The photosynthetic pigments of twigs in five tree and shrub species possessing chlorenchyma under a well developed, stomata-less, and highly photon absorptive periderm were analysed and compared to those of the corresponding canopy leaves. We asked whether the unavoidable shade acclimation of corticular chlorenchyma results in photosynthetic pigment complements typically found in shade leaves. As expected, chlorophyll (Chl) a/b ratios in twigs were consistently low. However, carotenoid (Car) analysis did not confirm the initial hypothesis, since twigs generally contained increased Chl-based pool sizes of the xanthophyll cycle components. The contents of photo-selective neoxanthin and lutein were high as well. Yet, β-carotene content was extraordinarily low. In addition, twigs retained high pre-dawn ratios of the deepoxidized antheraxanthin and zeaxanthin, although environmental conditions were not pre-disposing for such a state. The unexpected Car composition allows the conclusion that other micro-environmental conditions within twigs (hypoxia, increased red to blue photon ratios, and extremely high CO2 concentrations) are more important than shade in shaping the Car profiles. and E. Levizou, Y. Petropoulou, Y. Manetas.
Recent reports have indicated a considerably inactivated PSII in twig cortices, in spite of the low light transmittance of overlying periderms. Corresponding information for more deeply located and less illuminated tissues like xylem rays and pith are lacking. In this investigation we aimed to characterize the efficiency of PSII and its light sensitivity along twig depth, in conjunction with the prevailing light quantity and quality. To that aim, optical methods (spectral reflectance and transmittance, chlorophyll fluorescence imaging, low temperature fluorescence spectra) and photoinhibitory treatments were applied in cut twig sections of four tree species, while corresponding leaves served as controls. Compared to leaves, twig tissues displayed lower chlorophyll (Chl) levels and dark-adapted PSII efficiency, with strong decreasing gradients towards the twig center. The low PSII efficiencies in the inner stem were not an artifact due to an actinic effect of measuring beam or to an enhanced contribution of PSI fluorescence. In fact, the PSII/PSI ratios in cortices were higher and those in the xylem rays similar to that of leaves. Inner twig tissues were quite resistant to photoinhibitory treatments, tolerating irradiation levels several-fold higher than those encountered in their microenvironment. Moreover, the extent of high light tolerance was similar in naturally exposed and shaded twig sides. The results indicate an increasing, inherent and light-independent inactivation of PSII along twig depth. The findings are discussed on the basis of a recently proposed model for photosynthetic electron flow in twigs, taking into account the specific atmospheric and light microenvironment as well as the possible metabolic needs of such bulky organs. and C. Yiotis, Y. Petropoulou, Y. Manetas.
We checked the hypothesis that the transient presence of anthocyanins in young leaves serves a photoprotective function. For this purpose, Rosa sp. and Ricinus communis L., whose young leaves are red to become green upon maturation, were used. Thus, young leaves with high and mature leaves with low anthocyanin contents were analysed concerning their carotenoid (Car) composition and susceptibility to photoinhibition. Cars, including the components of the xanthophyll cycle, had similar contents in young and mature leaves, when expressed on a chlorophyll basis. Yet, when expressed on a leaf area basis or on the assumed photon absorptive capacity of leaves, Cars contents were considerably lower in anthocyanic young leaves. Although this may indicate a low photodissipative potential, red young leaves were considerably less susceptible to photoinhibitory damage. The results are compatible with a photoprotective function of anthocyanins, indicating also that their presence may compensate for a low capacity in the xanthophyll cycle-dependent harmless dissipation of excess excitation energy. and Y. Manetas, A. Drinia, Y. Petropoulou.
Limitations in photosystem function and photosynthetic electron flow were investigated during leaf senescence in two field-grown plants, i.e., Euphorbia dendroides L. and Morus alba L., a summer- and winter-deciduous, shrub and tree, respectively. Analysis of fast chlorophyll (Chl) a fluorescence transients and post-illumination fluorescence yield increase were used to assess photosynthetic properties at various stages of senescence, the latter judged from the extent of Chl loss. In both plants, the yield of primary photochemistry of PSII and the content of PSI remained quite stable up to the last stages of senescence, when leaves were almost yellow. However, the potential for linear electron flow along PSII was limited much earlier, especially in E. dendroides, by an apparent inactivation of the oxygen-evolving complex and a lower efficiency of electron transfer to intermediate carriers. On the contrary, the corresponding efficiency of electron transfer from intermediate carriers to final acceptors of PSI was increased. In addition, cyclic electron flow around PSI was accelerated with the progress of senescence in E. dendroides, while a corresponding trend in M. alba was not statistically significant. However, there was no decrease in PSI activity even at the last stages of senescence. We argue that a switch to cyclic electron flow around PSI during leaf senescence may have the dual role of replenishing the ATP and maintaining a satisfactory nonphotochemical energy quenching, since both are limited by hindered linear electron transfer., C. Kotakis, A. Kyzeridou, Y. Manetas., and Obsahuje bibliografii
Photochemical efficiency of photosystem 2 (PS2), assessed from in situ chlorophyll (Chl) fluorescence measurements, was seasonally monitored in five evergreen sclerophyll and five malacophyllous drought semi-deciduous species, co-occurring in the same Mediterranean field site. In evergreen sclerophylls, a considerable drop in the variable (Fv) to maximum (Fm) Chl fluorescence ratio coincided with the lowest winter temperatures, indicating low PS2 efficiency during this period. Summer drought caused a comparatively slight decrease in Fv/Fm and only in three of the five evergreen sclerophyll species tested. In drought semi-deciduous shrubs, the winter drop in Fv/Fm was much less conspicuous. During the summer, and in spite of the severe and prolonged desiccation of their malacophyllous leaves, Fv/Fm was maintained high and only in one species the PS2 efficiency was transiently suppressed, when the leaf relative water content became lower than 30 %. Thus evergreen sclerophylls are more prone to photoinhibition by low winter temperatures, while the sensitivity of drought semi-deciduals depends on the extent and duration of summer drought. and S. Karavatas, Y. Manetas.
Some photosynthetic attributes of leaves and stems were seasonally followed in the small-leaved, summer-deciduous, green-stemmed Mediterranean shrub Calicotome villosa. Both leaves and stems displayed similar photon energy-saturated photosystem 2 (PS2) efficiencies with a minimum during winter. A second minimum in stems during the leafless summer period could be ascribed to sustained photoinhibition. Yet, stems were slightly inferior in photon capture, resulting partly from lower chlorophyll (Chl) contents and partly from higher reflectance due to pubescence. As a result, photon energy-saturated linear electron transport rates were slightly higher in leaves. However, when the total leaf and stem areas were taken into account, this superiority was abolished during autumn and winter and more than overturned during spring. Given that during summer the stems were the only photosynthetic organs, the yearly photosynthetic contribution of stems was much higher. Chl contents in stems displayed a transient and considerable summer drop, accompanied by an increase in the carotenoid to Chl ratio, indicating a photo-protective adaptation to summer drought through a decrease of photo-selective capacity, typical for leaves of many Mediterranean plants. and C. Yiotis, G. K. Psaras, Y. Manetas.
nclusion of natural (glycerol, proline) or synthetic (polyethylene glycol) compatible solutes in the assay medium for phosphoenolpyruvate carboxylase (EC 4.1.1.31) activity at low substrate (phosphoenolpyruvate) levels enhanced the enzymic activity throughout the temperature range tested (11-42 °C). This effect was much more pronounced above 30 °C and, therefore, the optimum for activity was shifted from 27-29 °C to 37-39 °C, a temperature more consistent with the prevailing leaf temperatures during the growing season for C4-plants; there was also a better correlation between activity/temperature profile and the net photosynthetic rate of intact leaves of Cynodon dactylon (L.) Pers., at the temperature range tested.
Fully exposed, senescing leaves of Cornus sanguinea and Parthenocissus quinquefolia display during autumn considerable variation in both anthocyanin and chlorophyll (Chl) concentrations. They were used in this study to test the hypothesis that anthocyanins may have a photoprotective function against photosystem II (PSII) photoinhibitory damage. The hypothesis could not be confirmed with field sampled leaves since maximum photochemical efficiency (Fv/Fm) of PSII was negatively correlated to anthocyanin concentration and the possible effects of anthocyanins were also confounded by a decrease in Fv/Fm with Chl loss. However, after short-term laboratory photoinhibitory trials, the percent decrease of Fv/Fm was independent of Chl concentration. In this case, a slight alleviation of PSII damage with increasing anthocyanins was observed in P. quinquefolia, while a similar trend in C. sanguinea was not statistically significant. It is inferred that the assumed photoprotection, if addressed to PSII, may be of limited advantage and only under adverse environmental conditions. and Y. Manetas, C. Buschmann.
Total carotenoids assessed spectrophotometrically in crude extracts may be considerably overestimated when high contents of phenolic compounds are co-extracted. In this case, the absorbance tails of phenolics extend well into the blue part of the spectrum, interfering with carotenoid estimation. Extracts of phenolic-rich organs, with a low ratio of photosynthetic to heterotrophic and/or supportive cells (for example, stems or twigs) are vulnerable to such pitfalls and may need chromatographic separation of carotenoids. and E. Levizou, Y. Petropoulou, Y. Manetas.