Nicole L. Goebel

Abundance and distribution of major groups of diazotrophic cyanobacteria and their potential contribution to N2 fixation in the tropical Atlantic Ocean.

The abundances of six N₂-fixing cyanobacterial phylotypes were profiled at 22 stations across the tropical Atlantic Ocean during June 2006, and used to model the contribution of the diazotrophs to N₂ fixation. Diazotroph abundances were measured by targeting the nifH gene of Trichodesmium, unicellular groups A, B, C (UCYN-A, UCYN-B and UCYN-C), and diatom-cyanobiont symbioses Hemiaulus-Richelia, Rhizosolenia-Richelia and Chaetoceros-Calothrix. West to east gradients in temperature, salinity and nutrients [NO₃⁻ + NO₂⁻, PO₄³⁻, Si(OH)₄] showed the influence of the Amazon River plume and its effect on the distributions of the diazotrophs. Trichodesmium accounted for more than 93% of all nifH genes detected, dominated the warmer waters of the western Atlantic, and was the only diazotroph detected at the equatorial upwelling station. UCYN-A was the next most abundant (> 5% of all nifH genes) and dominated the cooler waters of the eastern Atlantic near the Cape Verde Islands. UCYN-C was found at a single depth (200 m) of high salinity and low temperature and nutrients, whereas UCYN-B cells were widespread but in very low abundance (6.1 × 10¹ ± 4.6 × 10² gene copies l⁻¹). The diatom-cyanobionts were observed primarily in the western Atlantic within or near the high Si(OH)₄ input of the Amazon River plume. Overall, highest diazotroph abundances were observed at the surface and declined with depth, except for some subsurface peaks in Trichodesmium, UCYN-B and UCYN-A. Modelled contributions of Trichodesmium, UCYN-B and UCYN-A to total N₂ fixation suggested that Trichodesmium had the largest input, except for the potential of UCYN-A at the Cape Verde Islands.

GROWTH AND CARBON CONTENT OF THREE DIFFERENT-SIZED DIAZOTROPHIC CYANOBACTERIA OBSERVED IN THE SUBTROPICAL NORTH PACIFIC1

To develop tools for modeling diazotrophic growth in the open ocean, we determined the maximum growth rate and carbon content for three diazotrophic cyanobacteria commonly observed at Station ALOHA (A Long‐term Oligotrophic Habitat Assessment) in the subtropical North Pacific: filamentous nonheterocyst‐forming Trichodesmium and unicellular Groups A and B. Growth‐irradiance responses of Trichodesmium erythraeum Ehrenb. strain IMS101 and Crocosphaera watsonii J. Waterbury strain WH8501 were measured in the laboratory. No significant differences were detected between their fitted parameters (±CI) for maximum growth rate (0.51 ± 0.09 vs. 0.49 ± 0.17 d−1), half‐light saturation (73 ± 29 vs. 66 ± 37 μmol quanta · m−2 · s−1), and photoinhibition (0 and 0.00043 ± 0.00087 [μmol quanta · m−2 · s−1]−1). Maximum growth rates and carbon contents of Trichodesmium and Crocosphaera cultures conformed to published allometric relationships, demonstrating that these relationships apply to oceanic diazotrophic microorganisms. This agreement promoted the use of allometric models to approximate unknown parameters of maximum growth rate (0.77 d−1) and carbon content (480 fg C · μm−3) for the uncultivated, unicellular Group A cyanobacteria. The size of Group A was characterized from samples from the North Pacific Ocean using fluorescence‐activated cell sorting and real‐time quantitative PCR techniques. Knowledge of growth and carbon content properties of these organisms facilitates the incorporation of different types of cyanobacteria in modeling efforts aimed at assessing the relative importance of filamentous and unicellular diazotrophs to carbon and nitrogen cycling in the open ocean.

Modeled contributions of three types of diazotrophs to nitrogen fixation at Station ALOHA.

A diagnostic model based on biomass and growth was used to assess the relative contributions of filamentous nonheterocystous Trichodesmium and unicellular cyanobacteria, termed Groups A and B, to nitrogen fixation at the North Pacific Station ALOHA over a 2-year period. Average (and 95% confidence interval, CI) annual rates of modeled monthly values for Trichodesmium, Group B and Group A were 92 (52), 14 (4) and 12 (8) mmol N per m2 per year, respectively. The fractional contribution to modeled instantaneous nitrogen fixation by each diazotroph fluctuated on interannual, seasonal and shorter time scales. Trichodesmium fixed substantially more nitrogen in year 1 (162) than year 2 (12). Group B fixed almost two times more nitrogen in year 1 (17) than year 2 (9). In contrast, Group A fixed two times more nitrogen in year 2 (16) than year 1 (8). When including uncertainties in our estimates using the bootstrap approach, the range of unicellular nitrogen fixation extended from 10% to 68% of the total annual rate of nitrogen fixation for all three diazotrophs. Furthermore, on a seasonal basis, the model demonstrated that unicellular diazotrophs fixed the majority (51%–97%) of nitrogen during winter and spring, whereas Trichodesmium dominated nitrogen fixation during summer and autumn (60%–96%). Sensitivity of the modeled rates to some parameters suggests that this unique attempt to quantify relative rates of nitrogen fixation by different diazotrophs may need to be reevaluated as additional information on cell size, variability in biomass and C:N, and growth characteristics of the different cyanobacterial diazotrophs become available.

ISOLATION OF CALOTHRIX RHIZOSOLENIAE (CYANOBACTERIA) STRAIN SC01 FROM CHAETOCEROS (BACILLARIOPHYTA) SPP. DIATOMS OF THE SUBTROPICAL NORTH PACIFIC OCEAN1

In November 2004, Chaetoceros spp. (diatom) cells were collected from 5 m at Station ALOHA (22º45′ N, 158º0′ W) in the subtropical North Pacific Ocean. Attached to the spines of several Chaetoceros spp. were symbiotic heterocystous cyanobacterial cells, identified as Calothrix rhizosoleniae Lemmerm. The symbiotic diatom cells were handpicked and placed in N‐deplete media. The host cells have been absent since isolation, and the cyanobacterial symbiont has persisted in a free‐living state in two different liquid and solid media: one made from artificial seawater, and another with natural seawater. All media are devoid of fixed inorganic nitrogen. Trichome length varied from three to four vegetative cells when growing colonially, to >10 vegetative cells when growing as single trichomes. The DNA and amino acid sequences of the narB, rbcL, and rnpB genes are most similar to those from other heterocystous cyanobacteria (Anabaena and Fischerella). Acetylene reduction (AR) assays were run in conjunction with multiplexing quantitative reverse transcription–PCR (qRT–PCR) assays. Gene transcription for rbcL and nifH was high, coincident with maximum AR, and occurred in the middle of the photoperiod. Eight irradiance curves of nitrogenase activity at varying biomass concentrations showed evidence of photoinhibition at high light intensities. Here, we report on the genetic identification and photophysiology for the first symbiotic isolate, Ca. rhizosoleniae SC01, of an open‐ocean diatom (Chaetoceros).

Primary production in Long Island sound

Daily and annual integrated rates of primary productivity and community respiration were calculated using physiological parameters measured in oxygen-based photosynthesis-irradiance (P-I) incubations at 8 stations throughout central and western Long Island Sound (cwLIS) during the summer and autumn of 2002 and 2003 and the late spring of 2003. Each calculation takes into account actual variations in incident irradiance over the day and underwater irradiance and standing stock with depth. Annual peak rates, ±95% confidence interval of propagated uncertainty in each measurement, of gross primary production (GPP, 1,730±610 mmol O2 m−2 d−1), community respiration (Rc, 1,660±270 mmol O2 m−2 d−1), and net community production (NCP, 1,160±1,100 mmol O2 m−2 d−1) occurred during summer at the western end of the Sound. Lowest rates of GPP (4±11 mmol O2 m−2 d−1), Rc (−50±300 mmol O2 m−2 d−1), and NCP (−1,250±270 mmol O2 m−2 d−1) occurred during late autumn-early winter at the outer sampled stations. These large ranges in rates of GPP, Rc, and NCP throughout the photic zone of cwLIS are attributed to seasonal and spatial variability. Algal respiration (Ra) was estimated to consume an average of 5% to 52% of GPP, using a literature-based ratio of Ra:Rc. From this range, we established that the estimated Ra accounts for approximately half of GPP, and was used to estimate daily net primary production (NPP), which ranged from 2 to 870 mmol O2 m−2 d−1 throughout cwLIS during the study. Annual NPP averaged 40±8 mol O2 m−2 yr−1 for all sampled stations, which more than doubled along the main axis of the Sound, from 32±14 mol O2 m−2 yr−1 at an eastern station to 82±25 mol O2 m−2 yr−1 at the western-most station. These spatial gradients in productivity parallel nitrogen loads along the main axis of the Sound. Daily integrals of productivity were used to test and formulate a simple, robust biomass-light model for the prediction of phytoplankton production in Long Island Sound, and the slope of the relationship was consistent with reports for other systems.

Air-sea CO2 fluxes in the California Current: Impacts of model resolution and coastal topography

The present study uses a suite of coupled physical-biogeochemical model simulations at 1/3°, 1/10°, and 1/30° to assess the impact of horizontal resolution on air-sea CO2 fluxes in the California Current System (CCS), a relevant issue for downscaling between coarser resolution global climate models and higher resolution regional models. The results demonstrate that horizontal resolution is important to reproduce the sharp transition between near-shore outgassing and offshore absorption, as well as to resolve regions of enhanced near-shore outgassing in the lee of capes. The width of the outgassing region is overestimated when horizontal resolution is not eddy resolving (i.e., 1/3°) but becomes more dependent on shelf topography for eddy-resolving simulations (i.e., 1/10° and 1/30°). Enhanced near-shore outgassing is associated with a local increase in wind-driven upwelling in the lee of capes (i.e., expansion fans), meaning that sufficient horizontal resolution is needed both in the ocean circulation model and in the wind field forcing the model. From a global carbon budget perspective, the model indicates that biological production generates sufficient absorption within a few hundred kilometers of the coast to offset near-shore outgassing, which is consistent with the notion that midlatitude eastern boundary current upwelling systems act both as a sink and source for atmospheric CO2. Based on the 1/30° solution, the CCS between 35 and 45 N and out to 600 km offshore is a net carbon sink of approximately 6 TgC yr−1, with the 1/10° solution underestimating this value by less than 10% and the 1/3° solution by a factor of 3.

Modeled diversity effects on microbial ecosystem functions of primary production, nutrient uptake, and remineralization.

Ecosystem-wide primary productivity generally increases with primary producer diversity, emphasizing the importance of diversity for ecosystem function. However, most studies that demonstrate this positive relationship have focused on terrestrial and aquatic benthic systems, with little attention to the diverse marine pelagic primary producers that play an important role in regulating global climate. Here we show how phytoplankton biodiversity enhances overall marine ecosystem primary productivity and other ecosystem functions using a self-organizing ecosystem model. Diversity manipulation numerical experiments reveal positive, asymptotically saturating relationships between ecosystem-wide phytoplankton diversity and functions of productivity, nutrient uptake, remineralization, and diversity metrics used to identify mechanisms shaping these relationships. Increase in productivity with increasing diversity improves modeled ecosystem stability and model robustness and leads to productivity rates that exceed expected yields primarily through niche complementarity and facilitative interactions between coexisting phytoplankton types; the composition of traits in assemblages determines the magnitude of complementarity and selection effects. While findings based on these aggregate measures of diversity effects parallel those from the majority of experimental outcomes of terrestrial and benthic biodiversity-ecosystem function studies, we combine analyses of community diversity effects and investigations of the underlying interactions among phytoplankton types to demonstrate how an increase in recycled production of non-diatoms through an increase in new production of diatoms drives this diversity-cosystem function response. We demonstrate the important role that facilitation plays in the modeled marine plankton and how this facilitative interaction could amplify future climate-driven changes in ocean ecosystem productivity.