The Loroxanthin cycle: A new type of xanthophyll cycle in green algae (Chlorophyta)

Xanthophyll cycles have proven to be major contributors to photoacclimation for many organisms. This work describes a light-driven xanthophyll cycle operating in the chlorophyte Chlamydomonas reinhardtii and involving the xanthophylls Lutein (L) and Loroxanthin (Lo). Pigments were quantified during a switch from high to low light and at different time points from cells grown in Day/night cycle. Trimeric LHCII was purified from cells acclimated to high or low light and their pigment content and spectroscopic properties were characterized. The Lo/(L+Lo) ratio in the cells varies by a factor of 10 between cells grown in low or high light leading to a change in the Lo/(L+Lo) ratio in trimeric LHCII from 0.5 in low light to 0.07 in high light. Trimeric LhcbMs binding Loroxanthin have 5+/-1% higher excitation energy transfer from carotenoid to Chlorophyll as well as higher thermo- and photostability than trimeric LhcbMs that only bind Lutein. The Loroxanthin cycle operates on long time scales (hours to days) and likely evolved as a shade adaptation. It has many similarities with the Lutein-epoxide - Lutein cycle of plants.

Phlorotannin and Pigment Content of Native Canopy-Forming Sargassaceae Species Living in Intertidal Rockpools in Brittany (France): Any Relationship with Their Vertical Distribution and Phenology?

Five native Sargassaceae species from Brittany (France) living in rockpools were surveyed over time to investigate photoprotective strategies according to their tidal position. We gave evidences for the existence of a species distribution between pools along the shore, with the most dense and smallest individuals in the highest pools. Pigment contents were higher in lower pools, suggesting a photo-adaptive process by which the decreasing light irradiance toward the low shore was compensated by a high production of pigments to ensure efficient photosynthesis. Conversely, no xanthophyll cycle-related photoprotective mechanism was highlighted because high levels of zeaxanthin rarely occurred in the upper shore. Phlorotannins were not involved in photoprotection either; only some lower-shore species exhibited a seasonal trend in phlorotannin levels. The structural complexity of phlorotannins appears more to be a taxonomic than an ecological feature: Ericaria produced simple phloroglucinol while Cystoseira and Gongolaria species exhibited polymers. Consequently, tide pools could be considered as light-protected areas on the intertidal zone, in comparison with the exposed emerged substrata where photoprotective mechanisms are essential.

The fine-tuning of NPQ in diatoms relies on the regulation of both xanthophyll cycle enzymes

Diatoms possess an efficient mechanism to dissipate photons as heat in conditions of excess light, which is visualized as the Non-Photochemical Quenching of chlorophyll a fluorescence (NPQ). In most diatom species, NPQ is proportional to the concentration of the xanthophyll cycle pigment diatoxanthin formed from diadinoxanthin by the diadinoxanthin de-epoxidase enzyme. The reverse reaction is performed by the diatoxanthin epoxidase. Despite the xanthophyll cycle’s central role in photoprotection, its regulation is not yet well understood. The proportionality between diatoxanthin and NPQ allowed us to calculate the activity of both xanthophyll cycle enzymes in the model diatom Phaeodactylum tricornutum from NPQ kinetics. From there, we explored the light-dependency of the activity of both enzymes. Our results demonstrate that a tight regulation of both enzymes is key to fine-tune NPQ: (i) the rate constant of diadinoxanthin de-epoxidation is low under a light-limiting regime but increases as photosynthesis saturates, probably due to the thylakoidal proton gradient ΔpH (ii) the rate constant of diatoxanthin epoxidation exhibits an optimum under low light and decreases in the dark due to an insufficiency of the co-factor NADPH as well as in higher light through an as yet unresolved inhibition mechanism, that is unlikely to be related to the ΔpH. We observed that the suppression of NPQ by an uncoupler was due to an accelerated diatoxanthin epoxidation enzyme rather than to the usually hypothesized inhibition of the diadinoxanthin de-epoxidation enzyme.

Autumn Warming Delays the Downregulation of Photosynthesis and Does Not Increase the Risk of Freezing Damage in Interior and Coastal Douglas-fir

During autumn, evergreen conifers utilize the decrease in daylength and temperature as environmental signals to trigger cold acclimation, a process that involves the downregulation of photosynthesis, upregulation of photoprotection, and development of cold hardiness. Global warming will delay the occurrence of autumn low temperatures while daylength remains unaffected. The impact of autumn warming on cold acclimation and the length of the carbon uptake period of species with ranges that encompass diverse climates, such as Douglas-fir (Pseudotsuga menziesii), remains unclear. Our study investigated intraspecific variation in the effects of autumn warming on photosynthetic activity, photosynthetic pigments, and freezing tolerance in two interior (var. glauca) and two coastal (var. menziesii) Douglas-fir provenances. Following growth under simulated summer conditions with long days (16 h photoperiod) and summer temperatures (22/13°C day/night), Douglas-fir seedlings were acclimated to simulated autumn conditions with short days (8 h photoperiod) and either low temperatures (cool autumn, CA; 4/−4°C day/night) or elevated temperatures (warm autumn, WA; 19/11°C day/night). Exposure to low temperatures in the CA treatment induced the downregulation of photosynthetic carbon assimilation and photosystem II efficiency, increased the size and de-epoxidation of the xanthophyll cycle pigment pool, and caused the development of sustained nonphotochemical quenching (NPQ). Seedlings in the WA treatment exhibited no downregulation of photosynthesis, no change in xanthophyll cycle pigment de-epoxidation, and no development of sustained NPQ. Albeit these changes, freezing tolerance was not impaired under WA conditions compared with CA conditions. Interior Douglas-fir seedlings developed greater freezing tolerance than coastal seedlings. Our findings suggest that autumn warming, i.e., short photoperiod alone, does not induce the downregulation of photosynthesis in Douglas-fir. Although autumn warming delays the downregulation of photosynthesis, the prolonged period of photosynthetic activity does not bear a trade-off of impaired freezing tolerance.

Photosynthetic and Photoprotective Responses to Steady-State and Fluctuating Light in the Shade-Demanding Crop Amorphophallus xiei Grown in Intercropping and Monoculture Systems

Photosynthetic and photoprotective responses to simulated sunflecks were examined in the shade-demanding crop Amorphophallus xiei intercropped with maize (intercropping condition) or grown in an adjacent open site (monoculture condition). Both intercropping leaves and monoculture leaves exhibited very fast induction responses. The times taken to achieve 90% maximum net photosynthetic rate in intercropping leaves and monoculture leaves were 198.3 ± 27.4 s and 223.7 ± 20.5 s during the photosynthetic induction, respectively. During an 8-min simulated sunfleck, the proportion of excess excited energy dissipated through the xanthophyll cycle-dependent pathway (ΦNPQ) and dissipated through constitutive thermal dissipation and the fluorescence (Φf, d) pathway increased quickly to its maximum, and then plateaued slowly to a steady state in both intercropping and monoculture leaves. When the illumination was gradually increased within photosystem II (PSII), ΦNPQ increased quicker and to a higher level in monoculture leaves than in intercropping leaves. Relative to their monoculture counterparts, intercropping leaves exhibited a significantly lower accumulation of oxygen free radicals, a significantly higher content of chlorophyll, and a similar content of malondialdehyde. Although monoculture leaves exhibited a larger mass-based pool size of xanthophyll cycle [V (violaxanthin) + A (antheraxanthin) + Z (zeaxanthin)] than intercropping leaves, intercropping leaves had a higher ratio of (Z + A)/(V + Z + A) than monoculture leaves. intercropping leaves had markedly higher glutathione content and ascorbate-peroxidase activity than their monoculture counterparts. Similar activities of catalase, peroxidase, dehydroascorbate reductase, and monodehydroascorbate were found in both systems. Only superoxide dismutase activity and ascorbate content were lower in the intercropping leaves than in their monoculture counterparts. Overall, the xanthophyll cycle-dependent energy dissipation and the enzymatic antioxidant defense system are important for protecting plants from photooxidation in an intercropping system with intense sunflecks.

Synergistic adaptations: freezing tolerance is associated with desiccation tolerance and activation of violaxanthin de-epoxidase in wintergreen ferns

This article comments on: Fernández-Marín B, Arzac MI, López-Pozo M, Laza JM, Roach T, Stegner M, Neuner G, García-Plazaola JI. 2021. Frozen in the dark: interplay of night-time activity of xanthophyll cycle, xylem attributes, and desiccation tolerance in fern resistance to winter. Journal of Experimental Botany 72, 3168–3184.