叶黄素循环和D1蛋白周转在毛竹叶片光保护中的作用

利用叶绿素荧光技术,对强光胁迫下以及叶黄素循环抑制剂-二硫苏糖醇(DTT)和D1蛋白合成抑制剂-硫酸链霉素(SM)处理后毛竹(Phyllostachys edulis (Carr.) Lehaie)的光抑制特征进行研究。结果显示:在夏季中午强光或人为强光胁迫下,毛竹叶片最大光化学效率Fv/Fm均显著降低;在下午光强减弱或黑暗、弱光条件下,Fv/Fm可有效恢复。DTT和SM均可抑制毛竹叶片非光化学淬灭(NPQ),且DTT效果明显优于SM。另外,在强光下,DTT和SM处理均能使毛竹叶片Fv/Fm、实际光化学效率Y(Ⅱ)和光化学淬灭qP等荧光参数下降幅度增大。研究结果表明毛竹叶片具有完善的光破坏防御机制,NPQ与叶黄素循环和D1蛋白周转紧密关联,在叶片光保护机制中具有重要作用。     

依赖于叶黄素循环的热耗散对茶树叶片防御强光破坏的相关试验研究

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

濒危植物长序榆(Ulmus elongata)幼苗叶绿素荧光特性的初步研究

用Li-6400XT便携式光合作用仪对濒危植物长序榆幼苗的各叶绿素荧光参数的日变化和快速光响应曲线进行了测定。结果发现,光系统Ⅱ(PSⅡ)的实际光化学效率(ΦPSⅡ)、电子传递速率(ETR)在整个白天阶段较稳定,下午18:00显著下降。光化学淬灭(qP)先增大后减小。非光化学淬灭(NPQ)呈现出与光化学淬灭(qP)相反的变化趋势,中午最低,说明长序榆幼苗光能利用率较高。快速光曲线表明实际光化学效率(ΦPSⅡ)和光化学淬灭(qP)随着光合有效辐射(PAR)的增大而减小,电子传递速率(ETR)和非光化学淬灭(NPQ)随着光合有效辐射(PAR)的增大而增大。使用幂函数能够很好的拟合实际光化学效率(ΦPSⅡ)和电子传递速率(ETR)随光强的变化,而对数函数能较好的拟合实际光化学淬灭(qP)和非光化学淬灭(NPQ)随光强的变化。     

光强转换对不同生长环境下桑树叶片光化学效率的影响

以桑树品种‘蒙古桑’为试验材料,利用叶绿素荧光技术研究了光强转换对生长在不同光强下的桑树叶片实际光化学效率(ΦPSⅡ)、电子传递速率(ETR)和非光化学淬灭(NPQ)的影响,分析了非光化学淬灭(NPQ)3个组分的变化.结果表明:当光强从黑暗或弱光转换到自然光条件下,自然光桑树叶片的光量子转化效率高于弱光叶片,ΦPSⅡ、ETR诱导平衡较快,NPQ诱导呈先升后降趋势.自然光叶片在强光下状态转换淬灭组分(qT)占NPQ的18%,而弱光叶片qT仅占NPQ的7%.与弱光桑树叶片相比,自然光桑树叶片可以通过较高的光量子转化效率和较强的调节激发能在PSⅠ和PSⅡ之间的分配能力来适应光强的变化.     

植物叶片和冠层光化学反射指数与叶黄素循环的关系

以一串红(Salvia splendens Ker-Gawl.)和白车轴草(Trifolium repens Linn.)为材料,使用光谱反射技术测定了这两种植物叶片水平和冠层水平的光化学反射指数(PRI)的日变化,同时使用高效液相色谱分析法测定了这两种植物叶黄素循环的日变化,分析了单叶、群体冠层的PRI与叶黄素循环和叶片实际光化学效率(ΦPSⅡ)和非光化学淬灭(NPQ)之间的关系.结果表明,不论是叶片水平还是冠层水平,两种植物PRI的变化均与叶黄素的脱环氧化程度和NPQ之间呈显著的负相关、与ΦPSⅡ呈显著的正相关.研究结果表明无论叶片水平还是冠层水平上的光谱反射指数均能非常好地反映植物光合机构对光能的利用效率.     

木棉叶片叶绿素荧光参数和SPAD值对干旱胁迫的响应

采用盆栽控水干旱法模拟干旱逆境,对1年生木棉(Bombax ceiba)盆栽苗进行不同程度的胁迫处理,测定干旱胁迫时其叶片叶绿素SPAD值和荧光参数动态变化。结果表明,干旱胁迫下,木棉叶片叶绿素相对含量逐渐降低;初始荧光(Fo)随着干旱程度的加剧而呈上升趋势,而最大荧光(Fm)、潜在光化学效率(Fv /Fo)、最大光化学效率(Fv /Fm)、光化学淬灭系数(qP)和表观光合电子传递速率(ETR)下降,在中度和重度干旱胁迫时差异显著(P＜0.05);非光化学淬灭系数(NPQ)变化不显著。干旱胁迫导致木棉植株出现光抑制,植株可通过降低光化学淬灭、增加热耗散的形式增强对干旱的适应能力。     

EFFECTS OF TEMPERATURE AND LIGHT TREATMENT ON VIOLAXANTHIN DE-EPOXIDASE ACTIVITY AND XANTHOPHYLL CYCLE-DEPENDENT ENERGY DISSIPATION IN WHEAT LEAVES

 为了探讨温度和光强是如何影响离体紫黄质脱环氧化酶(VDE)活性, 阐明依赖叶黄素循环的热耗散与VDE活性关系, 该文以小麦(Triticum aestivum)为材料, 研究了不同光强(200、500、900和1 200 μmol&;#8226;m^–2&;#8226;s^–1)和不同温度(4、25、38和45 ℃) 交叉处理对小麦叶片VDE活性以及依赖叶黄素循环热耗散能力的影响。结果表明: 小麦叶片VDE活性在30 ℃最高, 说明30 ℃是小麦叶片VDE体外条件下的最适温度; 不同光强处理下小麦叶片VDE活性基本一致。与室温(25 ℃)处理的叶片相比, 低温(4 ℃)处理的叶片VDE活力没有明显下降, 而高温(45 ℃)处理则导致了叶片VDE活性急剧下降。小麦叶片热耗散(NPQ)以及依赖叶黄素循环的热耗散(qE)均随着处理光强的增加不断上升, 而qE/NPQ则随光强增加略微下降, 在1 200 μmol&;#8226;m^–2&;#8226;s^–1光强条件下qE/NPQ则急剧下降。该研究揭示VDE活性与依赖叶黄素循环热耗散能力的指标qE/NPQ的变化有一定的相关性, 但不完全一致。并针对此问题进行了讨论。     

复苏被子植物牛耳草脱水和复水期间叶黄素循环的保护作用

研究了复苏被子植物牛耳草(Boea hygrometrica (Bunge) R.Br.)离体叶片在微弱光强下(3 μmol photons*m-2*s-1)和黑暗中叶黄素循环组分及叶绿素荧光随脱水复水的变化.结果发现:脱水期间随着光系统Ⅱ光化学效率(Fv/Fm)、实际量子产率(ΦPSⅡ)、光化学淬灭(qP)和非光化学淬灭(NPQ)值的降低,微弱光强下的对照叶片玉米黄素含量显著增加,而微弱光强下DTT处理的叶片和黑暗中的叶片都没有玉米黄素的积累.经过3 d复水后,微弱光强下对照叶片的Fv/Fm, ΦPSⅡ, qP 和 NPQ值能完全恢复,但是微弱光强下DTT处理的叶片和黑暗中的叶片其Fv/Fm、ΦPSⅡ、qP 和 NPQ值只有部分恢复.说明脱水的牛耳草离体叶片光系统Ⅱ的光化学活性的恢复明显受到DTT处理和黑暗的影响,因此玉米黄素可能对微弱光强下脱水的牛耳草叶片具有重要的保护作用.     

缺锰降低大豆叶片叶绿素荧光的高能态猝灭

缺锰大豆叶片光合速率比对照低 5 0 %左右。缺锰叶片的Fv/Fm较对照低约 30 %～ 40 % ,而Fo比对照高 2～ 3倍左右。强光下 ,缺锰叶片的qP为对照的5 0 %。缺锰降低了叶片的高能态猝灭 (qE) ,仅占其NPQ的 6 0 %左右 ;而对照 qE组分占NPQ的 90 %左右。对照和缺锰叶片叶黄素库的脱环氧化程度分别为36 %和 2 1%左右。认为缺锰叶片 qE的降低可能是由于叶黄素脱环氧化程度减少所致     

低氧胁迫下黄瓜植株热耗散途径

采用营养液栽培,研究了低氧(营养液溶氧浓度为0.9～1.1 mg·L-1)胁迫下黄瓜幼苗光合作用热耗散与叶黄素循环的关系.结果表明:低氧胁迫下,黄瓜叶片PSⅡ的实际光化学效率(φPSⅡ)、饱和光强下的净光合速率(Pn)、表观量子效率(AQY)和PSⅡ的最大光化学效率(Fv/Fm)均显著降低,表明黄瓜植株的光合作用受到了光抑制;同时,光化学猝灭系数(qp)降低,而热耗散(NPQ)和天线耗散能量(D)的比值显著升高,说明黄瓜叶片热耗散增强;NPQ与叶黄素脱环氧化状态(DEPS)呈显著正相关,且两者均被抗坏血酸(AsA)所促进,被二硫苏糖醇(DTT)所抑制,说明低氧胁迫下,叶黄素循环是黄瓜植株光合作用热耗散的主要途径.     

田间大豆叶片成长过程中的光合特性及光破坏防御机制

田间大豆叶片在成长进程中光饱和光合速率持续提高,但气孔导度的增加明显滞后.尽管叶片在成长初期就具有较高的最大光化学效率,但是仍略低于发育成熟的叶片.随着叶片的成长,光下叶片光系统Ⅱ实际效率增加;非光化学猝灭下降.幼叶叶黄素总量与叶绿素之比较高,随着叶面积的增加该比值下降,在光下,幼叶的脱环氧化程度较高.因此认为大豆叶片成长初期就能够有效地进行光化学调节;在叶片生长过程中依赖叶黄素循环的热耗散机制迅速建立起来有效抵御强光的破坏.     

Xanthophyll Cycle and Inactivation of Photosystem Ⅱ Reaction Centers Alleviating Reducing Pressure to Photosystem Ⅰ in Morning Glory Leaves under Short-term High Irradiance

investigated through chlorophyll fluorescence parameters in morning glory (Ipomoea setosa) leaves, which were dipped into water, dithiothreitol (DTT) and lincomycin (LM), respectively. During the stress, both the xanthophyll cycle and D1 protein turnover could protect PSI from photoinhibition. In DTT leaves, non-photochemical quenching (NPQ) was inhibited greatly and the oxidation level of P700 (P700+) was the lowest one. However, the maximal photochemical efficiency of PSII (Fv/Fm) in DTT leaves was higher than that of LM leaves and was lower than that of control leaves. These results suggested that PSI was more sensitive to the loss of the xanthophyll cycle than PSII under high irradiance. In LM leaves, NPQ was partly inhibited, Fv/Fm was the lowest one among three treatments under high irradiance and P700+ was at a similar level as that of control leaves. These results implied that inactivation of PSII reaction centers could protect PSI from further photoinhibition. Additionally, the lowest of the number of active reaction centers to one inactive reaction center for a PSII cross-section (RC/CSo), maximal trapping rate in a PSII cross-section (TRo/CSo), electron transport in a PSII cross-section (ETo/CSo) and the highest of 1-qP in LM leaves further indicated that severe photoinhibition of PSII in LM leaves was mainly induced by inactivation of PSII reaction centers, which limited electrons transporting to PSI. However, relative to the LM leaves the higher level of RC/CSo, TRo/CSo, Fv/Fm and the lower level of 1-qP in DTT leaves indicated that PSI photoinhibition was mainly induced by the electron accumulation at the PSI acceptor side, which induced the decrease of P700+ under high irradiance.     

Xanthophyll Cycle and Inactivation of Photosystem Ⅱ Reaction Centers Alleviating Reducing Pressure to Photosystem Ⅰ in Morning Glory Leaves under Short-term High Irradiance

Under 30-min high irradiance （1500μmol m^-2 s^-1）, the roles of the xanthophyll cycle and D1 protein turnover were investigated through chlorophyll fluorescence parameters in morning glory （Ipomoea setosa） leaves, which were dipped into water, dithiothreitol （DTT） and lincomycin （LM）, respectively. During the stress, both the xanthophyll cycle and D1 protein turnover could protect PSI from photoinhibition. In DTT leaves, non-photochemical quenching （NPQ） was inhibited greatly and the oxidation level of P700 （P700^＋） was the lowest one. However, the maximal photochemical efficiency of PSII （Fv/Fm） in DTT leaves was higher than that of LM leaves and was lower than that of control leaves. These results suggested that PSI was more sensitive to the loss of the xanthophyll cycle than PSII under high irradiance. In LM leaves, NPQ was partly inhibited, Fv/Fm was the lowest one among three treatments under high irradiance and P700^＋ was at a similar level as that of control leaves. These results implied that inactivation of PSII reaction centers could protect PSI from further photoinhibition. Additionally, the lowest of the number of active reaction centers to one inactive reaction center for a PSII cross-section （RC/CSo）, maximal trapping rate in a PSll cross-section （TRo/CSo）, electron transport in a PSll cross-section （ETo/CSo） and the highest of 1-qP in LM leaves further indicated that severe photoinhibition of PSII in LM leaves was mainly induced by inactivation of PSII reaction centers, which limited electrons transporting to PSh However, relative to the LM leaves the higher level of RC/CSo, TRo/CSo, Fv/Fm and the lower level of 1-qP in DTT leaves indicated that PSI photoinhibition was mainly induced by the electron accumulation at the PSI acceptor side, which induced the decrease of P700^＋ under high irradiance.     

Roles of xanthophylls and exogenous ABA in protection against NaCl-induced photodamage in rice (Oryza sativa L) and cabbage (Brassica campestris)

Changes in actual efficiency of PS II photochemistry, non-photochemical quenching (NPQ), content of xanthophylls and kinetics of de-epoxidation were studied in ABA-fed and non-ABA-fed leaves of rice and cabbage under NaCl stress. Salt stress induced more progressive decrease in actual efficiency of PS II photochemistry (ФPS II), higher reduction state of PS II, and a small significant increase in NPQ in NaCl-sensitive rice plants as compared with NaCl-tolerant cabbage plants, whereas exogenously supplied ABA alleviated the decrease in actual efficiency of PS II photochemistry (ФPS II), induced a lower reduction state of PS II, and caused higher capacity of NPQ in ABA-fed plants than in non-ABA-fed plants. As a result, there were higher activities of photosynthetic electron transport, higher capacity of energy dissipation, and lower cumulation of excess light in cabbage than in rice plants, and in ABA-fed leaves than in non-ABA-fed leaves. The effect of ABA was more efficient in cabbage than in rice plants. Addition of exogenous ABA resulted in enhancement of the size of the xanthophyll cycle pool, promotion of de-epoxidation of the xanthophyll cycle components, and a rise in the level of NPQ by altering the kinetics of de-epoxidation of the xanthophyll cycle. Protection from photodamage appears to be achieved by coordinated contributions by exogenous ABA and xanthophyll cycle-mediated NPQ. This variety of photoprotective mechanisms may be essential for conferring photodamage tolerance under NaCl stress.     

Xanthophyll cycle components and capacity for non-radiative energy dissipation in sun and shade leaves ofLigustrum ovalifolium exposed to conditions limiting photosynthesis

The relationships between photosynthetic efficiency, non-radiative energy dissipation and carotenoid composition were studied in leaves ofLigustrum ovalifolium developed either under full sunlight or in the shade. Sun leaves contained a much greater pool of xanthophyll cycle components than shade leaves. The rate of non-radiative energy dissipation, measured as non-photochemical fluorescence quenching (NPQ), was strictly related to the deepoxidation state (DPS) of xanthophyll cycle components in both sun and shade leaves, indicating that zeaxanthin (Z) and antheraxanthin (A) are involved in the development of NPQ. Under extreme conditions of excessive energy, sun leaves showed higher maximum DPS than shade leaves. Therefore, sun leaves contained not only a greater pool of xanthophyll cycle components but also a higher proportion of violaxanthin (V) actually photoconvertible to A and Z, compared to shade leaves. Both these effects contributed to the higher NPQ in sun versus shade leaves. The amount of photoconvertible V was strongly related to chla/b ratio and inversely to leaf neoxanthin content. This evidence indicates that the amount of photoconvertible V may be dependent on the degree of thylakoid membrane appression and on the organization of chlorophyll-protein complexes, and possible explanations are discussed. Exposure to chilling temperatures caused a strong decline in the photon yield of photosynthesis and in the intrinsic efficiency of PS II photochemistry in sun leaves, but little effects in shade leaves. These effects were accompanied by increases in the pool of xanthophyll cycle components and in DPS, more pronounced in sun than in shade leaves. This corroborates the view that Z and A may play a photoprotective role under unfavorable conditions. In addition to the xanthophyll-related non-radiative energy dissipation, a slow relaxing component of NPQ, independent from A and Z concentrations, has been found in leaves exposed to low temperature and high light. This quenching component may be attributed either to other regulatory mechanism of PS II efficiency or to photoinactivation.Research supported by National Research Council of Italy, Special Project RAISA, Sub-Project 2, Paper N. 1587.     

Changes in activity of energy dissipating mechanisms in wheat flag leaves during senescence

Abstract: Excitation energy dissipation, including the xanthophyll cycle, during senescence in wheat flag leaves grown in the field was investigated at midday and in the morning. With progress of senescence, photosynthesis (Pn) and actual PSII photochemical efficiency (ΦPSII) decreased markedly at midday. The decrease in extent of Pn was greater than that of ΦPSII. However, there was no significant decline in Pn and ΦPSII observed in the morning, except in leaves 60 days after anthesis. The kinetics of xanthophyll cycle activity, thermal dissipation (NPQ), and q_f observed at midday during senescence exhibited two distinct phases. The first phase was characterized by an increase of xanthophyll cycle activity, NPQ, and q_f during the first 45 days after anthesis. The second phase took place 45 days after anthesis, characterized by a dramatic decline in the above parameters. However, the qI, observed both at midday and in the morning, always increased along with senescence. A larger proportion of NPQ insensitive to DTT (an inhibitor of the de-epoxidation of V to Z) was also observed in severely senescent leaves. In the morning, only severely senescent leaves showed higher xanthophyll cycle activity, NPQ, q_f, and qI. It was demonstrated that, at the beginning of senescence or under low light, wheat leaves were able to dissipate excess light energy via NPQ, depending on the xanthophyll cycle. However, the xanthophyll cycle was insufficient to protect leaves against photodamage under high light, when leaves became severely senescent. The ratio of (Fj - Fo)/(Fp - Fo) increased gradually during the first 45 days after anthesis, but dramatically increased 45 days after anthesis. We propose that another photoprotection mechanism might exist around reaction centres, activated in severely senescent leaves to protect leaves from photodamage.     

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.     

Increase in Resistance to Low Temperature Photoinhibition Following Ascorbate Feeding is Attributable to an Enhanced Xanthophyll Cycle Activity in Rice (Oryza sativa L.) Leaves

The mechanistic basis for protection of exogenous ascorbate against photoinhibition at low temperature was examined in leaves of rice (Oryza sativa L.). Exposure of intact leaves to chilling temperature resulted in a drastic decrease in the speed of development of non-photochemical fluorescence quenching (NPQ). This was related to the low temperature-imposed restriction on the formation of the fast relaxing component of NPQ (q_f). Feeding with 20 mM ascorbate markedly increased the rate of q_f development at chilling temperature due primarily to the enhanced rate of zeaxanthin (Z) formation. On the other hand, ascorbate feeding had no influence on photosystem 2 (PS2)-driven electron flow. The reduced state of the PS2 primary electron acceptor Q_A decreased in ascorbate-fed leaves exposed to high irradiance at chilling temperature owing to the increased Z-associated thermal energy dissipation in the light-harvesting antenna system of PS2. Furthermore, ascorbate feeding increased the photosynthetic apparatus of rice leaves to resist photoinhibition at low temperature. The protective effect of exogenous ascorbate was fully accounted for by the enhanced xanthophyll cycle activity.     

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

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