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deep-sea environment is from any other that ecologists have visited.

The discovery of high diversity in the deep sea was critically important to the evolution and maturation of biological oceanography because it provided scientific respectability to this expensive research. Deep-sea animals were found to be interesting and sometimes weird, as National Geographic articles frequently reminded us, but of what relevance was deep-sea ecology? The discussion of diversity thrust deep-sea research into a mainstream ecology debate that was important and exciting. This development was pivotal because NSF is most comfortable supporting hypothesis-driven research on questions that are significant to mainstream science. After Howie Sanders and Bob Hessler published on deep-sea diversity (Sanders, 1967; Hessler and Sanders, 1969; Sanders and Hessler, 1967; Dayton and Hessler, 1972; Grassle and Sanders, 1973), there were abundant hypotheses to be tested, and tested they were. The Biological Oceanography Program at NSF was, and still is, the major supporter of this work.

Deep-Sea Diversity References

1967 Hessler, R.R., and H.L. Sanders. 1967. Faunal diversity in the deep-sea. Deep-Sea Res. 14:65-78.

1968 Sanders, H.L. 1968. Marine benthic diversity: A comparative study. Am. Natur. 102:243-282.

1969 Rowe, G.T., and R.J. Menzies. 1969. Zonation of large benthic invertebrates in the deep-sea off the Carolinas. Deep-Sea Res. 16:531-537.

1969 Sanders, H.L., and R.R. Hessler. 1969. Ecology of the deep-sea benthos. Science 163:1419-1424.

1972 Dayton, P.K., and R.R. Hessler. 1972. Role of biological disturbance in maintaining diversity in the deep sea. Deep-Sea Res. 19:199-208.

1973 Grassle, J.F., and H.L. Sanders. 1973. Life histories and the role of disturbance. Deep-Sea Res. 20:643-659.

1976 Jumars, P.A. 1976. Deep-sea species diversity: Does it have a characteristic scale? J. Mar. Res. 34:217-246.

New and Regenerated Productivity

This landmark achievement had its origin in a Limnology and Oceanography publication by Dugdale and Goering (1967) that introduced a deceptively simple notion: primary productivity in the ocean can be divided into the portion that uses locally recycled nutrients (regenerated production) and the portion that uses nutrients newly transported into the euphotic zone (new production), usually by the physical processes of mixing and upwelling. Dugdale and Goering's exciting and powerful concept was presented in very basic terms and specifically included ''new" nutrients entering from the atmosphere, a process that was not considered important in 1967 but is now known to be significant.

The new production concept, together with the Dugdale (1967) paper on nutrient uptake dynamics in the same issue of Limnology and Oceanography , provided biological oceanography with the mathematical formalism needed for rigorous, quantitative modeling of ocean productivity and biogeochemical fluxes. (See also Eppley et al., 1969; MacIsaac and Dugdale, 1969.) This formalism fueled the explosive growth of modeling described in the modeling section later in this paper.

Eppley and Peterson (1979) further developed the concept by arguing that at steady state the magnitude of new production is equal to the export flux of particulate organic matter out of the euphotic zone to the ocean interior. Together, the Dugdale and Goering (1967) and Eppley and Peterson (1979) papers have impressive citation index scores. At the ages of 31 and 19 years, respectively, they are cited more now than they were in their first decades. They are like fine wines. Significantly, Eppley and Peterson (1979) estimated global new production to be about 4 petagrams per year and suggested for the first time that this number approximates the sinking flux of organic carbon and, hence, the rate at which the deep sea sequesters atmospheric carbon dioxide. This number has proved very durable; it is still used in global biogeochemical budgets.

As a consequence of the work of Dugdale and Goering (1967) and Eppley and Peterson (1979), a link was forged between physical and biological oceanography. The new concept required that physical processes of mixing and upwelling be an integral part of ecosystem models dealing with new production, fish production, or export of organic material from the surface layer. Ocean physics and biology were formally wed by this landmark achievement.

The technological advance that made progress on new production possible was the use of a stable isotope tracer 15N and a mass spectrometer to measure it precisely. The 15N tracer method was a logical development of Steemann-Nielsen's (1952) breakthrough use of 14C as a tracer of carbon fixation.

Dugdale and Goering's work was supported by the NSF International Indian Ocean Expedition and its successor program, the Southeastern Pacific Expedition, using the NSF ship Anton Bruun for focused biological oceanography. The NSF decision to fund this vessel specifically for biological oceanography was a decision that had positive long-range consequences for the field. In addition to expeditionary support from NSF, laboratory support came from the Atomic Energy Commission (AEC), now the Department of Energy (DOE). In the 1960s and 1970s, NSF and AEC had a productive partnership, with NSF providing focused investigator and expedition awards and AEC providing block grants to support research groups. Dugdale and Goering went to sea with John Ryther's AEC-supported research group at the Woods Hole Oceanographic Institution. Eppley was a member of J.D.H. Strickland's AEC group at Scripps Institution of Oceanography and headed this group later during its most productive years. However, NSF was the lead agency responsible for this breakthrough and the agency should take great pride in this landmark achievement.



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