Comparison of Pelagic and Nepheloid Layer Marine Snow: Implications for Carbon Cycling
Total Page:16
File Type:pdf, Size:1020Kb
ELSEVIER Marine Geology 150 (1998) 39±50 Comparison of pelagic and nepheloid layer marine snow: implications for carbon cycling Barbara Ransom a,Ł,KevinF.Sheab, Patti Jo Burkett b, Richard H. Bennett c, Roy Baerwald d a Scripps Institution of Oceanography, University of California at San Diego, La Jolla, CA 92093-0220, USA b Naval Research Laboratory, Stennis Space Center, MS 39529, USA c SEAPROBE Inc., 01 Pine Street, Picayune, MS 39566, USA d Department of Biology, University of New Orleans, New Orleans, LO 70148, USA Received 2 June 1997; accepted 8 May 1998 Abstract Marine snow from upper and mid-water (i.e., pelagic) depths on the California margin is texturally and compositionally different from that traveling in the nepheloid layer. Transmission electron microscopy shows that pelagic marine snow consists primarily of bioclasts (e.g., diatom frustules, foram tests), organic matter, and microbes. These components are entrained as discrete particles or small aggregates .Ä 10 mm in diameter) in a loose network of exocellular, muco-polysaccharide material. Clays are infrequent but, when present, are constituents of comparatively compact organic-rich microaggregates. Microbes are abundant and appear to decrease in number with increasing water depth. In contrast, marine snow aggregates collected from just above the sea ¯oor in the nepheloid layer are assemblages of clay particles, clay ¯ocs, and relatively dense clay±organic-rich microaggregates in an exocellular organic matrix. Bioclasts and microorganisms occur only rarely. The prevalence of clay±organic-rich aggregates in the nepheloid layer suggests that, prior to ®nal deposition and burial, marine snow from the pelagic zone is subject to disaggregation and recombination with terrigenous detrital material near or at the sea ¯oor. Results have signi®cant implications for the accumulation and burial rates of organic carbon on continental margins and the aging and bioavailability of sedimentary organic matter. Samples examined were collected offshore of northern and central California. 1998 Elsevier Science B.V. All rights reserved. Keywords: nepheloid layer; marine snow; organic carbon; continental margin; TEM 1. Introduction has focused on surface to mid-water column ag- gregate production and the rates and variability of Marine snow, the population of marine aggre- organic carbon transport to the sea ¯oor (see review gates with diameters greater than 5 mm, is the in Alldredge and Silver, 1988). Recent research now primary means by which organic matter is trans- suggests that marine snow aggregates may also play ported down through the water column (Fowler and an important role in the development of sediment Knauer, 1986). In large part, marine snow research geotechnical properties (Syvitski, 1991) and the ul- timate distribution of organic matter in continental Ł Corresponding author. Fax: C1 (619) 534-0784; E-mail: margin sediments (Ransom et al., 1997). Below we [email protected] present the results of a comparative study of the com- 0025-3227/98/$19.00 1998 Elsevier Science B.V. All rights reserved. PII S0025-3227(98)00052-8 40 B. Ransom et al. / Marine Geology 150 (1998) 39±50 ponents of marine snow from various water depths fabric and sediment chemical and physical properties offshore California. Our objectives were to identify across the California continental slope (Kim, 1996; the components and textures of aggregates traveling Ransom et al., 1997, 1998; Sawyer et al., 1997). in the nepheloid layer, improve our understanding Mendocino samples were retrieved from an anchored of how processes and conditions in the nepheloid array deployed for 64 days in ¾950 m of water with layer affect marine aggregates that originate in the collectors at water depths of 250, 750, and 930 m upper water column (pelagic zone), and examine the at 40ë1.990N, 124ë31.60W. San Luis Obispo samples possible impact these interactions may have on the were recovered from a ¯oating array deployed for 24 distribution of organic matter in the sediment. h at 35ë27.50N and 121ë31.00W. The ¯oating array Marine snow is a complex assemblage of or- had ®ve evenly spaced collectors at water depths ganisms, bioclasts (e.g., tests and frustules), organic from 50 to 250 m. matter, minerals, and fecal material entrained as dis- Trap depths were selected on the basis of trans- crete particles or small aggregates in a loose network missometer data which indicated that the water col- of exocellular muco-polysaccharide slime and ®brils umn in the sampling areas could be divided into (e.g., Alldredge, 1989; Decho, 1990; Heissenberger three zones. The surface layer, shown in representa- et al., 1996). The composition of marine snow is tive transmissometer traces in Fig. 2, coincides with a function of local particle abundance, particle en- the turbid photic zone which is characterized by high counter rates, and the ease with which different com- particle concentrations resulting from biological pro- ponents can be incorporated into larger aggregates. ductivity and sea-surface aggregation processes. The Through aggregation and disaggregation, as well as composition, size, and number of aggregates formed grazing by bacterial and planktonic heterotrophs, the in this layer are episodically affected by suspended architecture, particle composition and organic matter material from rivers during periods of heavy conti- in marine snow evolves through time (see review in nental runoff. The pelagic layer, a zone of reduced Wakehan and Lee, 1993). particle concentration, through which marine snow The direct observation of marine snow con- formed in the surface layer descends, occurs at the stituents and their textural relations requires use base of the surface layer and extends to the top of of a transmission electron microscope (TEM). To the nepheloid layer. The nepheloid layer stretches date, such studies have concentrated on fecal pel- from the sediment±water interface up to the base lets and=or aggregates generated in surface waters of the pelagic zone. This layer directly overlies the and pelagic regions of the water column (e.g., Silver sea ¯oor and experiences episodic turbidity from the and Bruland, 1981; Gowing and Silver, 1983, 1985; lateral transport and resuspension of detrital clastic Leppard et al., 1996). Only one study contains im- material as a result of oceanographic, sedimentolog- ages of nepheloid aggregates (Ransom et al., 1997). ical, and biological processes (e.g., bottom currents, The potential of this class of aggregates to in¯uence storms, sediment gravity ¯ows, slope failure, organ- sea ¯oor sediment microfabric and the distribution ism burrowing, etc.). To some extent, the boundaries and bioavailability of organic matter warrants further between the three layers shown in Fig. 2 are tem- investigation. porally and spatially variable, being dependent on water depth and prevailing biological, physical, and oceanographic conditions. 2. Sampling locations Marine snow samples were collected with Martin- 3. Methods Knauer multi-collector (eight tubes per water depth) sediment traps (Knauer et al., 1979) from transects Samples were collected and prepared to ensure that near Mendocino and San Luis Obispo on the Califor- aggregates represented the in-situ textures and parti- nia continental margin (Fig. 1) from June 13 to Au- cles present in the water column. Two of the eight col- gust 16, 1993. These samples form a companion data lectors at each water depth were used to obtain sam- set to a high-resolution study of the sediment micro- ples for TEM study; the six others were used for an B. Ransom et al. / Marine Geology 150 (1998) 39±50 41 Fig. 1. Location map of the Mendocino (northern) and San Luis Obispo (southern) locations sampled off the coast of California (see inset). Bathymetric contours are in meters. Sediment trap locations are indicated by crosses. investigation of organic carbon ¯ux and spiked with Prior to processing the carbon ¯ux material, formalin to kill planktonic scavengers. The TEM-des- planktonic scavengers were removed from the ®lter ignated collectors were ®lled with pH-buffered sea- papers so as not to affect the mass of organic carbon water containing 2 wt% glutaraldehyde. Sucrose was corresponding to that of the water column particles. added to these to increase solution density to ¾1.07 Filter papers were then dried and weighed to obtain gcm3 to minimize washout during trap recovery. the mass of marine aggregate material collected. The Upon retrieval, liquid in the upper three quarters of dried material was analyzed for total carbon using the TEM collectors was slowly siphoned off. All but a Perkin Elmer CHN analyzer (precision Dš0:3%, 50 ml of the remaining liquid was then slowly drained based on repeated measurements of a cystine stan- through the bottom of the collector. The resulting sus- dard) and inorganic carbon using acid-dissolution pension, which contained marine snow, was gently coulometry (precision Dš0:5%, based on repeated decanted into a jar spiked with glutaraldehyde and analysis of a carbonate standard). Organic carbon osmium tetroxide and refrigerated. In the carbon ¯ux was determined by difference. For further discus- collectors, ¯uid in each column was drained through sion of carbon analytical procedures and errors see the bottom and all entrained particulate matter was Ransom et al. (1998). deposited on a tared ®lter paper. In the laboratory, marine snow which was sus- 42 B. Ransom et al. / Marine Geology 150 (1998) 39±50 Fig. 2. Transmissometer plots from the Mendocino and San Luis Obispo sediment trap transects. The distinct gradient changes at the base of the surface and pelagic layers were used to distinguish boundaries between layers and shaded zones and indicate the relative extent of each layer. pended in the TEM-designated collectors was gently clastic particles (pieces of diatom frustules), clay separated with forceps into pieces 1±3 mm in di- minerals, organic matter, and microbes. Identi®ca- ameter in preparation for embedding. Aggregates tion of speci®c components carries a number of were then quickly ethanol-exchanged (¾10 min per caveats and the reliability of such determinations is ¯uid) and embedded in Spurr's resin (Hayat, 1970).