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Micromonas strains of small prasinophyte green algae are found throughout the world's oceans, exploiting widely different niches. We grew arctic and temperate strains of Micromonas and compared their susceptibilities to photoinactivation of Photosystem II, their counteracting Photosystem II repair capacities, their Photosystem II content, and their induction and relaxation of non-photochemical quenching. In the arctic strain Micromonas NCMA 2099, the cellular content of active Photosystem II represents only about 50 % of total Photosystem II protein, as a slow rate constant for clearance of PsbA protein limits instantaneous repair. In contrast, the temperate strain NCMA 1646 shows a faster clearance of PsbA protein which allows it to maintain active Photosystem II content equivalent to total Photosystem II protein. Under growth at 2 °C, the arctic Micromonas maintains a constitutive induction of xanthophyll deepoxidation, shown by second-derivative whole-cell spectra, which supports strong induction of non-photochemical quenching under low to moderate light, even if xanthophyll cycling is blocked. This non-photochemical quenching, however, relaxes during subsequent darkness with kinetics nearly comparable to the temperate Micromonas NCMA 1646, thereby limiting the opportunity cost of sustained downregulation of PSII function after a decrease in light.

When growth irradiance changes, phytoplank- ton acclimates by changing allocations to cellular compo- nents to re-balance their capacity to absorb photons versus their capacity to use the electrons from the oxidation of water at photosystem II. Published changes in the cellular allocations resulting from photoacclimation across algal groups highlight that algae adopt different strategies. We examined the photoacclimation of the photosynthetic ap- paratus of six marine phytoplankters under near-natural diel irradiance patterns. For most of the phytoplankters, Chl a per structural photosystem II unit decreased with increasing growth irradiance, but a parallel decline in op- tical packaging effect allowed cells to maintain their functional absorption cross section serving active photo- system II units (rPSII). Furthermore, no significant changes were observed in the ratio of Chl a per photosystem I. The diatom Skeletonema marinoi proved an exception to this pattern as Chl a per photosystem II is stable and Chl a per photosystem I slightly decreased with light intensity. A clear decrease in the photosystem content per cell was observed for all species except for Thalassiosira oceanica and S. marinoi. Rubisco content per cell showed little variation with irradiance for most algae, except for a 3-fold increase in S. marinoi .A *700 % increase in the Rubis- co:photosystem ratio across species with increasing growth irradiance indicates this is a key cellular stoichiometric adjustment to balance photon absorption capacity and the carbon reduction capacity. Increasing the Rubisco:photo- system ratio occurs through a decrease in the photosystems per cell for most of the phytoplankters in this study, except in the case of S. marinoi where Rubisco per cell increased.

The diazotrophic cyanobacterium Trichodesmium is a major contributor to marine nitrogen fixation. We analyzed how light acclimation influences the photophysiological performance of Trichodesmium IMS101 during exponential growth in semi-continuous nitrogen fixing cultures under light levels of 70, 150, 250, and 400 μmol photons m−2 s−1, across diel cycles. There were close correlations between growth rate, trichome length, particulate organic carbon and nitrogen assimilation, and cellular absorbance, which all peaked at 150 μmol photons m−2 s−1. Growth rate was light saturated by about 100 μmol photons m−2 s−1 and was photoinhibited above 150 μmol photons m−2 s−1. In contrast, the light level (I k) to saturate PSII electron transport (e − PSII−1 s−1) was much higher, in the range of 450–550 μmol photons m−2 s−1, and increased with growth light. Growth rate correlates with the absorption cross section as well as with absorbed photons per cell, but not to electron transport per PSII; this disparity suggests that numbers of PSII in a cell, along with the energy allocation between two photosystems and the state transition mechanism underlie the changes in growth rates. The rate of state transitions after a transfer to darkness increased with growth light, indicating faster respiratory input into the intersystem electron transport chain.

Changing light environments force photoautotroph cells, including coral symbionts, to acclimate to maintain photosynthesis. Photosystem II (PSII) is subjected to photoinactivation at a rate proportional to the incident light, and cells must adjust their rates of protein repair to counter this photoinactivation. We examined PSII function in the coral symbiont Symbiodinium to determine the effect of photoacclimation on their capacity for PSII repair. Colonies of the coral Stylophora pistillata were collected from moderate light environments on the Lizard Island reef (Queensland, Australia) and transported to a local field station, where they were assigned to lower or higher light regimes and allowed to acclimate for 2 weeks. Following this photoacclimation period, the low-light acclimated corals showed greater symbiont density, higher chlorophyll per symbiont cell, and higher photosystem II protein than high-light acclimated corals did. Subsequently, we treated the corals with lincomycin, an inhibitor of chloroplastic protein synthesis, and exposed them to a high-light treatment to separate the effect of de novo protein synthesis in PSII repair from intrinsic susceptibility to photoinactivation. Low-light acclimated corals showed a sharp initial drop in PSII function but inhibition of PSII repair provoked only a modest additional drop in PSII function, compared to uninhibited corals. In high-light acclimated corals inhibition of PSII repair provoked a larger drop in PSII function, compared to uninhibited high-light corals. The greater lincomycin effects in the corals pre-acclimated to high-light show that high-light leads to an increased reliance on the PSII repair cycle.

All oxygenic photoautotrophs suffer photoinactivation of their Photosystem II complexes, at a rate driven by the instantaneous light level. To maintain photosynthesis, PsbA subunits are proteolytically removed from photoinactivated Photosystem II complexes, primarily by a membrane-bound FtsH protease. Diatoms thrive in environments with fluctuating light, such as coastal regions, in part because they enjoy a low susceptibility to photoinactivation of Photosystem II. In a coastal strain of the diatom Thalassiosira pseudonana growing across a range of light levels, active Photosystem II represents only about 42 % of the total Photosystem II protein, with the remainder attributable to photoinactivated Photosystem II awaiting recycling. The rate constant for removal of PsbA protein increases with growth light, in parallel with an increasing content of the FtsH protease relative to the substrate PsbA. An offshore strain of Thalassiosira pseudonana, originating from a more stable light environment, had a lower content of FtsH and slower rate constants for removal of PsbA. We used this data to generate the first estimates for in vivo proteolytic degradation of photoinactivated PsbA per FtsH(6) protease, at similar to 3.9 x 10(-2) s(-1), which proved consistent across growth lights and across the onshore and offshore strains.

We studied the interactions of the CO2-concentrating mechanism and variable light in the filamentous cyanobacterium Leptolyngbya sp. CPCC 696 acclimated to low light (15 μmol m−2 s−1 PPFD) and low inorganic carbon (50 μM Ci). Mass spectrometric and polarographic analysis revealed that mediated CO2 uptake along with both active Na+-independent and Na+-dependent HCO3 − transport, likely through Na+/HCO3 − symport, were employed to concentrate Ci internally. Combined transport of CO2 and HCO3 − required about 30 kJ mol−1 of energy from photosynthetic electron transport to support an intracellular Ci accumulation 550-fold greater than the external Ci. Initially, Leptolyngbya rapidly induced oxygen evolution and Ci transport to reach 40–50% of maximum values by 50 μmol m−2 s−1 PPFD. Thereafter, photosynthesis and Ci transport increased gradually to saturation around 1,800 μmol m−2 s−1 PPFD. Leptolyngbya showed a low intrinsic susceptibility to photoinhibition of oxygen evolution up to PPFD of 3,000 μmol m−2 s−1. Intracellular Ci accumulation showed a lag under low light but then peaked at about 500 μmol photons m−2 s−1 and remained high thereafter. Ci influx was accompanied by a simultaneous, light-dependent, outward flux of CO2 and by internal CO2/HCO3 − cycling. The high-affinity and high-capacity CCM of Leptolyngbya responded dynamically to fluctuating PPFD and used excitation energy in excess of the needs of CO2 fixation by increasing Ci transport, accumulation and Ci cycling. This capacity may allow Leptolyngbya to tolerate periodic exposure to excess high light by consuming electron equivalents and keeping PSII open.

In this minireview we discuss effects of excitation stress on the molecular organization and function of PS II as induced by high light or low temperature in the cyanobacterium Synechococcus sp. PCC 7942. Synechococcus displays PS II plasticity by transiently replacing the constitutive D1 form (D1:1) with another form (D1:2) upon exposure to excitation stress. The cells thereby counteract photoinhibition by increasing D1 turn over and modulating PS II function. A comparison between the cyanobacterium Synechococcus and plants shows that in cyanobacteria, with their large phycobilisomes, resistance to photoinhibition is mainly through the dynamic properties (D1 turnover and quenching) of the reaction centre. In contrast, plants use antenna quenching in the light-harvesting complex as an important means to protect the reaction center from excessive excitation.

Cyanobacteria acclimate to environmental inorganic carbon (C(i)) concentrations through re-organisations of photosynthetic function and the induction of carbon concentrating mechanisms (CCMs), which alter and constrain their subsequent acclimation to changing light. We grew cells acclimated to high C(i) (4 mM) or low C(i) (0.02 mM), shifted them from 50 micromol m(-2) s(-1) to 500 micromol m(-2) s(-1), and quantified their photosynthetic performance in parallel with quantitation of allocations to key indicator macromolecules. Pigments cell(-1) declined, PsbA (PS II), AtpB (ATP Synthase), RbcL (Rubisco) and GlnA (Glutamine Synthetase) increased, and PsaC (PS I) remained stable through the light shift. The increase in these protein pools was slower and smaller in low C(i) cells, but acted in both cell types to re-normalise the electron fluxes through the catalytic complexes back toward values before the light shift (for PsbA and GlnA) or even below the initial flux per complex (for RbcL). In contrast, an increased electron flux per PsaC was sustained for at least 6 h after the increase in light. Initially, high levels of PS II cell(-1) and PS II connectivity in high C(i) cells caused a more rapid net photoinactivation of PS II in high C(i) cells than in low C(i) cells, depressing the rate of PS II-specific electron transport (PS II ETR) to levels similar to linear ETR (net O(2) evolution minus respiration). In low C(i) cells, PS II ETR remained in excess of linear ETR and may have helped maintain CCM activity. The pool sizes of PsbA, AtpB and GlnA correlated with cellular growth rate, and changed at similar rates in high C(i) and low C(i) cells when expressed on a generational rather than chronological timescale, which has implications for differing ecology of high and low C(i) cells under variable natural light.

Synechococcus sp. PCC 7942 (Anacystis nidulans R2) contains two forms of the Photosystem II reaction centre protein D1, which differ in 25 of 360 amino acids. D1: 1 predominates under low light but is transiently replaced by D1:2 upon shifts to higher light. Mutant cells containing only D1:1 have lower photochemical energy capture efficiency and decreased resistance to photoinhibition, compared to cells containing D1:2. We show that when dark-adapted or under low to moderate light, cells with D1:1 have higher non-photochemical quenching of PS II fluorescence (higher qN) than do cells with D1:2. This is reflected in the 77 K chlorophyll emission spectra, with lower Photosystem II fluorescence at 697–698 nm in cells containing D1:1 than in cells with D1:2. This difference in quenching of Photosystem II fluorescence occurs upon excitation of both chlorophyll at 435 nm and phycobilisomes at 570 nm. Measurement of time-resolved room temperature fluorescence shows that Photosystem II fluorescence related to charge stabilization is quenched more rapidly in cells containing D1:1 than in those with D1:2. Cells containing D1:1 appear generally shifted towards State II, with PS II down-regulated, while cells with D1:2 tend towards State I. In these cyanobacteria electron transport away from PS II remains non-saturated even under photoinhibitory levels of light. Therefore, the higher activity of D1:2 Photosystem II centres may allow more rapid photochemical dissipation of excess energy into the electron transport chain. D1:1 confers capacity for extreme State II which may be of benefit under low and variable light.

In Table 2 of the original publication, all instances of krec in the Parameter and Equation columns should read krecinact.

Among marine phytoplankton groups, diatoms span the widest range of cell size, with resulting effects upon their nitrogen uptake, photosynthesis and growth responses to light. We grew two strains of marine centric diatoms differing by ~4 orders of magnitude in cell biovolume in high (enriched artificial seawater with ~500 µmol L−1 µmol L−1 NO3 −) and lower-nitrogen (enriched artificial seawater with