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Not Et Al 2005.Pdf Limnol. Oceanogr., 50(5), 2005, 1677±1686 q 2005, by the American Society of Limnology and Oceanography, Inc. Late summer community composition and abundance of photosynthetic picoeukaryotes in Norwegian and Barents Seas Fabrice Not1 Station Biologique, UMR7144 Centre National de la Recherche Scienti®que (CNRS), Institut National des Sciences de l'Univers (INSU) et Universite Pierre et Marie Curie, Place George Teissier, 29680 Roscoff, France Ramon Massana and Mikel Latasa Institut de CieÁncies del Mar, Consejo Superior de Investigaciones Cientõ®cas (CSIC), Passeig MarõÂtim de la Barceloneta 37-49, 08003 Barcelona, Spain Dominique Marie and CeÂline Colson Station Biologique, UMR7144 CNRS, INSU et Universite Pierre et Marie Curie, Place George Teissier, 29680 Roscoff, France Wenche Eikrem Department of Biology, University of Oslo, P.O. Box 1069 Blindern, 0316 Oslo, Norway Carlos PedroÂs-Alio Institut de CieÁncies del Mar, CSIC, Passeig MarõÂtim de la Barceloneta 37-49, 08003 Barcelona, Spain Daniel Vaulot and Nathalie Simon Station Biologique, UMR7144 CNRS, INSU et Universite Pierre et Marie Curie, Place George Teissier, 29680 Roscoff, France Abstract We investigated marine picoeukaryotic diversity (cells ,3 mm) in samples collected in late summer 2002 at the boundary between the Norwegian, Greenland, and Barents Seas. The two main Arctic and Atlantic water masses in this region are separated by the polar front. We combined total counts of picoeukaryotes assemblages by ¯ow cytometry and epi¯uorescence microscopy with taxa detection by tyramide signal ampli®cation±¯uorescent in situ hybridization (TSA- FISH) and high performance liquid chromatography (HPLC) pigment analyses. The picoeukaryotic community was primarily composed of photoautotrophs (75% of the cells on average). Members of the division Chlorophyta, in particular the species Micromonas pusilla (Butcher) Manton and Parke, were the major components in truly Arctic waters (32% of the picoeukaryotes, maximum 3,200 cells ml21). M. pusilla was also well represented in coastal waters and at the polar front (25% of the picoeukaryotes, maximum 9,100 cells ml21). Haptophyta were prominent in more typical Atlantic waters (up to 35% of the picoeukaryotes, maximum 4,500 cells ml21). Quanti®cation of haptophyte biomass by HPLC pigment analyses and CHEMTAX, and haptophyte abundances by TSA-FISH were in good agreement. This con®rms previous studies, which suggested that M. pusilla is a dominant contributor of picoeukaryotic communities in both coastal and nutrient rich environments, whereas haptophytes seem to be more important in open seawaters. The Arctic Ocean and adjacent seas are areas of major Atlantic circulation (e.g., deep water formation) and their oceanographic interest because of their contribution to the role in carbon sequestration (Anderson et al. 1998). The area investigated (Fig. 1), at the intersection of the Barents, Nor- 1 To whom correspondence should be addressed. Present address: wegian, and Greenland Seas is hydrologically complex and Marine Microbial Ecology group, Rosenstiel School of Marine and is one of the areas north of 708N that becomes ice free for Atmospheric Science, University of Miami, 4600 Rickenbacker some part of the year. Warm and saline Atlantic waters enter Causeway, Miami, Florida 33149-1098 ([email protected]). from the Norwegian Sea. This North Atlantic current ¯ows Acknowledgments northward parallel to the Norwegian coast and then divides We wish to thank the F/F Johan Hjort crew for ef®cient assis- into two main branches. One follows the continental slope tance with the sampling, as well as the Norwegian Marine Research west of the Spitsbergen archipelago and the other ¯ows east- Institute for providing ship time. Thanks are also due to Francisco ward into the Barents Sea (Fig. 1). These waters meet less Rodriguez for help with CHEMTAX analysis and helpful discus- saline and colder Arctic waters ¯owing southward along the sion. We thank Connie Lovejoy and Alexandra Worden for critically east coast of the Spitsbergen archipelago. The encounter of reading this manuscript and two anonymous reviewers for helpful Atlantic and Arctic water masses around Bear Island forms comments and suggestions. This work was funded by the following programs: PICODIV supported by the European Union (EVK3-CT- 1999-00021), PICMANCHE supported by the ReÂgion Bretagne, and the Biogeochemistry and Optic South Paci®c Experiment (BIO- Flux (PROOF), CNRS. F.N. was supported by a doctoral fellowship SOPE) supported by Processus BiogeÂochimiques dans l'OceÂan et from the French Research Ministry. 1677 1678 Not et al. sen (1970) noted that green ¯agellates related to the class Prasinophyceae were signi®cantly represented and that the prasinophyte species Micromonas pusilla was very abundant in this area. Here we examined the diversity and abundance of eu- karyotic picophytoplankton in relation to hydrological fea- tures along two transects through the complex region around the polar front located between the Norwegian and Barents Seas. Samples were collected in late summer, when the over- all contribution of picophytoplankton to total phytoplankton was expected to be large. The combined use of pigment anal- yses, tyramide signal ampli®cation±¯uorescent in situ hy- bridization (TSA-FISH) with oligonucleotide probes, epi¯u- orescence microscopy, and ¯ow cytometry (FCM) allowed us to investigate the importance of eukaryotic picoplankton and to identify the main phylogenetic groups within this complex assemblage. Material and methods Sampling proceduresÐSampling was done during an oceanographic cruise on board the F/F Johan Hjort (Nor- wegian Institute of Marine Research). This cruise took place in an area between the Norwegian, Greenland, and Barents Seas (Fig. 1). Ten stations were sampled from 20 August to 8 September 2002 along two transects (south/north, S/N and east/west, E/W) south of the Spitsbergen archipelago (Fig. 1 and Table 1). Samples were collected at ®ve depths with a conductivity±temperature±depth (CTD) rosette system equipped with Niskin bottles (5 liters). Chlorophyll ¯uores- cence provided by a SD200W ¯uorometer was used as a Fig. 1. Map of the cruise area. The two transects S/N (south/ guide to choose the ®ve sampling depths: in the mixed layer north) and E/W (east/west) are indicated. Arrows represent surface (5 m), just above the chlorophyll maximum (ca. 10 m), chlo- circulation and origin of waters encountered, adapted from Mar- kowski and Wiktor (1998). rophyll maximum, just below the chlorophyll maximum (ca. 10 m), and below the euphotic zone (60 m). Water was pre- ®ltered through a 200-mm mesh to remove zooplankton, a marked polar front. The exact position of this front varies large phytoplankton, and particles before further ®ltrations. with time. The Barents Sea shelf is one of the most biologically pro- DAPI stainingÐMicrobial counts were performed on ductive regions of the world. In the waters of Atlantic origin, board with a Zeiss epi¯uorescence microscope. Glutaralde- a phytoplankton bloom develops in May±June while post- hyde (Sigma) ®xed samples (20 ml, 1% glutaraldehyde ®nal bloom communities are observed in summer (Kogeler and concentration) were stained with 496-diamidino-2-phenylin- Rey 1999). Although large interannual variations in the tim- dole (DAPI, 5 mgml21 ®nal concentration) and ®ltered ing of the bloom are observed in the northern Barents Sea through 0.6-mm black, 25-mm diameter polycarbonate ®lters (Arctic waters), it usually starts earlier, in April±May im- (Poretics). Flagellates were identi®ed and counted by stan- mediately after the ice starts melting (Kogeler and Rey dard epi¯uorescence microscopy (Porter and Feig 1980) 1999). In the region we investigated, diatoms make up typ- based on their blue ¯uorescence under ultraviolet (UV) ex- ically most of the microphytoplankton species inventories, citation (360 nm). The presence or absence of chlorophyll while dino¯agellates as well as ¯agellates such as Phaeo- in ¯agellates was determined based on their red ¯uorescence cystis pouchetii (Hariot) Lagerheim and cryptophytes have under blue light excitation (490 nm). The size of each in- also been reported in signi®cant numbers (Markowski and dividual cell was estimated using an ocular grid in order to Wiktor 1998; Rat'kova et al. 1998). In contrast, there have obtain the abundances of cells for three size classes (2, 3± been few studies focused on the picophytoplankton (cells ,3 4, and 5 mm). Total ¯agellate counts were grouped into cat- mm) in Arctic waters. Murphy and Haugen (1985) found that egories based on size and presence or absence of chloro- the cyanobacterium Synechococcus NaÈgeli decreases in plasts. abundance with increasing latitude. Verity et al. (1999) and Rat'kova and Wassmann (2002) observed that total chloro- Flow cytometryÐSamples for ¯ow cytometry were pre- phyll biomass is dominated by small ¯agellates and monads ®ltered through a 3-mm Nuclepore ®lter (Whatman). Then during periods outside microphytoplankton blooms (respec- 1.5 ml of seawater was ®xed with a mixture of 1% parafor- tively in the Barents Sea and off northern Norway). Thrond- maldehyde (PFA) and 0.1% glutaraldehyde (®nal concentra- Picoplankton diversity in Arctic seas 1679 Table 1. Position, bottom depth, and hydrology for each sampling station. Bottom Chlorophyll Depth of Surface depth max* chlorophyll temperature Surface Station Latitude Longitude (m) (ng L21) max (m) (8C) salinity Z01 708309 208049 155 650 13 12.6 33.26 Z07 718309 198489 234 1,320 25 11.9 34.41 Z11 728319 198369 390 1,249 25 11.0 34.56 Z15 738299 198189 479 1,323 16 8.9 34.86 Z18 738599 198149 128 1,701 6 4.5 34.31 M09 768199 238459 67 1,835 20 4.5 34.39 Z68 768209 188479 268 1,393 5 7.1 34.11 Z65 768209 148539 354 1,674 16 7.1 34.88 Z61 768209 78609 2,170 891 16 8.2 35.05 Z59 768209 38599 3,231 737 36 6.0 33.63 * Chlorophyll max, maximum of Chlorophyll a concentration. tion), frozen in liquid nitrogen, and stored at 2808C.
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