A. B. Demidov

Spatial Variability of the Primary Production and Chlorophyll a Concentration in the Drake Passage in the Austral Spring

The spatial distribution of the primary production (PP) and the chlorophyll a concentration (Chl) were investigated during two research cruises in the Drake Passage area in October–November of 2007 and 2008. The algorithm evaluating the integral PP (PPint) for the water column in this area was developed based on the data on the surface chlorophyll (Chls) and the incident solar irradiance obtained in 2004–2008 in the Atlantic Sector of the Southern Ocean. The results obtained both by the experimental and model approaches suggested that the Polar Front (PF) region of the Drake Passage was characterized by low values of both the PPint (<100 mg C/m2 per day) and Chls (0.08–0.20 mg/m3) in October–November. Low values of the Chls and relatively high phaeophytine a concentrations indicated the winter succession state of the phytoplankton community in the Antarctic Ocean and the southern Polar Frontal Zone (PFZ). The seasonal warming of the surface water layers and the developing pycnocline resulted in a phytoplankton bloom and a Chls concentration of more than 1 mg/m3 in mid-November in this area and the Subantarctic waters.

Vertical distribution of primary production and chlorophyll a in the Kara Sea

On the basis of data obtained during three ecosystem expeditions in the Kara Sea, the vertical variability of the primary production (PP) and chlorophyll a (Chl) in autumn was studied. The Chl maximum was detected mainly on the surface (Chl0). A homogenous Chl distribution in the euphotic layer (1% photo-synthetically available radiation) and a nearly linear decrease in the Chl concentration below this layer were observed in waters with Chl0 values of 0.1–0.5 mg/m3. In waters with Chl0 > 0.5 mg/m3, the Chl concentration in the studied layer decreased linearly or exponentially. The subsurface Chl maximum (SCM) was registered weekly and was detected mostly in waters with a Chl0 content of 0.1–0.5 mg/m3. The SCM formation in the Kara Sea was consistent with the general patterns for the World Ocean. Water-column stability, the content of biogenic elements, and the level of subsurface irradiance had an approximately equal effect on SCM formation. The contribution of the SCM to the depth-integrated PP varied from 1 to 27%. The parameterization of vertical profiles of Chl was performed in order to be used in depth-integrated PP models. The Chl maximum on the surface and the negligible SCM facilitate the estimation of depth-integrated PP on the basis of satellite data and the use of vertical-resolution models.

The structure of phytoplankton communities in the eastern part of the Laptev Sea

Studies have been performed on a transect along 130°30′ E from the Lena River delta (71°60′ N) to the continental slope and adjacent deepwater area (78°22′ N) of the Laptev Sea in September 2015. The structure of phytoplankton communities has distinct latitudinal zoning. The southern part of the shelf (southward of 73°10′ N), the most desalinated by riverine discharge, houses a phytoplankton community with a biomass of 175–840 mg/m2, domination of freshwater Aulacoseira diatoms, and significant contribution of green algae (both in abundance and biomass). The northern border for the distribution range of the southern complex of phytoplankton species lies between the 8 and 18 psu isohalines (~73°10′ N). The continental slope and deepwater areas of the Laptev Sea (north of 77°30′ N), with a salinity of >27 psu in the upper mixed layer, are populated by the community prevalently composed of Chaetoceros and Rhizosolenia diatoms, very abundant in the Arctic, and dinoflagellates. The phytoplankton number in this area fall in the range of 430–1100 × 106 cell/m2, and the biomass, in the range of 3600 mg/m2. A moderate desalinating impact of the Lena River discharge is observed in the outer shelf area between 73°20′ and 77°30′ N; the salinity in the upper mixed layer is 18–24 psu. The phytocenosis in this area has a mosaic spatial structure with between-station variation in the shares of different alga groups in the community, cell number of 117–1200 × 106 cells/m2, and a biomass of 1600–3600 mg/m2. As is shown, local inflow of “fresh” nutrients to the euphotic layer in the fall season leads to mass growth of diatoms.

Meridional asymmetric distribution of the primary production in the Atlantic Sector of the Southern Ocean in the austral spring and summer

The spatial distribution of the phytoplankton productivity was investigated in the Atlantic Sector of the Southern Ocean in the austral summer of 2009–2010 and the spring of 2010 on the basis of field measurements. In October–November, the average integrated primary production and the concentration of surface and integrated (photosynthetic layer) chlorophyll “a” in the Subantarctic waters of the Drake Passage exceeded the similar values along the Greenwich meridian by 3, 2, and 1.5 times, respectively. Similar primary production was observed in December–January for the water column in the eastern and western sectors of the studied area. The chlorophyll “a” concentration in the surface water layer was higher by 1.7 times for the Greenwich meridian area compared to the deep layers, but the concentrations of this pigment in the deeper layers did not differ. During the spring, the average primary production in the water column (the chlorophyll “a” concentration in the surface and in the photosynthetic layer) differed in the Drake Passage and along the Greenwich meridian by 2.3, 1.6, and 1.7 times, respectively. The opposite pattern was observed during the summer period, when the parameters listed above were higher for the Greenwich meridian area by 1.9, 2.5, and 1.7 times, respectively. Therefore, the western Antarctic areas in the spring are characterized by higher production than the eastern ones, and an opposite pattern is observed in the summer. The possible reasons for the meridional zonation of the chlorophyll “a” and the primary production are discussed in regard to different seasons.

Seasonal variability of the surface chlorophyll “a” in the Drake Passage

The seasonal variability of the surface chlorophyll “a” (Chl-s) was studied for five different hydrological areas in the Drake Passage. The data were collected both in the field (December 2001–March 2002, and November 2007) and by satellite observations. One maximum of Chl-s was registered for the area northward of the Antarctic Polar Front in November 2007. This maximum moves southwards to the Antarctic and Continental Antarctic regions in December and January, respectively. The major factors affecting the phytoplankton growth were analyzed, namely, the decrease of the mixed water layer’s depth due to jogging during the austral late spring and summer and seasonal water temperature increase. The comparison of the field and satellite data allows us to conclude that the standard OC4v4 algorithm usually underreports the Chl-s concentration when it exceed 0.2 mg m−3.

Phytoplankton production characteristics in the Eastern Atlantic and Atlantic sector of the Southern Ocean in October–November 2004

Production parameters of surface phytoplankton were measured along three transects: La Manche-Cape Town (I); Cape Town-54°S (II); 0°-49°W (along 54°S) (III). The Canary upwelling waters were most productive along transect I, where the surface chlorophyll a (Chl0) and the surface primary production (PP0) were as high as 4.3 mg/m3 and 173 mg C/m3 per day, respectively. Mosaic patterns in the distribution of these parameters were recorded in the northeastern regions of the South Subtropical Anticyclonic Gyre (Chl0 = 0.03–0.35 mg/m3; PP0 = 1.6–12.6 mg C/m3 per day). Along transect II, the average twofold southward increase in Chl0 (from 0.2 to 0.4 mg/m3) and the concurrent decline of the phytoplankton assimilation activity ( AN0) resulted in deviations from typical latitudinal changes inPP0. At most sites, PP0 values varied between 6 and 15 mg C/m3 per day. Negligible changes in Chl0 (0.36–0.85 mg/m3), PP0 (8–19 mg C/m3 per day), and AN0 (0.7–1.6 mg C/mg chl a per hour) were registered for the oceanic waters along transect III. Along all the transects, PP0 depended on Chl0 to a greater extent than AN0. The values of the latter parameter were largely determined by the water temperature and showed a slight correlation with the insolation. Along transect II, the integrated primary production (PPint) and the layer-integrated chlorophyll a in the upper 200 m (Chl0–200) generally varied from 180 to 360 mg C/m2 per day and from 30 to 70 mg/m2, respectively. In the Polar Front region, an increase in Chl0–200, PPint, Chl0, and PP0 up to respective values of 190 mg/m2, 520 mg C/m2 per day, 1.2 mg/m3, and 32 mg C/m3 per day was observed. A comparison of the water column (0–100 m) stability with the vertical distribution of the primary production and chlorophyll content along transect II implies that the thick (>100 m) upper mixed layer (UML) formed in response to the strong water cooling and wind forcing was largely responsible for the limited primary production in the Subantarctic and Antarctic regions. The large UML thickness resulted in an intense removal of plant cells from the photosynthetic layer and light starvation of a significant (up to 60%) part of the phytoplankton community.

Seasonal variation of the satellite-derived phytoplankton primary production in the Kara Sea

Seasonal variation of the integrated primary production (IPP) and surface chlorophyll (Chl0) in different regions of the Kara Sea was studied from satellite data obtained by the MODIS-Aqua colour scanner and averaged for 2003–2015. The minimum variation of Chl0 concentration during the growing season (from April to October) was 1.5 times in southwestern region and 2 times in the northern region of the sea. It was found that the Chl0 concentration increased slightly in all regions by the end of the growing season. The maximum IPP value recorded in June coincided with the peak level of photosynthetically active radiation (PAR) and maximum river discharge. The IPP value varied in a wider range compared with the Chl0 concentration. The ratio of the maximum and minimum monthly average IPP values varied from 8.9 times in Southwestern region to 11.7 times in the Northern region of the sea. The average increase in the Chl0 concentration was 1.7 times (from 0.78 mg/m3 in April to 1.29 mg/m3 in October). The IPP value varied by a factor of 10.7 (from 26 mg C/m2 per day in October to 279 mg C/m2 per day in June). The article also discusses the influence of water column stratification, the concentration of nutrients, the PAR level, and river discharge on the seasonal IPP dynamics in the Kara Sea.

Peculiarities of the primary production process in the Kara Sea at the end of the vegetation season

Research was implemented from September 15 through October 4, 2011 in the Kara Sea along transects located southeastwards Novaya Zemlya, in the St. Anna Trough, the Yenisei River estuary, and the adjacent shelf. The concentration of chlorophyll a was the highest in the photic zone (0.05–2.30 mg/m3, on average, 0.80 ± 0.37 mg/m3). The maximal concentration of Chl a at most of the stations located in the water layer of 7–30 m. Integral primary production in the water column varied from 3.0 to 151.0 mg C/m2 per day, on average, 37.2 ± 36.6 mg C/m2 per day. The maximal rate of primary production at most of the stations has been observed for the surface layer of the water column. Within the upper mixed water layer, relative primary production was from 31 to 100% (on average, 77 ± 20%). The most productive zone was the waters along Yenisei transect. In the estuary and at the adjacent shelf, primary production was 50 mg C/m2 per day, exceeding the range observed for other areas by 1.5–2.0 times. The concentrations of silica and nitrogen together with light regime and water temperature were the major limiting factors affecting the primary production rate in the Kara Sea in autumn.

Evaluation of the influence of abiotic and biotic factors on primary production in the Kara Sea in autumn

A regression analysis of the parameters of primary production versus environmental factors was performed on the basis of data of three complex expeditions performed in the Kara Sea in September to October 1993, 2007, and 2011. The analysis of the dependence of the depth-integrated primary production (PPint) on the value of surface chlorophyll a (Chl0) and assimilation activity (ANm) showed that only 12% of the variability of the integrated PPint was determined by the variability of Chl0, whereas the correlation between PPint and ANm was strong (R2 = 0.635). Thus, in autumn, PPint values in the Kara Sea depended primarily on the activity of phytoplankton assimilation. At the end of the vegetative season, high (close to or above 1 mg/m3) Chl0 values did not reflect phytoplankton production within the entire photosynthetic layer, where organic matter was synthesized at a low rate. In turn, PPint and ANm depended primarily on the intensity of insolation and was weakly related to the content of dissolved forms of nitrogen and phosphorus. In autumn, at the end of the vegetative season, insolation apparently is the main factor in the determination of the formation of primary production in the Kara Sea.

Verification of Kara Sea primary production models with field and satellite observations

The depth-integrated model (Ψ-Mod) and depth-resolved Kara Sea model (KDRSM) of primary production in the water column were verified using field (2013–2015) and satellite (MODIS-Aqua scanner, 2007, 2011, 2013–2015) observations. The KSDRM and Ψ-Mod over- or underestimate the values of integrated primary production (IPP) in autumn by a factor of 2 and 2.5 with shipboard data as input parameters; the rootmean-square difference (RMSD) was 0.29 and 0.39, respectively. In summer, the efficiency of Ψ-Mod decreased by a factor of 1.5 (RMSD = 0.57), while the predictive capacity of the KSDRM remained the same (RMSD = 0.31). In the Laptev Sea in autumn, the KSDRM performed better than Ψ-Mod (the RMSD was 0.24 and 0.41, respectively). There was no sufficient decrease in the predictive skill of either algorithm when MODIS-Aqua data were used as input parameters. Thus, Ψ-Mod, being a simple and precise algorithm, can be recommended for evaluating the annual IPP in the Kara Sea and for studying its long-term variability using satellite data.