Phytoplankton in high-light, low-nutrient ocean environments are challenged with maintaining high photosynthetic efficiency and simultaneously preventing photodamage that results from low levels of electron acceptors downstream of photosystem II (PSII). Here, we identify a process in open ocean picophytoplankton that preserves PSII activity by diverting electrons from the photosystem I (PSI) complex-mediated carbon assimilation to oxygen via a propyl gallate-sensitive oxidase associated with the photosynthetic electron transport chain. This process stabilizes diel photochemical efficiency of PSII, despite midday photoinhibition, by maintaining oxidized PSII reaction centers. Although measurements of the maximum photochemical efficiency of PSII, Fv: Fmshow midday photoinhibition, midday CO2fixation is not depressed. Moreover, CO2fixation saturates at low irradiances even though PSII electron flow is not saturated at irradiances of 1,985 μmol photons m-2s-1. This disparity between PSII fluorescence and CO2fixation is consistent with the activity of an oxidase that serves as a terminal electron acceptor, maintaining oxidized PSII reaction centers even when CO2fixation has saturated and the total number of functional reaction centers decreases because of photoinhibition (reflected in lower midday Fv: Fmvalues). This phenomenon is less apparent in coastal phytoplankton populations, suggesting that it is a strategy particularly distinctive of phytoplankton in the oligotrophic ocean. Spatial variability in features of photosynthetic electron flow could explain biogeographical differences in productivity throughout the ocean and should be represented in models that use empirical photosynthesis and chlorophyll fluorescence measurements from a limited number of ocean sites to estimate the productivity of the entire ocean. © 2008, by the American Society of Limnology and Oceanography, Inc.

The spring bloom in seasonally stratified seas is often characterized by a rapid increase in photosynthetic biomass. To clarify how the combined effects of nutrient and light availability influence phytoplankton composition in the oligotrophic Gulf of Aqaba, Red Sea, phytoplankton growth and acclimation responses to various nutrient and light regimes were recorded in three independent bioassays and during a naturally-occurring bloom. We show that picoeukaryotes and Synechococcus maintained a "bloomer" growth strategy, which allowed them to grow quickly when nutrient and light limitation were reversed. During the bloom picoeukaryotes and Synechococcus appeared to have higher P requirements relative to N, and were responsible for the majority of photosynthetic biomass accumulation. Following stratification events, populations limited by light showed rapid photoacclimation (based on analysis of cellular fluorescence levels and photosystem II photosynthetic efficiency) and community composition shifts without substantial changes in photosynthetic biomass. The traditional interpretation of "bloom" dynamics (i.e., as an increase in photosynthetic biomass) may therefore be confined to the upper euphotic zone where light is not limiting, while other acclimation processes are more ecologically relevant at depth. Characterizing acclimation processes and growth strategies is important if we are to clarify mechanisms that underlie productivity in oligotrophic regions, which account for approximately half of the global primary production in the ocean. This information is also important for predicting how phytoplankton may respond to global warming-induced oligotrophic ocean expansion. © 2009 Springer-Verlag.

Mitigating further anthropogenic changes to the global climate will require reducing greenhouse-gas emissions ("abatement"), or else removing carbon dioxide from the atmosphere and/or diminishing solar input ("climate engineering"). Here, we develop and apply criteria to measure technical, economic, ecological, institutional, and ethical dimensions of, and public acceptance for, climate engineering strategies; provide a relative rating for each dimension; and offer a new interdisciplinary framework for comparing abatement and climate engineering options. While abatement remains the most desirable policy, certain climate engineering strategies, including forest and soil management for carbon sequestration, merit broad-scale application. Other proposed strategies, such as biochar production and geological carbon capture and storage, are rated somewhat lower, but deserve further research and development. Iron fertilization of the oceans and solar radiation management, although cost-effective, received the lowest ratings on most criteria. We conclude that although abatement should remain the central climate-change response, some low-risk, cost-effective climate engineering approaches should be applied as complements. The framework presented here aims to guide and prioritize further research and analysis, leading to improvements in climate engineering strategies. © The Ecological Society of America.

Two alternative mechanisms are suggested for nitrite accumulation in the oxygenated oligotrophic water column: (1) excretion by phytoplankton or (2) microbial oxidation of ammonium (nitrification). This study assessed the role of these 2 mechanisms, based on seasonal and high-resolution diurnal depth profiles of the dissolved inorganic nitrogen (DIN) species (nitrite, ammonium, nitrate) and chlorophyll a in the Gulf of Aqaba, Red Sea. Both mechanisms operated in the water column, but in different seasons; nitrification was the prime process responsible for nitrite accumulation during the stratified summer season and phytoplankton nitrite excretion operated during winter mixing. At the onset of summer stratification two N peaks developed below the photic zone, an ammonium maximum (AM) and below it the primary nitrite maximum (PNM). Both peaks were located at a depth range where phytoplankton are thought to be inactive and not excreting nitrite. During summer stratification, the water column deep chlorophyll maximum (DCM), AM, PNM and the nitracline were ordered by a downward increase in N oxidation state similar to the temporal order of the N-species during nitrification. This similarity, together with the diurnal stability of the PNM and its co-existence with oscillating chlorophyll profiles above the DCM, is consistent with nitrification as the key process forming the PNM. We suggest that transport and reaction control the vertical order and separation of N-species in the water column. The ratios between the rate constants for ammonification, ammonium oxidation, nitrite oxidation and nitrate assimilation were estimated by a simple box model to be 1:3:1.5:0.15, respectively. These field estimates are similar to the ratios between the rate constants measured in laboratory experiments.

Cyanobacteria dominate the world's oceans where iron is often barely detectable. One manifestation of low iron adaptation in the oligotrophic marine environment is a decrease in levels of iron-rich photosynthetic components, including the reaction center of photosystem I and the cytochrome b6f complex [R.F. Strzepek and P.J. Harrison, Photosynthetic architecture differs in coastal and oceanic diatoms, Nature 431 (2004) 689-692.]. These thylakoid membrane components have well characterised roles in linear and cyclic photosynthetic electron transport and their low abundance creates potential impediments to photosynthetic function. Here we show that the marine cyanobacterium Synechococcus WH8102 exhibits significant alternative electron flow to O2, a potential adaptation to the low iron environment in oligotrophic oceans. This alternative electron flow appears to extract electrons from the intersystem electron transport chain, prior to photosystem I. Inhibitor studies demonstrate that a propyl gallate-sensitive oxidase mediates this flow of electrons to oxygen, which in turn alleviates excessive photosystem II excitation pressure that can often occur even at relatively low irradiance. These findings are also discussed in the context of satisfying the energetic requirements of the cell when photosystem I abundance is low. © 2008 Elsevier B.V. All rights reserved.