Philip R. van Hasselt

Photoinhibition of photosystem II in vivo is preceded by down-regulation through light-induced acidification of the lumen: Consequences for the mechanism of photoinhibition in vivo

The mechanism of photoinhibition of photosystem II (PSII) was studied in intact leaf discs of Spinacia oleracea L. and detached leaves of Vigna unguiculata L. The leaf material was exposed to different photon flux densities (PFDs) for 100 min, while non-photochemical (qN) and photochemical quenching (qp) of chlorophyll fluorescence were monitored. The ‘energy’ and redox state of PSII were manipulated quite independently of the PFD by application of different temperatures (5–20° C), [CO2] and [O2] at different PFDs. A linear or curvilinear relationship between qp and photoinhibition of PSII was observed. When [CO2] and [O2] were both low (30 μl · l−1 and 2%, respectively), PSII was less susceptible at a given qp than at ambient or higher [CO2] and photoinhibition became only substantial when qp decreased below 0.3. When high levels of energy-dependent quenching (qE) (between 0.6 and 0.8) were reached, a further increase of the PFD or a further decrease of the metabolic demand for ATP and NADPH led to a shift from qE to photoinhibitory quenching (qI). This shift indicated that photoinhibition was preceded by down-regulation through light-induced acidification of the lumen. We propose that photoinhibition took place in the centers down-regulated by qE. The shift from qE to qI occurred concomitant with qP decreasing to zero. The results clearly show that photoinhibition does not primarily depend on the photon density in the antenna, but that photoinhibition depends on the energy state of the membrane in combination with the redox balance of PSII. The results are discussed with regard to the mechanism of photoinhibition of PSII, considering, in particular, effects of light-induced acidification on the donor side of PSII. Interestingly, cold-acclimation of spinach leaves did not significantly affect the relationship between qP, qE and photoinhibition of PSII at low temperature.

Differences in acclimation potential of photosynthesis in seven isolates of the tropical to warm temperate macrophyte Valonia utricularis (Chlorophyta)

A. Eggert, R.J.W. Visser, P.R. Van Hasselt and A.M. Breeman. 2006. Differences in acclimation potential of photosynthesis in seven isolates of the tropical to warm temperate macrophyte Valonia utricularis (Chlorophyta). Phycologia 45: 546–556. DOI: 10.2216/05-03.1 The potential to acclimate photosynthesis to sub- and supra-optimal temperatures was investigated in seven isolates of Valonia utricularis (Roth) C. Agardh, a green macrophyte with a tropical to warm-temperate distribution. Photosynthesis–light response curves were obtained by measuring chlorophyll a fluorescence characteristics of algae grown at optimal (25°C), sub- and supra-optimal temperatures. Suboptimal temperatures were chosen to support 30% of the maximum relative growth rate in each isolate. Thermal acclimation was investigated by comparing short-term and long-term temperature effects on the initial rate of increase of the relative electron transport rate (rETR) and the maximum rETR under light-saturating conditions. Isolates from the northeast Atlantic and the Mediterranean all showed a strong potential to acclimate maximum rETR to suboptimal growth temperatures, i.e. short-term temperature effects were diminished after acclimation. However, photoinhibition, measured as a decrease of the maximal quantum yield (Fv/Fm), was found when plants were grown at 30°C. The isolates reduced light harvesting at 30°C by decreasing total chlorophyll content and by increasing the chlorophyll a/b ratio. Up-regulation of photoprotective processes by the xanthophyll cycle pigments was not observed. In contrast, isolates from the Indo-west Pacific were unable to acclimate photosynthesis to suboptimal growth temperatures and these temperatures were strongly photoinhibiting, even though adjustments on the pigment level were observed. All Indo-west Pacific isolates reached comparable maximum rETR values at 30° and 25°C. Thus, the Atlantic/Mediterranean isolates had a stronger potential to acclimate photosynthetic rates at suboptimal growth temperatures compared to the Indo-west Pacific isolates, which was accompanied by losses at 30°C. The results are discussed in a biogeographical context.

Chilling-induced photoinhibition in nine isolates of Valonia utricularis (Chlorophyta) from different climate regions.

Chilling induced inhibition of photosynthesis was studied in nine isolates of the marine tropical to warm-temperate green macrophyte Valonia utricularis (Roth) C. Agardh. According to their temperature requirements for growth and survival, the isolates belong to a cold-tolerant Atlantic/Mediterranean group and a cold-sensitive Indo-west Pacific group. After 5 hours exposure to 5 degrees C under moderate light, all isolates experienced similar substantial photoinhibition, which approached steady state levels after a decline in Fv/Fm to about 40% of the initial values. After return to optimal temperature and dim light conditions, Fv/Fm values increased with biphasic kinetics. A fast phase with half-life times of less than 30 minutes (dynamic photoinhibition) was followed by a slow phase lasting a few hours, indicating repair of photodamaged PSII reaction centres (chronic photoinhibition). In the Atlantic/Mediterranean isolates the fast phase accounted for more than 80 % of the recovery response, showing that these isolates were able to cope with the applied low temperature stress by down-regulating their PSII reaction centres. In contrast, the two isolates from the Seychelles were predominantly photodamaged. In a second experiment, three isolates (Corsica, Seychelles, Japan) were exposed to a similar relative amount of cold stress (0, 10, 15 degrees C, respectively). The Japanese isolate and the isolate from the Seychelles showed significantly less inhibition compared to 5 degrees C exposure, but no significant difference was found in the Corsican isolate. However, the degree of low temperature stress had no significant influence on the relative contributions of dynamic and chronic photoinhibition. Only two of the seven investigated isolates had a lower final inhibition level when grown at sub-optimal temperatures than at optimal temperatures. However, all sub-optimally grown Atlantic/Mediterranean isolates exhibited faster recovery kinetics from chilling-induced photoinhibition than optimally grown plants. This is related to a faster recovery from chronic photoinhibition than to a higher relative contribution of dynamic photoinhibition. A specific role of the photoprotective pigments of the xanthophyll cycle, leading to an acclimation response in the Atlantic/Mediterranean isolates may be involved. We conclude that ecotypic differentiation in V. utricularis is mirrored in different degrees of susceptibility to low temperature stress.

Low-temperature-related growth and photosynthetic performance of alloplasmic tomato (Lycopersicon esculentum Mill.) with chloroplasts from L. hirsutum Humb. & Bonpl.

Growth and photosynthetic performance were analyzed in alloplasmic tomato at a high- (25/17 °C; HTR) and low-temperature regime (12/6 °C; LTR) in order to establish the role of cytoplasmic variation on low-temperature tolerance of tomato (Lycopersicon esculentum Mill.). Four alloplasmic tomato lines, containing the nuclear genome of tomato and the plastome of L. hirsutum LA 1777 Humb. & Bonpl., an accession collected at high-altitude in Peru, were reciprocally crossed with 11 tomato entries with a high inbreeding level and a wide genetic variation, resulting in a set of 44 reciprocal crosses. Irrespective of growth temperature, alloplasmic families with alien chloroplasts of L. hirsutum (h) were on average characterized by a high shoot biomass, a large leaf area, and a low specific leaf area in comparison with their euplasmic counterparts. These results do not directly point to an advantageous effect of h-chloroplasts on biomass accumulation at low temperature but rather towards a small general beneficial effect on growth and/or distribution of assimilates. Significant chloroplast-related differences in photosynthetic performance, however, were not detected at both temperature regimes, indicating that h-chloroplasts can properly function in a variable nuclear background of L. esculentum. It is concluded that chloroplast substitution is not an effective method for breeding tomato plants with improved low-temperature tolerance.