Dynamics of violaxanthin and lutein epoxide xanthophyll cycles in Lauraceae tree species under field conditions.

Two xanthophyll cycles have been described in higher plants: the violaxanthin xanthophyll (V or VAZ) cycle, which is present in all species, and the taxonomically restricted lutein epoxide xanthophyll (Lx) cycle, which involves the light-induced de-epoxidation of Lx to lutein (L) and its epoxidation back to Lx in low light. Laboratory experiments indicate that the first reaction occurs quickly, but the second reaction is much slower. We investigated the Lx cycle under field conditions in several tree species of the Lauraceae family to determine its relationship with the ubiquitous V cycle. The field study was conducted in two natural laurel forests: one in the Canary Islands, where Laurus azorica (Seub.) Franco, Ocotea foetens (Aiton.) Benth, Apollonias barbujana (Cav.) Bornm. and Persea indica (L.) Spreng were studied; and one in the Basque Atlantic coast where Laurus nobilis L. was studied. The results were complemented by a taxonomic study. The presence of Lx was widespread among Lauraceae species, but its concentration varied even among closely related species. The V pool size correlated positively with growth irradiance, whereas the relationship between Lx pool size and growth irradiance varied with species. A functional Lx cycle was confirmed under field conditions only in O. foetens and L. nobilis. Furthermore, in O. foetens, a correlation between Lx de-epoxidation and photoinhibition suggested a protective role for this cycle. We conclude that, unlike the V cycle, which is normally correlated with irradiance, the operation and light dependence of the Lx cycle is species-dependent.

Functional role of red (retro)-carotenoids as passive light filters in the leaves of Buxus sempervirens L.: increased protection of photosynthetic tissues?

Red (retro)-carotenoids accumulate in chloroplasts of Buxus sempervirens leaves during the process of winter leaf acclimation. As a result of their irregular presence, different leaf colour phenotypes can be found simultaneously in the same location. Five different colour phenotypes (green, brown, red, orange, and yellow), with a distinct pattern of pigment distribution and concentration, have been characterized. Leaf reddening due to the presence of anthocyanins or carotenoids, is a process frequently observed in plant species under photoinhibitory situations. Two main hypotheses have been proposed to explain the function of such colour change: antioxidative protection exerted by red-coloured molecules, and green light filtering. The potential photoprotective role of red (retro-) carotenoids as light filters was tested in Buxus sempervirens leaves. In shade leaves of this species the upper (adaxial) mesophyll of the lamina was replaced by the equivalent upper part of a different colour phenotype. These hybrid leaves were exposed to a photoinhibitory treatment in order to compare the photoprotective effect exerted by adaxial parts of phenotypes with a different proportion of red (retro)-carotenoids in the lower mesophyll of a shade leaf. The results indicated that the presence of red (retro)-carotenoids in the upper mesophyll did not increase photoprotection of the lower mesophyll when compared with chlorophyll, and the best protection was achieved by an upper green layer. This was due to the fact that the extent of photoinhibition was proportional to the amount of red light transmitted by the upper mesophyll and/or to the chlorophyll pool located above. These results do not exclude a protective function of carotenoids in the upper leaf layer, but imply that, at least under the conditions of this experiment, the accumulation of red pigments in the outer leaf layers does not increase photoprotection in the lower mesophyll.

The lutein epoxide cycle in vegetative buds of woody plants.

Recent works have shown that two xanthophyll cycles operate simultaneously in several plant species: the ubiquitous violaxanthin + anteraxanthin + zeaxanthin (VAZ) cycle and the lutein epoxide (Lx) cycle. In the present work we tested for the presence of the Lx cycle in vegetative buds of woody plants. After an extensive screening of 130 species, we have shown that Lx is present in buds of many different plant species and leaf primordia are enriched in Lx compared with leaves. As a result, the Lx pool was higher than the violaxanthin (V) pool in several species. Although Lx can be potentially de-epoxidised in buds, light attenuation by scales inhibited the daily operation of the Lx cycle. This finding would imply that the Lx cycle is not involved in short-term reversible photoprotection in buds. However, a light-induced decrease in Lx was observed through the winter. An extensive screening of 130 species in 49 families conducted to elucidate the taxonomic extension of this cycle showed a widespread presence of the Lx cycle. The presence of Lx has a high fidelity at the family level, but its presence in unrelated taxa suggests that this character has appeared independently in several different groups of plants.