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Young leaves in some plants are transiently red due to the presence of anthocyanins, which disappear upon maturation. We investigated the hypothesis that light attenuation by anthocyanins may lead to a shade acclimation of the photosynthetic machinery in red leaves. We took advantage of the intra-species variation in anthocyanin levels of young, exposed leaves of Quercus coccifera. Thus, photosynthetic and photoprotective characteristics were compared in young green and red leaves of the same age, sampled from the corresponding phenotypes occupying the same habitat. Red leaves displayed several shade attributes like thinner laminae, lower Chl a/b ratios and lower levels of the xanthophyll cycle components and β-carotene. In addition, although both leaf kinds had the same area based levels of chlorophylls, these pigments were excluded from the sub-epidermic anthocyanic cell layers, leading to a further reduction of effective mesophyll thickness and an increase in chlorophyll density. Accordingly, red leaves had higher absolute chlorophyll fluorescence signals. In spite of these apparent shade characters, red leaves were less prone to photoinhibition under mild laboratory conditions and displayed slightly but significantly higher PS II photochemical efficiencies at pre-dawn in the field. No differences in all the above measured parameters were found in mature green leaves of the two phenotypes. The results confirm the light acclimation hypothesis and are also compatible with a photoprotective function of anthocyanins.

Under apparently similar field conditions individual plants of Cistus creticus turn transiently red during winter, while neighbouring plants remain green. These two phenotypes provide a suitable system for comparing basic photosynthetic parameters and assessing critically two hypotheses, i.e. anthocyanins afford photoprotection and anthocyanins induce shade characteristics on otherwise exposed leaves. With that aim, pigment levels and in vivo chlorophyll fluorescence parameters were monitored in dark-acclimated (JIP-test) and light-acclimated (saturation pulse method) leaves during both the green and the red period of the year. No evidence for actual photoprotection by anthocyanins was obtained. On the contrary, all fluorescence parameters related to yields and probabilities of photochemical energy conversion and electron flow, from initial light trapping to final reduction of ultimate electron acceptors in PSI, declined in the red phenotype after leaf reddening. Moreover, the pool sizes of final electron acceptors of PSII diminished, indicating that both photosystems were negatively affected. Vulnerability to winter stress was also indicated by sustained chlorophyll loss, inability to increase the levels of photoprotective xanthophylls and increased quantum yield of non-regulated energy loss during reddening. However, during the same period, the relative PSII antenna size increased, indicating an apparent shade acclimation after anthocyanin accumulation, while changes in the photosynthetic pigment ratios were also compatible to the shade acclimation hypothesis. All parameters recovered to pre-reddening values upon re-greening. It is concluded that the photosynthetic machinery of the red leaf phenotype has an inherently low capacity for winter stress tolerance, which is not alleviated by anthocyanin accumulation.

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