[1] Univ Helsinki, Dept Microbiol, POB 56, Helsinki 00014, Finland
[2] Univ Helsinki, Dept Biosci, POB 65, FIN-00014 Helsinki, Finland
[3] Eawag, Dept Aquat Ecol, CH-8600 Zurich, Switzerland
[4] Swiss Fed Inst Technol, Dept Environm Syst Sci, CH-8092 Zurich, Switzerland