PSⅡ Photochemistry and Xanthophyll Cycle in Two Superhigh yield Rice Hybrids,Liangyoupeijiu and Hua an 3 During Photoinhibition and Subsequent Restoration

PSⅡ photochemistry and xanthophyll cycle during photoinhibition (exposed to strong light of 2 000 μmol photons·m-2·s-1) and the subsequent restoration were compared between two superhigh yield rice hybrids (Liangyoupeijiu and Hua an 3, the newly developed rice hybrids from two parental lines) and the traditional rice hybrid Shanyou 63 developed from three parental lines. The results showed that the maximal efficiency of PSⅡ photochemistry (Fv/Fm), the efficiency of excitation energy capture by open PSⅡ centers (Fv′/Fm′), and the yield of PSⅡ electron transport (ΦPSⅡ) of the three rice hybrids decreased during photoinhibition. However, a greater decrease in Fv/Fm, Fv′/Fm′, and ΦPSⅡ was observed in Shanyou 63 than in Liangyoupeijiu and Hua an 3. At the same time, the components of xanthophyll cycle, antherxanthin (A) and zeathanxin (Z) increased rapidly while violaxanthin (V) decreased considerably. Both the rate of accumulation and the amount of A and Z in the two superhigh yield rice hybrids were higher than that in Shanyou 63. The de epoxidation state (DES) of xanthophyll cycle increased rapidly with the fast accumulation of A and Z, and reached the maximal level after first 30 min during photoinhibition. Of the three hybrids, the increasing rate of DES in Liangyoupeijiu and Hua an 3 was higher than that in Shanyou 63. After photoinhibition treatment, the plant materials were transferred to a dim light (70 μmol photons·m-2·s-1) for restoration. During restoration, both chlorophyll fluorescence parameters and xanthophyll cycle relaxed gradually, but the rate and level of restoration in the two superhigh yield rice hybrids were higher than those in Shanyou 63. Our results suggest that Liangyoupeijiu and Hua an 3 had higher resistance to photoinhibition and higher capacity of non radiative energy dissipation associated with xanthophyll cycle, as well as higher rate of restoration after photoinhibition, than Shanyou 63 when subjected to strong light.     

Photosynthetic light and CO2 utilization and C4 traits of two novel super-rice hybrids

Characteristics of photosynthetic light and CO2 use efficiency from seedling to heading stage, and C4 pathway enzyme activities in both flag leaves and lemma were compared between two newly developed super-rice hybrids (Oryza sativa L.), Liangyoupeijiu and Hua-an 3, and a traditional rice hybrid, Shanyou 63. At seedling and tillering stages, Liangyoupeijiu and Hua-an 3 had higher net photosynthetic rates (Pn) and light saturated assimilation rates (Asat) than did Shanyou 63, at both normal (360 micromol mol(-1)) and doubled (720 micromol mol(-1)) CO2 concentrations. At the heading stage, the flag leaves of all three rice hybrids had similar Pn and Asat. However, the two super-rice hybrids had higher apparent quantum yield (AQY) and carboxylation efficiency (CE) during all three typical developmental stages, and higher quantum yield of CO2 fixation (PhiCO2) at the tillering and heading stages. In addition, Liangyoupeijiu showed significantly higher activities of the C(4) pathway enzymes in both flag leaves and lemmas than did Shanyou 63. As a result, flag leaves of the two super-rice hybrids had higher Pn at morning, noontime and late afternoon during the daily cycle. Since most of the grain yield of rice comes from the photosynthesis of flag leaves, the similar Asat and much higher AQY, CE and PhiCO2 at heading stage of the two super-rice hybrids indicates that higher photosynthetic efficiency rather than higher photosynthetic capacity may be the primary factor contributing to their higher grain yields.     

Characterization of photosynthesis, photoinhibition and the activities of C4 pathway enzymes in a superhigh-yield rice, Liangyoupeijiu

Characteristics of photosynthetic gas exchange, photoinhibition and C₄ pathway enzyme activities in both flag leaves and lemma were compared between a superhigh-yield rice (Oryza sativa L.) hybrid, Liangyoupeijiu and a traditional rice hybrid, Shanyou63. Liangyoupeijiu had a similar light saturated assimilation rate (Asat) to Shanyou63, but a much higher apparent quantum yield (AQY), carboxylation efficiency (CE) and quantum yield of CO₂ fixation (ΦCO₂). Liangyoupeijiu also showed a higher resistance to photoinhibition and higher non-radiative energy dissipation associated with the xanthophyll cycle than Shanyou63 when subjected to strong light. In addition, Liangyoupeijiu had higher activities of the C₄ pathway enzymes in both flag leaves and lemmas than Shanyou63. These results indicate that higher light and CO₂ use efficiency, higher resistance to photoinhibition and C₄ pathway in both flag leaf and lemma may contribute to the higher yield of the superhigh-yield rice hybrid, Liangyoupeijiu.     

超高产杂交稻不同生育期的光合色素含量、净光合速率和水分利用效率

 较为系统地研究了两个超高产杂交稻‘两优培九’、‘华安3号’和多年来大面积推广的常规杂交稻‘汕优63’不同生育期的光合色素含量、净光合速率和水分利用效率。结果表明,在苗期,3个杂交稻的单位叶面积的叶绿素（Chl）含量差别不大,类胡萝卜素（Car）的含量以‘汕优63’为最高。然而,随着发育阶段的推进,到分蘖期,尤其是抽穗期剑叶的单位叶面积Chl和Car含量,两个超高产杂交稻高于‘汕优63’。从苗期到抽穗期,超高产杂交稻‘两优培九’和‘华安3号’的净光合速率（Pn）都比‘汕优63’高,而在苗期的午间强光条件下和分蘖期的早晨以及抽穗期的早晚相对弱光条件下其Pn的差别尤为显著。在苗期,‘汕优63’的水分利用效率（WUE）略高于‘两优培九’和‘华安3号’;到分蘖期和抽穗期,在上午10时之前和下午14时以后的时间段,则是两个超高产杂交稻明显高于‘汕优63’。这些结果说明,超高产杂交稻‘两优培九’和‘华安3号’不仅有较高的Pn和较强的抗光抑制能力,而且还能充分利用早晨和傍晚较弱的光强进行光合作用,这些是它们之所以高产的重要生理基础。此外,超高产杂交稻在生长发育的中后期（分蘖期和抽穗期）具有较高的WUE,有利于节约农业用水。     

Diurnal variation of gas exchange, chlorophyll fluorescence, and xanthophyll cycle components of maize hybrids released in different years

Diurnal variation of gas exchange, chlorophyll (Chl) fluorescence, and xanthophyll cycle components of three maize (Zea mays L.) hybrids released in different years, i.e. Baimaya (1950s), Zhongdan2 (1970s), and Nongda108 (1990s), were compared. On cloudless days, the newer hybrids always had higher net photosynthetic rate (P _N), especially at noon, than the older ones. At noon, all the hybrids decreased their maximal yield of photosystem 2 (PS2) photochemistry (F_v/F_m) and actual quantum yield of PS2 (Φ_PS2), the newer ones always showing higher values. Generally, the newer hybrids displayed higher photochemical quenching of Chl (q_P) and lower non-photochemical quenching (NPQ). The interhybrid differences in P _N may be owing to their differential photochemical efficiency. A midday depression in P _N occurred in all hybrids, which might be caused by serious photoinhibition or by decreased stomatal conductance. However, midday depression in P _N was more obvious in the older hybrids, especially when leaves were senescent. The higher de-epoxidation state of the xanthophylls was noted in older hybrids, which was confirmed by their larger NPQ. The newer maize hybrids did not need a strong de-epoxidation state since they had a better photosynthetic quantum conversion rate and a lower NPQ.     

Characteristics of Chlorophyll Fluorescence and Membrane-Lipid Peroxidation During Senescence of Flag Leaf in Different Cultivars of Rice

With japonica rice 98-08, indica hybrids Shanyou 63, Gangyou 881, and X07S/Zihui 100, and sub-species hybrid Peiai 64S/9311 as materials, chlorophyll (Chl) content, Chl a fluorescence parameters, and membrane lipid peroxidation in flag leaf were measured at late developmental stages under natural conditions. F_v/F_m, q_P, _PS2, and electron transport rate gradually decreased while q_N increased conversely. Excessive photon energy led to the accumulation of active oxygen (O₂ ^–), H₂O, malonyldialdehyde, and products of membrane lipid peroxidation, and resulted in reduced Chl content and early ageing subsequent to the photooxidation during flag leaf senescence. There was obvious diversification of these parameters among rice cultivars. In comparison with japonica cv. 98-08 (tolerant to photooxidation), F_v/F_m decreased in indica cv. Shanyou 63 (susceptible to photooxidation) with greater accumulation of active oxygen and a sharp drop in Chl content, which resulted in yellowish early ageing, and affected the filling and setting of rice grains. The mechanism for premature ageing in indica rice was related to irradiance and temperature at filling stages. On a sunny day at above 25 °C, the reaction centre of photosystem 2 (PS2) exhibited a dynamic change on reversible inactivation. Under the intense irradiance at noon, PS2 function in indica rice exhibited obvious down-regulation and photoinhibition. Under intense irradiance with lowered temperatures, PS2 resulted in photo-damage and early ageing, related to the degradation of PS2-D1 protein and the inhibition of endogenous protection systems such as the xanthophyll cycle and enzymes scavenging active oxygen. Hence for high-yield breeding, based on a good plant-type and utilising heterosis and tolerance of photooxidation, the selection of japonica rice or a sterile line with the japonica genotype as female is a strategy worthy of consideration.     

Characterization of photosynthesis, photoinhibition and the activities of C4 pathway enzymes in a superhigh-yield rice, Liangyoupeijiu

Characteristics of photosynthetic gas exchange, photoinhibition and C4 pathway enzyme activities in both flag leaves and lemma were compared between a superhigh-yield rice (Oryza sativa L.) hybrid, Liangyoupeijiu and a traditional rice hybrid, Shanyou63. Liangyoupeijiu had a similar light saturated assimilation rate (Asat) to Shanyou63, but a much higher apparent quantum yield (AQY), carboxylation efficiency (CE) and quantum yield of CO2 fixation (φCO2). Liangyoupeijiu also showed a higher resistance to photoinhibition and higher non-radiative energy dissipation associated with the xanthophyll cycle than Shanyou63 when subjected to strong light. In addition, Liangyoupeijiu had higher activities of the C4 pathway enzymes in both flag leaves and lemmas than Shanyou63. These results indicate that higher light and CO2 use efficiency, higher resistance to photoinhibition and C4 pathway in both flag leaf and lemma may contribute to the higher yield of the superhigh-yield rice hybrid, Liangyoupeijiu.     

超高产杂交稻光合特性的研究

比较了超高产杂交稻(Oryza sativa L.)X07S/紫恢100和两优培九与多年来大面积推广的杂交稻汕优63的光合功能和抗光胁迫能力.结果表明,超高产杂交稻X07S/紫恢100和两优培九的净光合速率(Pn)分别比汕优63高9.1%和11.9%,而其蒸腾速率(Tr)分别比汕优63低37.46%和31.42%,此外,其水分利用效率(WUE)分别比汕优63高出74.2%和63.5%;经强光(2 000μmol photons@m-2@s-1)处理2 h之后,X07S/紫恢100和两优培九的光系统Ⅱ光化学量子效率分别上升28.3%和37.0%,荧光光化学猝灭系数分别升高46.2%和18.0%,而汕优63的这两项参数值却变化很小;同时,X07S/紫恢100和两优培九的非光化学猝灭系数均下降50%左右,而汕优63则上升近50%,表明超高产杂交稻较高的光合能力、水分利用效率以及较强的抗光抑制能力可能是其高产的生理基础.     

Photoinhibition and Recovery of photosystem 2 in Grapevine (Vitis vinifera L.) Leaves Grown Under Field Conditions

Photoinhibition of photosynthesis was investigated in grapevine (Vitis vinifera L.) exposed to 2 or 4h of high irradiance (HI) (1 700–1 800 mol m^–2 s^–1) leaves under field conditions at different sampling time in a day. The degree of photoinhibition was determined by means of the ratio of variable to maximum chlorophyll fluorescence (F_v/F_m) and photosynthetic electron transport measurements. When the photochemical efficiency of photosystem 2 (PS2), F_v/F_m, markedly declined, F₀ increased in both 2 (HI2) and 4 h (HI4) HI leaves sampled at midday. When various photosynthetic activities were followed on isolated thylakoids, HI4 leaves showed significantly higher inhibition of whole chain and PS2 activity than the HI2 leaves sampled at midday. Later, the leaves reached maximum PS2 efficiencies similar to those observed early in the morning during sampling at evening. The artificial exogenous electron donor Mn²⁺ failed to restore PS2 activity in both variants of leaves, while DPC and NH²OH significantly restored PS2 activity in HI4 midday leaf samples. Quantification of the PS2 reaction centre protein D1 and 33 kDa protein of water splitting complex following midday exposure of leaves showed pronounced differences between HI2 and HI4 leaves. The marked loss of PS2 activity noticed in midday samples was mainly due to the marked loss of D1 protein in HI2, while in HI4 it was mainly 33-kDa protein.     

Characteristics of Photosynthetic Apparatus in Mn-Starved Maize Leaves

The effects of Mn-deficiency on CO₂ assimilation and excitation energy distribution were studied using Mn-starved maize leaves. Mn-deficiency caused about 70 % loss in the photon-saturated net photosynthetic rate (P _N) compared to control leaves. The loss of P _N was associated with a strong decrease in the activity of oxygen evolution complex (OEC) and the linear electron transport driven by photosystem 2 (PS2) in Mn-deficienct leaves. The photochemical quenching of PS2 (q_P) and the maximum efficiency of PS2 photochemistry (F_v/F_m) decreased significantly in Mn-starved leaves under high irradiance, implicating that serious photoinhibition took place. However, the high-energy fluorescence quenching (q_E) decreased, which was associated with xanthophyll cycle. The results showed that the pool of de-epoxidation components of the xanthophyll cycle was lowered markedly owing to Mn deficiency. Linear electron transport driven by PS2 de-creased significantly and was approximately 70 % lower in Mn-deficient leaves than that in control, indicating less trans-thylakoid pH gradient was built in Mn deficient leaves. We suggest that the decrease of non-radiative dissipation depending on xanthophyll cycle in Mn-starved leaves is a result of the deficiency of trans-thylakoid pH gradient.     

Diurnal Changes of Gas Exchange,Chlorophyll Fluorescence,and Stomatal Aperture of Hybrid Poplar Clones Subjected to Midday Light Stress

Diurnal changes in net photosynthetic rate (P _N), chlorophyll (Chl) fluorescence, and stomatal aperture of several hybrid poplar clones subjected to midday light stress were measured in July and August of 1996. Midday depression of P _N, photosystem 2 (PS2) efficiency, stomatal conductance (g _s), and stomatal aperture was observed in all clones, though at differing rates among them. Non-uniform stomatal closure occurred at noon and at other times, requiring a modification of intercellular CO₂ concentration (C ₁). A linear relationship was found between g _s and stomatal aperture. More than half of the photons absorbed by PS2 centre dissipated thermally when subjected to light stress at noon. There was a linear relationship between the rate of PS2 photochemical electron transport (PxPFD) and P _N. There was a consensus for two fluorescence indicators (1 – q_P/q_N and (F_m' – F)/F_m') in assessment of susceptibility of photoinhibition in the clones. According to P _N, Chl fluorescence, and stomatal aperture, we conclude that midday depression of photosynthesis can be attributed to both stomatal and non-stomatal limitations.     

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.     

Involvement of betacyanin in chilling-induced photoinhibition in leaves of <Emphasis Type="Italic">Suaeda salsa</Emphasis>

Seeds of Suaeda salsa were cultured in dark for 3 d and betacyanin accumulation in seedlings was promoted significantly. Then the seedlings with accumulated betacyanin (C+B) were transferred to 14/10 h light/dark and used for chilling treatment 15 d later. Photosystem 2 (PS2) photochemistry, D1 protein content, and xanthophyll cycle during the chilling-induced photoinhibition (exposed to 5 °C at a moderate photon flux density of 500 μmol m⁻² s⁻¹ for 3 h) and the subsequent restoration were compared between the C+B seedlings and the control (C) ones. The maximal efficiency of PS2 photochemistry (F_v/F_m), the efficiency of excitation energy capture by open PS2 centres (F_v′/F_m′), and the yield of PS2 electron transport (Φ_PS2) of the C+B and C leaves both decreased during photoinhibition. However, smaller decreases in F_v/F_m, F_v′/F_m′, and Φ_PS2 were observed in the C+B leaves than in C ones. At the same time, the deepoxidation state of xanthophyll cycle, indicated by (A+Z)/(V+A+Z) ratio, increased rapidly but the D1 protein content decreased considerably during the photoinhibition. The increase in rate of (A+Z)/(V+A+Z) was higher but the D1 protein turnover was slower in C+B than C leaves. After photoinhibition treatment, the plants were transferred to a dim irradiation (10 μmol m⁻² s⁻¹) at 25 °C for restoration. During restoration, the chlorophyll (Chl) fluorescence parameters, D1 protein content, and xanthophyll cycle components relaxed gradually, but the rate and level of restoration in the C+B leaves was greater than those in the C leaves. The addition of betacyanins to the thylakoid solution in vitro resulted in similar changes of F_v/F_m, D1 protein content, and (A+Z)/(V+A+Z) ratio during the chilling process. Therefore, betacyanin accumulation in S. salsa seedlings may result in higher resistance to photoinhibition, larger slowing down of D1 protein turnover, and enhancement of non-radiative energy dissipation associated with xanthophyll cycle, as well as in greater restoration after photoinhibition than in the control when subjected to chilling at moderate irradiance.     

Comparison of Some Photosynthetic Characters Between Two Hybrid Rice Combinations Differing in Yield Potential

Photosynthetic characteristics of two hybrid rice combinations, Peiai 64S/E32 and Shanyou 63, were compared at the panicle differentiation stage. As compared with Shanyou 63, the new combination Peiai 64S/E32 showed a significantly higher net photosynthetic rate (P _N), apparent quantum yield of carbon assimilation (_c), carboxylation efficiency (CE), and photorespiratory rate (R _P) as well as leaf chlorophyll content, but a significantly lower dark respiration rate (R _D) and compensation irradiance (I _c). It also showed a slightly higher photochemical efficiency (F_v/F_m and F/F_m) of photosystem 2, a lower non-photochemical quenching (q_N), and a similar CO₂ compensation concentration () as compared to Shanyou 63.     

Relationship Between PSⅡExcitation Pressure and Content of Super High-Yielding Hybrid Rice

The changes in photochemical features of PhotosystemⅡ(PSⅡ) and contents of Rubisco large subunit (RLS) and small subunit (RSS) in flag leaf from 75DAS to 113DAS (from filling to harvesting stages) were investigated in two hybrid rices (Oryza sativa L.) cv. Liangyoupeijiu and cv. Shanyou 63 grown in the field. Liangyoupeijiu is a super high-yielding rice and Shanyou 63 has widely been planted in China in these years. The results indicate that soluble protein and chlorophyll in both cultivars degraded slowly at first and dramatically thereafter. The degradation speed of soluble protein in Shanyou 63 was faster than that in Liangyoupeijiu. Both Fv/Fm and qP decreased in parallel with leaf senescence, whereas qN fell at first and then rose. No significant change in excitation pressure (1-qP ) was found before 89DAS but a sharply increase in both cultivars after it occurred. Excitation pressure rose more rapidly in Shanyou 63 than that in Liangyoupeijiu. The changes of RLS and RSS content exhibited the same trend as that of soluble protein content. A better linear correlation between RLS, RSS degradation and elevation of (1-qP ) were shown in both cultivars. We suggest that the increase in PSⅡ excitation pressure possibly induced the quick senescence process in rice flag leaf. The high-yielding of Liangyoupeijiu may be due to its maintenance of stronger photosynthetic capacity, longer and more stable photosynthetic functional du-ration than that of Shanyou 63.     

Response of xanthophyll cycle and chloroplastic antioxidant enzymes to chilling stress in tomato over-expressing glycerol-3-phosphate acyltransferase gene

Over-expression of chloroplastic glycerol-3-phosphate acyltransferase gene (LeGPAT) increased unsaturated fatty acid contents in phosphatidylglycerol (PG) of thylakoid membrane in tomato. The effect of this increase on the xanthophyll cycle and chloroplast antioxidant enzymes was examined by comparing wild type (WT) tomato with the transgenic (TG) lines at chilling temperature (4 °C) under low irradiance (100 μmol m⁻² s⁻¹). Net photosynthetic rate and the maximal photochemical efficiency of photosystem (PS) 2 (F_v/F_m) in TG plants decreased more slowly during chilling stress and F_v/F_m recovered faster than that in WT plants under optimal conditions. The oxidizable P700 in both WT and TG plants decreased during chilling stress under low irradiance, but recovered faster in TG plants than in the WT ones. During chilling stress, non-photochemical quenching (NPQ) and the de-epoxidized ratio of xanthophyll cycle in WT plants were lower than those of TG tomatoes. The higher activities of superoxide dismutase (SOD) and ascorbate peroxidase (APX) in TG plants resulted in the reduction of O₂ ^−· and H₂O₂ contents during chilling stress. Hence the increase in content of unsaturated fatty acids in PG by the over-expression of LeGPAT could alleviate photoinhibition of PS2 and PS1 by improving the de-epoxidized ratio of xanthophyll cycle and activities of SOD and APX in chloroplast.     

Photoinhibition of photosynthesis in leaves of grapevine (<Emphasis Type="Italic">Vitis vinifera</Emphasis> L. cv. Riesling). Effect of chilling nights

Photoinhibition of photosynthesis was investigated in control (C) and chilling night (CN) leaves of grapevine under natural photoperiod at different sampling time in a day. The degree of photoinhibition was determined by means of the ratio of variable to maximum chlorophyll fluorescence (F_v/F_m) and photosynthetic electron transport measurements. When the potential efficiency of photosystem (PS) 2, F_v/F_m was measured at midday, it markedly declined with significant increase of F₀ in CN leaves. In isolated thylakoids, the rate of whole chain and PS2 activity were markedly decreased in CN leaves than control leaves at midday. A smaller inhibition of PS1 activity was also observed in both leaf types. Later, the leaves reached maximum PS2 efficiencies similar to those observed in the morning during sampling at evening. The artificial exogenous electron donors diphenyl carbazide, NH₂OH, and Mn²⁺ failed to restore the PS2 activity in both leaf types at midday. Thus CN enhanced inactivation on the acceptor side of PS2 in grapevine leaves. Quantification of the PS2 reaction centre protein D1 following midday exposure of leaves showed pronounced differences between C and CN leaves. The marked loss of PS2 activity in CN leaves noticed in midday samples was mainly due to the marked loss of D1 protein of the PS2 reaction centre.     

Responses of Tobacco Plantlets to Change of Irradiance During Transfer from in vitro to ex vitro Conditions

Chlorophyll a fluorescence kinetics, net photosynthetic rate (P _N), water relations, and photosynthetic pigment contents were studied during acclimation of in vitro grown tobacco to higher irradiance (HL; 700 mol m^–2 s^–1). Plantlets were grown on medium containing sucrose in glass vessels (G-plants) or in Magenta boxes (M-plants) with better CO₂ supply in the latter ones. The effect of HL was studied either (1) in plantlets grown under original in vitro conditions (closed vessels), (2) in in vitro plantlets exposed to ambient CO₂ concentration (covers removed), or (3) in plantlets transplanted to ex vitro into pots with sand and nutrient solution. Higher P _N, and fraction of closed photosystem 2 (PS2) centres (1 – q_P), and lower content of xanthophyll cycle pigments were found in M-plants compared to G-plants. HL treatment caused photoinhibition particularly in plants kept in closed vessels. This was indicated by the decrease in the ratio of F_v/F_m and by the increase in non-photochemical quenching, 1 – qp, and content of xanthophyll cycle pigments. Better CO₂ supply ensured by the removal of closure lead to the moderate reduction of symptoms of photoinhibition, although stomatal conductance (g _s), transpiration rate (E), and P _N were negatively affected. The main reason was the decrease in relative air humidity, which caused similar reduction of P _N, E, and g _s after the transfer of plantlets to ex vitro. Nevertheless, plant response to HL seemed not to be affected by any possible root injury caused by transfer to ex vitro. The differences in contents of xanthophyll cycle pigments, degree of de-epoxidation, P _N, and quenching parameters between M- and G-plantlets were still significant 7 d after ex vitro transfer and HL acclimation.     

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.     

Photosynthetic acclimation and photoinhibition on exposure to high light in shade-developed leaves of Fagus crenata seedlings

The physiological response of leaves developed in low light (L) on Fagus crenata seedlings exposed to different levels of high light (H: high light, M: medium light) was studied. Measurements were conducted on potted seedlings in the F. crenata forest understory. The seedlings with leaves developed in L were transferred to H (L–H) and M (L–M) in summer. On exposure to high light, the photochemical efficiency of dark-adapted PSII (F_v/F_m) immediately decreased and was followed by a subsequent recovery in both L–H and L–M leaves. The mean value of F_v/F_m in L–H leaves was lower than that in L–M leaves through experiments, indicating that the degree of photoinhibition in L–H leaves was greater than that in L–M leaves. About 1 month after transfer, 37% and 5% of leaves had fallen in L–H and L–M seedlings, respectively. This result also indicated the greater photoinhibition in L–H leaves. Moreover, the photosynthetic capacity (P_Nmax) of L–H leaves decreased. In contrast, the P_Nmax of L–M leaves increased, although the P_Nmax was lower than that of M control leaves. An increase in the xanthophyll cycle pool (VAZ), indicating an increase of the photoprotective function, was found in both L–H and L–M leaves. Especially, the VAZ pool in L–M leaves was higher than that in M leaves by the end of experiments. L–M leaves may avoid photoinhibition effectively by the decrease in excess light with the increase of the P_Nmax or VAZ pool, compared to L–H leaves. Thus, the physiological acclimation on exposure to high light depended on the degree of high light. To achieve successful photosynthetic acclimation with slight photoinhibition, the variation of light intensity before and after exposure to high light would be an important factor because of the difference in excess light.