[1] Univ Southampton, Natl Oceanog Ctr, Ocean & Earth Sci, Southampton, Hants, England
[2] Univ Southampton, Inst Life Sci, Southampton, Hants, England
[3] Weizmann Inst Sci, Dept Plant & Environm Sci, Rehovot, Israel
[4] Woods Hole Oceanog Inst, Dept Marine Chem & Geochem, Woods Hole, MA 02543 USA