Drought Tolerance in Two Mosses: Correlated with Enzymatic Defence Against Lipid Peroxidation

Drought-induced changes in the activities of superoxide dismutase(SOD) and catalase, level of lipid peroxidation, and membranepermeability (solute leakage) have been studied in two mosses,the drought-tolerant Tortula ruralis and the drought-sensitiveCratoneuron filicinum. In T. ruralis the activities of SOD andcatalase increase during slow drying. The level of lipid peroxidationconsequently declines. On subsequent rehydration the enzymeactivities decline and the level of lipid peroxidation risesgradually to normal levels. The leakage of preloaded ⁸⁶Rb onrehydration of slowly dried T. ruralis is similar to that inturgid moss, i.e. leakage of about 20% of tissue ⁸⁶Rb. WhenT. ruralis is subjected to rapid drying there is no change inthe enzyme activities or in lipid peroxidation. However, whenthis moss is rehydrated there is a large immediate increasein lipid peroxidation. Half of the tissue ⁸⁶Rb is leaked intothe bathing medium during the first hour of rehydration. Butwithin the next hour, when SOD and catalase activities haveincreased to high levels, lipid peroxidation quickly declinesto a level lower than that in the turgid control moss, and the⁸⁶Rb leaked earlier is partly reabsorbed indicating that membranerepair is well underway. On prolonged rehydration the enzymeactivities decline and the level of lipid peroxidation risesgradually to reach normal levels found in control turgid moss.In the case of drought-sensitive C. filicinum the activitiesof SOD and catalase decline during drying as well as duringsubsequent rehydration. There is a rapid increase in lipid peroxidationduring rehydration and most of the preloaded ⁸⁶Rb leaks intothe bathing medium irreversibly. The changes in lipid peroxidationduring drying and subsequent rehydration of both the mossesappear to coincide in time with the reported changes in O₂ uptake,indicating that the drought-induced membrane damage may be dueto free radical-induced lipid peroxidation which is known torequire active O₂ uptake. Furthermore, there appears to be agood correlation between an ability of the tissue to controllipid peroxidation and its ability to retain solutes. It issuggested that ability of plant tissues to mobilize enzymaticdefence against uncontrolled lipid peroxidation may be an importantfacet of their drought tolerance.     

Leaf senescence and lipid peroxidation: Effects of some phytohormones, and scavengers of free radicals and singlet oxygen

During in vitro senescence (chlorophyll loss) of oat ( Avena sativa L. cv. Victory) leaf segments and of leaf discs of Rumex obtusifolius L, the activity of catalase decreases and lipid peroxidation increases. The activity of superoxide dismutase (SOD) decreases in Rumex leaf discs but changes little in oat leaf segments. Kinetin treatment of oat leaf segments, and GA₃ treatment of Rumex leaf discs, inhibit decline in the enzyme activities and increase in the level of lipid peroxidation and strongly inhibit senescence. In either leaf tissue a treatment with ethanol or vitamin E (scavengers of free radicals) or with diphenylisobenzofuran (scavenger of singlet oxygen) results in a strong inhibition of lipid peroxidation and senescence, but does not affect much the decline in the SOD and catalase activities. It is concluded that, i) senscence-associated lipid peroxidation is induced by free radicals and singlet oxygen; and, ii) kinetin and GA₃ inhibit senescence mainly by a modulation of lipid peroxidation through maintaining high levels of such cellular scavengers as SOD and catalase.     

The Effect of Light and Hormones on Leaf Senescence

With wheat leaves as material, the changes of superoxide dismutase (SOD), lipid peroxi-dation and membrane permeability during leaf senescence in light or dark, and treated withphytohormones (KT or ABA) have been studied. The changes of chlorophyll content, lipidperoxidation and fine structure of spinach chloroplasts senescing in light or dark have alsobeen studied. When leaves senesce in light, the activity of SOD increased at first then decreased. The increase of SOD activity was able to result from the synthesis of new protein. Lightwas found to delay the leaf senescence obviously but also accelerate leaf senescence by causinglipid peroxidation when prolonged the illumination time. The delay or acceleration of leafsenescence by exogenous hormones were observed, it may be due to the control of lipid peroxi-dation by adjusting the activity of SOD. O2-participated the chlorophyll decomposition andlipid peroxidation during chloroplasts senesce in light. A favourable role of light in mainta-lng the fine structure of isolated chloroplasts was clear.     

杂交稻及其三系叶片衰老过程中SOD、CAT活性和MDA含量的变化

对杂交水稻及其三系主茎第11叶叶片自然衰老过程中超氧物歧化酶(SOD)、过氧化氢酶(CAT)和丙二醛(MDA)含量的变化进行了研究,结果表明:叶片衰老过程中,SOD和CAT活性下降,MDA的含量增加,可作为衰老特征的叶绿素和可溶性蛋白质含量明显下降;SOD的活性和MDA的含量变化相对应;CAT活性大幅度下降与SOD之间的不平衡,致使O_2~-代谢中间产物累积而引起膜的损伤。不育系的衰老进程比杂交水稻、恢复系和保持系慢,其SOD和CAT活性明显高于其它三者,可能是不育系不易早衰的原因之一。     

Superoxide Dismutase Activity and Lipid Peroxidation in Relation to Senescence of Rica Leaves

Superoxide dismutase (SOD), lipid peroxidation, ribulose 1, 5-bisphosphate earboxylasc and chlorophyll content in the leaves of rice were investigated during bloomJug to ripening stages. The results indicated that leaf senescence was associated with the decreascs of SOD and RuBP earboxylase activities and chlorophyll content. A marked increase of malondialdehyde(MDA)contcnt, a produet of lipid pcroxidation was observed during leaf senescence, However, the relative high activities of SOD and RuBP earboxylase and chlorophyll content, lower MDA content in hybrid rice senescenee leaves, compared with their three lines, suggest the presence of physiological hybrid vigor. The analysis of polyacrylamide gel eleetrophoresis showed that three SOD isozymes in leaf extract, which were Cu-Zn SOD. Changes of SOD isozymes were observed during leaf agling and at different time of storage of its extract. The destroying effect of oxygen free radical on enzymes and membrane in the course of leaf senescence was discussed.     

水稻叶片的衰老与超氧物歧化酶活性及脂质过氧化作用的关系

研究了从抽穗开花到籽粒成熟过程中,水稻植株顶部三片叶子的超氧物歧化酶(SOD),脂质过氧化产物丙二醛(MDA)含量及二磷酸核酮糖羧化酶活性的变化。实验结果表明:叶片的衰老伴随着 SOD 活性、RuBP 羧化酶活性及叶绿素含量的降低、丙二醛含量显著增高。分离了三个 SOD 的同工酶,证明为 Cu—Zn SOD。观察了 SOD 同工酶在叶片老化及酶液存放不同时间中的变化。讨论了叶片衰老过程中氧自由基对酶及质膜的损伤影响。     

化学催熟剂对油菜角果叶绿素含量及抗氧化酶系统的影响

采用大田试验研究了两种化学催熟剂（敌草快和农达）对生长后期油菜角果的叶绿素含量、抗氧化酶系统（CAT、SOD、POD活性）、细胞膜透性及MDA含量的影响.结果表明：采用敌草快催熟,油菜角果皮叶绿素含量下降,SOD、POD、CAT活性及细胞膜透性和MDA含量显著提高,导致角果膜脂过氧化,且作用强度随处理浓度的增加而增加;采用农达催熟,油菜角果皮叶绿素含量所受影响较小,SOD、POD和CAT活性上升缓慢,细胞膜透性和MDA含量增加不明显.随着催熟时间的推移,油菜角果保护酶活性受到不同程度的抑制,这可能与催熟剂干扰酶系统分子结构有关.     

Changes in the Levels of Chlorophyll and Light-Harvesting Chlorophyll a/b Protein of PS II in Rice Leaves Aged under Different Irradiances from Full Expansion through Senescence

Effects of irradiance on changes in the amounts of chlorophyll(Chl) and light-harvesting chlorophyll a/b protein of PS II(LHCII) were examined in senescing leaves of rice (Oryza sativaL.). Results of treatments at two irradiances (100% and 20%natural sunlight) were examined after the full expansion ofthe 13th leaf throughout the course of senescence. With 20%sunlight, the Chl content decreased only a little during leafsenescence, while with 100% sunlight it decreased appreciably.Similarly, the amount of LHCII protein during treatment with20% sunlight remained almost constant. However, the ratio ofChl a/b during the shade treatment decreased significantly andthe rate of decrease was greater than during the full-sunlighttreatment. The ratio of Chl a/b for Chl a and b bound to LHCIIwas about 1.2, irrespective of leaf age or irradiance treatment.When the amounts of Chl bound to LHCII were calculated fromthe total leaf content of Chl and the ratio of Chl a/b, assuminga ratio of Chl a/b bound to LHCII of 1.2, they were well correlatedwith the amounts of LHCII protein. Changes in the amounts of LHCII synthesized during the two irradiancetreatments were examined using an ¹⁵ tracer. Incorporation of¹⁵N into LHCII declined dramatically during both treatmentsfrom full expansion through senescence, suggesting that therewas little synthesis of LHCII protein during that time. In addition,the amount of LHCII synthesized during senescence was lowerduring the shade treatment than during the 100% sunlight treatment.These results indicate that the absence of an apparent changein levels of LHCII with shade treatment during senescence wascaused by the very low rate of turnover of LHCII protein. (Received June 17, 1992; Accepted September 28, 1992)     

Antioxidant Defences under Hyperoxygenic and Hyperosmotic Conditions in Leaves of Two Lines of Maize with Differential Sensitivity to Drought

The effects of hyperoxygenic and hyperosmotic stress on severalaspects of antioxidant defences were studied in the leaves ofa drought-sensitive (LG11) and a drought-tolerant (LIZA) lineof maize (Zea mays L.). When leaf disks were subjected to theseverest stress conditions (100% O₂ and 0.5 M mannitol), theactivities of antioxidant enzymes, such as superoxide dismutase(SOD), catalase (Cat), ascorbate peroxidase (Asc-Px) and glutathionereductase (GSSG-Red), remained higher in disks of LIZA thanin disks of LG11. The ratios of activities of SOD to Cat, SODto Asc-Px and SOD to GSSG-Red were much higher in leaf disksfrom LG11 than in those from LIZA. Damage, as indicated by,for example, the extend of lipid peroxidation, the destructionof chlorophyll and carotenoids, the decrease in levels of proteinsulfhydryl groups and the leakage of electrolytes from cellswas apparent in leaf disks of both LIZA and LG11 as consequenceof the applied stresses. However, the damage was less markedin LIZA than in LG11. (Received September 2, 1992; Accepted July 20, 1993)     

Retardation and Reversal of Senescence in Bean Leaves by Benzyladenine and Decapitation

During the early stages of growth when the primary leaves ofbeans (Phaseolus vulgaris L., cv. Kinghorn) are expanding, thereis an increase in chlorophylls, carotenoids and protein levelsas well as superoxide dismutase (SOD), catalase and peroxidaseactivities. However, following the initiation of flowering onday (d) 21 there is a steady decrease in all of these parametersexcept levels of peroxidase activity, and by d 35 the primaryleaves are also showing pronounced morphological symptoms ofsenescence. Following the onset of senescence, primary leaveswere also leaking and showed increased levels of malondialdehyde(MDA). In addition, there was an increase in the lipid phasetransition temperature of a membrane fraction consisting predominantlyof chloroplast membranes, which reflected the formation of gelphase lipid. By d 56, the leaves had abscinded. A single applicationof benzyladenine (BA) to the primary leaves or decapitationof the shoot above the primary leaf on d 9 retarded all of thesesenescence symptoms, with decapitation being more effectivethan BA. On d 35 when the primary leaves were pale green-yellow, a singleapplication of BA to the leaves or decapitation of the shootabove them reversed the symptoms of senescence. Decapitationwas more effective than BA as a means of reversal, causing chlorophyll,carotenoid and protein levels to rise to values higher thanthose present in the control leaves. Decapitation also reversedthe senescence-related changes in membrane phase properties,decreased membrane leakiness and reduced lipid peroxidation. (Received August 22, 1983; Accepted January 26, 1984)     

萝卜离体子叶衰老与膜脂过氧化的关系

萝卜离体子叶在光下或暗中衰老及激素调节衰老过程中,作为叶片衰老指标的叶绿素和蛋白质含量的降低,发生在MDA含量增高之前,更早于SOD活性的下降。表明由SOD活性降低所导致的膜脂过氧化的增强,并非衰老的原初反应,而是叶片衰老到一定程度的生理变化。因此,至少在萝卜离体子叶上,不能将其衰老的启动归因于受SOD控制的膜脂过氧化作用导致的膜累积性质变。     

骆驼蓬提取物对豌豆幼苗生长的影响

用不同浓度的骆驼蓬提取物处理豌豆种子,明显导致幼苗根系活力及叶绿素和叶片可溶性蛋白质含量下降,膜透性增加。分析表明,骆驼蓬提取物处理下幼苗叶片膜脂过氧化作用增强;过氧化物酶(POD)活性提高;而超氧化物歧化酶(SOD)和过氧化氢酶(CAT)活性降低。     

Cr6+对菹草叶绿素荧光参数、抗氧化系统和超微结构的胁迫影响

研究了不同浓度的Cr⁶⁺(0、1、10、30、50 mg·L^-1)对菹草叶片叶绿素、叶绿素荧光参数、抗氧化系统、可溶性蛋白含量、膜脂过氧化产物、可溶性糖含量、超氧阴离子(O₂^—·)产生以及细胞超微结构的胁迫影响。结果表明,随着Cr⁶⁺浓度的增加,总叶绿素、叶绿素a、叶绿素b和叶绿素a/b均呈下降趋势,F_o先升后降,F_v/F_m、F_m及F_v/F_o均逐渐降低;可溶性蛋白和谷胱甘肽(GSH)含量先升后降,抗坏血酸(AsA)含量逐渐下降;超氧化物歧化酶(SOD)活性逐渐下降,而过氧化物酶(POD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)和谷胱甘肽还原酶(GR)活性均呈先升后降趋势; O₂^—·产生速率、丙二醛(MDA)和可溶性糖含量均表现先降后升趋势;电镜观察发现：随着Cr⁶⁺浓度的增加,对细胞超微结构的损伤也加剧,表现为叶绿体膨胀,被膜破裂,类囊体片层解体;线粒体嵴数目减少,呈空泡状。可见,Cr⁶⁺破坏了菹草正常生理活动的结构基础,造成菹草生理功能紊乱。     

燕麦叶片衰老与活性氧代谢的关系

燕麦连体叶片与高体叶片衰老中,过氧化氢酶和超氧物歧化酶(SOD)活性下降,脂类过氧化产物丙二醛(MDA)迅速积累,组织自动氧化速率显著加快。植物激素BA,GA_3,2,4—D及光、亚胺环己酮(CH),EDTA处理均不同程度地延缓离体叶片的衰老过程,同时抑制过氧化氢酶和SOD活性下降,阻止MDA的积累和组织自动氧化速率的提高.推测叶片衰老中活性氧起着重要的作用。     

NaCl-induced Senescence in Leaves of Rice (Oryza sativa L.) Cultivars Differing in Salinity Resistance

The influence of NaCl on senescence-related parameters (proteinand chlorophyll concentrations, membrane permeability and chlorophyllfluorescence) was investigated in young and old leaves of fiverice cultivars differing in salt resistance. NaCl hastened thenaturally-occurring senescence of rice leaves which normallyappears during leaf ontogeny: it decreased chlorophyll and proteinconcentrations and increased membrane permeability and malondialdehydesynthesis. Such an acceleration of deteriorative processes affectedall leaves in salt-sensitive cultivars while it was more markedin oldest than in youngest leaves of salt-resistant genotypes.NaCl-induced senescence also involved specific modifications,such as an increase in basal non-variable chlorophyll fluorescence(F ₀) recorded in all cultivars or a transient increase in solubleprotein concentration recorded in salt-resistant genotypes only.Alteration of membrane permeability appeared as one of the firstsymptoms of senescence in rice leaves and allowed discriminationamong cultivars after only 7 d of stress. In contrast, F _v/F _mratio (variable fluorescence/maximal fluorescence) was thesame for all cultivars during the first 18 d of stress and thuscould not be used for identifying salt-resistant rice exposedto normal light conditions. Relationships between parametersinvolved in leaf senescence are discussed in relation to salinityresistance of rice cultivars. Chlorophyll concentration; chlorophyll fluorescence; electrolyte leakage; magnesium; malondialdehyde; membrane permeability; NaCl; Oryza sativa L.; protein; rice; salinity resistance; senescence; UV absorbing substances     

The Senescence Process in Oat Leaves and its Regulation by Oxygen Concentration and Light Irradiance

The regulation of senescence by oxygen-concentration, lightirradiance and H₂O₂ has been studied in leaf segments of Avenasativa L. cv. Suregrain. The development of the components of the senescence process,for example chlorophyll breakdown, proteolysis (as soluble aminoacids), hydroperoxides (as malondi-aldehyde) and permeability(as conductivity) is accelerated in light as the O₂-tensionincreases. In darkness, 0.3% O₂ accelerates increases in hydroperoxides,permeability and proteolysis and delays the chlorophyll break-down,but 0.0005% O₂ delays all the components studied. In every casethe hydroperoxide content, permeability and proteolysis areclosely related. Any treatment inducing an increase in membranepermeability causes chlorophyll bleaching (photo-oxidation)if leaf segments are then treated with light in an atmospherecontaining oxygen. Light has a modulating effect on the senescenceprocess. An irradiance lower or higher than 40 W m^–2 hasan accelerating effect on the senescence process. (Received September 7, 1985; Accepted July 30, 1985)     

Roles for redox regulation in leaf senescence of pea plants grown on different sources of nitrogen nutrition

Leaf senescence and associated changes in redox components were monitored in commercial pea (Pisum sativum L. cv. Phoenix) plants grown under different nitrogen regimes for 12 weeks until both nodules and leaves had fully senesced. One group of plants was inoculated with Rhizobium leguminosarum and grown with nutrient solution without nitrogen. A second group was not inoculated and these were grown on complete nutrient solution containing nitrogen. Leaf senescence was evident at 11 weeks in both sets of plants as determined by decreases in leaf chlorophyll and protein. However, a marked decrease in photosynthesis was observed in nodulated plants at 9 weeks. Losses in the leaf ascorbate pool preceded leaf senescence, but leaf glutathione decreased only during the senescence phase. Large decreases in dehydroascorbate reductase and catalase activities were observed after 9 weeks, but the activities of other antioxidant enzymes remained high even at 11 weeks. The extent of lipid peroxidation, the number of protein carbonyl groups and the level of H(2)O(2) in the leaves of both nitrate-fed and nodulated plants were highest at the later stages of senescence. At 12 weeks, the leaves of nodulated plants had more protein carbonyl groups and greater lipid peroxidation than the nitrate-fed controls. These results demonstrate that the leaves of nodulated plants undergo an earlier inhibition of photosynthesis and suffer enhanced oxidation during the senescence phase than those from nitrate-fed plants.     

低温胁迫对麻竹叶片和根系抗性生理指标的影响

采用室内人工低温处理,研究了麻竹(Dendrocalamus latiflorus)叶片及根部质膜透性、丙二醛、可溶性蛋白、可溶性糖含量,超氧化物歧化酶(SOD)、过氧化物酶(POD)活性和膜脂脂肪酸组成的变化,寻找与竹类植物耐寒性关系最密切的抗性生理指标。结果表明:低温预处理(8℃)15d后,麻竹叶片中可溶性糖、可溶性蛋白含量、POD活性显著提高,根部可溶性糖含量、POD活性显著升高;低温胁迫处理(-2℃)72h后,经低温预处理的麻竹叶片可溶性糖、可溶性蛋白质含量、SOD、POD活性显著高于未经低温预处理,而质膜透性显著低于未经低温预处理,但膜脂过氧化程度显著高于-2℃处理前;经低温预处理的麻竹根部SOD、POD活性及膜脂不饱和脂肪酸相对含量显著高于未经低温预处理,而质膜透性、膜脂过氧化程度较-2℃处理前无显著差异。说明剧烈降温对麻竹生理特征造成严重影响,叶片通过提高可溶性蛋白、可溶性糖含量,维持较高的POD活性以减轻低温伤害;根系则通过维持较高的SOD、POD活性以减轻低温下膜脂过氧化水平,并通过提高膜脂不饱和脂肪酸比例降低质膜透性来抵御低温对膜的伤害。     

腐殖酸对小麦抗旱性的生理效应

在干旱气候条件下,喷施腐殖酸钠（ＨＡ）、黄腐酸钠（ＦＡ）可降低土壤水分损耗,提高小麦叶片持水能力,叶片细胞超氧物歧化酶（ＳＯＤ）、过氧化氢酶（ＣＡＴ）活性明显提高,丙二醛（ＭＤＡ）含量和电解质渗出率明显降低,可减缓叶绿素降解,增强光合速率和光合产物积累,延缓植株衰老,小麦抗旱性增强,千粒重增加．     

汞对玉米幼苗膜脂过氧化及体内保护系统的影响

随着处理ＨｇＣｌ２浓度的升高,细胞膜脂质过氧化水平升高,细胞膜透性增大,ＣＡＴ活性降低,ＳＯＤ、ＰＯＤ活性升高,组织可溶性蛋白质含量升高。