Mesoscale iron enrichment experiments 1993-2005: synthesis and future directions.

Since the mid-1980s, our understanding of nutrient limitation of oceanic primary production has radically changed. Mesoscale iron addition experiments (FeAXs) have unequivocally shown that iron supply limits production in one-third of the world ocean, where surface macronutrient concentrations are perennially high. The findings of these 12 FeAXs also reveal that iron supply exerts controls on the dynamics of plankton blooms, which in turn affect the biogeochemical cycles of carbon, nitrogen, silicon, and sulfur and ultimately influence the Earth climate system. However, extrapolation of the key results of FeAXs to regional and seasonal scales in some cases is limited because of differing modes of iron supply in FeAXs and in the modern and paleo-oceans. New research directions include quantification of the coupling of oceanic iron and carbon biogeochemistry.

Biogenic carbon cycling in the upper ocean: effects of microbial respiration.

Food-web processes are important controls of oceanic biogenic carbon flux and ocean-atmosphere carbon dioxide exchange. Two key controlling parameters are the growth efficiencies of the principal trophic components and the rate of carbon remineralization. We report that bacterial growth efficiency is an inverse function of temperature. This relationship permits bacterial respiration in the euphotic zone to be computed from temperature and bacterial production. Using the temperature-growth efficiency relationship, we show that bacterial respiration generally accounts for most community respiration. This implies that a larger fraction of assimilated carbon is respired at low than at high latitudes, so a greater proportion of production can be exported in polar than in tropical regions. Because bacterial production is also a function of temperature, it should be possible to compute euphotic zone heterotrophic respiration at large scales using remotely sensed information.

Diel periodicity of photosynthesis in polar phytoplankton: influence on primary production.

In the Southern Ocean, primary production estimated from seasonal chemical and geochemical changes is two to four times greater than the value calculated from carbon-14 uptake. Since carbon uptake had typically been measured only during midday incubations, the influence of diel periodicity of photosynthesis on daily productions was not considered. Phytoplankton from McMurdo Sound, Antarctica, exhibited distinct, but seasonally variable diel patterns of light-saturated and light-limited photosynthesis. Maximum photosynthetic capacity occurred about noon in early September, and its occurrence progressively shifted to about midnight by late October. This shift was accompanied by a concomitant phase shift in the occurrence of minimum photosynthetic capacity from midnight to midday. Daily production estimated from time-of-day corrected photosynthetic characteristics and from 24-hour incubations was 2.5 to 4 times greater than that predicted from 6-hour midday incubations. If similar diel periodicity in photosynthesis occurs in other polar oceans, primary production would be significantly higher than previously estimated from carbon-14 uptake measurements.

Effects of flow cytometric analysis and cell sorting on photosynthetic carbon uptake by phytoplankton in cultures and from natural populations.

Photosynthetic rates of phytoplankton were significantly lower after analysis by flow cytometry (FCM) than before. Exposure to the laser beam during the sorting process caused significant physiological damage. The cellular content of a radiolabel accumulated prior to FCM was not affected by FCM. Although it may not be possible to use FCM to preconcentrate cells for further physiological studies, samples may be incubated with stable or radioactive isotopes and then analyzed by FCM.

Bacterivory: a novel feeding mode for asteroid larvae.

Planktotrophic larvae that occur beneath the annual sea ice in the Antarctic assimilate organic solutes and preferentially ingest bacteria, whereas they actively exclude phytoplankton. In regions where phytoplankton biomass is temporally limited by light or nutrient concentrations, the growth and development of planktotrophic larvae may not be directly coupled to phytoplankton production.

Radioisotopic method for measuring cell division rates of individual species of diatoms from natural populations.

Silicon is an essential element for diatom frustule synthesis and is usually taken up only by dividing cells. With Ge, a radioactive analog of Si, the cell cycle marker event of frustule formation was identified for individual species of diatom. The frequency of cells within a population undergoing this division event was estimated, and the cell division rate was calculated. In laboratory cultures, these rates of cell division and those calculated from changes in cell numbers were similar. By dual labeling with Ge(OH)(4) and NaHCO(3), rates of cell division and photosynthesis were coincidently measured for diatoms both in laboratory cultures and when isolated from natural populations in estuarine, offshore, and polar environments. These techniques permit the coupling between photosynthesis and cell division to be examined in situ for individual species of diatom.

Phytoplankton division rates in light-limited environments: two adaptations.

Red tide-forming dinoflagellates maximize cell numbers during periods of low light intensities in two ways. For short-term exposures to suboptimal light intensities such as might occur during recirculation in frontal convergences, cell division rates can be maintained at the expense of stored carbon for up to two generation times. During longer periods, corresponding to subsurface transport below a pycnocline, cell division rates eventually decrease as a portion of the fixed carbon is diverted to replenishing stored carbon. As a result, maximum rates of cell division can be resumed rapidly upon advection into surface waters where light intensities are optimal for growth.