Phytoplankton sinking rates in the Ross Sea fileChlorophyll was determined fluorometri-cally after...

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Phytoplankton sinking rates in the Ross Sea WALKER 0. SMITH, JR., HOLLY P. KELLY, and DALE VOGELIN Graduate Program in Ecology University of Tennessee Knoxville, Tennessee 37996 ANN R. CLOSE Bermuda Biological Station Ferry Reach, Bermuda As part of a coordinated, interdisciplinary study of the pro- duction of biogenic material at the surface, its flux and reminer- alization through the water column, and its accumulation in the sediments, we measured the sinking rates of suspended particulate matter in the Ross Sea in January and February 1990. Substantial deposits of diatoms occur in the sediments of the Ross Sea (Ledford-Hoffman, DeMaster, and Nittrouer 1986), and these deposits generally are composed of intact phy- toplankton cells. One of our hypotheses in this program was that the vertical flux of phytoplankton from the ice-edge bloom was large relative to less productive regions, and we wanted to know whether this flux was continual or episodic in time (i.e., a large pulse of intact cells sink near the end of the growing season). If the latter were true, it would have significant impli- cations for the biogeochemical cycles of biogenic elements (car- bon, nitrogen, silicon) as well as food web dynamics. We measured sinking rates using settling columns originally described by Bienfang (1981) and modified for use at sea (figure; Johnson and Smith 1986). Samples were collected from two depths, and the sinking rates of each were determined simul- taneously. Generally, sinking rates were determined from the surface and the depth of the chlorophyll maximum, as deter- mined by a continuous fluorescence trace collected as part of the routine conductivity-temperature-depth cast. Samples for chlorophyll and particulate carbon were taken before and after incubation. No correction for growth during incubation was made because growth rates over the short incubation intervals (2 hours) and at the low incubation temperatures are within analytical precision. Chlorophyll was determined fluorometri- cally after filtration and extraction, and particulate carbon was determined by high-temperature pyrolysis. Sinking rates were calculated from the differences between initial and final partic- ulate biomass concentrations (Johnson and Smith 1986). Sinking rates were moderate for the stations sampled (table 1). Sinking rates based on chlorophyll averaged 0.24 and 0.17 meters per day at the surface and depth of chlorophyll maxi- mum, respectively. These rates are slightly greater than those found in the Arctic (Culver and Smith 1989), but less than those observed in the Weddell Sea in 1983 (Johnson and Smith 1986), which averaged 0.89 meters per day. The rates at the chloro- phyll maximum tended to be lower than those at the surface, although the means were not statistically different due to the variability in sinking rates between stations. We also compared the sinking rates of the two locations where we had deployed sediment traps (site A: 76°30'S 167°30'E; site B: 76°30'S 175°W) (table 2). Sinking rates were Schematic diagram of the gimballed settling column used at sea to measure particulate matter sinking rates. greater at the eastern location for both depths. We also ob- served that the species composition was greatly different be- tween the two locations, with the site A being dominated by the diatom Nitzschia curta and site B being a mixed diatom- Phaeocystis assemblage. Vertical flux data suggested that there was a considerable temporal difference within one location, as well as a significant spatial difference (table 2). The contents of the material collected by the sediment traps at the two locations also differed, with the easternmost location being dominated by small, ellipsoidal fecal pellets, whereas site B was dominated by loose aggregates. Based on a comparison of calculated sink- ing rates and vertical flux information, passive particle sinking did not contribute significantly to the total vertical flux. Accel- erated removal of particulate material from the euphotic zone must be mediated by either micro- or macrozooplankton graz- Table 1. Statistics for chlorophyll-based sinking rates observed in the Ross Sea during January and February 1990 Parameter Surface Depth of chlorophyll maximum Mean sinking rate 0.24 0.17 Standard deviation 0.27 0.22 Number of observations 21 20 Range 0.0-1.10 0.0-0.85 a In meters per day. 1991 REVIEW 151

Transcript of Phytoplankton sinking rates in the Ross Sea fileChlorophyll was determined fluorometri-cally after...

Phytoplankton sinking ratesin the Ross Sea

WALKER 0. SMITH, JR., HOLLY P. KELLY, and DALE VOGELIN

Graduate Program in EcologyUniversity of Tennessee

Knoxville, Tennessee 37996

ANN R. CLOSE

Bermuda Biological StationFerry Reach, Bermuda

As part of a coordinated, interdisciplinary study of the pro-duction of biogenic material at the surface, its flux and reminer-alization through the water column, and its accumulation inthe sediments, we measured the sinking rates of suspendedparticulate matter in the Ross Sea in January and February1990. Substantial deposits of diatoms occur in the sedimentsof the Ross Sea (Ledford-Hoffman, DeMaster, and Nittrouer1986), and these deposits generally are composed of intact phy-toplankton cells. One of our hypotheses in this program wasthat the vertical flux of phytoplankton from the ice-edge bloomwas large relative to less productive regions, and we wanted toknow whether this flux was continual or episodic in time (i.e.,a large pulse of intact cells sink near the end of the growingseason). If the latter were true, it would have significant impli-cations for the biogeochemical cycles of biogenic elements (car-bon, nitrogen, silicon) as well as food web dynamics.

We measured sinking rates using settling columns originallydescribed by Bienfang (1981) and modified for use at sea (figure;Johnson and Smith 1986). Samples were collected from twodepths, and the sinking rates of each were determined simul-taneously. Generally, sinking rates were determined from thesurface and the depth of the chlorophyll maximum, as deter-mined by a continuous fluorescence trace collected as part ofthe routine conductivity-temperature-depth cast. Samples forchlorophyll and particulate carbon were taken before and afterincubation. No correction for growth during incubation wasmade because growth rates over the short incubation intervals(2 hours) and at the low incubation temperatures are withinanalytical precision. Chlorophyll was determined fluorometri-cally after filtration and extraction, and particulate carbon wasdetermined by high-temperature pyrolysis. Sinking rates werecalculated from the differences between initial and final partic-ulate biomass concentrations (Johnson and Smith 1986).

Sinking rates were moderate for the stations sampled (table1). Sinking rates based on chlorophyll averaged 0.24 and 0.17meters per day at the surface and depth of chlorophyll maxi-mum, respectively. These rates are slightly greater than thosefound in the Arctic (Culver and Smith 1989), but less than thoseobserved in the Weddell Sea in 1983 (Johnson and Smith 1986),which averaged 0.89 meters per day. The rates at the chloro-phyll maximum tended to be lower than those at the surface,although the means were not statistically different due to thevariability in sinking rates between stations.

We also compared the sinking rates of the two locationswhere we had deployed sediment traps (site A: 76°30'S167°30'E; site B: 76°30'S 175°W) (table 2). Sinking rates were

Schematic diagram of the gimballed settling column used at seato measure particulate matter sinking rates.

greater at the eastern location for both depths. We also ob-served that the species composition was greatly different be-tween the two locations, with the site A being dominated bythe diatom Nitzschia curta and site B being a mixed diatom-Phaeocystis assemblage. Vertical flux data suggested that therewas a considerable temporal difference within one location, aswell as a significant spatial difference (table 2). The contents ofthe material collected by the sediment traps at the two locationsalso differed, with the easternmost location being dominatedby small, ellipsoidal fecal pellets, whereas site B was dominatedby loose aggregates. Based on a comparison of calculated sink-ing rates and vertical flux information, passive particle sinkingdid not contribute significantly to the total vertical flux. Accel-erated removal of particulate material from the euphotic zonemust be mediated by either micro- or macrozooplankton graz-

Table 1. Statistics for chlorophyll-based sinking ratesobserved in the Ross Sea during January and February 1990

Parameter SurfaceDepth of chlorophyll maximum

Mean sinking rate 0.24 0.17Standard deviation0.27 0.22Number of observations21 20Range 0.0-1.10 0.0-0.85

a In meters per day.

1991 REVIEW 151

ing and fecal pellet production, or by aggregate formation andtheir associated accelerated sinking rates.

This research was supported by National Science Foundationgrant DPP 88-17070. We thank Rob Dunbar and Amy Leventerfor use of the vertical flux data.

Table 2. Chlorophyll-based sinking rates and vertical fluxat sites A (76030'S 167°30'W) and B (76 030 1 S 1750E).

Vertical flux data from floating sediment trapsdeployed at 50 meters for approximately 24 hours.

ReferencesParameter Site A Site B

Surface sinking rate 0.32 0.23Sinking rate at depth of

chlorophyll maximum a 0.25 0.09Total carbon fluxb 25.6c 254d

24.9° 92.71

a In meters per day.In milligrams of carbon per square meter per day.Sediment trap deployment 12 January 1990.

ci Sediment trap deployment 4 February 1990.Sediment trap deployment 16 January 1990.Sediment trap deployment 31 January 1990.

Bienfang, P.K. 1981. SETCOL: A technologically simple and reliablemethod for measuring phytoplankton sinking rates. Journal of Plank-ton Research, 3, 235-253.

Culver, ME., and WO. Smith, Jr. 1989. Effects of environmental vari-ation on sinking rates of marine phytoplankton. Journal of Phycology,25,262-270.

Johnson, T.O., and WO. Smith, Jr. 1986. Sinking rates of natural phy-toplankton populations from the Weddell Sea marginal ice zone.Marine Ecology Progress Series, 33, 131-137

Ledford-Hoffman, P.A., D.J. DeMaster, and C.J. Nittrouer. 1986. Bio-genic-silica accumulation in the Ross Sea and the importance of Ant-arctic continental-shelf deposits in the marine silica budget. Geochim-ica et Cosmochimica Acta, 50, 2099-2110.

Massive prasinophyte bloomin northern Gerlache Strait

D.F. BIRD

Department de Sciences BiologiquesUniversite du Quebec a Montreal

Montreal, QuebecCanada

D.M. KARL

School of Ocean and Earth Science and TechnologyUniversity of Hawaii

Honolulu, Hawaii 96822

Algal blooms observed in the southern ocean have beendominated either by diatoms or by the colonial prymnesio-phyte, Phaeocystis (Kopczynska, Weber, and El-Sayed 1986;Priddle, Hawes, and Ellis-Evans 1986). These same algae uni-formly dominate ice algal communities (Homer 1976; Fryxelland Kendrick 1988). The greater the biomass of the bloom, thegreater the proportion of diatoms or Phaeocystis (Sakshaug andHolm-Hansen 1984). Therefore, we were surprised to discoverthat one of the densest blooms ever reported in the Antarctic(Holm-Hansen and Mitchell in press; Huntley et al. in press)consisted almost entirely of unicellular green flagellates of thegenus Pyramirnonas.

The objectives and sampling strategy of the Research on Ant-arctic Coastal Ecosystem Rates (RACER) program have beendescribed previously (Huntley, Niiler, and Holm-Hansen 1987;Huntley et al. 1991). Briefly, 69 stations placed regularly over a

25,000-square-kilometer area of southern Drake Passage,Bransfield Strait and northern Gerlache Strait were sampledrapidly (within 5 days) four times during the 1986-1987 australspring and summer (December, January, February, and March).More intensive studies were carried out at selected stationsbetween these monthly regional surveys.

The RACER program uncovered striking spatial and seasonalheterogeneity in both biomass and productivity of the planktonwithin the study area. The greatest primary productivity oc-curred in December (Holm-Hansen and Mitchell 1991) whereasthe biomass peaked a month later (Karl et al. 1991). During thisperiod, the size structure of the plankton shifted from net- tonanoplankton. The stations in Gerlache Strait and adjoiningstations north of Brabant Island were the site of a massivebloom, reaching 25 micrograms chlorophyll a per liter (Holm-Hansen and Mitchell 1991).

Surface samples (100 milliliters) for enumeration and floristicidentification were collected at each station. The samples werepassed through a 20-micrometer Nitex mesh, fixed immedi-ately with glutaraldehyde, concentrated onto 25-millimeter di-ameter Nuclepore filters (0.8 micrometer porosity), stainedwith proflavine (Haas 1982), mounted, and stored frozen untilexamined using epifluorescence microscopy. Cells were iden-tified and counted in transects at 1,250 x magnification.

The January bloom consisted almost entirely of an unicellularquadriflagellate of the genus Pyramimonas, subgenus Trichocystis(McFadden, Hill, Wetherbee 1986; figures 1 and 2). The identi-fication as Pyramimonas was unmistakable based on the cell'scolor, morphology (prominent pyrenoid, flagellar pit, chioro-plast shape), flagellation, the presence of numerous tricho-cysts, and high-pressure liquid chromatography confirmationof chlorophyll b (Head personal communication). Chlorophyllb has been detected previously in pigment extracts from south-ern ocean habitats (Bidigare et al. 1986; Buma et al. 1990), butit is typically only a minor constituent compared to chlorophylls

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