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MARINE ECOLOGY PROGRESS SERIES Vol. 180: 23-36, 1999 Published May 3 Mar Ecol Prog Ser Microbial dynamics in coastal waters of East Antarctica: plankton production and respiration Carol R~binson'~~~*.**,Stephen D. ~rcher2r314r*, Peter J. le B. ~illiams' 'University of Wales Bangor, School of Ocean Sciences. Menai Bridge, Anglesey LL59 5EY. United Kingdom 'British Antarctic Survey, High Cross, Madingley Road, Cambridge CB3 OET, United Kingdom 3Department of Biology. University of Southampton. Medical and Biological Sciences Building, Bassett Crescent East, Southampton S016 7PX, United Kingdom 'In collaboration with the Australian Antarctic Division, Channel Highway, Kingston, Tasmania 7050, Australia ABSTRACT The rates of plankton community production and respiration were determined from in vitro changes in dissolved inorganic carbon and dissolved oxygen and the incorporation of NaH14C03 at a coastal site in East Antarctica between 16 December 1993 and 12 February 1994. The breakout of seasonal fast ice was associated with a succession of dominant phytoplankton from Cryptomonas to Phaeocystis to a d~atomassemblage. Gross production reached 33 mm01 C m-3 d-l and I4C incorpora- tion peaked at 24 mm01 C m-3 d-' on 23 January 1994, at the time of the chlorophyll a maximum (22 mg chl a m-3).Dark community respiration reached its maximum (13 mm01 C m-3 d-' ) 4 d later. Photosyn- thetic rates calculated from I4C incorporation were s~gnificantlylower (17 to 59%) than rates of gross production. The derivation of plankton processes from changes in both dissolved oxygen and dissolved inorganic carbon allowed the direct measurement of photosynthetic and respiratory quotients. A linear regression of all data gave a photosynthetic quotient of 1.33 + 0.23 and a respiratory quotient of 0.88 * 0.14. Concurrent determination$ of bacterial, heterotrophic dinoflagellate, nanoflagellate and ciliate respiration could account for 15 to 58% of measured dark community respiration. This study has improved the sparse data set of plankton respiration measurements, confirmed that heterotrophic res- piration is a significant process in the carbon flux of coastal Antarctic waters and achieved a first appor- tionment of community respiration to the major microbial groups in this region KEY WORDS: Photosynthesis . Respiration . Plankton . Antarctic INTRODUCTION of community respiration are often lacking. The devel- opment of the coulometric titration system for the pre- The biological conversion of dissolved inorganic car- cise measurement of DIC (Johnson et al. 1987, Robin- bon (DIC) to particulate inorganic and dissolved and son & Williams 1991) means that microbiologically particulate organic carbon in the euphotic zone of the induced changes in DIC concentration can now be ocean has a major influence on atmospheric CO2 con- directly measured and plankton respiration can be centrations and the amount of organic carbon available determined concurrently with plankton photosyn- to higher marine organisms. The accurate determina- thesis. tion of this DIC conversion is therefore of critical im- Plankton processes in the Southern Ocean are cur- portance and is equal to the excess of DIC consumed rently receiving much scientific attention, due in part during photosynthesis over that produced during to the recognition that this region may respond dra- plankton community respiration. Estimates of plankton matically to global change (Codispoti et al. 1991). photosynthesis derived from the incorporation of I4C Because of the short duration of oceanographic re- are now routine; however, concurrent measurements search cruises it is not possible to characterise and explain changes in oceanic biogeochemical and eco- 'Present address: Plymouth Marine Laboratory, Prospect logical variables over longer time periods or changes Place, West Hoe, Plymouth PL1 3DH, United Kingdom due to episodic environmental events. Seasonal and "E-mail: [email protected] annual time series studies provide invaluable data for 0 Inter-Research 1999 Resale of full article not permitted 24 Mar Ecol Prog Ser 180: 23-36, 1999 calibration and validation of biological and biogeo- approximately weekly intervals between 9 December chemical models, and if continued over the longer term 1993 and 12 February 1994 for determination of the (5 to 10+ yr), may detect changes in the ecological and depth distribution of temperature, salinity, dissolved chemical state of the ocean brought about by global oxygen and DIC. At approximately twice weekly inter- climate change (JGOFS 1990). vals, water samples were collected from a depth of 5 m The present study formed part of a co-ordinated pro- at O'Gorman Rocks for estimation of gross and net gramme which aimed to measure the temporal pro- community production and dark community respira- gression of microbial processes in Antarctic coastal tion from in vitro changes in dissolved oxygen and waters. Concurrent studies within the programme carbon dioxide, plankton photosynthesis from the included herbivory by heterotrophic dinoflagellates incorporation of NaH14C03,and chlorophyll a concen- (Archer et al. 1996a),bacterial production and nanofla- tration. gellate bacterivory (Leakey et al. 1996), the microbial CTD measurements. Conductivity, temperature and ecology of sea ice (Archer et al. 1996b) and ciliate her- depth were recorded with a handheld CTD (Platypus bivory (J. Grey & R. J.G. Leakey unpubl.). The com- Engineering, Tasmania) at weekly intervals at O'Gor- bined data of the co-ordinated programme allows the man Rocks. routes of carbon flow through the microbial food web Dissolved oxygen and inorganic carbon measure- to be examined (Robinson et al. 1996, Leakey et al. ments. Water was siphoned from the acid-washed unpubl.). Goflo bottle into borosilicate glass bottles using silicon This paper presents estimates of plankton photosyn- tubing. Dissolved oxygen concentration was measured thesis and respiration derived from 3 independent with an automated whole bottle Winkler titration sys- methods: changes in DIC and dissolved oxygen con- tem using a photometric endpoint (Williams & Jenkin- centrations and the incorporation of I4C-labelled bi- son 1982).Oxygen saturation was calculated using the carbonate at 1 site over a 60 d period. equations for the solubility of oxygen in seawater of This has enabled the determination of the temporal Benson & Krause (1984). DIC was measured by auto- change in both autotrophic and heterotrophic micro- mated coulometric titration (Robinson & Williams 1991, bial processes and the first direct measurements of DOE 1994). Analysis of seawater DIC reference mate- plankton respiration, production/respiration (P/R) rials certified at 1984.05 * 0.57 (A.G. Dickson) ratios and photosynthetic and respiratory quotients in throughout the fieldwork (mean = 1985.81 * 2.35; n = this region. 9) provided quality assessment of the precision and Due to an incomplete understanding of intracellular accuracy of the DIC measurements. carbon flow within the algal cell, the scientific commu- Measurement of gross production (GP), net commu- nity is still without consensus as to whether, and under nity production (NCP) and dark community respira- what controlling conditions, the I4C technique mea- tion (DCR). Bulk water samples (60 dm3) were col- sures net or gross photosynthesis (Ryther 1956, Stee- lected in an acid-washed opaque polypropylene mann Nielsen & Hansen 1959). Williams et al. (1996) aspirator using an acid-washed 5 dm3 Goflo sampling highlighted the need to accrue a database of I4C and bottle through a dedicated hole in the ice, or deployed DIC rate comparisons in order to establish the scale of from the side of a boat once the ice moved away. Sam- the problem of interpreting 14C field data. Although pling took place during hours of least sunlight (approx. not a primary aim of thls study, a data set of concurrent 23:OO h),and precautions were taken to minimise light 14C and DIC rate measurements was obtained.. shocking of the sample during sample collection and Finally, and to our knowledge uniquely, concurrent manipulation. Water was siphoned from the 60 dm3 measurements of bacterial, ciliate, nanoflagellate and opaque aspirator into borosilicate glass bottles through heterotrophic dinoflagellate biomass and activity have darkened silicon tubing, overflowing each sample been used to apportion community respiration to these bottle by 4 times its volume. Four 150 cm3 dissolved major trophic groups. oxygen sample bottles and three 250 cm"1C sample bottles were fixed immediately for measurement of zero time concentrations. DIC samples were fixed with METHODS a saturated mercuric chloride solution (250 mm3 per 250 cm3 sample). A further 8 dissolved oxygen and 6 This study was undertaken at a coastal location in DIC sample bottles were clamped to perspex bottle the vicinity of the Australian Antarctic station of Davis racks to enable a light/dark bottle incubation. Sample (68"35'S, 77'58'E) during the austral summer of bottles were incubated in sifu whenever possible; how- 1993-1994 ever, when logistics demanded, simulated in situ incu- Sampling. The study site O'Gorman Rocks, Antarc- bations were undertaken at 1 m depth alongside the tica (Fig. 1, 1 km offshore, 22 m depth),was sampled at wharf. Samples were fixed at the end of the 24 h incu- Robinson et al.: Plankton production and respiration 25 - Depths In metres Fig. 1 Study area showing location of sampling site (@) at O'Gorman Rocks in the vicinity of the Australian Antarctic station of Davis bation and stored underwater until analysis (within samples were fixed immediately and the remainder 24 h) as described above. After analysis, DIC sample incubated in the dark at surface water temperature. bottles were acid-washed (10% HCl) in order to Triplicate samples were fixed after each of 2, 4, 6, 9, remove any trace of mercuric chloride which would 12 and 24 h. affect any subsequent incubation. NCP was calculated from the net flux of dissolved The remainder of the collected 60 dm3 water sample oxygen and DIC in the 'light/dark' bottles, DCR as the was used to assess the size distribution of community decrease in dissolved oxygen and increase in DIC in respiration.