Parietin in the tolerant lichen Xanthoria parietina (L.) Th. Fr. increases protection of Trebouxia photobionts from cadmium excess

Secondary metabolites of lichens can be involved in production of chelates with heavy metals. We hypothesized that parietin plays important role in protection of photobiont cells in Xanthoria parietina from an excess of cadmium ions. Two types of X. parietina lichen thalli, natural with presence of secondary metabolite parietin (p+) as well as without parietin (p−) were exposed to different doses of cadmium (up to 300 μmol g⁻¹ dw). Based on determination of the total and intracellular Cd-accumulation, ergosterol and thiobarbituric acid reactive substances (TBARS) content did not show statistically significant differences in the response of both types of thalli (p+ and p−). However, a stronger toxic effect of the highest Cd-dose on photosynthetic pigment content and chlorophyll a fluorescence was observed in the parietin-depleted thalli. The protective role of parietin against Cd excess was better supported and concluded from the differences observed in the production of non-protein thiol compounds (cysteine, glutathione and phytochelatins) involved in Cd detoxification. In the p+ thalli Cys content was stable but GSH content slightly decreased in the studied Cd range, while in the p− thalli these compounds were completely absent at high Cd doses. At Cd doses higher than 37.5 μmol Cd g⁻¹ dw, toxic to both types of X. parietina thalli, Cys and GSH contents were significantly higher in p+ than in p− thalli. Also, the photobiont partner in the p+ thalli was better protected of the metal exposition, and able to produce phytochelatins (PCs) over the whole range of metal, while in the p− thalli the production was completely inhibited at 75 μmol Cd g⁻¹ dw and higher concentrations, together with the inhibition of cysteine (Cys) and reduced glutathione (GSH) production. The obtained results indicate that the parietin layer is a natural barrier decreasing Cd access to algal cells in X. parietina. Comparison of PCs production appeared to be the most sensitive marker for estimation of Cd availability to photobiont in the symbiotic system.     

The bromoperoxidase from the lichen Xanthoria parietina is a novel vanadium enzyme.

A novel bromoperoxidase was isolated from the lichen Xanthoria parietina. The enzyme contained vanadium, which is essential for enzymic activity. Under denaturating conditions the preparation showed a single protein band with an Mr of 65,000. Thermal-denaturation studies showed that this bromoperoxidase could tolerate high temperatures. The affinity of the enzyme for its substrate bromide is high; the Km for bromide was 29 microM. Excess halides (50 mM) inhibited enzymic activity considerably.     

Stomatal aperture,photosythesis and water fluxes in mesophyll cells as affected by the abscission of leaves. Simultaneous measurements of gas exchange,light scattering and chlorphyll fluorescence

Carbon dioxide exchange, transpiration, chlorophyll fluorescence and light scattering of leaves of Lycopersicom esculentum, Helianthus annuus and Arbutus unedo were measured simultaneously before and after abscission of leaves. Scattering of a weak green measuring beam was used to monitor water fluxes across the thylakoid membranes of the mesophyll. When leaves were cut under water, stomata initially closed partially and then occasionally exhibited distinct regulatory oscillations. As stomata closed, light scattering decreased indicating water influx into the mesophyll. Stomatal oscillations were accompanied, with small but noticeable phase shifts, by oscillations of water fluxes at the thylakoid level. These fluxes could be distinguished from the water fluxes accompanying light-dependent ion pumping across the thylakoids by the concomitant chlorophyll fluorescence signals. The latter record energy-dependent ion fluxes in addition to redox changes of the electron-transport chain. As stomata closed partially after cutting a leaf under water, photosynthesis decreased. In Arbutus unedo and Helianthus annuus leaves, transient stomatal closure was insufficient to account for transient inhibition of photosynthesis which appeared to be brought about by transfer of an inhibitory solute through the petiole into the mesophyll. This solute also stimulated respiration in the dark. When leaves were cut in air, stomata opened transiently (Iwanoff effect) before wilting enforced closure. Photosynthesis followed the stomatal responses, increasing during opening and decreasing during closure.Dedicated to Professor H. Ullrich on the occasion of his 85th birthday     

Synthesis and Antioxidant Evaluation of O-Methylated Emodacidamides: Starting from Parietin,a Secondary Metabolite of Lichen Xanthoria parietina

Polyhydroxy-anthraquinones bearing amino acids are found rather seldom in nature. Emodacidamides, isolated from a marine-derived fungus, Penicillium sp. SCSIO sof101 by Luo et al. (2017) are the first natural example of amino acid conjugated anthraquinone. In this study, O-methylated emodacidamides and emodinic acid-anilides were synthesized starting from parietin, extracted from the lichen Xanthoria parietina (L.) Th. Fr. The structural elucidations of prepared compounds were confirmed by 1D and 2D NMR analyses including HSQC and HMBC techniques. In addition, all newly synthesized compounds were evaluated for the antioxidant activities with free radical 1,1-diphenyl-2-picrylhydrazyl (DPPH) scavenging. The synthesized compounds showed low to moderate antioxidant and DPPH scavenging activities. The antioxidant activities were supported within quantum chemical calculations using the DFT−B3LYP/6-311++G(d,p) level of theory. It is observed that the antioxidant activity of emodacidamides mostly depends on the phenolic groups on anthraquinone ring. The phenolic groups on other substituents help to improve antioxidant activity and also the position of hydroxy group is a decisive factor for antioxidant ability.     

The effect of different night conditions on the CO2 fixation in a lichen Xanthoria parietina

CO₂ fixation was studied in a lichen, Xanthoria parietina, kept in continuous light, and with cyclic changes in light intensity, dark period or temperature. The diurnal and seasonal courses of CO₂ exchange were followed. The rate of net photosynthesis was observed to fall from morning to evening, and this decline was more pronounced in winter than in summer. The maximal net photosynthetic rate, 223 ng CO₂g^-1dws^-1, occured in winter and the minimum, 94 ng CO₂g^-1dws^-1, late in spring. The light compensation point in summer was four times as high as in winter. In continuous light (180 or 90 mol photons m^-2s^-1, 15°C) net photosynthesis decreased noticeably during one week, falling below the level maintained in a 12 h light: 12 h dark cycle. Photosynthetic activity did not decrease, however, in lichens held in continuous light (90 mol photons m^-2s^-1) with cyclic changes of temperature (12 h 20 °C: 12 h 5 °C). Active photosynthesis was also maintained in light of cyclically changing intensity (12 h: 12 h, 15 °C) when night-time light was at least 75% lower than illumination by day. A dark period of 4 hours in a 24-h light:dark cycle was sufficient to keep CO₂ fixation at the control level. It seems that plants need an unproductive period during the day to survive and this can be induced by fluctuations in light and/or temperature.     

Photosynthates stimulate the UV-B induced fungal anthraquinone synthesis in the foliose lichen Xanthoria parietina

Synthesis of the cortical anthraquinone pigment parietin (= physcion) was studied in acetone‐rinsed, parietin‐free Xanthoria parietina thalli. UV‐B induced the synthesis, which increased linearly with UV‐B (log‐transformed) to the highest applied UV‐B level (1.8 W m^?2). At natural UV‐B levels (0.75 W m^?2), parietin resynthesis occurred at a constant pace (106 mg m^?2 d^?1) during a 14‐d period at 220 µmol m^?2 s^?1 PAR. Under these conditions, 56% of the natural parietin content prior to extraction was resumed, accounting for 10% of total net carbon gain. In the presence of UV‐B, the remaining results were consistent with the hypothesis assuming that photosynthates regulate the pace at which parietin is synthesized by the mycobiont. Resynthesis was rapid when photosynthesis was activated by light, or when certain carbohydrates were added exogenously. Additions of ribitol, the carbohydrate delivered from the photobiont, increased the parietin resynthesis substantially. Mannitol, the main fungal polyol, was significantly less effective. Furthermore, parietin resynthesis in X. parietina was depressed at high and low hydration when net photosynthesis is depressed. Therefore, the photobiont regulates the parietin resynthesis pace in its mycobiont partner by the delivery of photosynthates. In conclusion, both lichen bionts play important roles in the synthesis of parietin, which probably acts as a PAR‐ rather than a UV‐B‐screen.     

Desiccation tolerance of the epilithic lichen Lecanora muralis (Schreb.) Rabenh. in the temperate climate

Lecanora muralis (Schreb.) Rabenh. is a ubiquitous epilithic crustose lichen of the temperate climate. It is well studied in terms of diel and annual carbon budget and productivity with continuous long-term observations in the field in 1995/96 by Otto L. Lange, Würzburg, and collaborators. However, these earlier studies left open the question to which extent the lichen is desiccation tolerant and if desiccation tolerance might possibly limit photosynthetic activity. In present study measurements of chlorophyll fluorescence parameters were performed to assess photosynthetic activity under various daily weather conditions throughout the year and recovery from desiccation after various periods of dryness in ambient air. Under any weather conditions, including strong frost for several days with night-temperatures around −15 °C and strong heat of several days with day-temperatures around 35 °C, the lichen was fully photosynthetically competent after wetting the samples for 15 min by submersion in water when they were dry in the field in the absence of actual incident precipitation. Chlorophyll fluorescence parameters were identical under all weather conditions sampled. A sample kept dry in ambient air for 37 days showed full recovery of chlorophyll fluorescence parameters after wetting for 30 min. Samples desiccated for longer periods up to 155 days took longer wetting times of about 300 min and recovered only partially but nevertheless showed active photosynthetic electron transport. Of 17 samples desiccated for 177–178 days only three recovered after rewetting for several days. It is concluded that the desiccation tolerance of L. muralis is sufficient to overcome dry spells of duration as it normally occurs in its natural environment. Desiccation tolerance is not likely to limit carbon budget and productivity.     

珍稀树种红花玉兰在华北地区的最适光环境

珍稀树种红花玉兰对其华南原产地的自然环境有良好的适应性, 但在华北地区却生长不良。通过对红花玉兰在华北地区一个生长季内对三种光照水平(100%、70%、40%全光照)的光合和生长响应分析, 结果表明:在70%全光照条件下, 红花玉兰幼苗的净光合速率、光饱和点、株高、基径、根生物量和茎生物量均达到最高水平。随着光照强度的减弱, 暗呼吸速率、光补偿点、比叶重量、叶片厚度和密度显著降低, 表观量子效率、最大荧光Fm、可变荧光Fv、Fm/Fo(Fo为初始荧光)、Fv/Fo、Fv/Fm、叶绿素含量、叶面积和叶柄角度均显著增大。说明70%全光照最适于一年生红花玉兰幼苗在华北地区的生长, 全光照和40%全光照条件下幼苗则因光量的过剩和不足而生长不良。因此建议将红花玉兰栽植在林缘或林窗地带, 可为这一珍稀濒危树种在华北地区的引种提供有利的适生光照环境。     

Light scattering,chlorophyll fluorescence and state of the adenylate system in illuminated spinach leaves

Scattering of green light and chlorophyll fluorescence by spinach leaves kept in a stream of air or nitrogen were compared with leaf adenylate levels during illumination with blue, red or far-red light. Energy charge and ATP-ADP ratios exhibited considerable variability in different leaves both in the dark and in the light. Variability is explained by different possible states of the reaction oxidizing triose phosphate or reducing 3-phosphoglycerate. Except when oxygen levels were low, there was an inverse relationship between light scattering and chlorophyll fluorescence during illumination with blue or red light. When CO₂ was added to a stream of CO₂-free air, chlorophyll fluorescence increased, sometimes after a transient decrease, and both light scattering and leaf $\text{ATP}\text{ADP}$ ratios decreased. Similar observations were made when air was replaced by nitrogen under blue or high-intensity red light. Under these conditions, over-reduction caused inhibition of electron transport and phosphorylation in chloroplasts. However, when air was replaced by nitrogen during illumination with low-intensity red light or far-red light, light scattering increased instead of decreasing. Under these light conditions, $\text{ATP}\text{ADP}$ ratios were maintained in the light. They decreased drastically only after darkening. Although $\text{ATP}\text{ADP}$ ratios responded faster than light scattering or the slow secondary decline of chlorophyll fluorescence due to illumination, it appeared that in the steady state, light scattering and chlorophyll fluorescence are useful indicators of the phosphorylation state of the leaf adenylate system at least under aerobic conditions, when chloroplast and extrachloroplast adenylate systems can effectively communicate.     

Pseudocyphellaria dissimilis: a desiccation-sensitive,highly shade-adapted lichen from New Zealand

Summary Pseudocyphellaria dissimilis, a foliose, cyanobacterial lichen, is shown not to fit into the normal ecological concept of lichens. This species is both extremely shade-tolerant and also more intolerant to drying than aquatic lichens previously thought to be the most desiccation-sensitive of lichens. Samples of P. dissimilis from a humid rain-forest site in New Zealand were transported in a moist state to Germany. Photosynthesis response curves were generated. The effect of desiccation was measured by comparing CO₂ exchange before and after a standard 20-h drying routine. Lichen thalli could be equilibrated at 15° C to relative humidities (RH) from 5% to almost 100%. Photosynthesis was saturated at a photosynthetically active radiation (PAR) level of 20 mol m^-2 s^-1 (350 bar CO₂) and PAR compensation was a very low 1 mol m^-2 s^-1. Photosynthesis did not saturate until 1500 bar CO₂. Net photosynthesis was relatively unaffected by temperature between 10° C and 30° C with upper compensation at over 40° C. Temporary depression of photosynthesis occurred after a drying period of 20 h with equilibration at 45–65% relative humidity (RH). Sustained damage occurred at 15–25% RH and many samples died after equilibration at 5–16% RH. Microclimate studies of the lichen habitat below the evergreen, broadleaf forest canopy revealed consistently low PAR (normally below 10–20 mol m^-2 s^-1) and high humidities (over 80% RH even during the day time). The species shows many features of an extremely deep shade-adapted plant including low PAR saturation and compensation, low photosynthetic and respiratory rates and low dry weight per unit area.     

Green light drives photosynthesis in mosses

Temperate forests are characterised by variable light quality (i.e. spectral composition of light) at or near the forest floor. These understory environments have a high concentration of green light, as red and blue light are preferentially absorbed by upper canopy leaves. Understory species may be well-adapted for using green light to drive photosynthesis. Angiosperms have been shown to use green light for photosynthesis, but this ability has not been demonstrated in shade-dwelling bryophytes. In this study, net photosynthetic rate (P_N) of three temperate understory species of moss (Dichodontium pellucidum (Hedw.) Schimp., Leucobryum albidum (Brid. ex P.Beauv) Lindb. and Amblystegium serpens (Hedw.) Schimp.) was measured under green, red?+?blue, and red?+?blue?+?green light to assess green light use efficiency. All three species were capable of photosynthesising beyond their respiratory demands using solely green light, with higher green light use efficiency measured in plants collected from areas with greater canopy cover, suggesting growth in a green light concentrated environment increases green light use efficiency. Each species was also collected from sites differing in their degree of canopy cover and grown under three light treatments (high light, low light, and green light). Photosynthetic efficiency (chlorophyll fluorescence), tissue nitrogen and carbon isotope concentrations were assessed after a short growth period. Growth conditions had little effect on leaf chemistry and monochromatic green light did not significantly degrade photosynthetic efficiency. This study provides the first evidence to date of positive net ‘green light photosynthesis’ in mosses.     

An improved method,using saturating light pulses,for the determination of photosystem I quantum yield via P700+-absorbance changes at 830 nm

An improved method is introduced for the determination of the quantum yield of photosystem I. The new method employs saturating light pulses with steep rise characteristics to distinguish, in a given physiological state, centers with an open acceptor side from centers with a reduced acceptor side. The latter do not contribute to PSI quantum yield (_I). Oxidation of P700 is measured by a rapid modulation technique using the absorbance change around 830 nm. The quantum yield _I is calculated from the amplitude of the rapid phase of absorbance change (A; 830 nm) upon application of a saturation pulse in a given state, divided by the maximal A (830 nm) which is induced by a saturation pulse with far-red background illumination. Using this technique, _I can be determined even under conditions of acceptor-side limitation, as for example in the course of a dark-light induction period or after elimination of CO₂ from the gas stream. Thus determined _I values display a close-to-linear relationship with those for the quantum yield of PSII (_II) calculated from chlorophyll fluorescence parameters. It is concluded that the proposed method may provide new information on the activity of the PSI acceptor side and thus help to separate the effects of acceptorside limitation from those of cyclic PSI, whenever a non-linear relationship between _II and the P700-reduction level is observed.Abbreviations and Symbols A absorbance change - _I quantum yield of photosystem I - _II quantum yield of photosystem II - PAR photosynthetically active radiation This work was supported by the Deutsche Forschungsgemeinschaft (SFB 176 Molekulare Grundlagen der Signalübertragung und des Stofftransportes in Membranen and SFB 251 Ökologie, Physiologie und Biochemie pflanzlicher Leistung unter Streß).     

天山北麓拟南芥表型特征的自然选择

本文研究了天山北麓干旱、湿润2种不同水分条件下拟南芥(Arabidopsis thaliana)光合生理、形态及发育特征的自然适应特点。结果表明：分枝数、株高、相对适合度（角果质量）、莲座叶面积等特征在不同水分下差异极显著;根质量与相对适合度、莲座叶面积、水分利用效率和光合速率之间表现了正相关,胞间二氧化碳浓度与光合速率、水分利用效率之间表现了负相关;选择梯度分析得出,干湿水分条件下分枝数、株高、莲座叶面积均表现定向选择;主成分分析发现,光合生理特征整合在处理间有一定的规律性,表现出拟南芥物种特征整体协调一致性较强,对环境响应具有较大的趋同性;通径分析发现,水分利用效率、枝质量、株高、角果数对相对适合度直接产生较大影响,因而在通径模型中设置为直接路径特征,将莲座叶面积通过分枝数、花期、角果数,花期通过角果数到达相对适合度的过程设置为间接路径,这些特征通过间接路径对相对适合度间接产生较大影响。     

不同光强下高锰对黄瓜光合作用特性的影响

采用营养液培养的方法,研究了不同光强下高锰对黄瓜植株生长、叶绿素含量、叶绿素荧光参数和光合作用的影响.结果表明,高锰处理抑制了黄瓜植株的生长,与弱光处理相比强光下抑制幅度更加显著.强光下,高锰处理显著降低叶绿素含量,但降低光强却增加其含量.强光下,高锰处理显著降低原初光能转换效率(F_v/F_m)、光合电子传递量子效率(ΦPSII)和光化学猝灭系数(qP);弱光下,高锰处理对F_v/F_m和qP无显著影响.高锰处理使净光合速率(P_n)和气孔导度(G_s)下降.尤其是在强光下下降幅度更大.高锰处理使细胞间CO₂浓度(C_i)在强光下升高,而在弱光下则下降.与C_i相反,高锰处理使气孔限制值(L_s)在强光下下降,而在弱光下上升.因此,强光下高锰胁迫使净光合速率下降可能是由非气孔限制引起的,而弱光下高锰处理使净光合速率下降可能是由气孔因子限制引起的.     

An assessment of the relationship between chlorophyll a fluorescence and CO2 gas exchange from field measurements on a moss and lichen

The relationship between CO₂ exchange and relative electron-transport rate through photosystem II (ETR, measured using chlorophyll a fluorescence) was determined for a moss and a green algal lichen, photobiont probably Trebouxia sp., in the field in Antarctica. Net photosynthesis (NP) and dark respiration (DR) were measured over temperatures from zero to 25 °C and gross photosynthesis (GP) calculated (GP = NP + DR). The strong response of DR to temperature in these organisms resulted in substantial changes in CO₂ exchange rates. The moss Bryum argenteum Hedw. showed a strong, linear relationship between GP and ETR. This was an unexpected result since mosses are C₃ plants and, in higher plants, this group normally has a curvilinear GP versus ETR relationship. It is suggested that suppression of DR in the light might be involved. The lichen, Umbilicaria aprina Nyl., had nonlinear relationships between ETR and GP that were different at each measurement temperature. In some cases the lowest ETR was at the higher CO₂ exchange rates. It is suggested that these relationships are the result of strong quenching mechanisms that are inversely proportional to GP. The results support a growing impression that the relationships between ETR and CO₂ exchange are complex in these organisms and different from those found for higher plants. Received: 24 November 1997 / Accepted 2 May 1998     

Field assessment of sub-epidermal leaf anthocyanin, PSII photochemistry,and the xanthophyll-cycle as photoprotective mechanisms in two morphs of Agave striata

We tested the hypothesis that leaf epidermal pigments screen light of particular wavelengths from reaching the photosynthetic machinery, reducing dependence on the xanthophyll-cycle as an energy dissipation process. Under field conditions, photosynthesis and water relations were studied in two morphs of Agave striata that differ in leaf coloration (green vs. reddish-purple). Titratable acidity, chlorophyll fluorescence, and internal and surface leaf temperatures were measured under low irradiance by shading (30%) and full sunlight (100%) for six days. We also measured the reflectance ratio (R_RED: R_GREEN), an index of anthocyanin content and the change in photochemical reflectance index (ΔPRI), an index of xanthophyll-cycle de-epoxidation state (xanthophyll conversion). Our results showed that both morphs expressed typical CAM-activity with no significant differences under sun vs. shade. However, shading did reduce titratable acids in both morphs. Both morphs were well hydrated, with the relative water content (RWC) being greater than 93%. Leaf surface temperature was found to be significantly higher during the day in the green morph compared to the red morph under sun and shade. Dark level fluorescence (Fo), photochemical efficiency of PSII (Fv/Fm), and the quantum yield of PSII electron transport (Φ_PSII) were higher in the red morph under sun compared to the green morph. The value of qN (non-photochemical quenching) was significantly higher during the day for the green morph compared to the red morph and this higher qN value was associated with a greater xanthophyll conversion and surface leaf temperature. However, sunlight did not predispose either of the morphs to photoinhibition. It is clear that the sub-epidermal anthocyanins serve as a photoprotective mechanism in the red morph, screening light energy from reaching the photosynthetic machinery and reducing dependence on the xanthophyll-cycle. We concluded that under natural light conditions the leaves of two morphs tested utilized differential photoprotective mechanisms.     

Photosynthetic performance in Sphagnum transplanted along a latitudinal nitrogen deposition gradient

Increased N deposition in Europe has affected mire ecosystems. However, knowledge on the physiological responses is poor. We measured photosynthetic responses to increasing N deposition in two peatmoss species (Sphagnum balticum and Sphagnum fuscum) from a 3-year, north–south transplant experiment in northern Europe, covering a latitudinal N deposition gradient ranging from 0.28 g N m⁻² year⁻¹ in the north, to 1.49 g N m⁻² year⁻¹ in the south. The maximum photosynthetic rate (NP_max) increased southwards, and was mainly explained by tissue N concentration, secondly by allocation of N to the photosynthesis, and to a lesser degree by modified photosystem II activity (variable fluorescence/maximum fluorescence yield). Although climatic factors may have contributed, these results were most likely attributable to an increase in N deposition southwards. For S. fuscum, photosynthetic rate continued to increase up to a deposition level of 1.49 g N m⁻² year⁻¹, but for S. balticum it seemed to level out at 1.14 g N m⁻² year⁻¹. The results for S. balticum suggested that transplants from different origin (with low or intermediate N deposition) respond differently to high N deposition. This indicates that Sphagnum species may be able to adapt or physiologically adjust to high N deposition. Our results also suggest that S. balticum might be more sensitive to N deposition than S. fuscum. Surprisingly, NP_max was not (S. balticum), or only weakly (S. fuscum) correlated with biomass production, indicating that production is to a great extent is governed by factors other than the photosynthetic capacity.