[1] Univ Utrecht, Dept Geosci, NL-3508 TA Utrecht, Netherlands
[2] Newcastle Univ, Sch Civil Engn & Geosci, Newcastle Upon Tyne NE1 7RU, Tyne & Wear, England
[3] Radboud Univ Nijmegen, Dept Microbiol, IWWR, NL-6525 AJ Nijmegen, Netherlands
[4] Lancaster Environm Ctr, Ctr Ecol & Hydrol, Lancaster LA1 4AP, England
[5] Royal Netherlands Inst Sea Res NIOZ, Dept Marine Organ Biogeochem, NL-1790 AB Den Burg, Netherlands