Xanthophyll Cycle Components Research Articles

Low-night temperature (LNT) induced changes of photosynthesis in grapevine ( Vitis vinifera L.) plants

Changes of leaf pigments, ribulose-1,5-bisphosphate carboxylase (RuBPC) and photosynthetic efficiency were examined in grapevine ( Vitis vinifera L.) plants grown under ambient irradiation (maximum daily PAR = 1500 μmol m −2 s −1) for 7 days to low night temperature (LNT) of 5 °C (daily from 18:00 to 06:00). The contents of chlorophyll (Chl) and carotenoids (Car) per fresh mass were lower in LNT leaves than in control leaves. The contents of α + β carotene and lutein-5,6-epoxide remained unaffected, but the de-epoxidation state involving the components of xanthophyll cycle increased. RuBPC activity and soluble proteins were also significantly reduced in LNT leaves. In isolated thylakoids, a marked inhibition of whole chain (PS I + PS II) and PS II activity were observed in LNT leaves. Smaller inhibition of PS I activity was observed in LNT leaves. The artificial exogenous electron donors, MnCl 2, DPC and NH 2OH did not restored the loss of PS II activity in LNT leaves. The same results were obtained when F v/ F m was evaluated by Chl fluorescence measurements. The marked loss of PS II activity in LNT leaves was due to the marked loss of D1 protein which was determined by immunological studies.

Midday photoinhibition of two newly developed super-rice hybrids

Super-rice hybrids are two-line hybrid rice cultivars with 15 to 20 % higher yields than the raditional three-line hybrid rice cultivars. Response of photosynthetic functions to midday photoinhibition was compared between seedlings of the traditional hybrid rice (Oryza sativa L.) Shanyou63 and two super-rice hybrids, Hua-an3 and Liangyoupeijiu. Under strong midday sunlight, in comparison with Shanyou63, the two super-rice hybrids were less photoinhibited, as indicated by the lower loss of the net photosynthetic rate (P N), the quantum yield of photosystem 2 (ΦPS2), and the maximum and effective quantum yield of PS2 photochemistry (Fv/Fm and Fv′/Fm′). They also had a much higher transpiration rate. Hence the super-rice hybrids could protect themselves against midday photoinhibition at the cost of water. The photoprotective de-epoxidized xanthophyll cycle components, antheraxanthin (A) and zeaxanthin (Z), were accumulated more in Hua-an3 and Liangyoupeijiu than in Shanyou63, but the size of xanthophyll cycle pool of the seedlings was not affected by midday photoinhibition. Compared to Shanyou63, the super-rice hybrids were better photoprotected under natural high irradiance stress and the accumulation of Z and A, not the size of the xanthophyll pool protected the rice hybrids against photoinhibition.

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.

Winter photoinhibition in the field involves different processes in four co-occurring Mediterranean tree species.

Photoinhibition was examined in four co-occurring Mediterranean evergreen tree species during two consecutive winters. In response to low temperatures and saturating light, Juniperus phoenicea L., Pinus halepensis Mill., Quercus coccifera L. and Q. ilex ssp. ballota (Desf.) Samp. exhibited marked chronic photoinhibition, indicated by low predawn maximal photochemical efficiency of photosystem II (PSII) (Fv/Fm). Low Fv/Fm values were correlated with high concentrations of xanthophyll cycle components (VAZ) and with the maintenance of high concentrations of zeaxanthin overnight (DPSpd). In all species, however, chronic photoinhibition was enhanced as the winter progressed in the absence of changes in DPSpd, suggesting cumulative damage toward the end of winter. Photoinhibition differed among species: P. halepensis always displayed significantly higher Fv/Fm values; and Q. coccifera had the lowest Fv/Fm values, showing a high sensitivity to the combination of high light and low temperatures. Differences among species were not fully explained by differences in the xanthophyll pool or its de-epoxidation state. Chronic photoinhibition overlapped with a dynamic photoinhibition as shown by the low values of photochemical efficiency of the open reaction centers of PSII at midday. Winter photoprotective strategies differed among species and may involve photoprotective mechanisms in addition to those associated with xanthophylls. The observed species-specific differences matched results obtained for the same species in summer; however, comparison of the two seasons suggests that the higher VAZ concentration observed in winter has an additional structural photoprotective role.

Exceptional Photosynthetic Performance of Capparis spinosa L. Under Adverse Conditions of Mediterranean summer

Diurnal and seasonal fluctuations in water potential (Ψ), stomatal conductance (gs), transpiration rate (E), and net photosynthetic rate (PN) were monitored in Capparis spinosa L., a Mediterranean plant growing during summer, i.e. at the period considered the most stressful for local plant life. In spite of the complete absence of rain, Ψ exhibited a modest drop at midday (-2.7 MPa), but was fully recovered overnight, indicating sufficient access to water sources. The stomata remained open throughout the day and season and the high E resulted in leaf temperatures up to 3.9 °C below air temperature. Additionally, PN of the fully exposed leaves was higher than 25 μmol m-2 s-1 for more than 10 h per day throughout the summer growth period. No symptoms of photooxidative stress were shown, as judged by maximum photosystem 2 photochemical efficiency (Fv/Fm) and the function of xanthophyll cycle. Indeed, diurnal inter-conversions of the xanthophyll cycle components were modest during the summer and a more intensive function of the cycle was only evident during leaf senescence in autumn. In comparison with a semi-deciduous and an evergreen sclerophyll co-existing in the same ecosystem, C. spinosa assimilated up to 3.4 times more CO2 per m2 during its growth period (May to October) and up to 1.8 times more on an annual basis.

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.

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.

Daily changes in components of xanthophyll cycle and antioxidant systems in leaves of rice at different developing stage

The daily changes in the behavior of xanthophyll cycle and antioxidant systems in flag leaves of superhigh-yield hybrid rice were investigated in relation to various developing stages. Dark-adapted Fv/Fm decreased with the increasing incident light intensity on leaf surface in the morning and then minimized at midday; Deepoxidation State showed an opposed daily pattern to Fv/Fm at different developing stage. As compared with increased deepoxidation state maximum value, the relative content of xanthophyll cycle pigments remained almost constant during development. The daily changes in activities of superoxide dismutase, ascorbate-peroxidase and glutathione reductase and the content of ascorbate and glutathione displayed a similar pattern, where they increased from 8:00 and reached maximum at midday, however, a lower daily fluctuation of superoxide dismutase activity was observed in senescent leaves. The enhanced contribution of xanthophyll cycle and Mehler-ascorbate peroxidase reaction to photoprotection in old leaves could be partially due to the altered leaf posture. In conclusion, daily changes of xanthophyll cycle and antioxidant systems in leaves of rice at various developing stages were dependent on leaf age, leaf angle and intensity of solar irridiance.

Response of senescing cotyledons of clusterbean to water stress in moderate and low light: Possible photoprotective role of beta-carotene.

Senescence of clusterbean (Cyamopsis tetragonoloba L.) cotyledons in moderate light (12 W m-2) brings about a loss in the pigments, enhanced lipid peroxidation and a decline in PS II photochemical activity without any loss either in Dl protein or in the level of beta-carotene. The senescence syndrome is aggravated in the cotyledons of water-stressed seedlings with an increase in thylakoid lipid peroxidation, a decline in the level of beta-carotene and a quantitative loss in the Dl protein. Loss of the protein, however, is arrested in the seedlings experiencing water stress at low light (3 W m-2) intensity that correlates with the stability in the level of beta-carotene and a slow rate of lipid peroxidation. Loss of the protein in moderate light is attributed to water-stress sensitized photoinhibitory damage. The data on changes in the components of xanthophyll cycle suggest the low activity of the cycle both during senescence and water stress. It is, therefore, concluded that beta-carotene may contribute to the assembly and stability of the Dl protein during senescence and water stress in clusterbean cotyledons.

Photoinhibition in Chilling Stressed Leguminosae: Comparison of Vicia Faba and Pisum Sativum

The concentrations of photosynthetic pigments decreased in both chilling stressed species but the ratios of chlorophyll (Chl) a/b and total carotenoids (Car)/Chls were depressed only in faba bean. The contents of α+β carotene and lutein+lutein-5,6-epoxide remained unaffected in both species, but the de-epoxidation state involving the components of xanthophyll cycle increased in pea. Under chilling stress the photosynthetic electron transport associated with photosystem 2, PS2 (with and without the water oxidising complex) decreased in both plant species, the inhibition being higher in faba bean. The intrachloroplast quinone pool also decreased in both stressed species, yet an opposite trend was found for cytochrome b559LP. Under stress an increasing peroxidation of thylakoid acyl lipids was detected in pea, but higher protein/Chl ratio was detected in faba bean. Thus the acceptor side of PS2 is mostly affected in both chilling stressed species, but faba bean is more sensitive.

Studies on Photosynthetic Characteristics and Adaption to Environment of Some Alpine plants in Qinghai-Xizang Plateau

Mainly the alpine plants in Qinghai?Xizang plateau including Salix gilashanica, Salix pilosomicrophylla, Rhododendron coriaceum, Rhododendron faucium, Rhododendron nivale, Quercus aquifolioides were compared with the same genera, non-alpine plants including Salix babyleuica, Salix matsudana, Rhododendron maoronulatum, Quercus liaotangensis for studying their photosynthetic characteristics and photosynthetic adaptability to high mountain environment. The results showed that chlorophyll content of the leaves and chloroplasts tended to be lower in alpine plants than in non-alpine, but the chlorophyll a to b ratio and the carotenoid relative content were higher in alpine plants than non-alpine plants. The decreased trends of Fv/Fm and Fv/Fo (parameter of kinetics of Chl a fluorescence induction) of detached leaves indicated that the potential photosynthetic capacity of alpine plants did not be higher than those non-alpine plants. But that the PSI/PSII ratio was low and related content of PSII in the chloroplasts was high from alpine plant as indicated by fluorescence spectra. Separation and quantification of the various carotenoids was completed by the high performance liquid chromatography. The results show that the chloroplast of S. gilashanica and Q. aquifolioides contained greater amounts of the components of the xanthophyll cycle than the same genera, non-alpine plants S. matsudana and Q. liaotangensis. But the content of the components of the xanthophyll cycle in the chloroplast of R. coriacenm and R. nivale were lower than the same genera non-alpine plants R. macronulatum. The results showed the xanthophyll cycle may be very important in adaptation of the partial alpine plants to high light radiation.

Response of Tradescantia albiflora to growth irradiance: Change versus changeability.

Most chloroplasts undergo changes in composition, function and structure in response to growth irradiance. However, Tradescantia albiflora, a facultative shade plant, is unable to modulate its light-harvesting components and has the same Chl a/Chl b ratios and number of functional PS II and PS I reaction centres on a Chl basis at all growth irradiances. With increasing growth irradiance, Tradescantia leaves have the same relative amount of chlorophyll-proteins of PS II and PS I, but increased xanthophyll cycle components and more zeaxanthin formation under high light. Despite high-light leaves having enhanced xanthophyll cycle content, all Tradescantia leaves acclimated to varying growth irradiances have similar non-photochemical quenching. These data strongly suggest that not all of the zeaxanthin formed under high light is necessarily non-covalently bound to major and minor light-harvesting proteins of both photosystems, but free zeaxanthin may be associated with LHC II and LHC I or located in the lipid bilayer. Under the unusual circumstances in light-acclimated Tradescantia where the numbers of functional PS II and PS I reaction centres and their antenna size are unaltered during growth under different irradiances, the extents of PS II photoinactivation by high irradiances are comparable. This is due to the extent of PS II photoinactivation being a light dosage effect that depends on the input (photon exposure, antenna size) and output (photosynthetic capacity, non-radiative dissipation) parameters, which in Tradescantia are not greatly varied by changes in growth irradiance.

Photosystem II photochemical efficiency, zeaxanthin and antioxidant contents in the poikilohydric Ramonda serbica during dehydration and rehydration.

Changes in photochemical efficiency, non-radiative energy dissipation (NRD), de-epoxidation state of xanthophyll cycle components (DPS) and contents of the antioxidants ascorbic acid and glutathione were studied in leaves of the poikilohydric Ramonda serbica Panc. (Gesneriaceae) during cycles of dehydration and subsequent rehydration. In drying leaves, the intrinsic efficiency of PS II photochemistry and the photon yield of PS II electron transport showed strong progressive decreases. Simultaneously, the fraction of excitation energy dissipated as heat in the PS II antenna increased markedly. The energy-dependent component of non-photochemical quenching (NPQ) showed an increase in dehydrating leaves down to relative water contents (RWC) values near 30%. Further decreases in RWC below these values caused a decrease in NPQ. Accordingly, DPS showed a similar behaviour, with a sharp increase and a subsequent decrease at very low RWC, although the maximum DPS was reached at slightly lower RWC than that for the maximum NPQ. The pools of reduced ascorbate and glutathione increased strongly when the RWC values fell below 40% and remained high in fully dehydrated leaves. When plants were re-watered photosynthetic efficiency, NRD, DPS and antioxidant contents recovered their initial control values. However, during rehydration, the zeaxanthin content showed a transient increase, as did NPQ, indicating an increasing demand for non-radiative dissipation. On the other hand, the contents of reduced ascorbate and reduced glutathione decreased but were still relatively high in the initial phase of rehydration, when the rate of photosynthetic electron transport, proton pumping and NRD were still relatively low. These results indicate that several photoprotective mechanisms are operating in R. serbica. Protection from photo-oxidation and photoinhibition appears to be achieved by co-ordinated contributions by ascorbate, glutathione and zeaxanthin-mediated NPQ. This variety of photoprotective mechanisms may be essential for conferring desiccation-tolerance.

Seasonal fluctuations in photoprotective (xanthophyll cycle) and photoselective (chlorophylls) capacity in eight Mediterranean plant species belonging to two different growth forms

Xanthophyll cycle components and their pool sizes were seasonally examined in eight Mediterranean plant species from two different growth forms (drought semi-deciduals and evergreen sclerophylls), in an attempt to correlate possible fluctuations in the above parameters with potentially photoinhibitory conditions. The xanthophyll cycle was common in all species examined and was functional throughout the year. In semi-deciduals, the maximum midday values for conversion state, defined as the ratio (zeaxanthin + antheraxanthin)/(violaxanthin + zeaxanthin + antheraxanthin), and the maximum xanthophyll pool sizes, expressed on a Chl basis, were recorded during the dry summer, indicating that the period of water shortage may be the most demanding for a high photoprotective potential. However, the data suggests that sclerophylls may also need a high photoprotective potential during winter, since the xanthophyll pool sizes were equally high during both summer and winter. Corresponding spring and autumn values were low. Furthermore, winter pre-dawn conversion state values were the highest recorded amongst all species studied, indicating that the mildly low winter temperatures may be a considerable stress factor for these plants. It is assumed that by preserving high concentrations of zeaxanthin and antheraxanthin overnight, sclerophylls retain high energy dissipation activity early in the morning, when minimum temperatures, coinciding with high photon fluence rates, are likely to drive the photosynthetic apparatus to potentially harmful overexcitation. Of significance may be the fact that almost all species show a minimum in chlorophyll content during the summer. In this way, overexcitation is alleviated and the photoprotective action of carotenoids per chlorophyll molecule is enhanced.

Antioxidative defense system, pigment composition, and photosynthetic efficiency in two wheat cultivars subjected to drought

We analyzed antioxidative defenses, photosynthesis, and pigments (especially xanthophyll-cycle components) in two wheat (Triticum durum Desf.) cultivars, Adamello and Ofanto, during dehydration and rehydration to determine the difference in their sensitivities to drought and to elucidate the role of different protective mechanisms against oxidative stress. Drought caused a more pronounced inhibition in growth and photosynthetic rates in the more sensitive cv Adamello compared with the relatively tolerant cv Ofanto. During dehydration the glutathione content decreased in both wheat cultivars, but only cv Adamello showed a significant increase in glutathione reductase and hydrogen peroxide-glutathione peroxidase activities. The activation states of two sulfhydryl-containing chloroplast enzymes, NADP+-dependent glyceraldehyde-3-phosphate dehydrogenase and fructose-1,6-bisphosphatase, were maintained at control levels during dehydration and rehydration in both cultivars. This indicates that the defense systems involved are efficient in the protection of sulfhydryl groups against oxidation. Drought did not cause significant effects on lipid peroxidation. Upon dehydration, a decline in chlorophyll a, lutein, neoxanthin, and beta-carotene contents, and an increase in the pool of de-epoxidized xanthophyll-cycle components (i.e. zeaxanthin and antheraxanthin), were evident only in cv Adamello. Accordingly, after exposure to drought, cv Adamello showed a larger reduction in the actual photosystem II photochemical efficiency and a higher increase in nonradiative energy dissipation than cv Ofanto. Although differences in zeaxanthin content were not sufficient to explain the difference in drought tolerance between the two cultivars, zeaxanthin formation may be relevant in avoiding irreversible damage to photosystem II in the more sensitive cultivar.

Zeaxanthin and non‐photochemical quenching in sun and shade leaves of C3 and C4 plants

The relationships between non‐radiative energy dissipation and the carotenoid content, especially the xanthophyll cycle components, were studied in sun and shade leaves of several plants possessing C3 (Hedera helix and Laurus nobilis) or C4 (Zea mays and Sorghum bicolor) photosynthetic pathways. Sun‐shade acclimation caused marked changes in the organisation and function of photosynthetic apparatus, including significant variation in carotenoid content and composition. The contents of zanthophyll cycle pigments were higher in sun than in shade leaves in all species, but this difference was considerably greater in C3 than in C4 plants. The proportion of photoconvertible violaxanthin, that is the amount of violaxanthin (V) which can actually be de‐epoxidised to zeaxanthin, was much greater in sun than in shade leaves. The amount of photoconvertible V was always linearly dependent on the chlorophyll a/b ratio, although the slope of the relationship varied especially between C3 and C4 species. The leaf zeaxanthin and antheraxanthin contents were correlated with non‐radiative energy dissipation in all species under different light environments. These relationships were curvilinear and variable between sun and shade leaves and between C3 and C4 species. Hence, the dissipation of excess energy does not appear to be univocally dependent on zeaxanthin content and other photoprotective mechanisms may be involved under high irradiance stress. Such mechanisms appear largely variable between C3 and C4 species according to their photosynthetic characteristics.

Photochemical efficiency of Photosystem II and xanthophyll cycle components in Zea mays leaves exposed to water stress and high light

The effects of two light treatments (photosynthetically active photon flux density of either 650 or 1950 µmol m−2 s−1) on the photochemical efficiency of Photosystem II (PS II) (measured as variable to maximum fluorescence ratio) and on the xanthophyll cycle components was studied in wilted Zea mays leaves. For comparison, these parameters were followed under the same light conditions in well-hydrated leaves maintained either in normal or CO2-free air. The net CO2 assimilation of dehydrated leaves declined rapidly as their relative water content (RWC) decreased from 100 to 60% while the PS II efficiency measured after a prolonged dark period of 16 h declined only when RWC leaves was lower than 60%. Furthermore, drought caused an increase in the pool size of the xanthophyll cycle pigments and the presence of a sustained elevated level of zeaxanthin and antheraxanthin at the end of the long dark period. The leaf water deficit enhanced the sensitivity of PS II efficiency to light exposure. During illumination, strong inhibition of PS II efficiency and large violaxanthin deepoxidation was observed in wilted leaves even under moderate photon flux density compared to control leaves in the same conditions. After 2 h of darkness following the light treatment, the PS II efficiency that is dependent on the previous PPFD, decreased with leaf water deficit. Moreover, zeaxanthin epoxidation led to an accumulation of antheraxanthin in dehydrated leaves. All these drought effects on PS II efficiency and xanthophyll cycle components were also obtained in well-hydrated leaves by short-term CO2 deprivation during illumination. We conclude that the increased susceptibility of PS II efficiency to light in wilted maize leaves is mainly explained by the decrease of CO2 availability and the resulting low net CO2 assimilation.

The transiently pubescent young leaves of plane (Platanus orientalis) are deficient in photodissipative capacity

We have analyzed reflectance changes and carotenoid composition of young and mature leaves of Platanus orientalis L. in order to test the hypothesis that the transient occurrence of highly absorptive and reflective leaf hairs of young leaves (M. Ntefidou and Y. Manetas 1996, Aust. J. Plant Physiol. 23: 535–538) may be correlated to a weakly developed photodissipative capacity in the chloroplast. Compared to mature leaves, young leaves showed negligible reflectance changes at 530 nm upon sudden illumination, possibly indicating a limited production of zeaxanthin. In addition, actual pigment analysis confirmed lower pools of xanthophyll cycle components and reduced capacity for violaxanthin photoconversion in young leaves. Accordingly, the epoxidation state at saturating photon fluence rates was particularly high. A notable feature of xanthophyll cycle interconversions in young leaves was the inability to drive the system to complete de‐epoxidation, as antheraxanthin in the light was always higher than zeaxanthin. Among the rest of the carotenoids, the levels of β‐carotene were particularly low. Moreover, most of the photosynthetic pigments were considerably bleached when young leaves were exposed to high light. The above results strongly suggest that young leaves possess a limited photodissipative capacity and therefore, the presence of leaf hairs affords protection against excess light. When the leaf has matured and presumably the concentrations of photoprotective compounds are adequate, the loss of hairs is not of consequence. In fact, their presence on mature leaves may reduce the photosynthetically active radiation to non‐saturating levels for photosynthesis.

Seasonal changes in xanthophyll composition and photosynthesis of cork oak (Quercus suberL.) leaves under mediterranean climate

Seasonal changes in pigment composition of sun and shade leaves of cork oak (Quercus suber) were studied under field conditions in Portugal. Expanding leaves showed a high concentration of xanthophyll cycle components, violaxanthin, antheraxanthin and zeaxanthin. The pool of violaxanthin plus antheraxanthin plus zeaxanthin (V + A + Z) varied greatly between the seasons, being higher at the end of summer and in winter when photosynthesis was limited by water stress and cold, respectively. The size of V + A + Z pool was associated to synthesis of zeaxanthin in response to an excess of light. In sun leaves, midday A + Z relative content was positively correlated with the V + A + Z pool, whereas in shade leaves A + Z decreased with leaf ageing. In both leaf types A + Z was positively correlated with the non-photochemical quenching (NPQ) of chlorophyll a fluorescence. However, in winter NPQ did not change significantly throughout the day, whereas the (A + Z)/(V + A + Z) increased following the typical daily trend observed in other seasons.

Xanthophyll cycle components and capacity for non-radiative energy dissipation in sun and shade leaves ofLigustrum ovalifolium exposed to conditions limiting photosynthesis

The relationships between photosynthetic efficiency, non-radiative energy dissipation and carotenoid composition were studied in leaves ofLigustrum ovalifolium developed either under full sunlight or in the shade. Sun leaves contained a much greater pool of xanthophyll cycle components than shade leaves. The rate of non-radiative energy dissipation, measured as non-photochemical fluorescence quenching (NPQ), was strictly related to the deepoxidation state (DPS) of xanthophyll cycle components in both sun and shade leaves, indicating that zeaxanthin (Z) and antheraxanthin (A) are involved in the development of NPQ. Under extreme conditions of excessive energy, sun leaves showed higher maximum DPS than shade leaves. Therefore, sun leaves contained not only a greater pool of xanthophyll cycle components but also a higher proportion of violaxanthin (V) actually photoconvertible to A and Z, compared to shade leaves. Both these effects contributed to the higher NPQ in sun versus shade leaves. The amount of photoconvertible V was strongly related to chla/b ratio and inversely to leaf neoxanthin content. This evidence indicates that the amount of photoconvertible V may be dependent on the degree of thylakoid membrane appression and on the organization of chlorophyll-protein complexes, and possible explanations are discussed. Exposure to chilling temperatures caused a strong decline in the photon yield of photosynthesis and in the intrinsic efficiency of PS II photochemistry in sun leaves, but little effects in shade leaves. These effects were accompanied by increases in the pool of xanthophyll cycle components and in DPS, more pronounced in sun than in shade leaves. This corroborates the view that Z and A may play a photoprotective role under unfavorable conditions. In addition to the xanthophyll-related non-radiative energy dissipation, a slow relaxing component of NPQ, independent from A and Z concentrations, has been found in leaves exposed to low temperature and high light. This quenching component may be attributed either to other regulatory mechanism of PS II efficiency or to photoinactivation.