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The Science of Nature -     

Evolution of extreme polyandry in the honeybee Apis mellifera L.

A number of hypotheses have been proposed to explain the evolution of multiple mating in the honeybee queen. In particular, the consequences of reduced intracolonial relatedness provide plausible explanations for multiple mating with up to ten drones, but fail to account for the much higher mating frequencies observed in nature. In this paper, we propose an alternative mechanism which builds on non-linear relationships between intracolonial frequencies in genotypic worker specialization and colony fitness. If genes for any worker specialization confer an advantage on colony fitness only when they are rare, this would require a stable mix of sperm from a few drones which contribute that trait, and many which do not. To ensure both specific, low within-colony proportions of “rare specialist” genes, and to reduce random variation of these proportions would require mating with high numbers of drones. The quantitative implementation shows that moderate to very high numbers of matings are required to exploit colony advantages from genotypic allocation of workers to rare tasks. Extreme polyandry thus could result from colony selection dependent on the intracolonial frequency of rare genetic specialists. Received: 30 January 1998 / Accepted after revision: 7 October 1998     

GVO-Pollenmonitoring zum Bt-Maisanbau im Bereich des NSG/FFH-Schutzgebietes Ruhlsdorfer Bruch

Aim and Background Precautionary regulations for the production of genetically modified crops (Gentechnik-Pflanzenerzeugungsverordnung, GenTPflEV 2008) provide isolation distances for the cultivation of genetically modified maize – Bt-maize – in the vicinity of conventional maize cultivation (150?m) as well as of organic maize cultivation (300?m). Nature reserves are not included. The aim of this study was to investigate whether maize pollen of surrounding fields were dispersed in the Flora-Fauna-Habitat (FFH) Ruhlsdorfer Bruch. In the vicinity of the nature reserve Bt-maize species MON 810 as well as conventional maize was grown. The survey should provide appropriate isolation distances for the cultivation of Bt-maize with regard to sustainable protection of non-target-organisms (NTO) in the Ruhlsdorfer Bruch. Materials and Methods The collection of maize pollen in the Ruhlsdorfer Bruch was carried out at three sampling sites in the FFH or rather in the immediate vicinity in July and August 2007 by means of twelve technical and three biological pollen samplers. The technical samplers Sigma-2/PMF enable point sampling. They are primarily influenced by wind and topography and provide information about the effective entry, the maize pollen flow and the maize pollen deposition at the location of the sampler. Honey bees roam over longer distances and are hence planar collectors. Thus, they also provide information about the plants blooming in a distinct area. Furthermore, the biological preferences during the collection are captured, whereas a technical sampler does not perform a species dependent selection. Hence, both the technical and the biological samplers complement one another in their scope of application. The pollen samples were identified microscopically, enumerated quantitatively, and the pollen-DNA was analysed by means of the PCR-method. Results All monitoring sites at the Ruhlsdorfer Bruch revealed maize pollen entries. The pollen deposition reached values of 1.75 million maize pollen/m² in the close-up range. The monitoring sites located 120?m within the protected area still featured 99,000 maize pollen/m². As shown by the statistical analysis, at a distance of 1,000?m still 28,000 maize pollen/m² must be expected. The results of the microscopic pollen analysis of the pollen pellets proved that the bees collect maize pollen at all three sites. Although maize pollen is not the main food source the high collection efficiency of the bees resulted in large amounts of introduced pollen. The biomolecular proof of pollen-DNA in the field samples was corroborated by analogue results for both the technical and biological pollen sampling by two independent laboratories. The results of these quality controlled analyses gave unambiguous evidence that under the cultivation conditions in 2007 an entry of Bt-maize pollen into the FFH Ruhlsdorfer Bruch was existent. Discussion The maize pollen deposition at the Ruhlsdorfer Bruch corresponds with additional supra-regional investigations which were conducted over several years with the same method. However, the pollen entries into the Ruhlsdorfer Bruch are above-average. Possible reasons are the size of the maize fields (>?10?ha), the geographical collocation between maize fields and protected area (lee location, through-shaped, thermal currents) as well as weather conditions and maize bloom during the sampling period July and August 2007. A distance of 1,000?m or more is necessary to avoid maize pollen deposition of more than 100,000/m² with a probability of 90?%. Conclusions The results of this investigation prove an entry of Bt-maize pollen into the FFH under the cultivation and weather conditions in summer 2007. According to the results of this and related studies and considering precautionary principle and the effect terms defined in the Federal Immission Protection Law, it becomes clear that reasonable isolation distances between Bt-maize fields and protected areas have to be introduced in order to minimise the entry of GM-maize pollen and to prevent adverse effects on protected NTOs. Recommendations Due to the fact that no toxicological investigations are available for the butterflies occurring in the FFH, highest protection standards should be implemented to avoid Bt-maize pollen entries into the Ruhlsdorfer Bruch. That implies that highest possible isolation distances for Bt-maize cultivation have to be considered, but at least a distance of 1,000?m is recommended. Following this a maize pollen deposition of more than 100,000 maize pollen/m² should be pevented with a certainty of 90?% as well as an exceeding exposure of foraging insects. Outlook By means of the pollen monitoring it becomes possible to survey empirically defined limits of pollen exposure. This should be performed in the nature reserve area at sites of expected maximum exposure, e.?g. at the boundaries of the FFH towards the maize fields and at exposed sites.