There has been considerable debate about both the mechanisms controlling primary production, and the interpretation of data from Fe-enrichment experiments conducted in high nitrate-lotv chlorophyll (HNLC) regions. This paper presents results of 3 in vitro Fe-enrichment experiments performed in May 1993 and May 1994 at Ocean Station Papa (OSP) in the NE subarctic Pacific. Expt 1 (May 1993) considered jointly the influence of both Fe supply and microzooplankton herbivory on algal stocks, while the second (May 1993) investigated the influence of Fe supply on the partitioning of C, N and Fe between algal size classes. The third study (May 1994) monitored changes in phytoplankton stocks in Fe-enriched carboys containing mesozooplankton Assuming simllar environmental conditions in May 1993 and 1994, then the comblned findings indicate that under ambient conditions autotrophic cells < S pm compose >70'% of algal biomass, primarily utilized ammonium, showed no Femediated enhancement of N-normalized nitrate or ammonium uptake, and were thus unlikely to be Felimited. Although the coupling between grazer mortality and algal growth of these cells ( g / p ) was ca 0.71 k 0.25, they grew at close to maximal rates but exhibited negligible net growth rates, suggesting strong grazer control. Dissolved iron (DFe) supply promoted a rapid increase in the abundance of large, initially rare, diatoms (mainly Njtzschla spp.) over 6 d, elevated diatom growth rates to close to their theoretical maximum and increased rates of incorporation of N-normalized nitrate and Fe by these cells over time. Unlike the small cells, stocks of large autotrophs also increased in the controls; DFe measurements indicate that this was probably due to inadvertent Fe contamination. The development of a phytoplankton population, of a cell size probably too large to be grazed at a significant rate by microzooplankton, resulted in a decoupling of herbivory and algal growth. In addition, mesozooplankton herbivory appeared unable to prevent the accumulation of these large cells; in May 1994, Fe supply prompted an 8-fold increase in algal stocks over 6 d in carboys where grazers equivalent to the maximum annual in situ pelagic abundance were present. The findings agree with the theories of others, that Fe supply rather than grazing provides the ultimate control over the phytoplankton community in the NE subarctic Pacific.
Although an extensive biological time series data set of phytoplankton and zooplankton standing stocks has been collected over a 30 yr period in the NE subarctic Pacific at Ocean Station Papa (OSP), the majority of these data were obtained before recent advances in our understanding of the structure and functioning of the manne microbial food web. In addition, recent studies did not obtain data during the winter penod. This paper provides the first contemporary biological description of the NE subarctic Pacific dunng winter Data from 2 winter cruises in the vicinity of OSP indicate that the abundance and composition of the autotrophic and heterotroph~c assemblage at these times are similar to those observed during late spring and summer by others. In contrast, winter mesozooplankton standing stocks are considerably less than observed In summer. These findings confirm the hypothesis that the relatively shallow winter mixed layer permits relatively high standing stocks of phytoplankton (20 mg C m-3) and consequently of microheterotrophs (7.1 to 13.1 mg C m-7 to subsist over the winter period. An assessment of autotrophic and heterotrophic biomass suggests that the requirements of the mesozooplankton (Stages IV to VI), microzooplankton and heterotrophic nanoflagellates may potentially be balanced by the standlng stocks of microplankton, nanoplankton/picophytoplankton, and picoplankton, respectively. However, the carbon requirements of heterotrophic bacteria cannot be balanced without invoking slower turnover times for thls pool relative to the turnover times for the dissolved organic carbon (DOC) pool.
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