兼性CAM植物长叶景天叶片在C3和CAM型时的光氧化作用

通过停止浇水产生水分胁迫使兼性CAM植物长叶景天 (SedumspectabileBoreau)叶片光合途径由C3 型转为CAM型。干旱 15d时观察到典型的CAM生理特征 ,且叶片的δ13 C值与含水量成线性相关。水分胁迫改变了叶绿素荧光参数和抗氧化能力 ,ΦPSⅡ 和qP 降低 5 0 %和 34% ,NPQ提高约 180 % ,SOD活性和清除DPPH·自由基能力也明显下降 ,但膜半透性变化不大。当将处于C3 (浇水 )和诱导为CAM(缺水 )型的叶圆片用外源甲基紫精 (MV)和强光作光氧化处理后 ,与C3 型叶片相比 ,诱导CAM型叶片的NPQ不能提高 ,qP 和ΦPSⅡ 降至很低水平 ,光系统处于高还原态 ,光能供给与消耗失衡 (1-qP=0 .86和 (1-qP) /NPQ >1) ,膜系统几乎失去完整性。这种严重的光氧化损伤表明 ,与我们以前报告的专性CAM植物不同 ,以兼性CAM植物诱导表达的CAM型未能显示比C3 型较强的耐光氧化优势。讨论了出现这种光氧化敏感性差别的可能原因     

长药景天的叶龄与CAM活性

用CO_2分析。含酸量和PEP羧化酶活性测定的方法对长药景天(Sedum spectabile)光合型进行研究,其成熟叶浇水时表现为C_3型,干旱条件下转变成CAM型,但极度的干旱又使其CAM活性降低。浇水植株的幼叶PEP羧化酶活性和含酸量低,成熟叶较高。但在干旱过程中,幼叶酶活性和含酸量的增加比成熟叶更为显著,表明幼叶对水分胁迫更敏感。水分胁迫中,叶最大PEP羧化酶活性和最大含酸量在植株由下而上各叶位转移。     

宝山堇菜(Viola baoshanensis)、紫花地丁(V. yedoensis)光合异质性比较

利用荧光成像技术,研究了特定和不同光合有效辐射下,宝山堇菜(Viola baoshanensis)、紫花地丁(V.yedoensis)不同叶龄(幼叶和成熟叶片)叶片纵向(叶基、叶中部、叶尖)间的光合异质性特征。特定光化光照射下,两种堇菜不同叶龄的Fv/Fm、ΦpsⅡ、qP、PS/50和Abs在叶尖、叶中部、叶基间呈依次降低趋势,NPQ/4和qN变化趋势与之相反。Fv/Fm、ΦpsⅡ和Abs在两种堇菜不同叶龄的叶片纵向间均没有显著性差异,NPQ/4和qN均显示宝山堇菜不同叶龄的叶片纵向间存在显著差异;但NPQ/4和qN分别显示紫花地丁成熟叶和幼叶叶片的叶尖和叶基处差异显著。qP的显著差异只存在于宝山堇菜幼叶的叶尖和叶基处,PS/50在两种植物幼叶纵向间均有显著差异。快速光曲线的变化中,两种堇菜α由叶尖向叶基下降幅度不明显,幼叶纵向间Pm差异显著,成熟叶叶尖处Pm显著高于叶中部和叶基。两种堇菜成熟叶叶尖处Ik显著高于叶中部和叶基,宝山堇菜幼叶纵向间Ik差异显著,而紫花地丁幼叶纵向间Ik差异不显著。以上结果反映出两种堇菜叶片纵向间Fv/Fm和ΦpsⅡ具有较高的均质性,Fv/Fm和ΦpsⅡ的下降受到NPQ/4、qN、qP、PS/50和Abs的综合影响,但Fv/Fm和ΦpsⅡ的变化与NPQ/4、qN、qP、PS/50的显著变化并不一致。叶龄对两种堇菜叶片纵向间α影响不显著,对Pm影响显著,α的小幅下降反映两种堇菜叶片纵向间捕光能力基本相同。叶片纵向间Ik的显著差异受叶龄和植物种类的综合影响。     

金钟藤叶绿素荧光特征初探

利用叶绿素荧光技术初步观测丁金钟藤在不同作用光下的光合生理特征,并对其离体叶片在低温、高温及脱水胁迫下的光能利用状况作了比较。结果显示,随着作用光强的增加,叶片Fv＇/Fm＇、qP和ΦPSⅡ呈下降趋势,qN和NPQ表现为上升。离体叶片在5℃处理6h后,其Fv＇/Fm＇、qP和ΦPSⅡ高于25℃和45℃处理的,而NPQ在5℃和45℃处理6h后明显低于对照（25℃）,qN则略有降低。离体叶片脱水胁迫12h后,仍有一定的光能转化能力,但脱水24h和36h后,qP、ΦPSⅡ、qN、NPQ都明显下降。这表明金钟藤对于环境因子,如光强、温度及水分的变化具有较强的适应性,这可能是其在森林中能快速蔓延扩散的一个重要原因。     

不同菠菜品种对海水胁迫的生理响应差异

以耐海水菠菜品种荷兰3号和海水敏感品种圆叶菠菜为材料,采用水培方法,研究了海水胁迫对菠菜叶片、根系质膜的伤害作用以及叶片光合作用、叶绿素荧光参数的影响.结果表明:(1)海水胁迫对荷兰3号单株干重影响不大,而显著降低圆叶菠菜的单株干重,并使2个品种植株的叶片和根系MDA含量增加,质膜透性增大,叶片光合色素含量降低,但荷兰3号的变化幅度(叶片MDA除外)小于圆叶菠菜.(2)海水胁迫下,短期内2个品种由于气孔限制引起叶片胞间CO2浓度(Ci)降低,净光合速率(Pn)下降;长期胁迫下,荷兰3号Pn恢复到对照水平,而圆叶菠菜同化力下降,Pn降低.(3)海水胁迫对荷兰3号光化学猝灭系数(qP)影响不大,实际光化学效率(ΦPSⅡ)明显升高,而圆叶菠菜的qP和ΦPSⅡ均下降;荷兰3号初始荧光(F0)的下降幅度和非光化学猝灭系数(qN)上升幅度比圆叶菠菜大;2个菠菜品种的最大光化学效率(Fv/Fm)均下降,但荷兰3号光抑制程度(1-qP/qN)的升高幅度比圆叶菠菜小.研究结果说明,海水胁迫下,2个耐性不同的菠菜品种植株都产生了光合作用的光抑制和光氧化伤害,使膜质过氧化和叶绿素含量降低;耐性强的品种能够较多地将光能用于光化学反应,热耗散能力较强,光抑制程度较低,膜系统和光合色素受到活性氧的破坏程度较低,保持了较高的净光合速率,最终可明显降低海水胁迫对植株生长的影响.     

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.     

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（3μmol 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3d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值能完全恢复。说明脱水的牛耳草主体叶片光系统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。     

夜间低温后日间光照对海桐和榕树叶片的光抑制以及光系统Ⅱ功能的影响

夜间低温导致海桐和榕树叶片光系统Ⅱ(PSⅡ)最大光化学效率(Fv/Fm)、PSⅡ光合电子传递量子效率(φPSⅡ)、天线转化效率(F'v'/Fm')、非光化学猝灭系数(NPQ)降低,其后日间光照先引起海桐叶片Fv//Fm、ΦPSⅡ、F'v'/Fm'稍微降低,其后又逐渐得到恢复,但NPQ却表现出相反趋势;夜间低温及随后的日间光照并未对海桐叶片光化学猝灭系数(qP)和初始荧光强度(Fo)产生影响.夜间低温后日间光照进一步引起榕树叶片Fv/Fm、ΦPSⅡ、Fv'/F'm、qp、NPQ下降,在午后光照减弱后仍不能得到恢复.     

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.     

自然条件下不同高产稻生育后期剑叶叶绿素荧光和膜脂过氧化的表现

以籼型 (OryzasativaL .)杂交组合汕优 6 3为对照 ,以中粳 95 16、两系亚种间杂交组合培矮 6 4S/E32、培矮6 4S/ 9311、亚种间三系杂交稻冈优 881和两系杂交组合X0 7S/紫恢 10 0为材料 ,研究其在生育后期 (抽穗 -成熟 )自然条件下剑叶的叶绿素衰减、CO2 交换、叶绿素荧光参数和膜脂过氧化表现。结果表明 :水稻在生育后期伴随叶绿素衰减 ,其叶内的原初光化学效率Fv/Fm、PSⅡ非环式电子传递效率ΦPSⅡ 、电子流传递速率ETR都有相应地下降 ,这种光能转化的障碍使多余的光能传递给PSⅡ的还原侧 ,产生O-·2 累积 ,发生膜脂过氧化和MDA的积累 ,引起光合色素及光合膜的破坏 ,发生光氧化早衰。这种现象在品种间有明显差异 ,耐光氧化的粳稻 95 16 ,其叶内的Fv/Fm、ΦPSⅡ 、ETR、qP下降较少 ,具有较稳定的光能转化能力 ,不易早衰 ,具有较高的结实率 ;而对光氧化敏感的籼稻汕优 6 3其叶内的Fv/Fm、ΦPSⅡ 、ETR ,光化学猝灭参数qP下降较多 ,易发生膜脂过氧化 ,导致叶片早衰 ,影响水稻灌浆结实和产量 ;而二系的和三系的杂交稻的耐光氧化特性和早衰表现居于中间。从水稻超高产育种的角度出发 ,在目前株型良好的基础上 ,兼顾杂种优势和防止早衰两方面考虑 ,在母本中利用粳型或带有粳型基因的不育系是育种上一个值得重     

Ecophysiology of Crassulacean Acid Metabolism (CAM)

BACKGROUND AND SCOPE: Crassulacean Acid Metabolism (CAM) as an ecophysiological modification of photosynthetic carbon acquisition has been reviewed extensively before. Cell biology, enzymology and the flow of carbon along various pathways and through various cellular compartments have been well documented and discussed. The present attempt at reviewing CAM once again tries to use a different approach, considering a wide range of inputs, receivers and outputs. INPUT: Input is given by a network of environmental parameters. Six major ones, CO(2), H(2)O, light, temperature, nutrients and salinity, are considered in detail, which allows discussion of the effects of these factors, and combinations thereof, at the individual plant level ('physiological aut-ecology'). RECEIVERS: Receivers of the environmental cues are the plant types genotypes and phenotypes, the latter including morphotypes and physiotypes. CAM genotypes largely remain 'black boxes', and research endeavours of genomics, producing mutants and following molecular phylogeny, are just beginning. There is no special development of CAM morphotypes except for a strong tendency for leaf or stem succulence with large cells with big vacuoles and often, but not always, special water storage tissues. Various CAM physiotypes with differing degrees of CAM expression are well characterized. OUTPUT: Output is the shaping of habitats, ecosystems and communities by CAM. A number of systems are briefly surveyed, namely aquatic systems, deserts, salinas, savannas, restingas, various types of forests, inselbergs and paramós. CONCLUSIONS: While quantitative census data for CAM diversity and biomass are largely missing, intuition suggests that the larger CAM domains are those systems which are governed by a network of interacting stress factors requiring versatile responses and not systems where a single stress factor strongly prevails. CAM is noted to be a strategy for variable, flexible and plastic niche occupation rather than lush productivity. 'Physiological syn-ecology' reveals that phenotypic plasticity constitutes the ecophysiological advantage of CAM.     

Phosphoenolpyruvate carboxylase genes in C3, crassulacean acid metabolism (CAM) and C3/CAM intermediate species of the genus Clusia: rapid reversible C3/CAM switches are based on the C3 housekeeping gene

The genus Clusia includes species that exhibit either the C3 or crassulacean acid metabolism (CAM) mode of photosynthesis, or those that are able to switch between both modes according to water availability. In order to screen for species-specific genetic variability, we investigated the key carboxylase for CAM, phosphoenolpyruvate carboxylase (PEPC). Sequence analysis of DNA isolated from the obligate CAM species, Clusia hilariana, the obligate C3 species, Clusia multiflora, and an intermediate species that can switch between C3 and CAM photosynthesis, Clusia minor, revealed three different isoforms for C. hilariana and one each for the other two species. Sequence alignments indicated that PEPC from the intermediate species had high homology with the C3 protein and with one of CAM plant proteins. These were assumed to constitute 'housekeeping' proteins, which can also support CAM in intermediate species. The other two isoforms of the CAM plant C. hilariana were either CAM-specific or showed homologies with PEPC from roots. Phylogenetic trees derived from neighbour-joining analysis of amino acid sequences from 13 different Clusia species resulted in two distinct groups of plants with either 'housekeeping' PEPC only, or additionally CAM-related isoforms. Only C. hilariana showed the third, probably root-specific isoform. The high homology of the PEPC from the intermediate species with the C3 protein indicates that for the reversible transition from the C3 to CAM mode of photosynthesis, the C3 type of PEPC is sufficient. Its expression, however, is strongly increased under CAM-inducing conditions. The use of the C3 isoform could have facilitated the evolution of CAM within the genus, which occurred independently for several times.     

NADP—苹果酸酶活性变化及其在CAM运行中的调节

ＮＡＤＰ－苹果酸酶是ＣＡＭ植物一种重要脱羧酶。实验结果表明,专一ＣＡＭ植物瓦松和兼性ＣＡＭ植物长药景天及露花其ＮＡＤＰ－苹果酸酶活性昼高夜低;５－８月,兼性ＣＡＭ植物长药景天和露花随着Ｃ３光合型向ＣＡＤ型转化,其中ＮＡＤＰ－苹果酸活活性逐渐升高。     

Cell organelles from crassulacean-acid-metabolism (CAM) plants

Cell organelles were isolated from the CAM plants Crassula lycopodioides Lam., Bryophyllum calycinum Salisb. and Sedum rubrotinctum R.T. Clausen by isopycnic centrifugation in sucrose gradients. The inclusion of 2.5% Ficoll in the grinding medium proved to be essential for a satisfactory separation of cell organelles during the subsequent centrifugation. Peroxisomes, mitochondria, and whole and broken chloroplasts were at least partially resolved as judged by marker-enzyme-activity profiles. The isolated peroxisomes contained activities of glycollate oxidase, catalase, hydroxypyruvate reductase, glycine aminotransferase, serine-glyoxylate aminotransferase, and aspartate aminotransferase, comparable to activities found in spinach (Spinacia oleracea L.) leaf peroxisomes. In contrast to spinach, however, only little, if any, particulate malate dehydrogenase activity could be attributed to isolated peroxisomes of the three CAM plants.     

新的C4及CAM光合途径植物

以稳定性碳同位素比(δ~(13)C)鉴别禾本科、莎草科、苋科和萝摩科共46种植物的光合作用途径,发现了36种新的C_4植物(δ~(13)C-10.43到-13.66‰)和1种CAM植物(δ~(13)C-15.24‰)。根据Hattersley区分C_4植物三种亚类型的不同δ~(13)C值,提出在36种C_4中有8种具δ_(13)C值-10.4到-10.9‰者是NADP-ME型,6种具δ~(13)C值-13‰左右的是NAD-ME型,其余种类可能是NADP-ME或PCK型。     

CAM induction by photoperiodism in green callus cultures from a CAM plant

Abstract Green calli obtained from leaves of the CAM-inducible plant Kalanchoe blossfeldiana cv. Montezuma were grown either under long-day or short-day regimes in the Phytotron of Gif-sur-Yvette. Callus cells were found to be CAM inducible by the short-day treatment, according to levels and diurnal oscillations of malate pools and phosphor-enolpyruvate (PEP) carboxylase (EC.4.1.1.31) capacity. De novo synthesis of PEP carboxylase was shown to occur under the short-day regime. In spite of continuous net CO² output, CAM-like patterns of CO₂ exchange by calli were obtained under the short-day treatment. After 2 months under both photoperiodic conditions, spontaneous organogenesis in callus tissues gave rise to numerous shootlets which were determined as photoperiod dependent for CAM; the same was so for plants originating from these shootlets.     

几种CAM植物PEP羧化酶同工酶的比较研究

比较研究几种兼性和专一性ＣＡＭ植物材料的ＰＥＰＣ同工酶表明：经自然干旱诱导,兼性ＣＡＭ植物露花（Ｍｅｓｅｍｂｒｙａｎｔｈｅｍｕｍｃｏｒｄｉｆｏｌｉｕｍ）、长药景天（Ｓｕｄｕｍｓｐｅｃｔａｂｉｌｅ）有新的ＰＥＰＣ同工酶的出现,诱导前后各同工酶的天然分子量变化不大;而土三七（Ｓｅｄｕｍａｉｚｏｏｎ）则没有新的ＰＥＰＣ同工酶出现,但诱导后其同工酶的天然分子量有所增大。以上几种兼性ＣＡＭ植物的ＰＥＰＣ同工酶酶谱无明显昼夜变化。专一性ＣＡＭ植物的ＰＥＰＣ酶谱和天然分子量均较一致,亦无昼夜差异。     

Kinetic changes with temperature of phosphoenolpyruvate carboxylase from a CAM plant

Abstract. Purified and crude phosphoenolpyruvate carboxylase from the CAM plant Kalanchoë daigremontiana Hamet et Perrier ( Bryophyllum diagremontianum ) was assayed at temperatures between 10 and 45° C. The optimum temperature of the enzyme activity changed with substrate availability and effector concentration in the assay. l -malate inhibited the enzyme activity and lowered the optimum temperature. Glucose-6-phosphate raised the optimum temperature to 43°C. K _m values for phosphoenolpyruvate increased with assay temperature from 0.12 mol m^-3 at 15° C to 0.36 mol^m−3 at 35° C. Inhibition by malate increased with temperature and acidity of the assay. In the crude enzyme 50% of control activity was inhibited by 1.65 mol m^-3 malate at 15° C and by 0.5 mol m^-3 at 35° C (at pH 7.0). With purification malate sensitivity was lost ( K _i values for malate at least 10 times higher). The shift in optimum temperatures for PEP-carboxylase activity thus results from changes in the kinetic parameters with temperature and allosteric effectors. The often low optimum temperatures for CO₂ fixation observed in nature may thus be the result of substrate and effector concentrations in the cytoplasm and the antagonistic effect of temperature on substrate affinity and effector efficiency on phosphoenolpyruvate carboxylase.