Y. Petropoulou

The effects of UV-B radiation on European heathland species

The effects of enhanced UV-B radiation on three examples of European shrub-dominated vegetation were studied in situ. The experiments were in High Arctic Greenland, northern Sweden and Greece, and at all sites investigated the interaction of enhanced UV-B radiation (simulating a 15% reduction in the ozone layer) with artificially increased precipitation. The Swedish experiment also involved a study of the interaction between enhanced UV-B radiation and elevated CO2 (600 ppm). These field studies were supported by an outdoor controlled environment study in the United Kingdom involving modulated enhancement of UV-B radiation in combination with elevated CO2 (700 ppm). Effects of the treatments on plant growth, morphology, phenology and physiology were measured. The effects observed were species specific, and included both positive and negative responses to the treatments. In general the negative responses to UV-B treatments of up to three growing seasons were small, but included reductions in shoot growth and premature leaf senescence. Positive responses included a marked increase in flowering in some species and a stimulation of some photosynthetic processes. UV-B treatment enhanced the drought tolerance of Pinus pinea and Pinus halepensis by increasing leaf cuticle thickness. In general, there were few interactions between the elevated CO2 and enhanced UV-B treatments. There was evidence to suggest that although the negative responses to the treatments were small, damage may be increasing with time in some long-lived woody perennials. There was also evidence in the third year of treatments for effects of UV-B on insect herbivory in Vaccinium species. The experiments point to the necessity for long-term field investigations to predict the likely ecological consequences of increasing UV-B radiation.

High Contents of Anthocyanins in Young Leaves are Correlated with Low Pools of Xanthophyll Cycle Components and Low Risk of Photoinhibition

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.

Beneficial effects of enhanced UV-B radiation under field conditions: improvement of needle water relations and survival capacity of Pinus pinea L. seedlings during the dry Mediterranean summer

The possible mechanism(s) by which supplemental UV-B radiation alleviates the adverse effects of summer drought in Mediterranean pines (Petropoulou et al. 1995) were investigated with seedlings of Pinus pinea. Plants received ambient or ambient plus supplemental UV-B radiation (biologically equivalent to a 15% ozone depletion over Patras, 38.3° N, 29.1° E) and natural precipitation or additional irrigation. Treatments started on 1 February, 1994 and lasted up to the end of the dry period (29 September). In well-watered plants, UV-B radiation had no influence on photosystem II photochemical efficiency and biomass accumulation. Water stressed plants suffered from needle loss and reduced photosystem II photochemical efficiency during the summer. These symptoms, however, were less pronounced in plants receiving supplemental UV-B radiation, resulting in higher total biomass at plant harvest. Laboratory tests showed that enhanced UV-B radiation did not improve the tolerance of photosystem II against drought, high light, high temperature and oxidative stress. Enhanced UV-B radiation, however, improved the water economy of water stressed plants, as judged by measurements of needle relative water content. In addition, it caused an almost two-fold increase of cuticle thickness. No such UV-B radiation effects were observed in well-watered pines. The results indicate that the combination of water stress and UV-B radiation may trigger specific responses, enabling the plants to avoid excessive water loss and, thereby, maintain a more efficient photosynthetic apparatus during the summer. The extent of this apparently positive UV-B radiation effect would depend on the amount of summer precipitation. Abbreviations: DW – dry weight, Fv/Fm – ratio of variable to maximum fluorescence, A 300 – absorbance at 300 nm, PAR – photosynthetically active radiation, PS II – photosystem II, RWC – relative water content, TCA – trichloroacetic acid, UV-BBE – biologically effective ultraviolet-B radiation

Exposed red (anthocyanic) leaves of Quercus coccifera display shade characteristics

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.

Transient winter leaf reddening in Cistus creticus characterizes weak (stress-sensitive) individuals, yet anthocyanins cannot alleviate the adverse effects on photosynthesis

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.

Effects of UV-B radiation on cuticle thickness and nutritional value of leaves in two mediterranean evergreen sclerophylls

Summary Seedlings of Laurus nobilis L. and Ceratonia siliqua L., grown for 1 year in the field under ambient radiation conditions were subsequendy allowed to produce their new spring growth in a glasshouse in the absence or presence of UV-B radiation. The UV-B radiation dose was biologically equivalent to that which would have being received by the plants in their natural environment. Plant height, number of leaves, total and mean leaf area, the biomass allocated to both above and below ground parts, photochemical efficiency of PS II and chlorophyll content were not affected. However, UV-B radiation resulted in slighdy thicker leaves in L. nobilis and considerably thicker cuticles in both plants. UV-B absorbing compounds were increased only in L. nobilis. In addition, the leaf nutritional quality of this plant was considerably reduced by UV-B radiation, since phenolics and tannins were increased but nitrogen was decreased. No such changes were observed in C. siliqua . We may conclude that L. nobilis and C. siliqua are not only resistant against UV-B radiation damage, but the observed changes may benefit the plants through their anti-transpirant and anti-herbivore functions.

Perturbations of the normal UV-B radiation environment alter leaf growth rates in Phlomis fruticosa L. seedlings

Abstract Phlomis fruticosa L. seedlings were raised in the field under ambient UV-B radiation for 18 months. On 15 March 1994, just before the spring growth burst), seedlings were transferred in a glasshouse under three regimes of artificial UV-B radiation, i.e. 0.06, 5.22 and 8.55 kJ m −2 biologically effective daily doses. The last two doses correspond to UV-B radiation reaching the Patras area (38.3° N, 29.1° E) under clear sky and with normal column ozone thickness at mid-April and mid-July. Leaf demography was monitored for 96 days. Growth rates new leaf formation and total leaf area increase) were significantly higher under 5.22 kJ m −2 day −1 , intermediate under 8.55 kJ m −2 day −1 and severely suppressed in the absence of UV-B radiation. In addition, shedding of older leaves after mid-May, was significantly more intense in the absence of UV-B radiation. The observed differences could not be attributed to UV-B radiation effects on CO 2 assimilation, photosynthetic electron flow or leaf water relations, since net photosynthetic rates, photochemical efficiency of photosystem II and leaf relative water contents showed no significant differences between treatments. Total dry matter measured at plant harvest was highest under 5.22 kJ m −2 day −1 and lowest in the absence of UV-B radiation. The results indicate that natural UV-B radiation may be an important factor in normal leaf development of Phlomis fruticosa .

Carotenoid composition of peridermal twigs does not fully conform to a shade acclimation hypothesis

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.

Induction of Ageotropic Response in Lettuce Radicle Growth by Epicuticular Flavonoid Aglycons of Dittrichia viscosa

Thirteen flavonoid aglycons, contained in the strongly allelopathic epicuticular exudates of Dittrichia viscosa, were investigated for their effects on lettuce seedling radicle growth. Concerning radicle length and mass, variable results were obtained, with most of the substances having no effect, some being inhibitory and some even promotive. Shoot mass was slightly reduced in four cases. Seed germination rates, root hair and lateral root formation were not affected either. Three of the compounds (namely quercetin 3,3-dimethylether, naringenin and eriodictyol) induced a strong ageotropic response in radicle growth.

Evidence for light-independent and steeply decreasing PSII efficiency along twig depth in four tree species

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.