Michael P. Lizotte

Primary production in Southern Ocean waters

The Southern Ocean forms a link between major ocean basins, is the site of deep and intermediate water ventilation, and is one of the few areas where macronutrients are underutilized by phytoplankton. Paradoxically, prior estimates of annual primary production are insufficient to support the Antarctic food web. Here we present results from a primary production algorithm based upon monthly climatological phytoplankton pigment concentrations from the coastal zone color scanner (CZCS). Phytoplankton production was forced using monthly temperature profiles and a radiative transfer model that computed changes in photosynthetically usable radiation at each CZCS pixel location. Average daily productivity (g C m−2 d−1) and total monthly production (Tg C month−1) were calculated for each of five geographic sectors (defined by longitude) and three ecological provinces (defined by sea ice coverage and bathymetry as the pelagic province, the marginal ice zone, and the shelf). Annual primary production in the Southern Ocean (south of 50°S) was calculated to be 4414 Tg C yr−1, 4–5 times higher than previous estimates made from in situ data. Primary production was greatest in the month of December (816 Tg C month−1) and in the pelagic province (contributing 88.6% of the annual primary production). Because of their small size the marginal ice zone (MIZ) and the shelf contributed only 9.5% and 1.8%, respectively, despite exhibiting higher daily production rates. The Ross Sea was the most productive region, accounting for 28% of annual production. The fourfold increase in the estimate of primary production for the Southern Ocean likely makes the notion of an “Antarctic paradox” (primary production insufficient to support the populations of Southern Ocean grazers, including krill, copepods, microzooplankton, etc.) obsolete.

Rapid and early export of Phaeocystis antarctica blooms in the Ross Sea, Antarctica.

The Southern Ocean is very important for the potential sequestration of carbon dioxide in the oceans and is expected to be vulnerable to changes in carbon export forced by anthropogenic climate warming. Annual phytoplankton blooms in seasonal ice zones are highly productive and are thought to contribute significantly to pCO2 drawdown in the Southern Ocean. Diatoms are assumed to be the most important phytoplankton class with respect to export production in the Southern Ocean; however, the colonial prymnesiophyte Phaeocystis antarctica regularly forms huge blooms in seasonal ice zones and coastal Antarctic waters. There is little evidence regarding the fate of carbon produced by P. antarctica in the Southern Ocean, although remineralization in the upper water column has been proposed to be the main pathway in polar waters. Here we present evidence for early and rapid carbon export from P. antarctica blooms to deep water and sediments in the Ross Sea. Carbon sequestration from P. antarctica blooms may influence the carbon cycle in the Southern Ocean, especially if projected climatic changes lead to an alteration in the structure of the phytoplankton community.

The Contributions of Sea Ice Algae to Antarctic Marine Primary Production1

SYNOPSIS. The seasonally ice-covered regions of the Southern Ocean have distinctive ecological systems due to the growth of microalgae in sea ice. Although sea ice microalgal production is exceeded by phytoplankton production on an annual basis in most offshore regions of the Southern Ocean, blooms of sea ice algae differ considerably from the phytoplankton in terms of timing and distribution. Thus sea ice algae provide food resources for higher trophic level organisms in seasons and regions where water column biological production is low or negligible. A flux of biogenic material from sea ice to the water column and benthos follows ice melt, and some of the algal species are known to occur in ensuing phytoplankton blooms. A review of algal species in pack ice and offshore plankton showed that dominance is common for three species: Phaeocystis antarctica, Fragilariopsis cylindrus and Fragilariopsis curta. The degree to which dominance by these species is a product of successional processes in sea ice communities could be an important in determining their biogeochemical contribution to the Southern Ocean and their ability to seed blooms in marginal ice zones.

Phytoplankton taxonomic variability in nutrient utilization and primary production in the Ross Sea

Patterns of nutrient utilization and primary productivity (PP) in late austral spring and early summer in the southwestern Ross Sea were characterized with respect to phytoplankton taxonomic composition, polynya dynamics, and upper ocean hydrography during the 1996–1997 oceanographic program Research on Ocean-Atmosphere Variability and Ecosystem Response in the Ross Sea. Phytoplankton biomass in the upper 150 m of the water column ranged from 40 to 540 mg chlorophyll a (Chl a) m−2, exceeding 200 mg Chl a m−2 everywhere except the extreme northern and eastern boundaries of the Ross Sea polynya. Diatom biomass was greatest in the shallow mixed layers of Terra Nova Bay, while the more deeply mixed waters of the Ross Sea polynya were dominated by Phaeocystis antarctica. Daily production computed from the disappearance of NO3 (1.14 g C m−2 d−1) and total dissolved inorganic carbon (TDIC, 1.29 g C m−2 d−1) is consistent with estimates made from an algorithm forced with satellite measurements of Chl a (1.25 g C m−2 d−1) and from measurements of 14C uptake (1.33 g C m−2 d−1). Phytoplankton PP in the Ross Sea averaged 100 g C m−2 yr−1 during 1996–1997. Despite the early formation of the Terra Nova Bay polynya the diatom bloom there did not reach its peak PP until middle to late January 1997 (most likely because of more intense wind mixing in November), ∼6 weeks after the P. antarctica bloom in the Ross Sea polynya had reached the same stage of development. From 70 to 100% of the C and N deficits in the upper 150 m could be accounted for by particulate organic matter, indicating that there had been little dissolved organic matter production or export of particulate material prior to our cruise. This suggests that early in the season, PP and zooplankton grazing are decoupled in the southwestern Ross Sea. The NO3∶PO4 disappearance ratio in waters dominated by P. antarctica (19.0±0.61) was significantly greater than in waters where diatoms were most common (9.52±0.33), and both were significantly different from the Redfield N∶P ratio of 16. Vertical profiles of TDIC suggest that P. antarctica took up 110% more CO2 per mole of PO4 removed than did diatoms, an important consideration for climate models that estimate C uptake from the removal of PO4.

Phytoplankton dynamics in the stratified water column of Lake Bonney, Antarctica I. Biomass and productivity during the winter-spring transition

Phytoplankton populations in perennially ice-covered Lake Bonney, Antarctica grow in a unique non-turbulent environment. The absence of turbulence generated by winds or major streams, combined with strong vertical gradients in temperature and nutrients, create vertically stratified environmental conditions that support three discrete phytoplankton populations in the east lobe of this lake. Phytoplankton biomass and photosynthesis were measured in the east lobe of Lake Bonney during the winter-spring transicion (September) to mid-summer (January). During this period, irradiance beneath the ice increased from 0.03 to 1.9 mol quanta m−2 d−1. Chlorophylla concentrations ranged from 0.03 to 3.8 μl−1 within the trophogenic zone (just beneath the permanent ice cover to 20 m) and photosynthesis ranged from below detection to 3.2 μg Cl−1 d−1. Our results indicate: (1) phytoplankton photosynthesis began in late winter (before 9 September, our earliest sampling date); (2) maxima for phytoplankton biomass and production developed sequentially in time from the top to the bottom of the trophogenic zone, following the seasoral increase in irradiance; and (3) the highest photosynthetic efficiencies occurred in early spring, then decreased over the remainder of the phytoplankton growth season. The spring decrease in photosynthetic rates for shallower phytoplankton appeared to be related to nutrient availability, while photosynthesis in the deeper populations was solely lightdependent.

Bio‐optical properties of the southwestern Ross Sea

The bio-optical properties of the southwestern Ross Sea were measured as part of the Antarctic research program Research on Atmospheric Variability and Atmospheric Response in the Ross Sea (ROAVERRS). The study area contained three distinct phytoplankton blooms, distinguishable by species composition. The largest in area was located to the north of the Ross Ice Shelf and was dominated by the prymnesiophyte Phaeocystis antarctica; chlorophyll a (Chl a) ranged from 0.45 to 8.2 mg m−3. Beam attenuation and particle absorption at 435 nm were as high as 3.4 m−1 and 0.35 m−1, respectively. A bloom of diatoms was more spatially restricted, located to the north and west of the P. antarctica bloom, with Chl a generally below 4 mg m−3. Neither diatoms nor P. antarctica exhibited evidence of the level of pigment packaging measured in waters near the Antarctic Peninsula during the Research on Antarctic Coastal Ecosystem Rates (RACER) program, possibly because of their smaller sizes. A much smaller cryptophyte bloom, located south of the Drygalski Ice Tongue, displayed a lower pigment-specific absorption spectra than did P. antarctica or diatoms, a sign of greater pigment packaging. Pigment-specific diffuse attenuation coefficients were consistent with the pigment-specific particle absorption coefficients (aph*), both being ∼3 times greater than similar measurements made during RACER. Spectral absorption by solutes determined through regression analysis of Kd against Chl a for the ROAVERRS data set was nearly identical to that measured during RACER. Total diffuse attenuation spectra at a given station could be reconstructed by summing the inherent optical properties of the major optical components (pure water, soluble material, detritus, phytoplankton) measured there. Differences in the absorption ratio of aph*(λ) at 490 nm to aph*(λ) at 555 nm among the three dominant phytoplankton taxa in the southwestern Ross Sea were responsible for most of the variability in the ratio of remote sensing reflectance (Rrs) at these same wavelengths. At a given concentration of Chl a, the ratio log [Rrs(490):Rrs(555)] was greatest in cryptophyte-dominated waters, which also possessed the lowest aph*(490):aph*(555) ratio, and lowest in P. antarctica–dominated waters. These bio-optical differences suggest that no simple empirical relationship between Chl a and log [Rrs(490):Rrs(555)] will apply to all three taxonomically distinct phytoplankton blooms in the southwestern Ross Sea.

Biochemical composition and photosynthate distribution in sea ice microalgae of McMurdo Sound, Antarctica: evidence for nutrient stress during the spring bloom

The nutrient status of microalgae inhabiting sea ice in McMurdo Sound, Antarctica was evaluated during the peak and decline of the spring bloom in November and December. Natural populations of microalgae were analysed for C, N, chlorophyll a , protein, lipid, polysaccharide, and low-molecular-weight carbohydrate content, and for the distribution of 14 C-labelled photosynthate into macromolecular fractions. Ratios of N:C and protein to carbohydrate (PR:CHO) were similar to values reported for nutrient-limited phytoplankton. Biochemical ratios and 14 C-photosynthate allocation patterns suggest that microalgae from congelation ice habitats may be more nutrient-stressed than those from underlying platelet ice habitats. This trend would be consistent with the presumed gradient of seawater nutrient influx through the platelet layer to the bottom congelation ice habitat. Microalgae from congelation ice subjected to an experimental depletion of nutrients (particularly nitrate) showed decreased N:C, PR:CHO, and allocation of 14 C-photosynthate to proteins. This evidence suggests that sea ice microalgae are nutrient-stressed during the peak and decline of the spring bloom in McMurdo Sound, which presumably begins when microalgal biomass concentrations and demands for growth reach or exceed the rate of nutrient supply from underlying seawater.

Photosynthetic capacity in microalgae associated with Antarctic pack ice

SummaryPrevious studies of primary production in Antarctic seas have concluded that microalgae associated with sea ice make only a minor contribution to the carbon budget; however, production estimates for sea ice algae have been based almost exclusively on microalgae from nearshore fast ice. We measured biomass and rates of photosynthesis (at saturating irradiances) in microalgae collected from offshore pack ice during four cruises to the Weddell-Scotia Sea and the region west of the Antarctic Peninsula. Chlorophyll a concentrations in pack ice (0.089 to 260 μg 1-1) were as high as reported from fast ice. Photosynthetic rates typically ranged (median 75%) from 0.3 to 3.6 μ C μg chl a-1 h-1 (n=127; arithmetic mean = 1.7, S D =1.9). These photosynthetic capacities are approximately an order of magnitude greater than previously reported for fast ice microalgae, but are similar to rates reported for Antarctic phytoplankton. Because pack ice constitutes more than 90% of the ice cover in Antarctic seas and indigenous microalgae have a higher photosynthetic capacity than previously realized, we raise the question: has the importance of sea ice algae to primary product: on in the southern ocean been underestimated?