Advances in Marine Biology 41

Series Contents for Last Ten Years* VOLUME 28, 1992. Heath, M. R. Field investigations of the early life stages of marine fish. pp. 1-174. James, M. A., Ansell, A. Q. D., Collins, M. J., Curry, G. B., Peck, L. S. and Rhodes, M. C. Biology of living brachiopods, pp. 175-387. Trueman, E. R. and Brown, A. C. The burrowing habit of marine gastro- pods. pp. 389-431. VOLUME 29, 1993. KiCrboe, T. Turbulence, phytoplankton cell size, and the structure of pelagic food webs. pp. 1-72. Kuparinen, K. and Kuosa, H. Autotrophic and heterotrophic picoplankton in the Baltic Sea. pp. 73-128. Subramoniam, T. Spermatophores and sperm transfer in marine crus- taceans, pp. 129-214. Horwood, J. The Bristol Channel sole (Solea solea (L.)): a fisheries case study, pp. 215-367. VOLUME 30, 1994. Vincx, M., Bett, B. J., Dinet, A., Ferrero, T., Gooday, A. J., Lambshead, P. J. D., Pfannktiche, O., Soltweddel, T. and Vanreusel, A. Meiobenthos of the deep Northeast Atlantic. pp. 1-88. Brown, A. C. and Odendaal, F. J. The biology of oniscid Isopoda of the genus Tylos. pp. 89-153. Ritz, D. A. Social aggregation in pelagic invertebrates, pp. 155-216. Ferron, A. and Legget, W. C. An appraisal of condition measures for marine fish larvae, pp. 217-303. Rogers, A. D. The biology of seamounts, pp. 305-350. VOLUME 31, 1997. Gardner, J. P. A. Hybridization in the sea. pp. 1-78. Egloff, D. A., Fofonoff, P. W. and Onbr, T. Reproductive behaviour of marine cladocerans, pp. 79-167. Dower, J. F., Miller, T. J. and Leggett, W. C. The role of microscale turbulence in the feeding ecology of larval fish. pp. 169-220. Brown, B. E. Adaptations of reef corals to physical environmental stress. pp. 221-299. Richardson, K. Harmful or exceptional phytoplankton blooms in the marine ecosystem, pp. 301-385. *The full list of contents for volumes 1-37 can be found in volume 38. ix X CONTENTS FOR LAST TEN YEARS VOLUME 32, 1997. Vinogradov, M. E. Some problems of vertical distribution of meso- and macroplankton in the ocean, pp. 1-92. Gebruk, A. K., Galkin, S. V., Vereshchaka, A. J., Moskalev, L. I. and Southward, A. J. Ecology and biogeography of the hydrothermal vent fauna of the Mid-Atlantic Ridge. pp. 93-144. Parin, N. V., Mironov, A. N. and Nesis, K. N. Biology of the Nazca and Sala y Gomez submarine ridges, an outpost of the Indo-West Pacific fauna in the eastern Pacific Ocean: composition and distribution of the fauna, its communities and history, pp. 145-242. Nesis, K. N. Goniatid squids in the subarctic North Pacific: ecology, bio- geography, niche diversity and role in the ecosystem, pp. 243-324. Vinogradova, N. G. Zoogeography of the abyssal and hadal zones, pp. 325-387. Zezina, O. N. Biogeography of the bathyal zone. pp. 389-426. Sokolova, M. N. Trophic structure of abyssal macrobenthos, pp. 427-525. Semina, H. J. An outline of the geographical distribution of oceanic phytoplankton, pp, 527-563. VOLUME 33, 1998. Mauchline, J. The biology of calanoid copepods, pp. 1--660. VOLUME 34, 1998. Davies, M. S. and Hawkins, S. J. Mucus from marine molluscs, pp. 1-71. Joyeux, J. C. and Ward, A. B. Constraints on coastal lagoon fisheries, pp. 73-199. Jennings, S. and Kaiser, M. J. The effects of fishing on marine ecosystems. pp. 201-352. Tunnicliffe, V., McArthur, A. G. and McHugh, D. A. Biogeographical perspective of the deep-sea hydrothermal vent fauna, pp. 353--442. VOLUME 35, 1999. Creasey, S. S. and Rogers, A. D. Population genetics of bathyal and abyssal organisms, pp. 1-151. Brey, T. Growth performance and mortality in aquatic macrobenthic invertebrates, pp. 153-223. VOLUME 36, 1999. Shulman, G. E. and Love, R. M. The biochemical ecology of marine fishes. pp. 1-325. CONTENTS FOR LAST TEN YEARS x i VOLUME 37, 1999. His, E., Beiras, R. and Seaman, M. N. L. The assessment of marine pollution - bioassays with bivalve embryos and larvae, pp. 1-178. Bailey, K. M., Quinn, T. J., Bentzen, P. and Grant, W. S. Population structure and dynamics of walleye pollock, Theragra chalcogramma. pp. 179-255. VOLUME 38, 2000. Blaxter, J. H. S. The enhancement of marine fish stocks, pp. 1-54. Bergstrrm, B. I. The biology of Pandalus. pp. 55-245. VOLUME 39, 2001. Peterson, C. H. The "Exxon Valdez" oil spill in Alaska: acute indirect and chronic effects on the ecosystem, pp. 1-103. Johnson, W. S., Stevens, M. and Watling, L. Reproduction and develop- ment of marine peracaridans, pp. 105-260. Rodhouse, P. G., Elvidge, C. D. and Trathan, P. N. Remote sensing of the global light-fishing fleet: an analysis of interactions with oceanography, other fisheries and predators, pp. 261-303. VOLUME 40, 2001. Hemmingsen, W. and MacKenzie, K. The parasite fauna of the Atlantic cod, Gadus morhua L. pp. 1-80. Kathiresan, K. and Bingham, B. L. Biology of mangroves and mangrove ecosystems, pp. 81-251. Zaccone, G., Kapoor, B. G., Fasulo, S. and Ainis, L, Structural, histo- chemical and functional aspects of the epidermis of fishes, pp. 253-348. CONTRIBUTORS TO VOLUME 41 C. CONAND, Universit~ de La Rdunion, Laboratoire d'l~cologie Marine, 15 Avenue Ren~ Cassin, Saint-Denis, Cedex 9, La R~union 97715, France J.-F. HAMEL, Society for the Exploration and Valuing of the Environment (SEVE), 655 rue de la Rivi~re, Katevale (Quebec), Canada JOB 1 WO A. MERCIER, Society for the Exploration and Valuing of the Environment (SEVE), 655 rue de la Rivi~re, Katevale (Qudbec), Canada JOB 1WO; International Center for Living Aquatic Resources Management (ICLARM), Coastal Aquaculture Centre, PO Box 438, Honiara, Solomon Islands and Institut des Sciences de La Mer de Rimouski (ISMER), 310 all~e des Ursulines, Rimouski (Quebec), Canada G5L 3,41 D. L. PAWSON, National Museum of Natural History, Smithsonian Institution, Mail Stop 163, Washington DC, 20560-0163, USA M. WHITFIELD, Marine Biological Association of the United Kingdom, The Laboratory, Citadel Hill, Plymouth PL1 2PB, UK V Interactions between Phytoplankton and Trace Metals in the Ocean Michael Whitfield Marine Biological Association of the United Kingdom, The Laboratory, Citadel Hill, Plymouth PL1 2PB, UK FAX: +44 (0)1752 633 102 e-mail: 2 MICHAEL WHITFIELD This review assesses the degree to which phytoplankton in the contemporary oceans interact with the essential trace metals in their chemical environment as exemplified by the cycling of iron, manganese, cobalt, nickel and zinc. The toxic element cadmium is also considered because of the extent to which it is taken up. The stage is set by a brief consideration of the overall geo- chemical controls on the composition of sea water and their implications for the milieu within which life evolved. The utilization of the elements within the cells is addressed with the consequent implications for optimizing the uptake of essential elements and controlling the ingress of potentially toxic elements. The impact of the change from an anoxic to an oxygenated atmosphere some 2 billion years ago on the availability o f the basic building blocks for living systems is considered. The essential elements have to be delivered to the centres of synthesis in the appropriate ratios. The optimal tailoring of the flows of material into and out o f the cell to meet the requirements for maintenance, growth and reproduction requires a carefully regulated internal economy. This internal economy sets the guidelines for the interaction of phyto- plankton with the chemistry of their aqueous environment. The uptake mechanisms are explored using data derived from culture experiments indicating a significant degree of biological influence on the chemistry of the essential elements close to the cell surface (the near-field chemistry). The perspective is then widened to consider the implications of these processes for the relative concentrations of the elements and their distri- bution throughout the world's oceans (the far-field chemistry). Case histories are followed for iron, manganese, copper, zinc, cobalt and nickel reviewing (a) their distribution in the oceans, (b) their biological availabil- ity, (c) their uptake and impact upon primary production. This external economy is intimately related to the feedback between the organisms and their environment. The extent o f recycling within the ocean system by the mutually dependent processes of photosynthesis and respiration provides a clear measure of the regulatory power of the biological system itself. This is analysed in the context o f the Gaia hypothesis. Although the biological processes, to a large degree, control the availability and distribution of the essential trace metals in the oceans, the system does not appear to be optimized. For example cadmium, generally considered to be a non-essential element, is recycled more vigorously than any other element. Zinc in contrast appears to be rendered less accessible as the result o f biological activity, and phyto- plankton cells in the open ocean are straining at the limit o f diffusive trans- port to obtain sufficient supplies.