The increases in atmospheric pCO 2 over the last century are accompanied by higher concentrations of CO 2 (aq) in the surface oceans. This acidification of the surface ocean is expected to influence aquatic primary productivity and may also affect cyanobacterial nitrogen (N)-fixers (diazotrophs). No data is currently available showing the response of diazotrophs to enhanced oceanic CO 2 (aq). We examined the influence of pCO 2 [preindustrial $ 250 ppmv (low), ambient $ 400, future $ 900 ppmv (high)] on the photosynthesis, N fixation, and growth of Trichodesmium IMS101. Trichodesmium spp. is a bloom-forming cyanobacterium contributing substantial inputs of 'new N' to the oligotrophic subtropical and tropical oceans. High pCO 2 enhanced N fixation, C : N ratios, filament length, and biomass of Trichodesmium in comparison with both ambient and low pCO 2 cultures. Photosynthesis and respiration did not change significantly between the treatments. We suggest that enhanced N fixation and growth in the high pCO 2 cultures occurs due to reallocation of energy and resources from carbon concentrating mechanisms (CCM) required under low and ambient pCO 2 . Thus, in oceanic regions, where light and nutrients such as P and Fe are not limiting, we expect the projected concentrations of CO 2 to increase N fixation and growth of Trichodesmium. Other diazotrophs may be similarly affected, thereby enhancing inputs of new N and increasing primary productivity in the oceans.
Summary• As iron (Fe) deficiency is a main limiting factor of ocean productivity, its effects were investigated on interactions between photosynthesis and nitrogen fixation in the marine nonheterocystous diazotrophic cyanobacterium Trichodesmium IMS101.• Biophysical methods such as fluorescence kinetic microscopy, fast repetition rate (FRR) fluorimetry, and in vivo and in vitro spectroscopy of pigment composition were used, and nitrogenase activity and the abundance of key proteins were measured.• Fe limitation caused a fast down-regulation of nitrogenase activity and protein levels. By contrast, the abundance of Fe-requiring photosystem I (PSI) components remained constant. Total levels of phycobiliproteins remained unchanged according to single-cell in vivo spectra. However, the regular 16-kDa phycoerythrin band decreased and finally disappeared 16-20 d after initiation of Fe limitation, concomitant with the accumulation of a 20-kDa protein cross-reacting with the phycoerythrin antibody. Concurrently, nitrogenase expression and activity increased. Fe limitation dampened the daily cycle of photosystem II (PSII) activity characteristic of diazotrophic Trichodesmium cells. Further, it increased the number and prolonged the time period of occurrence of cells with elevated basic fluorescence (F 0 ). Additionally, it increased the effective cross-section of PSII, probably as a result of enhanced coupling of phycobilisomes to PSII, and led to up-regulation of the Fe stress protein IsiA.• Trichodesmium survives short-term Fe limitation by selectively down-regulating nitrogen fixation while maintaining but re-arranging the photosynthetic apparatus. Abbreviations:Chl, chlorophyll; FKM, fluorescence kinetic microscopy/microscope (for twodimensional (imaging) measurements of fluorescence kinetics); FRRf, fast repetition rate fluorimeter; F 0 , basic fluorescence yield of a dark-adapted sample, fluorescence in nonactinic measuring light; F m , maximum fluorescence yield of a dark-adapted sample; , maximum fluorescence yield of a sample during exposure to actinic light, i.e. diminished by nonphotochemical quenching; , maximum fluorescence yield of a fully light-adapted sample at the end of the actinic light period of the measurement, diminished by nonphotochemical quenching; , fluorescence yield under actinic irradiance immediately before the measurement of ; F v , variable fluorescence yield; F v = F m − F 0 , i.e. response to a supersaturating flash in the dark-adapted state of photosystem II (PSII); F v /F m , maximal efficiency of dark-adapted PSII (In this study,
The in vivo substitution of Mg2+ in chlorophyll by heavy metals is an important damage mechanism in heavy metal-stressed plants that leads to an inhibition of photosynthesis. In photosynthetic organisms with LHC II antennae, the in vivo substitution of Mg2+ by Cu2+ occurs particularly readily under low irradiance with a dark phase — a phenomenon referred to as 'shade reaction'. In the present study the limiting steps of the shade reaction were investigated with synchronised cultures of the chlorococcal green alga Scenedesmus quadricauda (Turp.) Bréb. The rate of copper chlorophyll formation during shade reaction was shown to be controlled by several factors; firstly, in some phases of the cell cycle, especially at the end of the light period, Mg2+ in chlorophyll was not accessible to substitution. This pattern is likely to be caused by cell cycle-dependent changes in photosynthesis and thylakoid ultrastructure, which were published earlier and are reconsidered in the discussion of the present work. Secondly, prolonged culture in a medium containing 3 μM Cu2+ reversibly increased the resistance of the strain to Cu2+. Culturing without added Cu2+ lowered the threshold concentrations of various deleterious effects more than 10-fold within 8 months of de-adaptation. Adaptation to high Cu2+ levels is discussed in the context of studies of the regulation of metal transporter proteins. In addition, it was also observed that toxic Cu2+ levels impaired photosynthesis sooner than cell division.
Oxygen-evolving photosystem II particles (DT 20) isolated from pea chloroplasts by digitonin-Triton X-100 fractionation were photoinhibited with 150 W. m(-2) white light, at 20°C under three conditions: aerobic, anaerobic and strongly reducing (E(h) poised to approx. -250 mV with dithionite). Hill reaction rate (H (20) + BQ)and variable fluorescence (Fv) declined in parallel in all three cases with shortening half times: 30, 10 and 2.5 min, respectively. Light-induced absorbance changes at 685 nm characteristic of reversible photo accumulation of reduced pheophytin (& z -2.50 mV) remained essentially unchanged. We conclude that the three types of photoinhibitory treatment do not impair the separation of charges between chlorophyll P-680 and pheophytin in the photosystem II reaction center.
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