Effects of Thallus Temperature and Hydration on Photosynthetic Parameters of Cetraria Islandica from Contrasting Habitats

Two methods of induced in vivo chlorophyll (Chl) fluorescence were used to investigate the effects of varying thallus temperature and hydration on the performance of photosynthetic apparatus of a foliar lichen Cetraria islandica: slow Chl fluorescence induction kinetic with the analysis of quenching mechanisms, and rapid irradiance response curves of photosynthesis derived from quantum yield of photochemical reactions of photosystem 2 (₂) recorded at increasing irradiances. We compared responses of photosynthetic apparatus in populations of C. islandica growing in lower altitude (LAP: 1 350 m a.s.l.) and in higher altitude (HAP: 2 000 m a.s.l.). At each altitude, the samples were collected both in fully irradiated sites (HI) and in shade (LI). Temperature optimum of photosynthetic processes was the same for LAP and HAP thalli of LI populations (18 °C), while it was significantly lower for HI HAP (14 °C). Gradual dehydration of fully hydrated thalli led to initial increase (up to 20 % of water saturation deficit, WSD) in F_V/F_M and ₂, no change at 20–50 % WSD, and a dramatic decrease of the parameters within 50–80 % of WSD. LI HAP of C. islandica was the best adapted population to low temperature having higher rates of photochemical processes of photosynthesis than HI HAP within temperature range of –5 to +5 °C. The differences between populations were apparent also in Chl content and thallus morphology.     

Analysis of Qualitative Contribution of Assimilatory and Non-Assimilatory De-Excitation Processes to Adaptation of Photosynthetic Apparatus of Barley Plants to High Irradiance

The adaptation of barley (Hordeum vulgare L. cv. Akcent) plants to low (LI, 50 µmol m^–2 s^–1) and high (HI, 1000 µmol m^–2 s^–1) growth irradiances was studied using the simultaneous measurements of the photosynthetic oxygen evolution and chlorophyll a (Chl a) fluorescence at room temperature. If measured under ambient CO₂ concentration, neither increase of the oxygen evolution rate (P) nor enhancement of non-radiative dissipation of the absorbed excitation energy within photosystem 2 (PS2) (determined as non-photochemical quenching of Chl a fluorescence, NPQ) were observed for HI plants compared with LI plants. Nevertheless, the HI plants exhibited a significantly higher proportion of Q_A in oxidised state (estimated from photochemical quenching of Chl a fluorescence, q_P), by 49–102 % at irradiances above 200 µmol m^–2 s^–1 and an about 1.5 fold increase of irradiance-saturated PS2 electron transport rate (ETR) as compared to LI plants. At high CO₂ concentration the degree of P stimulation was approximately three times higher for HI than for LI plants, and the irradiance-saturated P values at irradiances of 2 440 and 2 900 µmol m^–2 s^–1 were by 130 and 150 % higher for HI plants than for LI plants. We suggest that non-assimilatory electron transport dominates in the adaptation of the photosynthetic apparatus of barley grown at high irradiances under ambient CO₂ rather than an increased NPQ or an enhancement of irradiance-saturated photosynthesis.     

Effects of different irradiances on the photosynthetic process during ex-vitro acclimation of Anoectochilus plantlets

Six months old in vitro-grown Anoectochilus formosanus plantlets were transferred to ex-vitro acclimation under low irradiance, LI [60 μmol(photon) m⁻² s⁻¹], intermediate irradiance, II [180 μmol(photon) m⁻² s⁻¹], and high irradiance, HI [300 μmol(photon) m⁻² s⁻¹] for 30 d. Imposition of II led to a significant increase of chlorophyll (Chl) b content, rates of net photosynthesis (P _N) and transpiration (E), stomatal conductance (g _s), electron transfer rate (ETR), quantum yield of electron transport from water through photosystem 2 (Φ_PS2), and activity of ribulose-1,5-bisphosphate carboxylase/ oxygenase (RuBPCO, EC 4.1.1.39). This indicates that Anoectochilus was better acclimated at II compared to LI treatment. On the other hand, HI acclimation led to a significant reduction of Chl a and b, P _N, E, g _s, photochemical quenching, dark-adapted quantum efficiency of open PS2 centres (F_v/F_m), probability of an absorbed photon reaching an open PS2 reaction centre (F_v′/F_m′), ETR, Φ_PS2, and energy efficiency of CO₂ fixation (Φ_CO2/Φ_PS2). This indicates that HI treatment considerably exceeded the photo-protective capacity and Anoectochilus suffered HI induced damage to the photosynthetic apparatus. Imposition of HI significantly increased the contents of antheraxanthin and zeaxanthin (ZEA), non-photochemical quenching, and conversion of violaxanthin to ZEA. Thus Anoectochilus modifies its system to dissipate excess excitation energy and to protect the photosynthetic machinery.     

Enhancement of susceptivity to photoinhibition and photooxidation in rice chlorophyll <Emphasis Type="Italic">b</Emphasis>-less mutants

Two rice chlorophyll (Chl) b-less mutants (VG28-1, VG30-5) and the respective wild type (WT) plant (cv. Zhonghua No. 11) were analyzed for the changes in Chl fluorescence parameters, xanthophyll cycle pool, and its de-epoxidation state under exposure to strong irradiance, SI (1 700 μmol m⁻² s⁻¹). We also examined alterations in the chloroplast ultrastructure of the mutants induced by methyl viologen (MV) photooxidation. During HI (0–3.5 h), the photoinactivation of photosystem 2 (PS2) appeared earlier and more severely in Chl b-less mutants than in the WT. The decreases in maximal photochemical efficiency of PS2 in the dark (F_v/F_m), quantum efficiency of PS2 electron transport (Φ_PS2), photochemical quenching (q_P), as well as rate of photochemistry (P_rate), and the increases in de-epoxidation state (DES) and rate of thermal dissipation of excitation energy (D_rate) were significantly greater in Chl b-mutants compared with the WT plant. A relatively larger xanthophyll pool and 78–83 % conversion of violaxanthin into antheraxanthin and zeaxanthin in the mutants after 3.5 h of HI was accompanied with a high ratio of inactive/total PS2 (0.55–0.73) and high 1–q_P (0.57–0.68) which showed that the activities of the xanthophyll cycle were probably insufficient to protect the photosynthetic apparatus against photoinhibition. No apparent difference of chloroplast ultrastructure was found between Chl b-less mutants and WT plants grown under low, LI (180 μmol m⁻² s⁻¹) and high, HI (700 μmol m⁻² s⁻¹) irradiance. However, swollen chloroplasts and slight dilation of thylakoids occurred in both mutants and the WT grown under LI followed by MV treatment. These typical symptoms of photooxidative damage were aggravated as plants were exposed to HI. Distorted and loose scattered thylakoids were observed in particular in the Chl b-less mutants. A greater extent of photoinhibition and photooxidation in these mutants indicated that the susceptibility to HI and oxidative stresses was enhanced in the photosynthetic apparatus without Chl b most likely as a consequence of a smaller antenna size.     

Role of calcium ion in protection against heat and high irradiance stress-induced oxidative damage to photosynthesis of wheat leaves

Cross stress of heat and high irradiance (HI) resulted in the accumulation of active oxygen species and photo-oxidative damage to photosynthetic apparatus of wheat leaves during grain development. Pre-treatment with calcium ion protected the photosynthetic system from oxidative damage by reducing O^-. ₂ production, inhibiting lipid peroxidation, and retarding electrolyte leakage from cell. Therefore, high F_v/F_m [maximal photochemical efficiency of photosystem 2 (PS2) while all PS2 reaction centres are open], F_m/F₀ (another expression for the maximal photochemical efficiency of PS2), Φ_PS2 (actual quantum yield of PS2 under actinic irradiation), q_P (photochemical quenching coefficient), and P _N (net photosynthetic rate) were maintained, and lower q_NP (non-photochemical quenching coefficient) of the leaves was kept under heat and HI stress. EGTA (a chelant of calcium ion) and LaCl₃ (a blocker of Ca²⁺ channel in cytoplasmic membrane) had the opposite effect. Thus Ca ion may help protect the photosynthetic system of wheat leaves from oxidative damage induced by the cross stress of heat and HI.     

Short-term responses of photosynthetic membrane lipids and photochemical efficiency in plants of <Emphasis Type="Italic">Phaseolus vulgaris</Emphasis> and <Emphasis Type="Italic">Vigna unguiculata</Emphasis> submitted to high irradiance

Primary leaves of young plants of common bean (Phaseolus vulgaris cv. Carioca and Negro Huasteco) and cowpea (Vigna unguiculata Walp cv. Epace 10) were exposed to high irradiance (HI) of 2 000 μmol m⁻² s⁻¹ for 10, 20, and 30 min. The initial fluorescence (F₀) was nearly constant in response to HI in each genotype except for Carioca. A distinct reduction of maximum fluorescence (F_m) was clearly observed in stressed genotypes of beans after 20 min followed by a slight recovery for the longer stress times. In common bean, the maximum quantum yield (F_v/F_m) was reduced slowly from 10 to 30 min of HI. In cowpea, only a slight reduction of F_v/F_m was observed at 20 min followed by recovery to normal values at 30 min. HI resulted in changes in the photochemical (q_P) and non-photochemical (q_N) quenching in both species, but to a different extent. In cowpea plants, more efficiency in the use of the absorbed energy under photoinhibitory conditions was related to increase in q_P and decrease in q_N. In addition, lipid peroxidation changed significantly in common bean genotypes with an evident increase after 20 min of HI. Hence the photosynthetic apparatus of cowpea was more tolerant to HI than that of common bean and the integrity of cowpea cell membranes was apparently maintained under HI.     

Chlorophyll thermofluorescence and thermoluminescence as complementary tools for the study of temperature stress in plants

The photosynthetic apparatus, especially the electron transport chain imbedded in the thylakoid membrane, is one of the main targets of cold and heat stress in plants. Prompt and delayed fluorescence emission originating from photosystem II have been used, most often separately, to monitor the changes induced in the photosynthetic membranes during progressive warming or cooling of a leaf sample. Thermofluorescence of F ₀ and F _M informs on the effects of heat on the chlorophyll antennae and the photochemical centers, thermoluminescence on the stabilization and movements of charges and Delayed Light Emission on the permeability of the thylakoid membranes to protons and ions. Considered together and operated simultaneously, these techniques constitute a powerful tool to characterize the effect of thermal stress on intact photosynthetic systems and to understand the mechanisms of constitutive or induced tolerance to temperature stresses.     

Large reallocations of carbon,nitrogen, and photosynthetic reductant among phycobilisomes,photosystems, and Rubisco during light acclimation in <Emphasis Type="Italic">Synechococcus elongatus</Emphasis> strain PCC7942 are constrained in cells under low environmental inorganic carbon

Synechococcus elongatus strain PCC7942 cells were grown in high or low environmental concentrations of inorganic C (high-C_i, low-C_i) and subjected to a light shift from 50 µmol m^–2 s^–1 to 500 µmol m^–2 s^–1. We quantified photosynthetic reductant (O₂ evolution) and molar cellular contents of phycobilisomes, PSII, PSI, and ribulose-1,5-bisphosphate carboxylase oxygenase (Rubisco) through the light shift. Upon the increase in light, small initial relative decreases in phycobilisomes per cell resulted from near cessation of phycobilisome synthesis and their dilution into daughter cells. Thus, allocation of reductant to phycobilisome synthesis dropped fivefold from pre- to post-light shift. The decrease in phycobilisome synthesis liberated enough material and reductant to allow a doubling of Rubisco and up to a sixfold increase in PSII complexes per cell. Low-C_i cells had smaller initial phycobilisome pools and upon increased light; their reallocation of reductant from phycobilisome synthesis may have limited the rate and extent of light acclimation, compared to high-C_i cells. Acclimation to increased light involved large reallocations of C, N, and reductant among different components of the photosynthetic apparatus, but total allocation to the apparatus was fairly stable at ca. 50% of cellular N, and drew 25–50% of reductant from photosynthesis.     

Excess irradiance causes early symptoms of senescence during leaf expansion in photoautotrophically <Emphasis Type="Italic">in vitro</Emphasis> grown tobacco plants

Photosynthetic parameters, growth, and pigment contents were determined during expansion of the fourth leaf of in vitro photoautotrophically cultured Nicotiana tabacum L. plants at three irradiances [photosynthetically active radiation (400–700 nm): low, LI 60 μmol m⁻² s⁻¹; middle, MI 180 μmol m⁻² s⁻¹; and high, HI 270 μmol m⁻² s⁻¹]. During leaf expansion, several symptoms usually accompanying leaf senescence appeared very early in HI and then in MI plants. Symptoms of senescence in developing leaves were: decreasing chlorophyll (Chl) a+b content and Chl a/b ratio, decreasing both maximum (F_V/F_M) and actual (Φ_PS2) photochemical efficiency of photosystem 2, and increasing non-photochemical quenching. Nevertheless, net photosynthetic oxygen evolution rate (P _N) did not decrease consistently with decrease in Chl content, but exhibited a typical ontogenetic course with gradual increase. P _N reached its maximum before full leaf expansion and then tended to decline. Thus excess irradiance during in vitro cultivation did not cause early start of leaf senescence, but impaired photosynthetic performance and Chl content in leaves and changed their typical ontogenetic course.     

Structural and functional integrity of Sulla carnosa photosynthetic apparatus under iron deficiency conditions

• The abundance of calcareous soils makes bicarbonate‐induced iron (Fe) deficiency a major problem for plant growth and crop yield. Therefore, Fe‐efficient plants may constitute a solution for use on calcareous soils.
• We investigated the ability of the forage legume Sulla carnosa (Desf.) to maintain integrity of its photosynthetic apparatus under Fe deficiency conditions. Three treatments were applied: control, direct Fe deficiency and bicarbonate‐induced Fe deficiency.
• At harvest, all organs of deficient plants showed severe growth inhibition, the effect being less pronounced under indirect Fe deficiency. Pigment analysis of fully expanded leaves revealed a reduction in concentrations of chlorophyll a, chlorophyll b and carotenoids under Fe deficiency. Electron transport rate, maximum and effective quantum yield of photosystem II (PSII), photochemical quenching (qP), non‐photochemical quenching (qN) as well as P700 activity also decreased significantly in plants exposed to direct Fe deficiency, while qN was not affected. The effects of indirect Fe deficiency on the same parameters were less pronounced in bicarbonate‐treated plants. The relative abundances of thylakoid proteins related to PSI (PsaA, Lhca1, Lhca2) and PSII (PsbA, Lhcb1) were also more affected under direct than indirect Fe deficiency.
• We conclude that S. carnosa can maintain the integrity of its photosynthetic apparatus under bicarbonate‐induced Fe deficiency, preventing harmful effects to both photosystems under direct Fe deficiency. This suggests a high capacity of this species not only to take up Fe in the presence of bicarbonate (HCO₃^?) but also to preferentially translocate absorbed Fe towards leaves and prevent its inactivation.

     

Effect of Iron Deficiency Induced Changes on Photosynthetic Pigments,Ribulose-1,5-Bisphosphate Carboxylase,and Photosystem Activities in Field Grown Grapevine (Vitis Vinifera L. cv. Pinot Noir) Leaves

The effect of iron deficiency on photosynthetic pigments, ribulose-1,5-bisphosphate carboxylase (RuBPC), and photosystem activities were investigated in field grown grapevine (Vitis vinifera L. cv. Pinot noir) leaves. The contents of chlorophyll (Chl) (a+b) and carotenoids per unit fresh mass showed a progressive decrease upon increase in iron deficiency. Similar results were also observed in content of total soluble proteins and RuBPC activity. The marked loss of large (55 kDa) and small (15 kDa) subunits of RuBPC was also observed in severely chlorotic leaves. However, when various photosynthetic electron transport activities were analysed in isolated thylakoids, a major decrease in the rate of whole chain (H₂O methyl viologen) electron transport was observed in iron deficient leaves. Such reduction was mainly due to the loss of photosystem 2 (PS2) activity. The same results were obtained when F_v/F_m was evaluated by Chl fluorescence measurements in leaves. Smaller inhibition of photosystem 1 (PS1) activity was also observed in both mild and severely chlorotic leaves. The artificial electron donors, diphenyl carbazide and NH₂OH, markedly restored the loss of PS2 activity in severely chlorotic leaves. The marked loss of PS2 activity was evidently due to the loss of 33, 23, 28-25, and 17 kDa polypeptides in iron deficient leaves.     

Photosynthetic electron transport is differentially affected during early stages of cultivar/race-specific interactions between potato andPhytophthora infestans

The influence of the early stages of fungal infection on chloroplast metabolism was studied in cultivar/race-specific interactions between potato (Solanum tuberosum L. cv. Datura) and the late-blight fungusPhytophthora infestans. The accumulation of several mRNAs encoding components of the photosynthetic apparatus was not affected, either in compatible or in incompatible interactions. However, within 3 h after inoculation of potato leaves with fungal spores, a change in the photochemistry of photosystem II was detectable by measuring chlorophylla fluorescence. Characteristic fluorescence parameters, such as maximum fluorescence yield (F_m), variable fluorescence yield (F_v) and photochemical efficiency (F_v/F_m), were specifically reduced in the compatible host/pathogen interaction. Analyses of photochemical and nonphotochemical fluorescence quenching showed an increase in the photochemical fraction. The amounts of two selected thylakoid membrane proteins and of total chlorophyll remained unchanged during this process, suggesting that the functional modification of the electron-transport system was not correlated with a change in the composition of the photosynthetic apparatus. The alterations of photosynthetic electron transport represent a rapidly detectable and sensitive physiological marker for compatible interactions in the potato/Phytophthora infestans pathosystem.     

Acclimation to irradiance in seedlings of three tropical rain forest Garcinia species after simulated gap formation

We investigated the acclimation of seedlings of three tropical rain forest sub-canopy Garcinia species (G. xanthochymus, G. cowa, and G. bracteata) after transfer from 4.5 (LI) to 40 % (HI) sunlight and 12.5 (MI) sunlight to HI (LH₁ and LH₂ denoting transfer from LI to HI and MI to HI transfer, respectively). The changes of chlorophyll (Chl) fluorescence, net photosynthetic rate (P _N), dark respiration rate (R _D), Chl content per unit area (Chl_area), leaf mass per unit area (LMA), and seedling mortality were monitored over two months after transfer. These parameters together with leaf anatomy of transferred and control seedlings (kept in LI, MI, and HI) were also examined after two months. No seedlings died during the two months. F_v/F_m, P _N, and Chl_area of the transferred seedlings decreased in the first 3 to 12 d. LH₁ leaves showed larger reduction in F_v/F_m (>23 % vs. <16 %) and slower recovery of F_v/F_m than LH₂ leaves. P _N started to recover after about one week of I transfer and approached higher values in all G. cowa seedlings and G. xanthochymus LH₁ seedlings than those before the transfer. However, P _N of G. bracteata seedlings approached the values before transfer. The final P _N values in leaves of transferred G. xanthochymus and G. cowa seedlings approached that of leaves kept in HI, while the final P _N values of transferred leaves of G. bracteata were significantly lower than that of leaves grown under HI (p<0.05). R _D of G. xanthochymus LH₁ seedlings and all G. cowa seedlings increased and approached the value of the seedlings in HI. The final Chl_area of both G. xanthochymus and G. cowa approached the values before transfer, but that of G. bracteata did not recover to the level before transfer. The final Chl_area of all transferred seedlings was not significantly different from that of seedlings in HI except that G. cowa LH₁ seedlings had higher Chl_area than that in HI. LMA decreased within 2 d and then increased continuously until about 30 d and approached the value under HI. Spongy/palisade mesophyll ratio decreased after transfer because of the increase in palisade thickness. Leaf thickness did not change, so LMA increase of transferred seedlings was mainly due to the increase of leaf density. Thus the mature leaves under LI and MI of G. xanthochymus and G. cowa are able to acclimate to HI by leaf physiological and anatomical adjustment, while G. bracteata had limited ability to acclimate to HI.     

Thermostability of photosynthesis in two new chlorophyllb-less rice mutants

Leaves of the two new chlorophyll b-less rice mutants VG28-1, VG30-5 and the wild type rice cv. Zhonghua 11 were subjected to temperatures 28, 36, 40, 44 and 48℃ in the dark for 30 min or gradually elevated temperature from 30℃ to 80℃ at 0.5℃/min. The thermostability of photosynthetic apparatus was estimated by the changes in chlorophyll fluorescence parameters, photosynthetic rate and pigment content, chloroplast ultrastructure and tissue location of H2O2 accumulation. There were different patterns of Fo-temperature curves between the Chl b-less mutants and the wild type plant, and the temperature of F_o rising threshold was shifted 3℃ lower in the Chl b-less mutants (48℃) than in the wild type (51℃). At temperature up to about 45℃, chloroplasts were swollen and thylakoid grana became misty accompanied with the complete loss of photosynthetic oxygen evolution in the two Chl b-less mutants, but chloroplast ultrastruc-ture in the wild type showed no obvious alteration. After 55℃ exposure, the disordered thylakoid and significant H₂O₂ accumulation in leaves were found in the two Chl _b-less mutants, whereas in the wild type plant, less H₂O₂ was accumulated and the swollen thylakoid still maintained a cer-tain extent of stacking. A large extent of the changes in qP, NPQ and F_v/F_m was consistent with the Pn decreasing rate in the Chl b-less mutants during high temperature treatment as compared with the wild type. The results indicated that the Chl b-less mutants showed a tendency for higher thermosensitivity, and loss of Chl b in LHC II could lead to less thermostability of PSII structure and function. Heat damage to photosynthetic apparatus might be partially attributed to the in-ternal oxidative stress produced at severely high temperature.     

Comparative Study of Thermal Degradation of Iron–Sulfur Proteins in Spinach Chloroplasts and Membranes of Thermophilic Cyanobacteria: Mössbauer Spectroscopy

Mössbauer spectra of chloroplasts isolated from spinach plants grown in a mineral medium enriched with ⁵⁷Fe and Mössbauer spectra of native membranes of the thermophilic cyanobacterium Synechococcus elongatus contain a broad asymmetric doublet typical of the iron–sulfur proteins of Photosystem (PS) I. Exposure of chloroplasts to temperatures of 20-70°C significantly modifies the central part of the spectra. This spectral change is evidence of decreased magnitude of the quadrupole splitting. However, the thermally induced doublet (Q = 3.10 mm/sec and = 1.28 mm/sec) typical of hydrated forms of reduced (divalent) inorganic iron is not observed in spinach chloroplasts. This doublet is usually associated with degradation of active centers of ferredoxin, a surface-exposed protein of PS I. The Mössbauer spectra of photosynthetic membranes of spinach chloroplasts and cyanobacteria were compared using the probability distribution function of quadrupole shift (1/2 quadrupole splitting Q) of trivalent iron. The results of calculation of these functions for the two preparations showed that upon increasing the heating temperature there was a decrease in the probability of the presence of native iron–sulfur centers F_X, F_A, and F_B (quadrupole shift range, 0.43-0.67 mm/sec) in heated preparations. This process was also accompanied by an increase in the probability of appearance of clusters of trivalent iron. This increase was found to be either gradual and continuous or abrupt and discrete in photosynthetic membranes of cyanobacteria or spinach chloroplasts, respectively. The probability of the presence of the iron–sulfur centers F_X, F_A, and F_B in chloroplasts abruptly decreases to virtually to zero within the temperature range critical for inhibition of electron transport through PS I to oxygen. In cyanobacteria, both thermal destruction of iron–sulfur centers of PS I and functional degradation of PS I are shifted toward a higher temperature. The results of this study suggest that the same mechanism of thermal destruction of the PS I core occurs in both thermophilic and mesophilic organisms: destruction of iron–sulfur centers F_X, F_A, and F_B, release of oxidized (trivalent) iron, and its accumulation in membrane-bound iron-oxo clusters.     

Thermostability of photosynthesis in two new chlorophyllb-less rice mutants

Leaves of the two new chlorophyllb-less rice mutants VG28-1, VG30-5 and the wild type rice cv. Zhonghua 11 were subjected to temperatures 28, 36, 40, 44 and 48°C in the dark for 30 min or gradually elevated temperature from 30°C to 80°C at 0.5°C/min. The thermostability of photosynthetic apparatus was estimated by the changes in chlorophyll fluorescence parameters, photosynthetic rate and pigment content, chloroplast ultrastructure and tissue location of H₂O₂ accumulation. There were different patterns of F_o-temperature curves between the Chlb-less mutants and the wild type plant, and the temperature of F_o rising threshold was shifted 3°C lower in the Chlb-less mutants (48°C) than in the wild type (51°C). At temperature up to about 45°C, chloroplasts were swollen and thylakoid grana became misty accompanied with the complete loss of photosynthetic oxygen evolution in the two Chlb-less mutants, but chloroplast ultrastructure in the wild type showed no obvious alteration. After 55°C exposure, the disordered thylakoid and significant H₂O₂ accumulation in leaves were found in the two Chlb-less mutants, whereas in the wild type plant, less H₂O₂ was accumulated and the swollen thylakoid still maintained a certain extent of stacking. A large extent of the changes in qP, NPQ and F_v/F_m was consistent with the Pn decreasing rate in the Chlb-less mutants during high temperature treatment as compared with the wild type. The results indicated that the Chlb-less mutants showed a tendency for higher thermosensitivity, and loss of Chlb in LHC II could lead to less thermostability of PSII structure and function. Heat damage to photosynthetic apparatus might be partially attributed to the internal oxidative stress produced at severely high temperature.     

Duration of irradiation rather than quantity and frequency of high irradiance inhibits photosynthetic processes in the lichen <Emphasis Type="Italic">Lasallia pustulata</Emphasis>

Lichen thalli were exposed to 4 regimes differing in irradiance and duration of irradiation. Photosynthetic efficiency of thalli was monitored by chlorophyll fluorescence parameters and xanthophyll cycle analysis. Maximal quantum yield of photosystem 2 (F_V/F_M) decreased gradually with time in long-term treatment. The effect of additional short-term high irradiance (HI) treatment applied each 24 h was not significant. Nevertheless, short-term HI applied repeatedly on thalli kept in the dark led to a significant decrease of F_V/F_M. Non-photochemical quenching recorded during the long-term treatment corresponded to the content of zeaxanthin (Z). In short-term treatment, however, proportion of Z (and antheraxanthin) to total amount of xanthophyll cycle pigments recovered to the initial values every 24 h after each repeated short-term HI event in thalli kept in dark. Thus duration of irradiation rather than irradiance and frequency of HI events is important for a decrease in primary photosynthetic processes in wet thalli of Lasallia pustulata. Rapidly responding photoprotective mechanisms, such as conversion of xanthophyll cycle pigments, are involved mainly in short-term irradiation events, even at HI.     

Photosynthetic performance of <Emphasis Type="Italic">Lycoris radiata</Emphasis> var. <Emphasis Type="Italic">radiata</Emphasis> to shade treatments

The effects of shade on the growth, leaf photosynthetic characteristics, and chlorophyll (Chl) fluorescence parameters of Lycoris radiata var. radiata were determined under differing irradiances (15, 65, and 100% of full irradiance) within pots. The HI plants exhibited a typical decline in net photosynthetic rate (P _N) during midday, which was not observed in MI- and LI plants. This indicated a possible photoinhibition in HI plants as the ratio of variable to maximum fluorescence (F_v/F_m) value was higher and the minimal fluorescence (F₀) was lower in the, and LI plants. Diurnal patterns of stomatal conductance (g _s) and transpiration rate (E) were remarkably similar to those of P _N at each shade treatments, and the intercellular CO₂ concentration (C _i) had the opposite change trend. Under both shading conditions, the light saturation point, light compensation point and photon-saturated photosynthetic rate (P _max) became lower than those under full sunlight, and it was the opposite for the apparent quantum yield (AQY). The higher the level of shade, the lower the integrated daytime carbon gain, stomatal and epidermis cell densities, specific leaf mass (SLM), bulb mass ratio (BMR), leaf thickness, and Chl a/b ratio. In contrast, contents of Chls per dry mass (DM), leaf area ratio (LAR), leaf mass ratio (LMR), leaf length, leaf area and total leaf area per plant increased under the same shade levels to promote photon absorption and to compensate for the lower radiant energy. Therefore, when the integrated daytime carbon gain, leaf area and total leaf area per plant, which are the main factors determining the productivity of L. radiata var. radiata plant, were taken into account together, this species may be cultivated at about 60∼70% of ambient irradiance to promote its growth.     

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 (F_m), photochemical efficiency of PS2 (F_v/F_m) and quantum yield of PS2 (Φ_PS2), and increased reduction state of Q_A (1-q_P) 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 (F_i-F₀)/(F_p-F₀) which is an expression of the proportion of the Q_B non-reducing PS2 centres, whereas a remarked decrease in the slope of F_i to F_p which represents the rate of Q_A 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 F_m, F_v/F_m, Φ_PS2, NPQ, slope of F_i to F_p, V+A+Z, and DES, and increased F₀, 1-q_P, ratio (F_i-F₀)/(F_p-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.     

Response of the Photosynthetic Apparatus of the Diatom Thallassiosira weisflogii to High Irradiance Light

The response of the photosynthetic apparatus to high irradiance illumination (440–2200 W/m²) was studied in the diatom Thallassiosira weisflogii by fluorescence methods. Changes in the photosynthetic apparatus were monitored by measuring characteristics of chlorophyll fluorescence F ₀, F _m, F _v/F _m, and qN for several hours after illumination of the alga with high-intensity light. Incubation of the alga with 2 mM DTT, an inhibitor of de-epoxidase of carotenoids in the diadinoxanthin cycle, led to a decrease in the nonphotochemical quenching of chlorophyll fluorescence and a drop in the F _v/F _m ratio, a characteristic that reflects the quantum efficiency of the functioning of the photosynthetic apparatus. Light-induced absorption changes associated with transformations of carotenoids of diadinoxanthin cycle were recorded in vivo in algal suspensions in the absence and in the presence of DTT. Using the microfluorometric method, we measured cell distribution over the efficiency of the primary processes of photosynthesis (F _v/F _m) after illumination. We found cells with a high tolerance of their photosynthetic apparatus to photooxidative damage. The relatively high tolerance of a portion of the cell population to high-light illumination can be related to light-induced transformation of carotenoids and to the functioning of other protective systems of the photosynthetic apparatus in diatoms.