposition; (2) during the competition phase, when dominant workers inhibit ovarian development of other workers. Between these stages there seems to be a temporal ``window'' of enhanced worker reproductive development. The queen's typical switch to haploid egg production was not associated with changes in worker ovarian development or JH biosynthesis rates. These findings suggest that regulation of worker reproduction in B. terrestris is not determined by simple changes in the queen's inhibition capacity or by the sex of offspring and that the worker's role is more important than previously believed.
1996
Bloch G, Borst DW, Huang ZY, Robinson GE, Hefetz A. Effects of social conditions on Juvenile Hormone mediated reproductive development in Bombus terrestris workers. PHYSIOLOGICAL ENTOMOLOGY. 1996;21 :257-267.Abstract
During the annual life cycle of the bumble bee Bombus terrestris (L.) colony, there is a stage characterized by worker reproduction in the presence of the queen. It has been proposed that this is a result of a decrease in queen inhibition. This hypothesis was examined by studying the effects of queens taken from colonies at different stages of development on several aspects of worker physiology and behaviour: rates of Juvenile Hormone (JH) release in vitro, ovary development, and behaviour associated with reproduction. After optimizing and validating the radiochemical assay for JH release for bumble bee workers, we found that queenless workers had significantly more developed ovaries and higher rates of release of JH than did queenright workers, confirming and extending previous findings that suggest that bumblebee ovarian development is under JH control. Mated queens, separated from their colony and brood, can have the same inhibitory effect on the reproductive development of callow workers. In contrast, workers confined with virgin queens or in queenless groups demonstrated a significantly higher rate of release of JH, overt aggression and threatening behaviours. However, there were no differences in rates of release of JH between workers confined in groups in the laboratory with queens taken from colonies either before or after the onset of worker reproduction. Furthermore, overt aggression and threatening behaviours were similar and low in both types of groups. These results gave no support to the hypothesis that a decrease in queen inhibition is associated with the onset of worker reproduction. We also show that young workers reared in colonies either before or after worker reproduction occurs, or in queenless colonies, all demonstrated similar, low rates of release of JH. These results suggest that older workers may inhibit the corpora allata of younger workers in queenless colonies.
1995
Bloch G, WOOL D. ESTERASE-ACTIVITY IN POPULATIONS OF THE WHITEFLY, BEMISIA-TABACI (HOMOPTERA, ALEYRODIDAE) - HERITABILITY AND ASSOCIATED ORGANOPHOSPHORUS INSECTICIDE RESISTANCE. BULLETIN OF ENTOMOLOGICAL RESEARCH. 1995;85 :11-19.Abstract
The association of esterase (EST) activity with resistance to the organophosphorus (OF) insecticide methidathion was investigated in field-collected populations of whitefly (Bemisia tabaci (Gennadius)) in Israel. The inheritance of EST activity was studied by controlled crosses in the laboratory. Among-family variance of EST activity was highly significant although all families were maintained in the same rearing room. This indicates that genetic or common environmental effects due to rearing each family on a different caged plant must have been important. Heritability estimated from son-mother regression was h(2)=0.98, but this estimate seems to be unrealistically high. Daughter-mother and daughter-mid-parent regressions produced lower heritability estimates (as expected). None of the regression coefficients of daughters on mother, however, were significantly different from zero and the regression on parent explained only very small amounts of the activity variation in the offspring. Estimates obtained from intraclass correlations among offspring were higher and outside the acceptable range, reflecting the variance component due to the common environment. The predictive value of the heritability estimates appears to be very low. The frequency distribution of activity among individuals sampled from one insecticide treated site (AM) was skewed to the right, as previously reported. But samples from another site (TZ) showed a symmetrical distribution, unlike the previous pattern. When field collected individuals from AM were released on clean plants in the laboratory, samples taken from the plants one or more days later showed symmetrical EST distribution. Electrophoretic paterns of field and laboratory samples were the same, but band intensity was stronger in samples from laboratory populations. Differences in mean EST activity between populations and sampling years were unrelated to methidathion resistance.
1994
Bloch G, WOOL D. METHIDATHION RESISTANCE IN THE SWEET-POTATO WHITEFLY (ALEYRODIDAE, HOMOPTERA) IN ISRAEL - SELECTION, HERITABILITY, AND CORRELATED CHANGES OF ESTERASE-ACTIVITY. JOURNAL OF ECONOMIC ENTOMOLOGY. 1994;87 :1147-1156.Abstract
Artificial selection for increased resistance to methidathion in two replicate lines of the whitefly, Bemisia tabaci (Gennadius) (originating from a cotton-field-collected greenhouse population) was successful; LC(50)s increased 7-8.6-fold in eight generations in the selected lines. This indicates the existence of additive genetic variance for resistance in the source population. Minimal realized heritability estimates, calculated from the response to selection, were h(2) = 0.49 after one generation (with minimal effects of common laboratory environment and inbreeding) and a mean value of h(2) = 0.344 after eight generations. Esterase activity (measured from the hydrolysis of beta-naphthyl butyrate) increased in the selected lines in correlation with resistance. We observed no change in mean esterase activity in the unselected (control) population. No consistent differences in fitness components between selected and control lines were detected during selection, but females exposed to sublethal doses of methidathion tended to have increased fecundity.

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