The xanthophyll cycle and energy dissipation in differently oriented faces of the cactus Opuntia macrorhiza

  Diurnal changes in titratable acidity, photosynthesis, energy dissipation activity, and the carotenoid composition of differently oriented cladodes of the cactus Opuntia macrorhiza were characterized during exposure to full sunlight in the field. Four cladode faces were chosen such that each was exposed to maximum photon flux densities (PFD) at different times of the day in addition to receiving different daily integrated PFDs. The sum of all carotenoids per chlorophyll was found to increase with increasing exposure to PFD, with the carotenoids of the xanthophyll cycle present in the most exposed face at more than twice the concentration found in the least exposed face. All faces exhibited large increases in xanthophyll cycle-dependent energy dissipation as the sun rose in the morning, even those receiving only minimal levels of diffuse radiation. The transient high levels of energy dissipation in those faces that did not receive direct sunlight in the morning may have been due to low temperature inhibition of photosynthesis (predawn low of 2°C). For the two faces receiving peak PFDs in the morning hours (north and east faces), the level of energy dissipation activity increased rapidly during exposure to direct sunlight in the early morning, gradually declining in the late morning under warm temperatures, and was negligible during the afternoon low light conditions. Changes in the xanthophyll cycle paralleled the changes in energy dissipation with the majority of the cycle present as violaxanthin (V) prior to sunrise, largely de-epoxidized to zeaxanthin (Z) and antheraxanthin (A) during exposure to direct sunlight, and reconverted to V during the afternoon. For the two faces receiving peak PFDs in the afternoon (south and west faces), energy dissipation activity increased dramatically during the early morning low light period, subsequently decreasing during midday as decarboxylation of malic acid proceeded maximally (providing a high concentration of CO₂ for photosynthesis), and then increased to the highest level in the late afternoon as the supply of malic acid was depleted and rates of photosynthetic electron transport declined. The xanthophyll cycle, largely present as Z and A prior to sunrise in the south and west faces, was de-epoxidized to the greatest extent in the late afternoon, followed by epoxidation back to the predawn level by sunset. In all cladode faces high levels of energy dissipation activity were accompanied by decreases in the intrinsic efficiency of photosystem II (PSII), indicative of a regulatory process that diverted the excess energy away from the reaction centers during periods of excess light. Furthermore, the overnight retention of Z and A by the south and west faces was accompanied by a sustained reduction in PSII efficiency (i.e., “photoinhibition”). We suggest that this “photoinhibition” represents the sustained engagement of nocturnally retained Z and A in the photoprotective down-regulation of PSII. Received: 8 May 1996 / Accepted: 9 September 1996     

Positive correlation between levels of retained zeaxanthin + antheraxanthin and degree of photoinhibition in shade leaves of Schefflera arboricola (Hayata) Merrill

Attached intact leaves of Schefflera arboricola grown at three different photon flux densities (PFDs) were subjected to 24-h exposures to a high PFD and subsequent recovery at a low PFD. While sun leaves showed virtually no sustained effects on photosystem II (PSII), shade-grown leaves exhibited pronounced photoinhibition of PSII that required several days at low PFD to recover. Upon transfer to high PFD, levels of nonphotochemical quenching in PSII as well as levels of zeaxanthin were initially low in shade leaves but continued to increase gradually during the 24-h exposure. The xanthophyll cycle pool size rose gradually during and also subsequent to the photoinhibitory treatment in shade leaves. Upon return to low PFD, a marked and extremely long-lasting retention of zeaxanthin and antheraxanthin was observed in shade but not sun leaves. During recovery, changes in the conversion state of the xanthophyll cycle therefore closely mirrored the slow increases in PSII efficiency. This novel report of a close association between zeaxanthin retention and lasting PSII depressions in these shade leaves clearly suggests a role for zeaxanthin in photoinhibition of shade leaves. In addition, there was a decrease in β-carotene levels, some decrease in chlorophyll, but no change in lutein and neoxanthin (all per leaf area) in the shade leaves during and subsequent to the photoinhibitory treatment. These data may be consistent with a degradation of a portion of core complexes but not of peripheral light-harvesting complexes. A possible conversion of β-carotene to form additional zeaxanthin is discussed. Received: 24 October 1997 / Accepted: 12 November 1997     

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.Research supported by National Research Council of Italy, Special Project RAISA, Sub-Project 2, Paper N. 1587.     

The dissipation of excess excitation energy in British plant species

The reversible dissipation of excitation energy in higher plants is believed to protect against light-induced damage to the photosynthetic apparatus. This dissipation is measured as the non-photochemical quenching of chlorophyll fluorescence. A method is described whereby the saturated capacity for rapidly reversible non-photochemical quenching can be compared between plant species. This method was applied to 22 common British plant species whose habitat was quantified using an index that describes shade tolerance. An association was found between occurrence in open habitats and a high capacity for non-photochemical quenching. It was found that, whilst this capacity was species dependent, it did not depend upon the conditions under which the plant was grown. The possible role of zeaxanthin as a determinant of quenching capacity was examined by measuring the contents of xanthophyll cycle carotenoids for each species. Comparing species, no correlation was seen between the saturated level of non-photochemical quenching and zeaxanthin content expressed relative to either total carotenoid or to chlorophyll. When zeaxanthin was expressed relative to the amount of xanthophyll cycle intermediates (zeaxanthin, antheraxanthin and violaxanthin), a weak correlation was seen.     

Arabidopsis mutants define a central role for the xanthophyll cycle in the regulation of photosynthetic energy conversion.

A conserved regulatory mechanism protects plants against the potentially damaging effects of excessive light. Nearly all photosynthetic eukaryotes are able to dissipate excess absorbed light energy in a process that involves xanthophyll pigments. To dissect the role of xanthophylls in photoprotective energy dissipation in vivo, we isolated Arabidopsis xanthophyll cycle mutants by screening for altered nonphotochemical quenching of chlorophyll fluorescence. The npq1 mutants are unable to convert violaxanthin to zeaxanthin in excessive light, whereas the npq2 mutants accumulate zeaxanthin constitutively. The npq2 mutants are new alleles of aba1, the zeaxanthin epoxidase gene. The high levels of zeaxanthin in npq2 affected the kinetics of induction and relaxation but not the extent of nonphotochemical quenching. Genetic mapping, DNA sequencing, and complementation of npq1 demonstrated that this mutation affects the structural gene encoding violaxanthin deepoxidase. The npq1 mutant exhibited greatly reduced nonphotochemical quenching, demonstrating that violaxanthin deepoxidation is required for the bulk of rapidly reversible nonphotochemical quenching in Arabidopsis. Altered regulation of photosynthetic energy conversion in npq1 was associated with increased sensitivity to photoinhibition. These results, in conjunction with the analysis of npq mutants of Chlamydomonas, suggest that the role of the xanthophyll cycle in nonphotochemical quenching has been conserved, although different photosynthetic eukaryotes rely on the xanthophyll cycle to different extents for the dissipation of excess absorbed light energy.     

Epoxidation of zeaxanthin and antheraxanthin reverses non-photochemical quenching of photosystem II chlorophyll a fluorescence in the presence of trans-thylakoid ΔpH

The xanthophyll cycle apparently aids the photoprotection of photosystem II by regulating the nonradiative dissipation of excess absorbed light energy as heat. However, it is a controversial question whether the resulting nonphotochemical quenching is soley dependent on xanthophyll cycle activity or not. The xanthophyll cycle consists of two enzymic reactions, namely deepoxidation of the diepoxide violaxanthin to the epoxide-free zeaxanthin and the much slower, reverse process of epoxidation. While deepoxidation requires a transthylakoid pH gradient (ΔpH), epoxidation can proceed irrespective of a ΔpH. Herein, we compared the extent and kinetics of deepoxidation and epoxidation to the changes in fluorescence in the presence of a light-induced thylakoid ΔpH. We show that epoxidation reverses fluorescence quenching without affecting thylakoid ΔpH. These results suggest that epoxidase activity reverses quenching by removing deepoxidized xanthophyll cycle pigments from quenching complexes and converting them to a nonquenching form. The transmembrane organization of the xanthophyll cycle influences the localization and the availability of deepoxidized xanthophylls is to support nonphotochemical quenching capacity. The results confirm the view that rapidly reversible nonphotochemical quenching is dependent on deepoxidized xanthophyll.     

The xanthophyll cycle and sustained thermal energy dissipation activity in Vinca minor and Euonymus kiautschovicus in winter

The influence of low temperature on the operation of the xanthophyll cycle and energy dissipation activity, as ascertained through measurements of chlorophyll fluorescence, was examined in two broad-leaved evergreen species, Vinca minor L. and Euonymus kiautschovicus Loessner. In leaves examined under laboratory conditions, energy dissipation activity developed more slowly at lower leaf temperatures, but the final, steady-state level of such activity was greater at lower temperatures where the rate of energy utilization (through photosynthetic electron transport) was much lower. The rate at which energy dissipation activity increased was similar to that of the de-epoxidation of violaxanthin to antheraxanthin and zea-xanthin at different temperatures. However, leaves in the field examined prior to sunrise on mornings following cold days and nights exhibited a retention of antheraxanthin and zeaxanthin that was associated with sustained decreases in photosystem II efficiency. We therefore suggest that this phenomenon of ‘photoinhibition’ in response to light and cold temperatures during the winter results from sustained photoprotective thermal energy dissipation associated with the xanthophyll cycle. Such retention of the de-epoxidized components of the xanthophyll cycle responded to day-to-day changes in temperature, being greatest on the coldest mornings (when photoprotective energy dissipation might be most required) and less on warmer mornings when photosynthesis could presumably proceed at higher rates.     

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.     

Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching

The photosynthetic apparatus in plants is protected against oxidative damage by processes that dissipate excess absorbed light energy as heat within the light-harvesting complexes. This dissipation of excitation energy is measured as nonphotochemical quenching of chlorophyll fluorescence. Nonphotochemical quenching depends primarily on the [delta]pH that is generated by photosynthetic electron transport, and it is also correlated with the amounts of zeaxanthin and antheraxanthin that are formed from violaxanthin by the operation of the xanthophyll cycle. To perform a genetic dissection of nonphotochemical quenching, we have isolated npq mutants of Chlamydomonas by using a digital video-imaging system. In excessive light, the npq1 mutant is unable to convert violaxanthin to antheraxanthin and zeaxanthin; this reaction is catalyzed by violaxanthin de-epoxidase. The npq2 mutant appears to be defective in zeaxanthin epoxidase activity, because it accumulates zeaxanthin and completely lacks antheraxanthin and violaxanthin under all light conditions. Characterization of these mutants demonstrates that a component of nonphotochemical quenching that develops in vivo in Chlamydomonas depends on the accumulation of zeaxanthin and antheraxanthin via the xanthophyll cycle. However, observation of substantial, rapid, [delta]pH-dependent nonphotochemical quenching in the npq1 mutant demonstrates that the formation of zeaxanthin and antheraxanthin via violaxanthin de-epoxidase activity is not required for all [delta]pH-dependent nonphotochemical quenching in this alga. Furthermore, the xanthophyll cycle is not required for survival of Chlamydomonas in excessive light.     

Photosynthetic Characteristics and the Ratios of Chlorophyll, β-Carotene,and the Components of the Xanthophyll Cycle Upon a Sudden Increase in Growth Light Regime in Several Plant Species*

Among three species, Gossypium hirsutum, Rhizophora mangle, and Monstera deliciosa, which were transferred from low to high growth PFD, only small decreases in the efficiency of photochemical energy conversion were observed in those plants which exhibited an increase in photosynthetic capacity. Leaves of plants which showed no increase in photosynthetic capacity experienced a continuous decrease in photochemical efficiency, accompanied by a more pronounced loss of chlorophyll than that observed in the former group. In all species marked increases in the xanthophyll/β-carotene ratio resulted from small increases in lutein, and several-fold increases in the sum of the three components of the xanthophyll cycle, zeaxanthin, antheraxanthin, and violaxanthin. A strong increase in the level of zeaxanthin was only partially matched by a decrease of violaxanthin to zero, and was further paralleled by a decrease in β-carotene. Antiparallel changes in the sum of zeaxanthin + antheraxanthin + violaxanthin and β-carotene between morning and evening were observed in all species. These diel changes were overlaid on a net increase in β-carotene as well as total carotenoid content in those plants in which photosynthetic capacity increased. In those, however, which exhibited no photosynthetic acclimation upon transfer to high light, a decrease in both β-carotene and total carotenoid content was observed. Rhizophora mangle grown at 100 % seawater exhibited a particularly high capacity for increasing the level of zeaxanthin in response to high light.     

Survey of Thermal Energy Dissipation and Pigment Composition in Sun and Shade Leaves

A survey was conducted of the magnitude of energy dissipationin photosystem II (expressed as nonphotochemi-cal quenchingof chlorophyll fluorescence, NPQ) as well as leaf carotenoidcomposition of a wide range of different plant species growingin deep shade and/or full sun. Consistently higher levels ofthe reversible component of NPQ as well as higher degrees ofrapidly attainable de-epoxida-tion of the xanthophyll cycle(VAZ) pool were observed in sun leaves compared to deep shadeleaves. It is concluded that these altered features of the xanthophyllcycle allowed sun leaves to dissipate excess energy more effectivelyover the short term. In addition to the rapid increase in reversibleNPQ, shade leaves exhibited a slow further, and sustained, increasein NPQ. In contrast to these deep shade leaves experimentallyexposed to high PFDs, understory leaves experiencing highlyvariable PFD in their natural environment appeared to be ableto dissipate excess excitation energy adequately via xanthophyllcycle-dependent thermal dissipation. Furthermore, very consistenttrends across plant species were observed for changes in carotenoidcomposition (pools of carotenes, VAZ, and other xantho-phylls)in response to light environment, as long as it is assumed thatin some species rß-carotene can be replaced by     

植株叶片的光合色素构成对遮阴的响应

叶绿素在植株体内负责光能的吸收、传递和转化, 类胡萝卜素则行使光能捕获和光破坏防御两大功能, 它们在光合作用中起着非常重要的作用。该文综述了几大主要光合色素的分布和功能, 以及不同物种的色素含量和构成差异。阳生植物的叶黄素库较大, 但脱环氧化水平不及阴生植物。黄体素与叶黄素库的比值与植物的耐阴性呈正相关关系。由不同的遮阴源造成的遮阴环境, 光强和光质有很大的差异, 总体来说对植物生长的影响, 建筑物遮阴<阔叶林遮阴<针叶林遮阴。光强的改变可诱导类胡萝卜素的两大循环——叶黄素循环和黄体素循环。由光强诱导的叶绿素含量和叶绿素a/b比值的改变与该物种的耐阴性无关。短时间的遮阴不会对植物的生长造成危害, 叶黄素库的大小不仅与每天接受的光量子有关, 更与光量子在一天的分布有关, 因为光照和温度是协同作用的。光合作用或色素构成是蓝光、红光和远红光共同作用的结果, 不是某一种单色光所能替代的。我们总结了影响植物色素构成的内因和外因, 指出植物主要通过调整光反应中心和捕光天线色素蛋白复合体的比例, 以及两个光系统的比值来调整色素含量和构成以适应不同的光照条件, 提出了现存研究中存在的一些问题, 旨在为今后的相关研究提供建议。     

Photosynthesis and Photoprotection in Overwintering Plants

Abstract: Seasonal differences in the capacity of photosynthetic electron transport, leaf pigment composition, xanthophyll cycle characteristics and chlorophyll fluorescence emission were investigated in two biennial mesophytes ( Malva neglecta and Verbascum thapsus ) that grow in full sunlight, and in leaves/needles of sun and shade populations of several broad-leafed evergreens and conifers (Vinca minor, Euonymus kiautschovicus, Mahonia repens, Pseudotsuga menziesii [Douglas fir], and Pinus ponderosa). Both mesophytic species maintained or upregulated photosynthetic capacity in the winter and exhibited no upregulation of photoprotection. In contrast, photosynthetic capacity was downregulated in sun leaves/needles of V. minor, Douglas fir, and Ponderosa pine, and even in shade needles of Douglas fir. Interestingly, photosynthetic capacity was upregulated during the winter in shade leaves/needles of V. minor, Ponderosa pine and Euonymus kiautschovicus. Nocturnal retention of zeaxanthin and antheraxanthin, and their sustained engagement in a state primed for energy dissipation, were observed largely in the leaves/needles of sun-exposed evergreen species during winter. Factors that may contribute to these differing responses to winter stress, including chloroplast redox state, the relative levels of source and sink activity at the whole plant level, and apoplastic versus symplastic phloem loading, are discussed.     

依赖于叶黄素循环的热耗散对茶树叶片防御强光破坏的相关试验研究

经叶黄素循环抑制剂——二硫苏糖醇(DIT)处理的茶树叶片,以850μmol．m^-2．s^-1的PFD照射120min后,福鼎大白茶的叶黄素循环组分中的环氧玉米黄素(A)和玉米黄素(Z)含量之和降低了76．5％,结果导致非光化学猝灭(NPQ)、光系统Ⅱ(PSⅡ)的光化学效率(Fv／Fm)、光化学猝灭系数(qP)、PSⅡ实际光化学量子效率(ψPSⅡR)和光合电子传递速率(ETR)明显下降,而F0显著上升,暗恢复后Fv／Fm恢复程度小于未经DIT处理的叶片。自然光强下,NPQ与与叶黄素循环的脱环氧化程度(A Z)/(V A Z)比值呈明显的正线性关系(R=0．9488^***)。这些结果充分证明依赖与叶黄素循环的热耗散是茶树叶片光合器官防御强光破坏的主要途径。     

Carotenoid composition in sun and shade leaves of plants with different life forms

The carotenoid composition of sun leaves of nine species of annual crop plants (some with several varieties) was compared with sun and shade leaves of several other groups of plants, among those sun and shade leaves of several species of perennial shrubs and vines and deep-shade leaves of seven rainforest species. All sun leaves contained considerably greater amounts of the components of the xanthophyll cycle violaxanthin, antheraxanthin and zeaxanthin as well as of β-carotene than the shade leaves, as had previously been reported for a variety of other species by Thayer & Björkman (Photosynthesis Research, 1990, 23, 331–343). Therefore, high light specifically stimulated β,β-carotenoid synthesis. The sun leaves of these crop species did not contain α-carotene which was, however, present in large amounts in all shade leaves and in smaller amounts in sun leaves of three of the four species of perennial shrubs and vines. There was no difference in neoxanthin content on a chlorophyll basis between sun and shade leaves, and there was no consistent general difference in the lutein content between all sun and all shade leaves. The zeaxanthin (and antheraxanthin) content at peak irradiance and the xanthophyll cycle pool size were compared for sun leaves from the different groups of plants with different life forms and different metabolic activities. When growing in full sunlight the annual crop species and a perennial mesophyte had high rates of photosynthesis whereas the perennial shrubs and vines had relatively low photosynthesis rates. More zeaxanthin (and antheraxanthin) were accumulated at noon in full sunlight in those species with the lower photosynthesis rates. However, it was not such that those species also possessed the larger pools of violaxanthin + antheraxanthin + zeaxanthin. Instead, the xanthophyll cycle pools of sun leaves of the annual crop species and the perennial mesophyte were not smaller, and were even possibly larger, than those of sun leaves of the perennial shrubs and vines with low photosynthesis rates. This was so in spite of the fact that the crop species experienced much lesser degrees of excessive light at full sun than the shrubs and vines. Thus, many of the crop species converted only about 30–50% of their xanthophyll cycle pool to zeaxanthin at noon, whereas the shrubs and vines typically converted more than 80% of their pool into zeaxanthin. The crop species also had larger pools of β-carotene than the shrubs and vines but smaller pools of lutein than the majority of the latter species.     

Carotenoid composition and metabolism in green and blue-green algal lichens in the field

The carotenoid composition of 33 species of green algal lichens and 5 species of blue-green algal lichens was examined and compared with that of the leaves of higher plants. As in higher plants, green algal lichen species which were found in both shade and full sunlight exhibited higher levels of the carotenoids involved in photoprotective thermal energy dissipation (zeaxanthin as well as the total xanthophyll cycle pool) in the sun than in the shade. This was particularly true when thalli were moist during exposure to high light, or presumably became desiccated in full sunlight. However, the reverse trend in the carotenoid composition of green algal lichens was also observed in those species which were found predominantly either in the shade or in full sunlight. In this case sun-exposed lichens often possessed lower levels of zeaxanthin and of the components of the xanthophyll cycle than lichens which were found in the shade. In contrast to higher plants, the lichens from all habitats exhibited a relatively high ratio of carotenoids to chlorophylls (more characteristic of sun leaves), very low levels of α-carotene (similar to that found in sun leaves), and a level of β-carotene similar to that found in shade leaves. Zeaxanthin, but not the expoxides of the xanthophyll cycle, was also frequently found in blue-green algal lichens. A trend for increasing levels of zeaxanthin with increasing growth light regime was observed inPeltigera rufescens, the species which was found to occur over the widest range of light environments. The level of zeaxanthin per chlorophylla in these blue-green algal lichens was in a range similar to that per chlorophylla+b in green algal lichens. However, zeaxanthin was also absent in one species,Collema cristatum, in full sunlight. Thus, the zeaxanthin content of the blue-green algal lichens can be similar to that of higher plants, or it can be rather dissimilar, as was also the case in the green algal lichen species. The presence of large amounts of ketocarotenoids in blue-green algal lichens is also noteworthy.     

Photoinhibition and photoprotection in senescent leaves of field-grown wheat plants

Photosynthesis, photosystem II (PSII) photochemistry, photoinhibition and the xanthophyll cycle in the senescent flag leaves of wheat (Triticum aestivum L.) plants grown in the field were investigated. Compared to the non-senescent leaves, photosynthetic capacity was significantly reduced in senescent flag leaves. The light intensity at which photosynthesis was saturated also declined significantly. The light response curves of PSII photochemistry indicate that a down-regulation of PSII photochemistry occurred in senescent leaves in particular at high light. The maximal efficiency of PSII photochemistry in senescent flag leaves decreased slightly when measured at predawn but substantially at midday, suggesting that PSII function was largely maintained and photoinhibition occurred in senescent leaves when exposed to high light. At midday, PSII efficiency, photochemical quenching and the efficiency of excitation capture by open PSII centers decreased considerably, while non-photochemical quenching increased significantly. Moreover, compared with the values at early morning, a greater decrease in CO₂ assimilation rate was observed at midday in senescent leaves than in control leaves. The levels of antheraxanthin and zeaxanthin via the de-epoxidation of violaxanthin increased in senescent flag leaves from predawn to midday. An increase in the xanthophyll cycle pigments relative to chlorophyll was observed in senescent flag leaves. The results suggest that the xanthophyll cycle was activated in senescent leaves due to the decrease in CO₂ assimilation capacity and the light intensity for saturation of photosynthesis and that the enhanced formation of antheraxanthin and zeaxanthin at high light may play an important role in the dissipation of excess light energy and help to protect photosynthetic apparatus from photodamage. Our results suggest that the well-known function of the xanthophyll cycle to safely dissipate excess excitation energy is also important for maintaining photosynthetic function during leaf senescence.     

Differences in the capacity for radiationless energy dissipation in the photochemical apparatus of green and blue-green algal lichens associated with differences in carotenoid composition

Green algal lichens, which were able to form zeaxanthin rapidly via the de-epoxidation of violaxanthin, exhibited a high capacity to dissipate excess excitation energy nonradiatively in the antenna chlorophyll as indicated by the development of strong nonphotochemical quenching of chlorophyll fluorescence (F_M, the maximum yield of fluorescence induced by pulses of saturating light) and, to a lesser extent, F_O (the yield of instantaneous fluorescence). Blue-green algal lichens which did not contain any zeaxanthin were incapable of such radiationless energy dissipation and were unable to maintain the acceptor of photosystem II in a low reduction state upon exposure to excessive photon flux densities (PFD). Furthermore, following treatment of the thalli with an inhibitor of the violaxanthin de-epoxidase, dithiothreitol, the response of green algal lichens to light became very similar to that of the blue-green algal lichens. Conversely, blue-green algal lichens which had accumulated some zeaxanthin following long-term exposure to higher PFDs exhibited a response to light which was intermediate between that of zeaxanthin-free blue-green algal lichens and zeaxanthin-containing green algal lichens. Zeaxanthin can apparently be formed in blue-green algal lichens (which lack the xanthophyll epoxides, i.e. violaxanthin and antheraxanthin) as part of the normal biosynthetic pathway which leads to a variety of oxygenated derivatives of β-carotene during exposure to high light over several days. We conclude that the pronounced difference in the capacity for photoprotective energy dissipation in the antenna chlorophyll between (zeaxanthin-containing0 green algal lichens and (zeaxanthin-free) blue-green algal lichens is related to the presence or absence of zeaxanthin, and that this difference can explain the greater susceptibility to high-light stress in lichens with blue-green phycobionts.     

Dynamic light use and protection from excess light in upper canopy and coppice leaves of Nothofagus cunninghamii in an old growth, cool temperate rainforest in Victoria, Australia

Responses to simulated sunflecks were examined in upper canopy and coppice leaves of Nothofagus cunninghamii growing in an old-growth rainforest gully in Victoria, Australia. Shaded leaves were exposed to a sudden increase in irradiance from 20 to 1500 micromol m(-2) s(-1). Gas exchange and chlorophyll fluorescence were measured during a 10 min simulated sunfleck and, in the ensuing dark treatment, we examined the recovery of PS II efficiency and the conversion state of xanthophyll cycle pigments. Photosynthetic induction was rapid compared with tropical and northern hemisphere species. Stomatal conductance was relatively high in the shade and stomata did not directly control photosynthetic induction under these conditions. During simulated sunflecks, zeaxanthin was formed rapidly and photochemical efficiency was reduced. These processes were reversed within 30 min in coppice leaves, but this took longer in upper canopy leaves. Poor drought tolerance and achieving a positive carbon balance in a shaded canopy may be functionally related to high stomatal conductance in the shade in N. cunninghamii. The more persistent reduction in photochemical efficiency of upper canopy leaves, which means less efficient light use in subsequent shade periods, but stronger protection from high light, may be related to the generally higher irradiance and longer duration of sunflecks in the upper canopy, but potentially reduces carbon gain during shade periods by 30%.     

Acclimation of leaf carotenoid composition and ascorbate levels to gradients in the light environment within an Australian rainforest

The influence of the growth photon flux density (PFD) on the size and composition of the carotenoid pool and the size of the reduced ascorbate pool was determined across a light gradient from the forest floor to the canopy and the forest edge of a sub-tropical rainforest in New South Wales, Australia. Nineteen plant species (most collected from multiple sites) representing a broad taxonomic range consistently possessed larger total carotenoid pools when found growing in more exposed sites. There was a significant positive correlation between β-carotene content and growth PFD and a significant negative correlation between α-carotene content and growth PFD. Neoxanthin content exhibited no significant trend while the trend in lutein content varied with mode of expression. The pigments of the xanthophyll cycle (violaxanthin, antheraxanthin and zeaxanthin) exhibited the most pronounced response to growth PFD; they comprised a much greater portion of the total carotenoid pool in high light-acclimated plants. The pool of reduced ascorbate was also several-fold greater in high light-acclimated plants. These acclimatory changes in carotenoid and ascorbate content are consistent with a need for a greater capacity to dissipate excessive absorbed light energy in high light-acclimated plants.