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Sanz-Martin, M, Vernet M, Cape MR, Mesa E, Delgado-Huertas A, Reigstad M, Wassmann P, Duarte CM.  2019.  Relationship between carbon- and oxygen-based primary productivity in the Arctic Ocean, Svalbard Archipelago. Frontiers in Marine Science. 6   10.3389/fmars.2019.00468   AbstractWebsite

Phytoplankton contribute half of the primary production (PP) in the biosphere and are the major source of energy for the Arctic Ocean ecosystem. While PP measurements are therefore fundamental to our understanding of marine biogeochemical cycling, the extent to which current methods provide a definitive estimate of this process remains uncertain given differences in their underlying approaches, and assumptions. This is especially the case in the Arctic Ocean, a region of the planet undergoing rapid evolution as a result of climate change, yet where PP measurements are sparse. In this study, we compared three common methods for estimating PP in the European Arctic Ocean: (1) production of O-18-labeled oxygen (GPP-O-18), (2) changes in dissolved oxygen (GPP-DO), and (3) incorporation rates of C-14-labeled carbon into particulate organic carbon (C-14-POC) and into total organic carbon (C-14-TOC, the sum of dissolved and particulate organic carbon). Results show that PP rates derived using oxygen methods showed good agreement across season and were strongly positively correlated. While also strongly correlated, higher scatter associated with seasonal changes was observed between C-14-POC and C-14-TOC. The C-14-TOC-derived rates were, on average, approximately 50% of the oxygen-based estimates. However, the relationship between these estimates changed seasonally. In May, during a spring bloom of Phaeocystis sp., C-14-TOC was 52% and 50% of GPP-DO, and GPP-O-18, respectively, while in August, during post-bloom conditions dominated by flagellates, C-14-TOC was 125% of GPP-DO, and C-14-TOC was 175% of GPP-O-18. Varying relationship between C and O rates may be the result of varying importance of respiration, where C-based rates estimate net primary production (NPP) and O-based rates estimate gross primary production (GPP). However, uncertainty remains in this comparison, given differing assumptions of the methods and the photosynthetic quotients. The median O:C ratio of 4.75 in May is within the range of that observed for other regions of the world's ocean. However, the median O:C ratio for August is <1, lower than in any other reported region. Our results suggest further research is needed to estimate O:C in Arctic waters, and at different times of the seasonal cycle.

Paulsen, ML, Seuthe L, Reigstad M, Larsen A, Cape MR, Vernet M.  2018.  Asynchronous accumulation of organic carbon and nitrogen in the Atlantic gateway to the Arctic Ocean. Frontiers in Marine Science. 5   10.3389/fmars.2018.00416   AbstractWebsite

Nitrogen (N) is the main limiting nutrient for biological production in the Arctic Ocean. While dissolved inorganic N (DIN) is well studied, the substantial pool of N bound in organic matter (OM) and its bioavailability in the system is rarely considered. Covering a full annual cycle, we here follow N and carbon (C) content in particulate (P) and dissolved (D) OM within the Atlantic water inflow to the Arctic Ocean. While particulate organic carbon (POC), particulate organic nitrogen (PON), and dissolved organic carbon (DOC) accumulated in the surface waters from January to May, the dissolved organic nitrogen (DON)-pool decreased substantially (Delta - 50 mu g N L-1). The DON reduction was greater than the simultaneous reduction in DIN (Delta - 30 mu g N L-1), demonstrating that DON is a valuable N-source supporting the growing biomass. While the accumulating POM had a C/N ratio close to Redfield, the asynchronous accumulation of C and N in the dissolved pool resulted in a drastic increase in the C/N ratio of dissolved organic molecules (DOM) during the spring bloom. This is likely due to a combination of the reduction in DON, and a high release of carbon-rich sugars from phytoplankton, as 32% of the spring primary production (PP) was dissolved. Our findings thus caution calculations of particulate PP from DIN drawdown. During post-bloom the DON pool increased threefold due to an enhanced microbial processing of OM and reduced phytoplankton production. The light absorption spectra of DOM revealed high absorption within the UV range during spring bloom indicating DOM with low molecular weight in this period. The absorption of DOM was generally lower in the winter months than in spring and summer. Our results demonstrate that the change in ecosystem function (i.e., phytoplankton species and activity, bacterial activity and grazing) in different seasons is associated with strong changes in the C/N ratios and optical character of DOM and underpin the essential role of DON for the production cycle in the Arctic.

Cape, MR, Vernet M, Kahru M, Spreen G.  2014.  Polynya dynamics drive primary production in the Larsen A and B embayments following ice shelf collapse. Journal of Geophysical Research-Oceans. 119:572-594.   10.1002/2013jc009441   AbstractWebsite

The climate-driven collapses of the Larsen A and B ice shelves have opened up new regions of the coastal Antarctic to the influence of sea ice resulting in increases in seasonal primary production. In this study, passive microwave remote sensing of sea ice concentration and satellite imagery of ocean color are employed to quantify the magnitude of and variability in open water area and net primary productivity (NPP) in the Larsen embayments between 1997 and 2011. Numerical model output provides context to analyze atmospheric forcing on the coastal ocean. Following ice shelf disintegration the embayments function as coastal, sensible heat polynyas. The Larsen A and B are as productive as other Antarctic shelf regions, with seasonally averaged daily NPP rates reaching 1232 and 1127 mg C m(-2) d(-1) and annual rates reaching 200 and 184 g C m(-2) yr(-1), respectively. A persistent cross-shelf gradient in NPP is present with higher productivity rates offshore, contrasting with patterns observed along the West Antarctic Peninsula. Embayment productivity is intimately tied to sea ice dynamics, with large interannual variability in NPP rates driven by open water area and the timing of embayment opening. Opening of the embayment is linked to periods of positive Southern Annular Mode and stronger westerlies, which lead to the vertical deflection of warm, maritime air over the peninsula and down the leeward side causing increases in surface air temperature and wind velocity. High productivity in these new polynyas is likely to have ramifications for organic matter export and marine ecosystem evolution. Key Points Primary production and sea ice dynamics after ice shelf disintegration Larsen embayments function as productive coastal sensible heat polynyas High sea ice interannual variability affects total production

Ducklow, HW, Schofield O, Vernet M, Stammerjohn S, Erickson M.  2012.  Multiscale control of bacterial production by phytoplankton dynamics and sea ice along the western Antarctic Peninsula: A regional and decadal investigation. Journal of Marine Systems. 98-99:26-39.   10.1016/j.jmarsys.2012.03.003   AbstractWebsite

We present results on phytoplankton and bacterial production and related hydrographic properties collected on nine annual summer cruises along the western Antarctic Peninsula. This region is strongly influenced by inter-annual variations in the duration and extent of sea ice cover, necessitating a decade-scale study. Our study area transitions from a nearshore region influenced by summer runoff from glaciers to an offshore, slope region dominated by the Antarctic Circumpolar Current. The summer bacterial assemblage is the product of seasonal warming and freshening following spring sea ice retreat and the plankton succession occurring in that evolving water mass. Bacterial production rates averaged 20 mg C m(-2) d(-1) and were a low (5%) fraction of the primary production (PP). There was significant variation in BP between regions and years, reflecting the variability in sea ice, chlorophyll and PP. Leucine incorporation was significantly correlated (r(2) ranging 0.2-0.7, p<0.001) with both chlorophyll and PP across depths, regions and years indicating strong phytoplankton-bacteria coupling. Relationships with temperature were variable, including positive, negative and insignificant relationships (r(2)<0.2 for regressions with p<0.05). Bacterial production is regulated indirectly by variations in sea ice cover within regions and over years, setting the levels of phytoplankton biomass accumulation and PP rates; these in turn fuel BP, to which PP is coupled via direct release from phytoplankton or other less direct pathways. (C) 2012 Elsevier B.V. All rights reserved.

Vernet, M, Kozlowski WA, Yarmey LR, Lowe AT, Ross RM, Quetin LB, Fritsen CH.  2012.  Primary production throughout austral fall, during a time of decreasing daylength in the western Antarctic Peninsula. Marine Ecology-Progress Series. 452:45-61.   10.3354/meps09704   AbstractWebsite

Antarctic phytoplankton is characterized by a pronounced seasonality in abundance, driven mainly by changes in sunlight. We combined measurements and modeling to describe the influence of changing daylength on fall and winter phytoplankton production in coastal waters of the western Antarctic Peninsula (wAP) in 2001 and 2002. The model was parameterized with field observations from the Palmer Long-Term Ecological program in the wAP during summer and early fall and from the Southern Ocean Global Ecosystems Dynamics program fall and winter cruises to Marguerite Bay and shelf waters. Shorter daylength and a deepening of the mixed layer account for most of the decrease in primary production during March, April, and May. At this time, biomass decreases by an order of magnitude and remains low and constant until the end of August. An additional loss rate was added to the primary production model to fit output to observations. This loss rate, estimated at similar to 0.1 to 0.15 d(-1), is due to physical, chemical, and biological processes such as scavenging by sea ice, zooplankton grazing, cell lysis, and cell sedimentation, which are expected to be high at this time of year. Growth and loss rates of phytoplankton populations are similar on 1 March, with growth decreasing rapidly over time while the loss rates remain constant. By the beginning of winter (1 June), growth is low, with minimum rates in July and increasing towards September. During a period of diminishing food supply, preliminary estimates of grazing indicate that fall biomass could support existing macrozooplankton populations, but the timing and concentration of food supply is variable and expected to affect health of zooplankton as they enter the winter.

Vernet, M, Sines K, Chakos D, Cefarelli AO, Ekern L.  2011.  Impacts on phytoplankton dynamics by free-drifting icebergs in the NW Weddell Sea. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 58:1422-1435.   10.1016/j.dsr2.2010.11.022   AbstractWebsite

Glacier ice released to the oceans through iceberg formation has a complex effect on the surrounding ocean waters. We hypothesized that phytoplankton communities would differ in abundance, composition and production around or close to an iceberg. This paper tests the influence of individual icebergs on scales of meters to kilometers, observed through shipboard oceanographic sampling on March-April 2009. Surface waters (integrated 0-100 m depth, within the euphotic zone) sampled close to the iceberg C-18a ( <1 km) were characterized by lower temperatures, more dissolved nitrate, less total chlorophyll a (chla) concentration, less picoplankton ( <3 mu m) cell abundance, and higher transparency than surface conditions 18 km upstream. However, enrichment of large cells, identified as diatoms, was the basis of an active food chain. Upward velocity of meltwater and dissolved Fe concentrations in excess of 1-2 nM are expected to facilitate diatom specific growth. The presence of diatoms close to the iceberg C-18a and the higher variable fluorescence (Fv/Fm) indicated healthy cells, consistent with Antarctic waters rich in micronutrients. Furthermore, chla increased significantly 2 km around the iceberg and 10 days after the iceberg's passage. We hypothesize that the lower biomass next to the iceberg was due to high loss rates. Underwater melting is expected to dilute phytoplankton near the iceberg by entraining deep water or by introducing meltwater. In addition, high zooplankton biomass within 2 km of the iceberg, mainly Antarctic krill Euphausia superba and salps Salpa thompsonii, are expected to exert heavy grazing pressure on phytoplankton, the krill on large cells >10 mu m and the salps on smaller cells, 3-10 mu m. The iceberg's main influence in the austral fall is measured not so much by phytoplankton accumulation but by reactivation of the classic Antarctic food chain, facilitating diatom growth and sustaining high Antarctic krill populations. (C) 2011 Elsevier Ltd. All rights reserved.

Vernet, M, Martinson D, Iannuzzi R, Stammerjohn S, Kozlowski W, Sines K, Smith R, Garibotti I.  2008.  Primary production within the sea-ice zone west of the Antarctic Peninsula: I-Sea ice, summer mixed layer, and irradiance. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 55:2068-2085.   10.1016/j.dsr2.2008.05.021   AbstractWebsite

in shelf waters of the western Antarctic Peninsula (wAP), with abundant macro- and micronutrients, water-column stability has been suggested as the main factor controlling primary production; freshwater input from sea-ice melting stabilizes the upper water column by forming a shallow summer mixed layer. Retreating sea ice in the spring and summer thus defines the area of influence, the sea-ice zone (SIZ) and the marginal ice zone (MIZ). A 12-year time series (1995-2006) was analyzed to address two main questions: (1) what are the spatial and temporal patterns in primary production; and (2) to what extent and in what ways is primary production related to sea-ice dynamics. Data were collected on cruises performed during January of each year, at the height of the growth season, within the region bounded by 64 degrees S and 64 degrees W to the north and 68 degrees S and 66 degrees W to the south. Average daily integrated primary production varied by an order of magnitude, from similar to 250 to similar to 1100mg cm(-2) d(-1), with an average cruise primary production of 745 mg C m(-2) d(-1). A strong onshore-offshore gradient was evident along the shelf with higher production observed inshore. Inter-annual regional production varied by a factor of 7: maximum rates were measured in 2006 (1788 mg cm(-2) d-(1)) and minimum in 1999 (248 Mg C m(-2) d(-1)). The results support the hypothesis that primary production in the wAP shelf is related to sea-ice dynamics. To first order, shallower summer mixed-layer depths in the shelf correlated with late sea retreat and primary production. Principal component analysis showed that high primary production in January was associated with enhanced shelf production toward the coast and in the south, explaining 63% of the variability in space and time. This first mode captured the inter-annual variability in regional production. Temporal variability in primary production (time series of anomalies defined for each location) showed spatial dependence: higher primary production correlated with shallow mixed-layer depths only at mid-shelf; in coastal and offshore waters, primary production correlated with deeper mixed layers. Thus, coastal primary production can show a non-linear relationship with summer mixed layers. Under conditions of large biomass (> 20 mg ch1 a m(-3)) and shallow mixed-layer depth (e.g., 5 m) phytoplankton production becomes light limited. This limitation is reduced with a deepening of the summer mixed layer (e.g., 20m). Dominance of diatoms and the ability to adapt and photosynthesize at higher light levels characterized the large phytoplankton blooms. No significant regional trend in primary production was detected within the 12-year series. We conclude that the regional average primary production on the wAP shelf is associated with shallow summer mixed layers in conjunction with late sea-ice retreat. An opposite relationship is observed for the highest production rates in coastal waters, associated with large biomass, where a deepening of the summer mixed layer relieves light limitation. (C) 2008 Elsevier Ltd. All rights reserved.

Matrai, P, Vernet M, Wassmann P.  2007.  Relating temporal and spatial patterns of DMSP in the Barents Sea to phytoplankton biomass and productivity. Journal of Marine Systems. 67:83-101.   10.1016/j.jmarsys.2006.10.001   AbstractWebsite

Dimethylsulfoniopropionate (DMSP), produced by many marine phytoplankton, is the main precursor of the climate relevant gas dimethylsulfide (DMS). Currently, it is generally accepted that the relationship between DMSP and phytoplankton biomass (as chlorophyll a), while not representative of the absolute magnitude of the DMSP pool, is a good indicator of ecosystem structure. In this study we test the strength of the relationships between DMSP and various phytoplankton parameters in Arctic shelf waters of the Barents Sea. Our objective is to assess the predictive value that traditional phytoplankton carbon parameters have on DMSP. We discuss C:DMSP-S variability as a function of seasonality, water masses, grazing and nutrient limitation. For this purpose we analyze data from 5 cruises including winter, spring and summer conditions and across the seasonal ice zone at the time of the study. Highest phytoplankton DMSP concentration was usually measured at the ice edge. Marked seasonal variability was observed in phytoplankton carbon biomass and production but not necessarily in the particulate fraction of DMSP (DMSPp), resulting in seasonally varying C:DMSP-S. High winter DMSPp concentrations, when chlorophyll a and primary production were lowest and flagellates dominant, suggest a heterotrophic source. The production of extracellular carbon and the pool of dissolved DMSP (DMSPd) followed similar seasonal trends, with enhanced concentrations in spring, and we suggest that high dissolved primary production induced by nutrient limiting conditions resulted in high DMSPd concentrations. Mesoscale changes in total DMSP (particulate + dissolved) may be modeled from basin-wide total phytoplankton primary production (rather than from phytoplankton biomass) at seasonal and interannual scales. We conclude there is predictive power of DMSP concentrations in the Barents Sea based on seasonality, the position of the ice edge and the distribution of phytoplankton variables. (c) 2006 Elsevier B.V. All rights reserved.

Dierssen, HM, Vernet M, Smith RC.  2000.  Optimizing models for remotely estimating primary production in Antarctic coastal waters. Antarctic Science. 12:20-32. AbstractWebsite

Primary productivity and associated biogeochemical fluxes within the Southern Ocean are globally significant, sensitive to change and poorly known compared to temperate marine ecosystems. We present seasonal time series data of chlorophyll a, primary productivity and in-water irradiance measured in the coastal waters of the Western Antarctica Peninsula and build upon existing models to provide a more optimum parameterization for the estimation of primary productivity in Antarctic coastal waters. These and other data provide strong evidence that bio-optical characteristics and phytoplankton productivity in Antarctic waters an different from temperate waters. For these waters we show that over 60% of the variability in primary production can be explained by the surface chlorophyll a concentration alone, a characteristic, which lends itself to remote sensing models. if chlorophyll a concentrations are accurately determined, then the largest source of error 13-18) results from estimates of the photoadaptive variable (P-opt(B)). Further, the overall magnitude of P-opt(B) is low (median 1.09 mg C mg chl(-1) h(-1)) for these data compared to other regions and generally fits that expected for a cold water system. However, the variability of P-opt(B) over the course of a season (0.4 to 3 mg C mg chl(-1) h(-1)) is not consistently correlated with other possible environmental parameters, such as chlorophyll, sea surface temperature, incident irradiance, day length, salinity, or taxonomic composition. Nonetheless, by tuning a standard depth- integrated primary productivity model to fit representative P-opt(B) values and the relatively uniform chlorophyll-normalized production profile found in these waters, we can improve the model to account for approximately 72-73% variability in primary production both for our data as well as for independent historic Antarctic data.