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We measured N2 fixation rates from oceanic zones that have traditionally been ignored as sources of biological N2 fixation; the aphotic, fully oxygenated, nitrate (NO−3)-rich, waters of the oligotrophic Levantine Basin (LB) and the Gulf of Aqaba (GA). N2 fixation rates measured from pelagic aphotic waters to depths up to 720 m, during the mixed and stratified periods, ranged from 0.01 nmol N L−1 d−1 to 0.38 nmol N L−1 d−1. N2 fixation rates correlated significantly with bacterial productivity and heterotrophic diazotrophs were identified from aphotic as well as photic depths. Dissolved free amino acid amendments to whole water from the GA enhanced bacterial productivity by 2–3.5 fold and N2 fixation rates by ~2-fold in samples collected from aphotic depths while in amendments to water from photic depths bacterial productivity increased 2–6 fold while N2 fixation rates increased by a factor of 2 to 4 illustrating that both BP and heterotrophic N2 fixation were carbon limited. Experimental manipulations of aphotic waters from the LB demonstrated a significant positive correlation between transparent exopolymeric particle (TEP) concentrations and N2 fixation rates. This suggests that sinking organic material and high carbon (C): nitrogen (N) micro-environments (such as TEP-based aggregates or marine snow) could support high heterotrophic N2 fixation rates in oxygenated surface waters and in the aphotic zones. Indeed, our calculations show that aphotic N2 fixation accounted for 37 to 75% of the total daily integrated N2 fixation rates at both locations in the Mediterranean and Red Seas with rates equal or greater to those measured from the photic layers. Moreover, our results indicate that that while N2 fixation may be limited in the surface waters, aphotic, pelagic N2 fixation may contribute significantly to new N inputs in other oligotrophic basins, yet it is currently not included in regional or global N budgets.

Transparent exopolymer particles (TEPs) are planktonic, organic microgels that are ubiquitous in aqueous environments. Increasing evidence indicates that TEPs play an active role in the process of aquatic biofilm formation. Frequently, TEPs are intensely colonized by bacteria and other microorganisms, thus serving as hot spots of intense microbial activity. We introduce the term "protobiofilm" to refer to TEPs with extensive microbial outgrowth and colonization. Such particles display most of the characteristics of developing biofilm, with the exception of being attached to a surface. In this study, coastal seawater was passed through custom-designed flow cells that enabled direct observation of TEPs and protobiofilm in the feedwater stream by bright-field and epifluorescence microscopy. Additionally, we could follow biofilm development on immersed surfaces inside the flow cells. Within minutes, we observed TEP and protobiofilm patches adhering to these surfaces. By 30 min, confocal laser-scanning microscopy (CLSM) revealed numerous patches of Con A and SYTO 9 staining structures covering the surfaces. Atomic force microscopy showed details of a thin, highly sticky, organic conditioning layer between these patches. Bright-field and epifluorescence microscopy and CLSM showed that biofilm development (observed until 24 h) was profoundly inhibited in flow cells with seawater prefiltered to remove most large TEPs and protobiofilm. We propose a revised paradigm for aquatic biofilm development that emphasizes the critical role of microgel particles such as TEPs and protobiofilm in facilitating this process. Recognition of the role of planktonic microgels in aquatic biofilm formation can have applied importance for the water industry.

We report N(2) fixation rates measured from two stations monitored monthly off the Mediterranean coast of Israel during 2006 and 2007, and along a transect from Israel to Crete in September 2008. Analyses of time-series data revealed expression of nifH genes from diazotrophs in nifH clusters I and II, including cyanobacterial bloom-formers Trichodesmium and diatom-Richelia intracellularis associations. However, nifH gene abundance and rates of N(2) fixation were very low in all size fractions measured (> 0.7 µm). Volumetric (15) N uptake ranged from below detection (∼ 36% of > 300 samples) to a high of 0.3 nmol N l(-1) d(-1) and did not vary distinctly with depth or season. Areal N(2) fixation averaged ∼ 1 to 4 µmol N m(-2) d(-1) and contributed only ∼ 1% and 2% of new production and ∼ 0.25% and 0.5% of primary production for the mixed (winter) and stratified (spring-fall) periods respectively. N(2) fixation rates along the 2008 east-west transect were also extremely low (0-0.04 nmol N l(-1) d(-1), integrated average 2.6 µmol N m(-2) d(-1) ) with 37% of samples below detection and no discernable difference between stations. We demonstrate that diazotrophy and N(2) fixation contribute only a minor amount of new N to the P impoverished eastern Mediterranean Sea.