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We provide here an overview of the remarkable life and outstanding research of David (Dave) Charles Fork (March 4, 1929–December 13, 2021) in oxygenic photosynthesis. In the words of the late Jack Edgar Myers, he was a top ‘photosynthetiker’. His research dealt with novel findings on light absorption, excitation energy distribution, and redistribution among the two photosystems, electron transfer, and their relation to dynamic membrane change as affected by environmental changes, especially temperature. David was an attentive listener and a creative designer of experiments and instruments, and he was also great fun to work with. He is remembered here by his family, coworkers, and friends from around the world including Australia, France, Germany, Japan, Sweden, Israel, and USA.

Main conclusion: Decreased PG constrains PSI activity due to inhibition of transcript and polypeptide abundance of light-harvesting and reaction center polypeptides generating a reversible, yellow phenotype during cold acclimation of pgp1.

Cold acclimation of the Arabidopsis pgp1 mutant at 5 °C resulted in a pale-yellow phenotype with abnormal chloroplast ultrastructure compared to its green phenotype upon growth at 20 °C despite a normal cold-acclimation response at the transcript level. In contrast, wild type maintained its normal green phenotype and chloroplast ultrastructure irrespective of growth temperature. In contrast to cold acclimation of WT, growth of pgp1 at 5 °C limited the accumulation of Lhcbs and Lhcas assessed by immunoblotting. However, a novel 43 kD polypeptide of Lhcb1 as well as a 29 kD polypeptide of Lhcb3 accumulated in the soluble fraction which was absent in the thylakoid membrane fraction of cold-acclimated pgp1 which was not observed in WT. Cold acclimation of pgp1 destabilized the Chl–protein complexes associated with PSI and predisposed energy distribution in favor of PSII rather than PSI compared to the WT. Functionally, in vivo PSI versus PSII photochemistry was inhibited in cold-acclimated pgp1 to a greater extent than in WT relative to controls. Greening of the pale-yellow pgp1 was induced when cold-acclimated pgp1 was shifted from 5 to 20 °C which resulted in a marked decrease in excitation pressure to a level comparable to WT. Concomitantly, Lhcbs and Lhcas accumulated with a simultaneous decrease in the novel 43 and 29kD polypeptides. We conclude that the reduced levels of phosphatidyldiacylglycerol in the pgp1 limit the capacity of the mutant to maintain the structure and function of its photosynthetic apparatus during cold acclimation. Thus, maintenance of normal thylakoid phosphatidyldiacylglycerol levels is essential to stabilize the photosynthetic apparatus during cold acclimation.

To finish this special issue, some friends, colleagues and students of Prof. Chow (Emeritus Professor, the Research School of Biology, the Australian National University) have written small tributes to acknowledge not only his eminent career but to describe his wonderful personality.

High latitude forests will experience large changes in temperature and CO₂ concentrations this century. We evaluated the effects of future climate conditions on 2 dominant boreal tree species, Pinus sylvestris L. and Picea abies (L.) H. Karst, exposing seedlings to 3 seasons of ambient (430 ppm) or elevated CO₂ (750 ppm) and ambient temperatures, a + 4 degrees C warming or a + 8 degrees C warming. Pinus sylvestris responded positively to warming: seedlings developed a larger canopy, maintained high net CO₂ assimilation rates (A_net), and acclimated dark respiration (R_dark). In contrast, carbon fluxes in Picea abies were negatively impacted by warming: maximum rates of A_net decreased, electron transport was redirected to alternative electron acceptors, and thermal acclimation of R_dark was weak. Elevated CO₂ tended to exacerbate these effects in warm-grown Picea abies, and by the end of the experiment Picea abies from the +8 degrees C, high CO₂ treatment produced fewer buds than they had 3 years earlier. Treatments had little effect on leaf and wood anatomy. Our results highlight that species within the same plant functional type may show opposite responses to warming and imply that Picea abies may be particularly vulnerable to warming due to low plasticity in photosynthetic and respiratory metabolism.

Bibliometric impact analyses show that Swedish research has less international visibility than Danish research. When taking a global view on all subject fields and selecting publications cited higher than the 90th percentile, i.e., the Top 10 %—publications, the Swedish Research Council shows that although Sweden ranks 15 % above world average, Denmark, the Netherlands and Switzerland rank 35–40 % above. To explain these different performances, The Royal Swedish Academy of Sciences asked us to compare the national research systems on three levels: priority setting at national level, governance of universities and direction and funding of research. There are of course many similarities between the Danish and Swedish research systems but there are still subtle differences that have developed over time, which may explain the different international visibility. First of all, it does not depend on different levels of public spending on research and development. However, the core funding of universities relative external funding is higher in Denmark than in Sweden. The academic leadership of Danish universities in terms of board, vice-chancellor, faculty dean and department chair is also more coherent and focused on priority setting, recruitment, organization and deployment of resources to establish research environments that operate at the forefront of international research. On all these points we see a weaker leadership in Sweden. Furthermore, over the last 20 years, public funding of research in Sweden has become more and more unpredictable and program oriented with many new actors, while the Danish funding system, although it also has developed over time, shows more consistency with strong actors to fund individuals with novel ideas. The research policy in Sweden has also developed multiple, sometimes even conflicting goals, which have undermined conditions for high-impact research, while in Denmark a policy to support excellence in research has been more coherent.

Exposure of control (non-hardened) Arabidopsis leaves to high light stress at 5 A degrees C resulted in a decrease of both photosystem II (PSII) (45 %) and Photosystem I (PSI) (35 %) photochemical efficiencies compared to non-treated plants. In contrast, cold-acclimated (CA) leaves exhibited only 35 and 22 % decrease of PSII and PSI photochemistry, respectively, under the same conditions. This was accompanied by an accelerated rate of P700(+) re-reduction, indicating an up-regulation of PSI-dependent cyclic electron transport (CET). Interestingly, the expression of the NDH-H gene and the relative abundance of the Ndh-H polypeptide, representing the NDH-complex, decreased as a result of exposure to low temperatures. This indicates that the NDH-dependent CET pathway cannot be involved and the overall stimulation of CET in CA plants is due to up-regulation of the ferredoxin-plastoquinone reductase, antimycin A-sensitive CET pathway. The lower abundance of NDH complex also implies lower activity of the chlororespiratory pathway in CA plants, although the expression level and overall abundance of the other well-characterized component involved in chlororespiration, the plastid terminal oxidase (PTOX), was up-regulated at low temperatures. This suggests increased PTOX-mediated alternative electron flow to oxygen in plants exposed to low temperatures. Indeed, the estimated proportion of O-2-dependent linear electron transport not utilized in carbon assimilation and not directed to photorespiration was twofold higher in CA Arabidopsis. The possible involvement of alternative electron transport pathways in inducing greater resistance of both PSII and PSI to high light stress in CA plants is discussed.

Given its unique function in light-induced water oxidation and its susceptibility to photoinactivation during photosynthesis, photosystem II (PS II) is often the focus of studies of photosynthetic structure and function, particularly in environmental stress conditions. Here we review four approaches for quantifying or monitoring PS II functionality or the stoichiometry of the two photosystems in leaf segments, scrutinizing the approximations in each approach. (1) Chlorophyll fluorescence parameters are convenient to derive, but the information-rich signal suffers from the localized nature of its detection in leaf tissue. (2) The gross O-2 yield per single-turnover flash in CO2-enriched air is a more direct measurement of the functional content, assuming that each functional PS II evolves one O-2 molecule after four flashes. However, the gross O-2 yield per single-turnover flash (multiplied by four) could over-estimate the content of functional PS II if mitochondrial respiration is lower in flash illumination than in darkness. (3) The cumulative delivery of electrons from PS II to P700(+) (oxidized primary donor in PS I) after a flash is added to steady background far-red light is a whole-tissue measurement, such that a single linear correlation with functional PS II applies to leaves of all plant species investigated so far. However, the magnitude obtained in a simple analysis (with the signal normalized to the maximum photo-oxidizable P700 signal), which should equal the ratio of PS II to PS I centers, was too small to match the independently-obtained photosystem stoichiometry. Further, an under-estimation of functional PS II content could occur if some electrons were intercepted before reaching PS I. (4) The electrochromic signal from leaf segments appears to reliably quantify the photosystem stoichiometry, either by progressively photoinactivating PS II or suppressing PS I via photo-oxidation of a known fraction of the P700 with steady far-red light. Together, these approaches have the potential for quantitatively probing PS II in vivo in leaf segments, with prospects for application of the latter two approaches in the field.

Exposure of wild type (WT) and plastocyanin coding petE gene deficient mutant (Delta petE) of Synechococcus cells to low iron growth conditions was accompanied by similar iron-stress induced blue-shift of the main red Chl a absorption peak and a gradual decrease of the Phc/Chl ratio, although Delta petE mutant was more sensitive when exposed to iron deficient conditions. Despite comparable iron stress induced phenotypic changes, the inactivation of petE gene expression was accompanied with a significant reduction of the growth rates compared to WT cells. To examine the photosynthetic electron fluxes in vivo, far-red light induced P700 redox state transients at 820 nm of WT and Delta petE mutant cells grown under iron sufficient and iron deficient conditions were compared. The extent of the absorbance change (Delta A(820)/A(820)) used for quantitative estimation of photooxidizable P700(+) indicated a 2-fold lower level of P700(+) in Delta petE compared to WT cells under control conditions. This was accompanied by a 2-fold slower re-reduction rate of P700(+) in the Delta petE indicating a lower capacity for cyclic electron flow around PSI in the cells lacking plastocyanin. Thermoluminescence (TL) measurements did not reveal significant differences in PSII photochemistry between control WT and Delta petE cells. However, exposure to iron stress induced a 4.5 times lower level of P700(+), 2-fold faster re-reduction rate of P700(+) and a temperature shift of the TL peak corresponding to S-2/S(3)Q(B)(-) charge recombination in WT cells. In contrast, the iron-stressed Delta petE mutant exhibited only a 40% decrease of P700(+) and no significant temperature shift in S-2/S(3)Q(B)(-) charge recombination. The role of mobile electron carriers in modulating the photosynthetic electron fluxes and physiological acclimation of cyanobacteria to low iron conditions is discussed. This article is part of a Special Issue entitled: Photosynthesis Research for Sustainability: from Natural to Artificial. (C) 2012 Elsevier B.V. All rights reserved.

Thermo luminescence (TL) of photosynthetic membranes was discovered by William Arnold and Helen Sherwood in 1957. In the last half century, several studies have elucidated the mechanism of TL emission, which showed that the recombination of different charge pairs generated and trapped during pre-illumination are responsible for the observed light emission. Since most of the TL bands originate within Photosystem II (PS II), the technique of TL has become a useful complementary tool to chlorophyll a fluorescence to probe subtle changes in PS II photochemistry. The technique is simple and non-invasive; it has been successfully used to study leaf, cells, thylakoids and even reaction center preparations. The TL technique provides quick information about the redox potential changes of the bound primary quinone (Q(A)) and the secondary quinone (Q(B)) acceptors of PS II; TL has been extensively used to study the effects of photoinhibition, mutations, stresses and myriad responses of the photosynthetic apparatus during acclimation and adaptation. This chapter reviews crucial evidence for the identification of charge pairs responsible for the generation of different TL bands; the relationship of these bands to the components of delayed light emission; responses to excitation pressure arising out of environmental factors; methodology, and instrumentation. A model based on the detailed analysis of the redox shifts of the PS II electron acceptors Q(A) and Q(B), explaining the possibility of non-radiative dissipation of excess light energy within the reaction center of PS II (reaction center quenching) and its physiological significance in photoprotection of the photosynthetic membranes has been suggested. Developments in the analysis of biophysical parameters and the non-adherence of photosynthetic TL to the analysis by the 1945 theory of J.T. Randall and M.H.F. Wilkins have been briefly reviewed.

Second year needles of Lodgepole pine (Pinus contorta L.) were exposed for 6 weeks to either simulated control summer [summer; 25C/250 photon flux denisty (PFD)], autumn (autumn; 15C/250 PFD) or winter conditions (winter; 5C/250 PFD). We report that the proportion of linear electron transport utilized in carbon assimilation (ETRCO2) was 40 lower in both autumn and winter pine when compared with the summer pine. In contrast, the proportion of excess photosynthetic linear electron transport (ETRexcess) not used for carbon assimilation within the total ETRJf increased by 30 in both autumn and winter pine. In autumn pine acclimated to 15C, the increased amounts of excess electrons were directed equally to 21kPa O-2-dependent and 2kPa O-2-dependent alternative electron transport pathways and the fractions of excitation light energy utilized by PSII photochemistry ((PSII)), thermally dissipated through (NPQ) and dissipated by additional quenching mechanism(s) ((f,D)) were similar to those in summer pine. In contrast, in winter needles acclimated to 5C, 60 of photosynthetically generated excess electrons were utilized through the 2kPa O-2-dependent electron sink and only 15 by the photorespiratory (21kPa O-2) electron pathway. Needles exposed to winter conditions led to a 3-fold lower (PSII), only a marginal increase in (NPQ) and a 2-fold higher (f,D), which was O-2 dependent compared with the summer and autumn pine. Our results demonstrate that the employment of a variety of alternative pathways for utilization of photosynthetically generated electrons by Lodgepole pine depends on the acclimation temperature. Furthermore, dissipation of excess light energy through constitutive non-photochemical quenching mechanisms is O-2 dependent.