Since the 1980s, the San Francisco Bay Delta ecosystem has experienced large declines in primary production. Hypothesized reasons for this decline include (1) suppression of nitrate (NO 3 − ) uptake, and thus phytoplankton growth, due to high concentrations of ammonium (NH 4 + ), and (2) wastewater NH 4 + -induced changes in phytoplankton community composition away from large-celled diatoms. These twin hypotheses implicate NH 4 + loading from the Sacramento Regional Wastewater Treatment Plant effluent outfall in explaining declines in primary production in the region. They have been controversial within the water resources management community and have stimulated a lengthy public scientific and regulatory debate. Here, in an effort to resolve this debate, we present results from a 48-h incubation experiment with surface water from both upstream and downstream of the Sacramento Regional Wastewater Treatment Plant effluent outfall, a major source of NH 4 + loading to the ecosystem. We amended this water with either NH 4 + , NO 3 − , or full wastewater effluent. All assays were incubated under high light (52% of incident irradiance) or low light (6% of incident irradiance). NO 3 − uptake rates were suppressed to near zero in all treatments with either added NH 4 + , added wastewater effluent, or high in situ NH 4 + concentrations. Yet, phytoplankton uniformly grew well on all dissolved inorganic nitrogen sources, including effluent and NH 4 + . Diatom species were the most abundant taxa at all stations, and diatom cell abundances increased at greater rates than all other taxa over the course of the experiment. Among all treatments, the light treatment had the greatest effects on chlorophyll a accumulation and phytoplankton growth rates. Our results suggest that high NH 4 + loading is not a driver of the lower productivity in the San Francisco Bay Delta. Although phytoplankton preferred NH 4 + to NO 3 − when both were available in our experiment, the form of dissolved inorganic nitrogen had no effect on growth rates or species composition.

The decline of sea-ice thickness, area, and volume due to the transition from multi-year to first-year sea ice has improved the under-ice light environment for pelagic Arctic ecosystems. One unexpected and direct consequence of this transition, the proliferation of under-ice phytoplankton blooms (UIBs), challenges the paradigm that waters beneath the ice pack harbor little planktonic life. Little is known about the diversity and spatial distribution of UIBs in the Arctic Ocean, or the environmental drivers behind their timing, magnitude, and taxonomic composition. Here, we compiled a unique and comprehensive dataset from seven major research projects in the Arctic Ocean (11 expeditions, covering the spring sea-ice-covered period to summer ice-free conditions) to identify the environmental drivers responsible for initiating and shaping the magnitude and assemblage structure of UIBs. The temporal dynamics behind UIB formation are related to the ways that snow and sea-ice conditions impact the under-ice light field. In particular, the onset of snowmelt significantly increased under-ice light availability (>0.1–0.2 mol photons m –2 d –1 ), marking the concomitant termination of the sea-ice algal bloom and initiation of UIBs. At the pan-Arctic scale, bloom magnitude (expressed as maximum chlorophyll a concentration) was predicted best by winter water Si(OH) 4 and PO 4 3– concentrations, as well as Si(OH) 4 :NO 3 – and PO 4 3– :NO 3 – drawdown ratios, but not NO 3 – concentration. Two main phytoplankton assemblages dominated UIBs (diatoms or Phaeocystis ), driven primarily by the winter nitrate:silicate (NO 3 – :Si(OH) 4 ) ratio and the under-ice light climate. Phaeocystis co-dominated in low Si(OH) 4 (i.e., NO 3 :Si(OH) 4 molar ratios >1) waters, while diatoms contributed the bulk of UIB biomass when Si(OH) 4 was high (i.e., NO 3 :Si(OH) 4 molar ratios <1). The implications of such differences in UIB composition could have important ramifications for Arctic biogeochemical cycles, and ultimately impact carbon flow to higher trophic levels and the deep ocean.

Sea ice is one the largest biomes on earth, yet it is poorly described by biogeochemical and climate models. In this paper, published and unpublished data on sympagic (ice-associated) algal biodiversity and productivity have been compiled from more than 300 sea-ice cores and organized into a systematic framework. Significant patterns in microalgal community structure emerged from this framework. Autotrophic flagellates characterize surface communities, interior communities consist of mixed microalgal populations and pennate diatoms dominate bottom communities. There is overlap between landfast and pack-ice communities, which supports the hypothesis that sympagic microalgae originate from the pelagic environment. Distribution in the Arctic is sometimes quite different compared to the Antarctic. This difference may be related to the time of sampling or lack of dedicated studies. Seasonality has a significant impact on species distribution, with a potentially greater role for flagellates and centric diatoms in early spring. The role of sea-ice algae in seeding pelagic blooms remains uncertain. Photosynthesis in sea ice is mainly controlled by environmental factors on a small scale and therefore cannot be linked to specific ice types. Overall, sea-ice communities show a high capacity for photoacclimation but low maximum productivity compared to pelagic phytoplankton. Low carbon assimilation rates probably result from adaptation to extreme conditions of reduced light and temperature in winter. We hypothesize that in the near future, bottom communities will develop earlier in the season and develop more biomass over a shorter period of time as light penetration increases due to the thinning of sea ice. The Arctic is already witnessing changes. The shift forward in time of the algal bloom can result in a mismatch in trophic relations, but the biogeochemical consequences are still hard to predict. With this paper we provide a number of parameters required to improve the reliability of sea-ice biogeochemical models.

Antarctic pack ice is inhabited by a diverse and active microbial community reliant on nutrients for growth. Seeking patterns and overlooked processes, we performed a large-scale compilation of macro-nutrient data (hereafter termed nutrients) in Antarctic pack ice (306 ice-cores collected from 19 research cruises). Dissolved inorganic nitrogen and silicic acid concentrations change with time, as expected from a seasonally productive ecosystem. In winter, salinity-normalized nitrate and silicic acid concentrations (C*) in sea ice are close to seawater concentrations (C w ), indicating little or no biological activity. In spring, nitrate and silicic acid concentrations become partially depleted with respect to seawater (C* < C w ), commensurate with the seasonal build-up of ice microalgae promoted by increased insolation. Stronger and earlier nitrate than silicic acid consumption suggests that a significant fraction of the primary productivity in sea ice is sustained by flagellates. By both consuming and producing ammonium and nitrite, the microbial community maintains these nutrients at relatively low concentrations in spring. With the decrease in insolation beginning in late summer, dissolved inorganic nitrogen and silicic acid concentrations increase, indicating imbalance between their production (increasing or unchanged) and consumption (decreasing) in sea ice. Unlike the depleted concentrations of both nitrate and silicic acid from spring to summer, phosphate accumulates in sea ice (C* > C w ). The phosphate excess could be explained by a greater allocation to phosphorus-rich biomolecules during ice algal blooms coupled with convective loss of excess dissolved nitrogen, preferential remineralization of phosphorus, and/or phosphate adsorption onto metal-organic complexes. Ammonium also appears to be efficiently adsorbed onto organic matter, with likely consequences to nitrogen mobility and availability. This dataset supports the view that the sea ice microbial community is highly efficient at processing nutrients but with a dynamic quite different from that in oceanic surface waters calling for focused future investigations.

To evaluate what drives phytoplankton photosynthesis rates in the Amundsen Sea Polynya (ASP), Antarctica, during the spring bloom, we studied phytoplankton biomass, photosynthesis rates, and water column productivity during a bloom of Phaeocystis antarctica (Haptophyceae) and tested effects of iron (Fe) and light availability on these parameters in bioassay experiments in deck incubators. Phytoplankton biomass and productivity were highest (20 µg chlorophyll a L −1 and 6.5 g C m −2 d −1 ) in the central ASP where sea ice melt water and surface warming enhanced stratification, reducing mixed layer depth and increasing light availability. In contrast, maximum photosynthesis rate ( P * max ), initial light-limited slope of the photosynthesis–irradiance curve (α * ), and maximum photochemical efficiency of photosystem II ( F v /F m ) were highest in the southern ASP near the potential Fe sources of the Dotson and Getz ice shelves. In the central ASP, P * max , α * , and F v /F m were all lower. Fe addition increased phytoplankton growth rates in three of twelve incubations, and at a significant level when all experiments were analyzed together, indicating Fe availability may be rate-limiting for phytoplankton growth in several regions of the ASP early in the season during build-up of the spring bloom. Moreover, Fe addition increased P * max , α * , and F v /F m in almost all experiments when compared to unamended controls. Incubation under high light also increased P * max , but decreased F v /F m and α * when compared to low light incubation. These results indicate that the lower values for P * max , α * , and F v /F m in the central ASP, compared to regions close to the ice shelves, are constrained by lower Fe availability rather than light availability. Our study suggests that higher Fe availability (e.g., from higher melt rates of ice shelves) would increase photosynthesis rates in the central ASP and potentially increase water column productivity 1.7-fold, making the ASP even more productive than it is today.

Sea ice covers approximately 5% of the ocean surface and is one of the most extensive ecosystems on the planet. The microbial communities that live in sea ice represent an important food source for numerous organisms at a time of year when phytoplankton in the water column are scarce. Here we describe the distributions and physiology of sea ice microalgae in the poorly studied Amundsen Sea sector of the Southern Ocean. Microalgal biomass was relatively high in sea ice in the Amundsen Sea, due primarily to well developed surface communities that would have been replenished with nutrients during seawater flooding of the surface as a result of heavy snow accumulation. Elevated biomass was also occasionally observed in slush, interior, and bottom ice microhabitats throughout the region. Sea ice microalgal photophysiology appeared to be controlled by the availability of both light and nutrients. Surface communities used an active xanthophyll cycle and effective pigment sunscreens to protect themselves from harmful ultraviolet and visible radiation. Acclimation to low light microhabitats in sea ice was facilitated by enhanced pigment content per cell, greater photosynthetic accessory pigments, and increased photosynthetic efficiency. Photoacclimation was especially effective in the bottom ice community, where ready access to nutrients would have allowed ice microalgae to synthesize a more efficient photosynthetic apparatus. Surprisingly, the pigment-detected prymnesiophyte Phaeocystis antarctica was an important component of surface communities (slush and surface ponds) where its acclimation to high light may precondition it to seed phytoplankton blooms after the sea ice melts in spring.