N. P. A. Huner

Effects of cold acclimation on the susceptibility of photosynthesis to photoinhibition in Scots pine and in winter and spring cereals : a fluorescence analysis

Winter and spring varieties of cereals and seedlings of Scots pine were exposed to a low temperature regime of 4-5-degrees-C for the induction of frost hardiness. The effect of cold acclimation on ...

Temperature/light dependent development of selective resistance to photoinhibition of photosystem I

Exposure of winter rye leaves grown at 20°C and an irradiance of either 50 or 250 μmol m−2 s−1 to high light stress (1600 μmol m−2 s−1, 4 h) at 5°C resulted in photoinhibition of PSI measured in vivo as a 34% and 31% decrease in ΔA820/A820 (P700+). The same effect was registered in plants grown at 5°C and 50 μmol m−2 s−1. This was accompanied by a parallel degradation of the PsaA/PsaB heterodimer, increase of the intersystem e− pool size as well as inhibition of PSII photochemistry measured as Fv/Fm. Surprisingly, plants acclimated to high light (800 μmol m−2 s−1) or to 5°C and moderate light (250 μmol m−2 s−1) were fully resistant to photoinhibition of PSI and did not exhibit any measurable changes at the level of PSI heterodimer abundance and intersystem e− pool size, although PSII photochemistry was reduced to 66% and 64% respectively. Thus, we show for the first time that PSI, unlike PSII, becomes completely resistant to photoinhibition when plants are acclimated to either 20°C/800 μmol m−2 s−1 or 5°C/250 μmol m−2 s−1 as a response to growth at elevated excitation pressure. The role of temperature/light dependent acclimation in the induction of selective tolerance to PSI photoinactivation is discussed.

Abscisic acid induced protection against photoinhibition of PSII correlates with enhanced activity of the xanthophyll cycle

The exogenous application of abscisic acid (ABA) to barley seedlings resulted in partial protection of the PSII photochemistry against photoinhibition at low temperature, the effect being most pronounced at 10−5 M ABA. This was accompanied by higher photochemical quenching (qP) in ABA‐treated leaves. A considerable increase (122%) in the amount of total carotenoids and xanthophylls (antheraxanthin, violaxanthin and zeaxanthin) was also found in the seedlings subjected to ABA. The activity of the xanthophyll cycle measured by the epoxidation state of xanthophylls under high‐light treatment was higher in ABA‐treated plants compared with the control. This corresponds to a higher value (0.411) of non‐photochemical quenching (qNP) observed in ABA‐treated than in control (0.306) leaves.

Photosynthetic electron transport adjustments in overwintering Scots pine (Pinus sylvestris L.)

Abstract. As shown before [C. Ottander et al. (1995) Planta 197:176–183], there is a severe inhibition of the photosystem (PS) II photochemical efficiency of Scots pine (Pinus sylvestris L.) during the winter. In contrast, the in vivo PSI photochemistry is less inhibited during winter as shown by in vivo measurements of ΔA820/A820 (P700+). There was also an enhanced cyclic electron transfer around PSI in winter-stressed needles as indicated by 4-fold faster reduction kinetics of P700+. The differential functional stability of PSII and PSI was accompanied by a 3.7-fold higher intersystem electron pool size, and a 5-fold increase in the stromal electron pool available for P700+ reduction. There was also a strong reduction of the QB band in the thermoluminescence glow curve and markedly slower QA– re-oxidation in needles of winter pine, indicating an inhibition of electron transfer between QA and QB. The data presented indicate that the plastoquinone pool is largely reduced in winter pine, and that this reduced state is likely to be of metabolic rather than photochemical origin. The retention of PSI photochemistry, and the suggested metabolic reduction of the plastoquinone pool in winter stressed needles of Scots pine are discussed in terms of the need for enhanced photoprotection of the needles during the winter and the role of metabolically supplied energy for the recovery of photosynthesis from winter stress in evergreens.

Iron stress restricts photosynthetic intersystem electron transport in Synechococcus sp. PCC 7942

Although exposure of Synechococcus sp. PCC 7942 to iron stress induced the accumulation of the isiA gene product (CP43′) compared with non‐stressed controls, immunodetection of the N‐terminus of cytochrome (Cyt) f indicated that iron stress not only reduced the content of the 40 kDa, heme‐binding, Cyt f polypeptide by 32% but it also specifically induced the accumulation of a new, 23 kDa, non‐heme‐binding, putative Cyt f polypeptide. Concomitantly, iron stress restricted intersystem electron transport based on the in vivo reduction of P700+, monitored as ΔA 820/A 820 in the presence and absence of electron transport inhibitors, as well as the inhibition of the Emerson enhancement effect on O2 evolution. However, iron stress appeared to be associated with enhanced rates of PS I cyclic electron transport, low rates of PS I‐driven photoreduction of NADP+ but comparable rates for PS II+PS I photoreduction of NADP+ relative to controls. We hypothesize that Synechococcus sp. PCC 7942 exhibits a dynamic capacity to uncouple PS II and PS I electron transport, which may allow for the higher than expected growth rates observed during iron stress.

Chloroplast biogenesis at cold-hardening temperatures. Kinetics of trans-Δ3-hexadecenoic acid accumulation and the assembly of LHCII.

Etiolated seedlings developed at cold-hardening temperatures (5°C) exhibited etioplasts with considerable vesiculation of internal membranes compared to etioplasts developed at 20°C regardless of the osmotic concentration employed during sample preparation. This vesiculation disappeared during exposure to continuous light at 5°C. This transformation of 5°C and 20°C etioplasts to chloroplasts under continuous light at 5° and 20°C respectively proceeded normally with the initial development of non-appressed lamellae and the subsequent appearance of granal stacks. However, chloroplasts developed at 5°C exhibited fewer lamellae per granum than chloroplasts developed at 20°C.Although the polypeptide complements of etioplasts and chloroplasts developed at 5° or 20°C were not significantly different, monomeric light harvesting complex (LHCII3) was assembled into oligomeric light harvesting complex (LHCII1) during chloroplast biogenesis at 20°C (oligomer:monomer =1.8) whereas monomeric LHCII predominated at 5°C (oligomer:monomer =0.3). Low temperature fluorescence emission spectra of isolated thylakoids indicated that both the F685/F735 and F695/F735 were significantly higher after greening at 5°C than at 20°C. In addition, chloroplast biogenesis at 5°C was associated with a low ratio of trans-Δ3-hexadecenoic acid (0.5) in phosphatidylglycerol whereas at 20°C biogenesis was associated with a high ratio (1.6). Comparative kinetics indicated that the maximization of the trans-Δ3-hexadecenoic acid level precedes the assembly of monomeric LHCII into oligomeric LHCII during biogenesis at 20°C. It is suggested that low developmental temperatures modulate the assembly of LHCII by reducing the trans-Δ3-hexadecenoic acid content of phosphatidylglycerol such that monomeric or some intermediate form of LHCII predominates.

Genetic decrease in fatty acid unsaturation of phosphatidylglycerol increased photoinhibition of photosystem I at low temperature in tobacco leaves.

Leaves of transgenic tobacco plants with decreased levels of fatty acid unsaturation in phosphatidylglycerol (PG) exhibited a slightly lower level of the steady state oxidation of the photosystem I (PSI) reaction center P700 (P700(+)) than wild-type plants. The PSI photochemistry of wild-type plants was only marginally affected by high light treatments. Surprisingly, all plants of transgenic lines exhibited much higher susceptibility to photoinhibition of PSI than wild-type plants. This was accompanied by a 2.5-fold faster re-reduction rate of P700(+) in the dark, indicating a higher capacity for cyclic electron flow around PSI in high light treated transgenic leaves. This was associated with a much higher intersystem electron pool size suggesting over-reduction of the PQ pool in tobacco transgenic lines with altered PG unsaturation compared to wild-type plants. The physiological role of PG unsaturation in PSI down-regulation and modulation of the capacity of PSI-dependent cyclic electron flows and distribution of excitation light energy in tobacco plants under photoinhibitory conditions at low temperatures is discussed. This article is part of a Special Issue entitled: Photosynthesis Research for Sustainability: from Natural to Artificial.

Chloroplast biogenesis at cold-hardening temperatures development of photosystem I and photosystem II activities in relation to pigment accumulation

Chloroplast biogenesis during continuous illumination at either low, cold-hardening temperatures (5°C) or non-hardening temperatures (20°C) was examined by monitoring the etioplast-chloroplast transformation with respect to pigment accumulation and the development of PSI- and PSII-associated electron transport activities in winter rye (Secale cereale L. cv Puma). Generally, chlorophyll and carotenoid accumulation during greening at 20°C were characterized by rapid initial rates in contrast to pronounced, initial lag times during biogenesis at 5°C. Although greening temperature had no effect on the sequential appearance of PSI relative to PSII, greening temperature significantly altered the pattern of appearance of PSI relative to chlorophyll accumulation. Thylakoid biogenesis under continuous illumination at 20°C imposed a pattern whereby the development of PSI activity was antiparallel to chlorophyll accumulation. In contrast, the development of PSI activity under continuous illumination at 5°C was paralllel to chlorophylll accumulation. These developmental patterns were independent of the temperature experienced during etiolation. However, rye seedlings etiolated at 20°C and subsequently subjected to continuous illumination at 5°C exhibited a 70% reduction in the maximum PSII activity (100 μmol DCPIP reduced.mg Chl-1.h-1) attained relative to that observed for similar etiolated seedlings greened at 20°C (300 μmol DCPIP reduced.mg Chl-1.h-1). This low temperature-induced inhibition could be alleviated by an initial 2 h exposure to continuous light at 20°C prior to greening to 5°C. Rye seedlings etiolated at 5°C attained similar maximal PSII activities (300 μmol DCPIP reduced.mg Chl-1.h-1) regardless of the greening temperature. We suggest that the altered kinetics for pigment accumulation, the low temperature-induced change in the pattern for the appearance of PSI activity relative to chlorophyll accumulation and the differential sensitivity of 20° and 5° etiolated seedlings to greening temperature reflect an alteration in membrane organization incurred as a consequence of thylakoid assembly at low temperature.

Restricted capacity for PSI-dependent cyclic electron flow in ΔpetE mutant compromises the ability for acclimation to iron stress in Synechococcus sp. PCC 7942 cells☆

Exposure of wild type (WT) and plastocyanin coding petE gene deficient mutant (ΔpetE) 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 ΔpetE 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 820nm of WT and ΔpetE mutant cells grown under iron sufficient and iron deficient conditions were compared. The extent of the absorbance change (ΔA(820)/A(820)) used for quantitative estimation of photooxidizable P700(+) indicated a 2-fold lower level of P700(+) in ΔpetE compared to WT cells under control conditions. This was accompanied by a 2-fold slower re-reduction rate of P700(+) in the ΔpetE 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 ΔpetE 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 ΔpetE 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.

The lack of LHCII proteins modulates excitation energy partitioning and PSII charge recombination in Chlorina F2 mutant of barley.

Analysis of the partitioning of absorbed light energy within PSII into fractions utilized by PSII photochemistry (ØPSII), thermally dissipated via ΔpH-and zeaxanthin-dependent energy quenching (ØNPQ) and constitutive non-photochemical energy losses (ØNO) was performed in wild type and F2 mutant of barley. The estimated energy partitioning of absorbed light to various pathways indicated that the fraction of ØPSII was slightly higher, while the proportion of thermally dissipated energy through ØNPQ was 38% lower in F2 mutant than in WT. In contrast, ØNO, i.e. the fraction of absorbed light energy dissipated by additional quenching mechanism(s) was 34% higher in F2 mutant. The increased proportion of ØNO correlated with narrowing the temperature gap (ΔTM) between S2/3QB− and S2QA− charge recombinations in F2 mutant as revealed by thermoluminescence measurements. We suggest that this would result in increased probability for an alternative non-radiative P680+QA− radical pair recombination pathway for energy dissipation within the reaction centre of PSII (reaction center quenching) and that this additional quenching mechanism might play an important role in photoprotection when the capacity for the primary, zeaxanthin-dependent non-photochemical quenching (ØNPQ) and state transitions pathways are restricted in the absence of LHCII polypeptides in F2 mutant.