The availability of nitrogen is important in regulating biological productivity in marine environments. Deepwater nitrate has long been considered the major source of new nitrogen supporting primary production in oligotrophic regions of the open ocean, but recent studies have showed that biological N2 fixation has a critical role in supporting oceanic new production. Large colonial cyanobacteria in the genus Trichodesmium and the heterocystous endosymbiont Richelia have traditionally been considered the dominant marine N2 fixers, but unicellular diazotrophic cyanobacteria and bacterioplankton have recently been found in the picoplankton and nanoplankton community of the North Pacific central gyre, and a variety of molecular and isotopic evidence suggests that these unicells could make a major contribution to the oceanic N budget. Here we report rates of N2 fixation by these small, previously overlooked diazotrophs that, although spatially variable, can equal or exceed the rate of N2 fixation reported for larger, more obvious organisms. Direct measurements of 15N2 fixation by small diazotrophs in various parts of the Pacific Ocean, including the waters off Hawaii where the unicellular diazotrophs were first characterized, show that N2 fixation by unicellular diazotrophs can support a significant fraction of total new production in oligotrophic waters.

Abstract. Marine N2 fixing microorganisms, termed diazotrophs, are a key functional group in marine pelagic ecosystems. The biological fixation of dinitrogen (N2) to bioavailable nitrogen provides an important new source of nitrogen for pelagic marine ecosystems and influences primary productivity and organic matter export to the deep ocean. As one of a series of efforts to collect biomass and rates specific to different phytoplankton functional groups, we have constructed a database on diazotrophic organisms in the global pelagic upper ocean by compiling about 12 000 direct field measurements of cyanobacterial diazotroph abundances (based on microscopic cell counts or qPCR assays targeting the nifH genes) and N2 fixation rates. Biomass conversion factors are estimated based on cell sizes to convert abundance data to diazotrophic biomass. The database is limited spatially, lacking large regions of the ocean especially in the Indian Ocean. The data are approximately log-normal distributed, and large variances exist in most sub-databases with non-zero values differing 5 to 8 orders of magnitude. Reporting the geometric mean and the range of one geometric standard error below and above the geometric mean, the pelagic N2 fixation rate in the global ocean is estimated to be 62 (52–73) Tg N yr−1 and the pelagic diazotrophic biomass in the global ocean is estimated to be 2.1 (1.4–3.1) Tg C from cell counts and to 89 (43–150) Tg C from nifH-based abundances. Reporting the arithmetic mean and one standard error instead, these three global estimates are 140 ± 9.2 Tg N yr−1, 18 ± 1.8 Tg C and 590 ± 70 Tg C, respectively. Uncertainties related to biomass conversion factors can change the estimate of geometric mean pelagic diazotrophic biomass in the global ocean by about ±70%. It was recently established that the most commonly applied method used to measure N2 fixation has underestimated the true rates. As a result, one can expect that future rate measurements will shift the mean N2 fixation rate upward and may result in significantly higher estimates for the global N2 fixation. The evolving database can nevertheless be used to study spatial and temporal distributions and variations of marine N2 fixation, to validate geochemical estimates and to parameterize and validate biogeochemical models, keeping in mind that future rate measurements may rise in the future. The database is stored in PANGAEA (doi:10.1594/PANGAEA.774851).

Independent data from the Gulf of Mexico are used to develop and test the hypothesis that the same sequence of physical and ecological events each year allows the toxic dinoflagellate Karenia brevis to become dominant. A phosphorus‐rich nutrient supply initiates phytoplankton succession, once deposition events of Saharan iron‐rich dust allow Trichodesmium blooms to utilize ubiquitous dissolved nitrogen gas within otherwise nitrogen‐poor sea water. They and the co‐occurring K. brevis are positioned within the bottom Ekman layers, as a consequence of their similar diel vertical migration patterns on the middle shelf. Upon onshore upwelling of these near‐bottom seed populations to CDOM‐rich surface waters of coastal regions, light‐inhibition of the small red tide of ∼1 ug chl l−1 of ichthytoxic K. brevis is alleviated. Thence, dead fish serve as a supplementary nutrient source, yielding large, self‐shaded red tides of ∼10 ug chl l−1.The source of phosphorus is mainly of fossil origin off west Florida, where past nutrient additions from the eutrophied Lake Okeechobee had minimal impact. In contrast, the P‐sources are of mainly anthropogenic origin off Texas, since both the nutrient loadings of Mississippi River and the spatial extent of the downstream red tides have increased over the last 100 years. During the past century and particularly within the last decade, previously cryptic Karenia spp. have caused toxic red tides in similar coastal habitats of other western boundary currents off Japan, China, New Zealand, Australia, and South Africa, downstream of the Gobi, Simpson, Great Western, and Kalahari Deserts, in a global response to both desertification and eutrophication.

Monospecific blooms of phytoplankton can disrupt pelagic communities and negatively affect human health and economies. Interspecific competition may play an important role in promoting blooms, and so we tested (1) whether the outcome of competition between the red tide dinoflagellate Karenia brevis (ex Gymnodinium breve) and 12 cooccurring phytoplankters could be explained by allelopathic effects of compounds released by K. brevis and (2) whether waterborne, lipophilic molecules, including brevetoxins, are involved. Nine of 12 phytoplankton species were suppressed when grown with live K. brevis at bloom concentrations. K. brevis extracellular filtrates or lipophilic extracts of filtrates inhibited six of these nine species, indicating allelopathy. However, these inhibitory effects were weaker than those experienced by competitors exposed to live K. brevis. Brevetoxins at ecologically reasonable waterborne concentrations accounted for the modest inhibition by K. brevis of only one competitor, Skeletonema costatum. The addition of brevetoxins also caused significant autoinhibition, reducing the maximum concentration of K. brevis. Allelopathy is one mechanism by which K. brevis appears to exhibit competitive advantage over some sympatric phytoplankters, although unidentified compounds other than brevetoxins must be involved, in most cases. K. brevis was also susceptible to competitive exclusion by several species, including Odontella aurita and Prorocentrum minimum, known to thrive during K. brevis blooms. Although field experiments are required to assess whether allelopathy plays a fundamental role in bloom dynamics, our results indicate that allelopathy occurs widely but with species-specific consequences.Competition is one of the dominant forces structuring communities, including marine pelagic communities (Hutchinson 1961). The production and release of compounds that inhibit competitors, a process known as allelopathy, is a mechanism of interference competition that is hypothesized to be important among phytoplankton (Smayda 1997), affecting species succession (Keating 1977), especially under eutrophic conditions (Maestrini and Bonin 1981). Allelopathy may be a successful strategy for phytoplankton species that occur in dense blooms, maximizing the concentration of allelopathic compound(s) exposed to competitors and mini-1 Corresponding author (julia.kubanek@biology.gatech.edu). AcknowledgmentsThis research was supported by NSF grant OCE-0134843 to J.K. M.K.H. was supported as a summer undergraduate research assistant by the Camille and Henry Dreyfus Foundation Environmental Chemistry Program. Nutrient analyses were funded by NOAA MERHAB grant MER 02-627A to T.A.V., and ELISA analyses were funded by COP-NOAA MERHAB, NIEHS PO1 ES10594-03, and the CDC-FDOH U50/CCU423360-01 to J.N. We thank P. Tester, R. Pierce, M. Hay, and E. Litchman for advice in planning experiments and two anonymous reviewers for suggestions that improved the manuscript. We also thank E. Prince, A. Prusak, E. John, D. Collins, and A....

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Diatoms with symbiotic N2-fixing cyanobacteria are often abundant in the oligotrophic open ocean gyres. The most abundant cyanobacterial symbionts form heterocysts (specialized cells for N2 fixation) and provide nitrogen (N) to their hosts, but their morphology, cellular locations and abundances differ depending on the host. Here we show that the location of the symbiont and its dependency on the host are linked to the evolution of the symbiont genome. The genome of Richelia (found inside the siliceous frustule of Hemiaulus) is reduced and lacks ammonium transporters, nitrate/nitrite reductases and glutamine:2-oxoglutarate aminotransferase. In contrast, the genome of the closely related Calothrix (found outside the frustule of Chaetoceros) is more similar to those of free-living heterocyst-forming cyanobacteria. The genome of Richelia is an example of metabolic streamlining that has implications for the evolution of N2-fixing symbiosis and potentially for manipulating plant–cyanobacterial interactions.

Diel protein and carbohydrate content in Trichodesmium thiebautii was measured to evaluate the relationship to buoyancy status. Carbohydrate:protein ratio was the best predictor of buoyancy and fit a cosine curve with increasing values during the day and decreasing values at night in cycles that paralleled observed diel buoyancy patterns. This ratio also increased in short-term experiments as a function of light and increased in parallel with decreasing positive buoyancy. We used changes in this ratio to estimate the potential for vertical migration. Whereas limited vertical excursions in the upper 70 m are possible, deeper migrations appear unlikely unless respiration rates decrease significantly. N:P ratios in sinking and floating colonies were used to test for the P acquisition at depth (vertical migration). We noted that pooled N:P ratios were not significantly different between sinking and ascending colonies (N:P = 65.6 and 66.3, respectively) collected along the northern Australian coast, much like published results from north of Hawaii. Highly significant differences (p <0.0001) were observed in the western Gulf of Mexico between sinking and ascending colonies (N:P = 87.0 and 43.5, respectively) and provide the best direct evidence to date of vertical migration for P acquisition. Our physiological data on compositional changes during buoyancy reversals suggest a complex relationship between light and nutrients. It appears likely that light and P metabolism interact to regulate the vertical extent of migrations, with deep vertical migration regulated by P metabolism superimposed on a mixed-layer light-driven migration. The variability in N:P ratios suggests that care should be taken in assuming buoyancy reversals always result in P acquisition in this oceanic cyanobacterium.

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