Xanthophyll Cycle Activity Research Articles

Wavelength-dependent xanthophyll cycle activity in marine microalgae exposed to natural ultraviolet radiation

The wavelength dependency of xanthophyll cycling in two marine microalgae (Thalassiosira weissflogii and Dunaliella tertiolecta) was studied by establishing biological weighting functions (BWFs) during exposure to natural ultraviolet radiation. High-(HL) and low-(LL) light-acclimated cultures of both species were exposed outdoors for up to 60 min under a series of UVR (280–400 nm) cut-off filters, after which the de-epoxidation state of xanthophyll cycle pigments, radiocarbon assimilation and photochemical quantum yield were measured. Exposures were repeated 4–8 times during the daily cycle to create exposure–response curves for each wavelength condition. UVR affected the three target processes significantly in both species and biological weights increased with decreasing wavelength, particularly in the UVBR region (280–315 nm). Minor wavelength dependency was observed between 315 and 360 nm. After BWF normalization to 300 nm, the LL cultures showed highly similar responses when comparing the three target processes, while the BWFs for the HL cultures differed significantly. The observed enhanced xanthophyll cycling activity in the UVR region implied that xanthophylls had an active role in diminishing UVR stress. However, this enhancement seems to be an indirect effect of damage within the dark reactions of photosynthesis. Hence, another vital target process further downstream in the photosynthetic process, possibly involved in the dark reactions, seems to be responsible for the high similarity in BWFs.

Short-term antioxidative responses of 15 microalgae exposed to excessive irradiance including ultraviolet radiation

Short-term photosensitivity and oxidative stress responses were compared for three groups of marine microalgae: Antarctic microalgae, temperate diatoms and temperate flagellates. In total, 15 low-light-acclimated species were exposed to simulated surface irradiance including ultraviolet radiation (SSI). Photosensitivity was assessed as the rate of recovery of Fv/Fm in the hours following SSI treatment. Before, during and after the SSI treatment, oxidative stress responses were assessed by following xanthophyll content and cycling, and activities of superoxide dismutase, ascorbate peroxidase and glutathione reductase, and glutathione redox status. When acclimated to low irradiance, antioxidant levels were not group specific. Superoxide dismutase activity was positively correlated with cell size, whereas in general, ascorbate peroxidase activity appeared to be lower and glutathione redox status appeared to be higher in the Antarctic than in the temperate species. After SSI exposure, the strong inhibition of PSII was followed by variable rates of recovery, although four species remained photosynthetically inactive. SSI tolerance appeared unrelated to geographic or taxonomic background, or to cell size. PSII recovery was enhanced in species with decreasing superoxide dismutase activity, glutathione redox status and increased xanthophyll cycle activity. We conclude that antioxidant responses are highly species specific and not related to the geographic or taxonomic background. Furthermore, xanthophyll cycling seems more important than antioxidants. Finally, it can be hypothesized that glutathione could function as a stress sensor and response regulator.

Does ultraviolet radiation affect the xanthophyll cycle in marine phytoplankton?

This Perspective summarizes the state of knowledge of the impact of ultraviolet radiation on the photoprotective xanthophyll cycle in marine phytoplankton. Excess photosynthetically active radiation (PAR; 400-700 nm) and ultraviolet radiation (UVR; 280-400 nm) affect various cellular processes and can potentially lead to reduced growth or viability loss in situ. Algae deploy photoprotective mechanisms that limit the hazardous effects of excess light exposure. Xanthophyll cycle pigments play a crucial role in photoprotection via the development of non-photochemical fluorescence quenching (NPQ) during excess radiation exposure. Research on the interacting effects of excess PAR and UVR exposure on xanthophyll cycle pigment synthesis and xanthophyll cycle activity has produced contrasting views. The current contribution summarizes research on photoprotection via photoregulation (xanthophyll cycle activity) and photoacclimation (adjustment of the xanthophyll cycle pigment pool) for marine phytoplankton. Subsequently, UVR effects on the xanthophyll cycle and on xanthophyll cycle pigment pools are discussed and results of supporting experiments with the common diatom Thalassiosira weissflogii are presented. We show that UVR exposure may enhance xanthophyll cycle pigment synthesis. This suggests that UVR-induced reduction in de-epoxidation state does not necessarily imply reduced energy dissipating potential.

Ultraviolet Radiation Reduces the Photoprotective Capacity of the Marine Diatom Phaeodactylum tricornutum (Bacillariophyceae, Heterokontophyta)

This study demonstrates that UV radiation (UVR) reduces the photoprotective capacity of the diatom Phaeodactylum tricornutum by affecting xanthophyll cycle (XC) activity. The short-term reduction of photosystem II (PSII) maximum efficiency of charge separation (F(v)/F(m)) in cells exposed to UVR could be explained mainly by a reduced photoprotective capacity under this condition. Phaeodactylum tricornutum cells acclimated to two different photosynthetically active radiation (PAR) intensities, high light (HL, 200 micromol quanta m(-2) s(-1)) and low light (LL, 50 micromol quanta m(-2) s(-1)), were exposed to saturating irradiance (1100 micromol quanta m(-2) s(-1)) in the presence (PAR + UVR) and absence of UVR (PAR). HL cells exhibited a greater reduction in F(v)/F(m) in PAR + UVR when compared with the PAR treatment that was related to a reduction in the de-epoxidation of XC pigments. In contrast, in LL cells, UVR did not considerably affect XC de-epoxidation even though the reduction in F(v)/F(m) was greater than in HL cells. The negative effect of UVR on photoprotection was more pronounced in HL cells because they synthesized more XC pigments than LL cells. This was confirmed when XC activity was blocked with dithiothreitol and when PSII repair was inhibited with chloramphenicol (CAP). The differential reduction of F(v)/F(m) between PAR + UVR and PAR treatments disappeared when XC was blocked in HL cells. A higher reduction and an incomplete recovery of F(v)/F(m) were observed in cells incubated with CAP in the presence of UVR. Such responses confirm that UVR had a negative effect on photoprotective mechanisms causing an enhancement of damage by PAR, especially in HL-acclimated cells in which heat dissipation is important for PSII regulation.

New transgenic line of Arabidopsis thaliana with partly disabled zeaxanthin epoxidase activity displays changed carotenoid composition, xanthophyll cycle activity and non-photochemical quenching kinetics

Zeaxanthin epoxidase (ZE, E.C. 1.14.13.90), an enzyme belonging to the lipocalin superfamily, catalyses the conversion of zeaxanthin to antheraxanthin and violaxanthin. These reactions are part of the xanthophyll biosynthetic pathway and the xanthophyll cycle. The role of carotenoids in the dissipation of excessive light energy has been widely studied using mutants with a disabled carotenoid biosynthetic pathway. In this paper, the transgenic line MaZEP7 with partially disabled ZE activity is described and compared with wild-type plants and npq2 mutant lacking active ZE. We examined the presence and the abundance of aba1 transcripts, measured pigment composition, xanthophyll cycle functioning and chlorophyll fluorescence in all three lines. The MaZEP7 line contains additional copies of the aba1 gene introduced by agroinfiltration, but no enhanced aba1 transcript level was observed. In addition, ZE activity in MaZEP7 was impaired, resulting in an altered xanthophyll profile. In dark-adapted plants, violaxanthin and neoxanthin levels were lower than in wild-type plants, whereas antheraxanthin and zeaxanthin levels were considerably higher. The presence of lutein epoxide was also observed. Violaxanthin levels changed only minimally during light exposition, whereas antheraxanthin was converted to zeaxanthin and there was no epoxidation during the course of the experiment indicating disturbed xanthophyll cycle functioning. The amounts of carotenoids and chlorophylls on a dry weight basis and chl a/chl b ratio were similar in all lines. The presence of epoxidated pigments in MaZEP7 plants indicates that epoxidation occurs, but it is likely very slow. Chlorophyll fluorescence measurements showed that the dependence of electron transport rates on light intensity for the MaZEP7 line resembled the npq2 mutant. Kinetic measurements showed that the MaZEP7 line exhibited very rapid induction and a high steady-state value of non-photochemical quenching.

The regulation of xanthophyll cycle activity and of non-photochemical fluorescence quenching by two alternative electron flows in the diatoms Phaeodactylum tricornutum and Cyclotella meneghiniana

Intact cells of diatoms are characterized by a rapid diatoxanthin epoxidation during low light periods following high light illumination while epoxidation is severely restricted in phases of complete darkness. The present study shows that rapid diatoxanthin epoxidation is dependent on the availability of the cofactor of diatoxanthin epoxidase, NADPH, which cannot be generated in darkness due to the inactivity of PSI. In the diatom Phaeodactylum tricornutum, NADPH production during low light is dependent on PSII activity, and addition of DCMU consequently abolishes diatoxanthin epoxidation. In contrast to P. tricornutum, DCMU does not affect diatoxanthin epoxidation in Cyclotella meneghiniana, which shows the same rapid epoxidation in low light both in the absence or presence of DCMU. Measurements of the reduction state of the PQ pool and PSI activity indicate that, in the presence of DCMU, NADPH production in C. meneghiniana occurs via alternative electron transport, which includes electron donation from the chloroplast stroma to the PQ pool and, in a second step, from PQ to PSI. Similar electron flow to PQ is also observed during high light illumination of DCMU-treated P. tricornutum cells. In contrast to C. meneghiniana, the electrons are not directed to PSI, but most likely to a plastoquinone oxidase. This chlororespiratory electron transport leads to the establishment of an uncoupler-sensitive proton gradient in the presence of DCMU, which induces diadinoxanthin de-epoxidation and NPQ. In C. meneghiniana, electron flow to the plastoquinone oxidase is restricted, and consequently, diadinoxanthin de-epoxidation and NPQ is not observed after addition of DCMU.

Biochemical constrains limit the potential of the photochemical reflectance index as a predictor of effective quantum efficiency of photosynthesis during the winter spring transition in Jack pine seedlings.

Leaf reflectance spectral measurements are an emerging non-invasive technique that can be used to derive the photochemical reflectance index (PRI) to assess the physiological state of plants from leaf to ecosystem level. Changes in PRI are associated with changes in the xanthophyll cycle activity and provide an estimate of changes in the effective photochemical quantum efficiency (ΦII) during the growing season. However, we hypothesised that the correlation between PRI and ΦII might be poor when the xanthophyll cycle is primed for sustained thermal dissipation of the light energy absorbed. To test our hypothesis, we studied the recovery of winter acclimated Jack pine (Pinus banksiana Lamb.) seedlings that were exposed to different simulated spring recovery treatments in controlled environments. Different growth temperatures and light intensities were used to dissect the effect of these two factors on chlorophyll fluorescence, pigment composition and leaf reflectance. ΦII showed a clear response to temperature whereas PRI was mostly affected by light intensity. In contrast, the de-epoxidation state of the xanthophyll cycle pigments was both temperature and light dependent. Our data suggest that zeaxanthin-independent non-photochemical quenching is employed to various degrees in the different treatments. As a result, within the limits of our experimental setup, PRI could not explain the variation in ΦII. This indicates that an improved understanding of the different energy quenching mechanisms is critical to accurately interpret the PRI signal under environmental conditions where the predominant mode of excess energy dissipation does not involve a dynamic operation of the xanthophyll cycle, but a sustained mechanism of energy dissipation.

Willow species (genus: Salix) with contrasting habitat affinities differ in their photoprotective responses to water stress.

Although many Mediterranean and xeric plant species enhance their xanthophyll-mediated thermal dissipation under drought conditions, there has been limited research on photoprotective mechanism in droughted plants from other habitats. To investigate whether wetland plants utilise this mechanism under drought conditions, and whether species differ in their responses depending on their habitat affinities, we investigated the response of six willow (Salix) species to a short-term drought. In a greenhouse, 40 individuals per species were dried down over 4 weeks. Periodically during the drought, predawn and midday chlorophyll fluorescence measurements were taken and leaf discs were collected for pigment analysis with HPLC. Predawn water potential was also monitored throughout the experiment. All six species increased xanthophyll cycle activity and their capacity to dissipate excess energy during the drought by increasing their total de-epoxidised xanthophyll concentration and the concentration of zeaxanthin in proportion to chlorophyll. In general, habitat generalists had greater photoprotective responses than wetland specialists, while the wetland specialists had higher pre-drought nonphotochemical quenching. These differences are consistent with their contrasting photosynthetic rates. The observed variation in species drought responses suggests that their photoprotective strategies vary with habitat affinity.

DETECTION OF DIATOM XANTHOPHYLL CYCLE USING SPECTRAL REFLECTANCE(1).

Analysis of reflectance spectra was used to monitor the conversion of diadinoxanthin (DD) into diatoxanthin (DT) in two benthic diatom species, Amphora coffeaeformis (C. Agardh) Kütz. and Cylindrotheca closterium (Ehrenb.) J. C. Lewin et Reiman, cultured at high light (HL, 400 μmol · m(-2) · s(-1) PAR) and low light (LL, 25 μmol · m(-2) · s(-1) PAR). Cultures were exposed to saturating light for 32 min. HL cultures of both species showed higher (DT + DD) content, whereas LL cultures exhibited higher chl a and fucoxanthin content. DD to DT conversion, measured by HPLC, occurred mainly in the first 2 min (LL) or 5 min (HL) after exposure to saturating light. Nonphotochemical quenching (NPQ), measured by PAM fluorescence, showed the same pattern as DT/(DD + DT), resulting in a linear relationship between these parameters. Addition of dithiothreitol (DTT) blocked the conversion of DD into DT and significantly reduced NPQ induction. Reflectance spectra showed no obvious change after light exposure. However, second derivative spectra (δδ) showed a shift in reflectance from 487 to 508 nm, which was not present for DTT-treated samples. Changes in δδ487 were strongly correlated with changes in DD (r = 0.76), while changes in δδ508 were strongly correlated with changes in DT (r = 0.94). The best index to estimate DD to DT conversion was δδ508 /δδ630 (r = 0.87). This index was very sensitive to minute changes that occurred immediately after exposure to light and was species insensitive. Good relationships were observed between indices for xanthophyll cycle activation (DD to DT conversion and NPQ induction) and the second derivative spectra. With further in situ validation, this index may prove to be highly useful for investigation into aquatic global photoregulation mechanisms in diatom-dominated samples.

Changes in photosynthesis, xanthophyll cycle, and sugar accumulation in two North Australia tropical species differing in leaf angles

Two tropical species of North Australia, Acacia crassicarpa and Eucalyptus pellita, have similar leaf size and leaf structure but different leaf angles. A. crassicarpa with near vertical leaf angle directly reduced photon absorption and leaf temperature (T l) and had relatively high photosynthetic activity (P max) and low xanthophyll cycle activity. In contrast, E. pellita with a small leaf angle exhibited high T l, low P max, and high activity of xanthophyll cycle which was useful for the dissipation of excessive energy and reduction of photoinhibition. In the dry season, contents of soluble sugars including pinitol, sucrose, fructose, and glucose in A. crassicarpa increased whereas larger amounts of only fructose and glucose were accumulated in E. pellita. Different sugar accumulation may be involved in osmotic adjustment of leaves during water stress that makes photosynthesis more efficient. The leaf angle may be critical for developing different protective mechanisms in these two tropical tree species that ensure optimal growth in the high irradiance and drought stress environment in North Australia.

Dynamics of the xanthophyll cycle and non-radiative dissipation of absorbed light energy during exposure of Norway spruce to high irradiance

The response of Norway spruce saplings (Picea abies [L.] Karst.) was monitored continuously during short-term exposure (10 days) to high irradiance (HI; 1000micromol m(-2)s(-1)). Compared with plants acclimated to low irradiance (100micromol m(-2)s(-1)), plants after HI exposure were characterized by a significantly reduced CO(2) assimilation rate throughout the light response curve. Pigment contents varied only slightly during HI exposure, but a rapid and strong response was observed in xanthophyll cycle activity, particularly within the first 3 days of the HI treatment. Both violaxanthin convertibility under HI and the amount of zeaxanthin pool sustained in darkness increased markedly under HI conditions. These changes were accompanied by an enhanced non-radiative dissipation of absorbed light energy (NRD) and the acceleration of induction of both NRD and de-epoxidation of the xanthophyll cycle pigments. We found a strong negative linear correlation between the amount of sustained de-epoxidized xanthophylls and the photosystem II (PSII) photochemical efficiency (F(V)/F(M)), indicating photoprotective down-regulation of the PSII function. Recovery of F(V)/F(M) at the end of the HI treatment revealed that Norway spruce was able to cope with a 10-fold elevated irradiance due particularly to an efficient NRD within the PSII antenna that was associated with enhanced violaxanthin convertibility and a light-induced accumulation of zeaxanthin that persisted in darkness.

The giant kelp Macrocystis pyrifera presents a different nonphotochemical quenching control than higher plants.

Here the mechanisms involved in excitation energy dissipation of Macrocystis pyrifera were characterized to explain the high nonphotochemical quenching of chlorophyll a (Chla) fluorescence (NPQ) capacity of this alga. We performed a comparative analysis of NPQ and xanthophyll cycle (XC) activity in blades collected at different depths. The responses of the blades to dithiothreitol (DTT) and to the uncoupler NH4Cl were also assayed. The degree of NPQ induction was related to the amount of zeaxanthin synthesized in high light. The inhibition of zeaxanthin synthesis with DTT blocked NPQ induction. A slow NPQ relaxation upon the addition of NH4Cl, which disrupts the transthylakoid proton gradient, was detected. The slow NPQ relaxation took place only in the presence of de-epoxidated XC pigments and was related to the epoxidation of zeaxanthin. These results indicate that in M. pyrifera, in contrast to higher plants, the transthylakoid proton gradient alone does not induce NPQ. The role of this gradient seems to be related only to the activation of the violaxanthin de-epoxidase enzyme.

Photoinhibition and xanthophyll cycle activity in bayberry (Myrica rubra) leaves induced by high irradiance

The effect of high irradiance (HI, photosynthetically active photon flux density of 1 300 µmol m−2 s−1) on net photosynthetic rate (P N), chlorophyll fluorescence parameters, and xanthophyll cycle components were studied in fruit tree bayberry leaves. HI induced the photoinhibition and inactivation of photosystem 2 (PS2) reaction centres (RCs), which was characterized by decreased P N, maximum yield of fluorescence after dark adaptation (Fm), photochemical efficiency of PS2 (Fv/Fm) and quantum yield of PS2 (ΦPS2), and increased reduction state of QA (1-qP) and non-photochemical quenching (NPQ). Initial fluorescence (F0) showed a decrease after the first 2 h, and subsequently increased from the third hour exposure to HI. Furthermore, a greater increase in the ratio (Fi-F0)/(Fp-F0) which is an expression of the proportion of the QB non-reducing PS2 centres, whereas a remarked decrease in the slope of Fi to Fp which represents the rate of QA reduction was observed in leaves after HI exposure. Additionally, HI caused an increase in the pool size of the xanthophyll cycle pigments and sustained elevated contents of zeaxanthin (Z), antheraxanthin (A), and de-epoxidation state (DES) at the end of the irradiation period. During HI, decreased Fm, Fv/Fm, ΦPS2, NPQ, slope of Fi to Fp, V+A+Z, and DES, and increased F0, 1-qP, ratio (Fi-F0)/(Fp-F0), and V were observed in dithiothreitol (DTT)-fed leaves compared to control ones under the same conditions. Hence photoinhibition caused by HI in bayberry was probably attributed to inactivation of PS2 RCs, and photoprotection from photodamage were mainly related to the xanthophyll cycle-dependent heat dissipation in excess photons.

The roles of specific xanthophylls in light utilization

To evaluate the role of specific xanthophylls in light utilization, wild-type and xanthophyll mutant plants (npq1, npq2, lut2, lut2npq1 and lut2npq2) from Arabidopsis thaliana were grown under three different light regimes: 30 (low light, LL), 150 (medium light, ML) and 450 (high light, HL) mumol photons m(-2) s(-1). We studied the pigment content, growth rate, xanthophyll cycle activity, chlorophyll fluorescence parameters and the response to photoinhibition. All genotypes differed strongly in the growth rates and the resistance against photoinhibition. In particular, replacement of lutein (Lut) by violaxanthin (Vx) in the lut2npq1 mutant did not affect the growth at non-saturating light intensities (LL and ML), but led to a pronounced reduction of growth under HL conditions, indicating an important photoprotective role of Lut. This was further supported by a much higher sensitivity of all Lut-deficient plants to photoinhibition in comparison with the wild type. In contrast, replacement of Lut by zeaxanthin (Zx) in lut2npq2 led to a pronounced reduction of growth under all light regimes, most likely related to the permanent non-photochemical dissipation of excitation energy by Zx at Vx-binding sites and the destabilization of antenna proteins by binding of Zx to Lut-binding sites. The high susceptibility of lut2npq2 to photoinhibition in comparison with npq2 further indicated that the photoprotective function of Zx is abolished in the absence of Lut. Thus, it can be concluded from our work that neither Vx nor Zx is able to fulfil the essential photoprotective function at Lut-binding sites under in vivo conditions.

The impact of shifts to elevated irradiance on the growth and photochemical activity of the harmful algae Chattonella subsalsa and Prorocentrum minimum from Delaware

The physiological impact of short and long-term shifts from low to high light intensity (30–600 μmol photons m −2 s −1) was assessed for the harmful blooming raphidophyte, Chattonella subsalsa and dinoflagellate, Prorocentrum minimum, isolated from the inland bays of Delaware. Analysis of photochemical activity by active chlorophyll fluorescence indicated a significant decline in the photochemical capacity of photosystem II (PSII), F v/ F m, in both algal species when shifted to high light, with a greater decline noted in P. minimum. While there was no significant difference in the effective quantum yield in the light acclimated state ( Δ F / F m ′ ) under high light between these algae, there were differences in how light energy was utilized between the two species. P. minimum displayed a typical upregulation in light energy dissipation via nonphotochemical pathways by significant elevated nonphotochemical fluorescence quenching and a rapid reduction in electron transport with increasing immediate light exposure. Low light acclimated C. subsalsa maintained greater electron transport during short term high light exposure, and showed minimal change in nonphotochemical energy dissipation despite higher excitation pressure placed upon PSII when shifted to high light for several days. C. subsalsa displayed a significantly higher growth rate than P. minimum following the shift of unialgal cultures from low to high light, while both species grew at equivalent rates when cultured together and shifted to high light. Results from these experiments and analysis of loss and subsequent recovery of PSII in the presence and absence of chloroplast protein translation suggest that during shifts to high light P. minimum relies heavily on photosystem turnover and repair as well as nonphotochemical dissipation most likely from xanthophyll cycle activity. Meanwhile, C. subsalsa appears to down-regulate the number of functional PSII reaction centers yet maintains photochemical activity through lowered electron transport to avoid potential photoinhibition.

The xanthophyll cycle activity in kidney bean and cabbage leaves under salinity stress

Seven-day-old kidney bean and cabbage seedlings were treated with 0.1–0.3 M NaCl solutions for 3 days. Chlorophyll content decreased in NaCl-treated Phaseolus seedlings, but did not significantly decrease in Brassica seedlings. Photochemical efficiency of photosystem II at dark-adapted state was similar in both Phaseolus and Brassica. The de-epoxidation state of violaxanthin increased more than sixfold in Phaseolus but showed no significant change in Brassica seedlings during NaCl treatment under low light. Maximum de-epoxidation state of violaxanthin in vivo tested in high light (2000 μmol quanta/(m2 s) increased in salt-stressed Phaseolus but decreased in Brassica seedlings. The nonphotochemical quenching (NPQ) also increased in Phaseolus but decreased in Brassica. This suggests that xanthophyll cycle pigments influence the NPQ in both Phaseolus and Brassica, but in an opposite way. The increase in the de-epoxidation state of violaxanthin in salt-stressed Phaseolus even under low light may be considered an early light signal to protect the pigment-protein complexes from salt-stress induced photodamage. It is proposed that in salt-stressed Brassica, the de-epoxidation is retarded and/or the epoxidation is accelerated leading to the accumulation of violaxanthin and a lower de-epoxidation state. Thus, light-induced violoxanthin cycle operation largely controls the photoprotection of photosynthetic apparatus in kidney bean leaves.

Temperature and Light Modulate the trans-Δ3-Hexadecenoic Acid Content of Phosphatidylglycerol: Light-harvesting Complex II Organization and Non-photochemical Quenching

The interaction of light and temperature in the modulation of the trans-delta3-hexadecenoic acid (trans-16:1) content of phosphatidylglycerol (PG) in winter rye (Secale cereale L.) was assessed and related to the organization of light-harvesting complex II (LHCII). Increasing the growth irradiance from 50 to 800 micromol m(-2) s(-1) at 20 degrees C resulted in a 1.8-fold increase in the trans-16:1 content in PG which favoured a greater preponderance of oligomeric LHCII, measured in vitro as the ratio of oligomer : monomer. Similar irradiance-dependent increases were observed during growth at 5 degrees C; however, 1.4-fold lower trans-16:1 contents and lower LHCII oligomer : monomer ratios were observed compared with growth at 20 degrees C and the same irradiance. These trends were also observed under natural field conditions. Thus, the accumulation of trans-16:1, as well as the organization of LHCII are modulated by both growth irradiance and growth temperature in an independent but additive manner. We also examined how changes in the supramolecular organization of LHCII affected the capacity for non-photochemical quenching (q(N)) and photoprotection via antenna quenching (q(O)). While q(O) was positively correlated with q(N), there was no correlation with either LHCII organization or xanthophyll cycle activity under the steady-state growth conditions examined.

Rapid light curves: A powerful tool to assess photosynthetic activity

Rapid light curves provide detailed information on the saturation characteristics of electron transport, as well as the overall photosynthetic performance of a plant. Rapid light curves were collected from samples of Zostera marina grown under low and high-light conditions (50 and 300 μmol photons m −2 s −1) and the distinctive patterns of RLC parameters are discussed, in terms of differential sink capacity and PSII reaction centre closure. Derived cardinal points of a rapid light curve ( α, E k and rETR max) describe the photosynthetic capacity of a seagrass leaf, its light adaptation state and its capacity to tolerate short-term changes in light. The shapes of the corresponding F and F ′ m curves also provide information on the development of the trans-thylakoid proton gradient and thermal energy dissipation. Low-light leaves showed limited photosynthetic capacity and reduced activity of non-photochemical quenching pathways, whereas photosynthesis of high light leaves were not limited and showed an elevated level of non-photochemical quenching, possibly associated with xanthophyll cycle activity. Light-dark kinetics are also discussed in relation to relaxation of non-photochemical quenching and its various components. A curve fitting model is recommended based on the double exponential decay function. In this paper, we explain the fundamental aspects of a RLC, describe how it reflects the response to light exposure of a leaf, how to interpret these curves, and how to quantitatively describe and compare RLCs.

Xanthophyll cycle activity in detached barley leaves senescing under dark and light

Senescence-induced changes in the xanthophyll cycle activity and chlorophyll (Chl) fluorescence parameters were compared in detached barley (Hordeum vulgare L.) leaf segments kept for 6 d in darkness or under continuous “ white light” (90 μmol m−2 s−1). Before detachment of the leaf segments, the plants were grown at periodic regime [12 h light (90 μmol m−2 s−1)/12 h dark]. The de-epoxidation state of the xanthophyll cycle pigments (DEPS) in the leaf samples was determined immediately (the actual DEPS), after 1 h of dark-adaptation (the residual DEPS), and during 14 min of a high-irradiance (HI) exposure (500 μmol m−2 s−1) (HI-induced DEPS). In the light-senescing segments, senescence was delayed pronouncedly compared to dark-senescing ones as the Chl content, the photosystem 2 photochemistry, and electron transport processes were highly maintained. Further, the actual DEPS increased, probably due to the increased mean photon dose. The HI-induced increase in the DEPS was stimulated in the light-senescing segments, whereas it was slowed down in the dark-senescing ones. However, after the 14 min HI-exposure of the dark-senescing segments the HI-induced DEPS was not markedly lower than in the mature leaves, which indicated the maintenance of the xanthophyll cycle operation.

The super-excess energy dissipation in diatom algae: comparative analysis with higher plants

When grown at intermittent light regime, diatom alga Phaeodactylum tricornutum is able to form photoprotective non-photochemical chlorophyll fluorescence quenching (NPQ) three to five times larger than that observed in the higher plants. This quenching is sustained in the dark for 5 to 10 min, reverses completely within approximately 1 h and seems to be very tightly related to the presence of the zeaxanthin analogue, diatoxanthin. Addition of the uncoupler NH4Cl before illumination can completely abolish formation of NPQ, revealing the DeltapH-dependency of the xanthophyll cycle activity. Once established, NPQ can also be almost completely reversed by the uncoupler. However, the higher NPQ is formed the more time is required for its reversal. At the point when the fluorescence was approximately 90% recovered the level of illumination-induced diatoxanthin was found to be only partially reduced. This indicates that the proton gradient is a key triggering factor of NPQ. It was also noticed that NPQ in Phaeodactylum cells was absent even when majority of reaction centers were closed and the plastoquinone pool was significantly reduced. The absence of NPQ at these conditions could be due to very low levels of DeltapH. It is likely that in diatoms alternative sources of protons such as the PS I cyclic electron transfer and/or chlororespiration are important in generating the proton gradient sufficient to trigger NPQ. Absorption changes associated with the xanthophyll cycle activity were found to be larger than those for higher plants. The position of the positive maximum in the difference spectrum illuminated-minus-dark was 512-514 nm in comparison to the 505-508 nm for leaves. The 535 nm band associated with NPQ in plants is absent in Phaeodactylum. An uncoupler-sensitive absorption change at 522 nm was discovered. Kinetics of NPQ showed linear correlation with the 522 nm absorption change.