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铝是地壳中含量最高的金属元素,约占土壤矿物质总量的7%,仅次于氧元素和硅元素,通常以难溶性的硅酸盐和氧化物的形式存在于长石、云母、高岭石等矿石中,对植物和环境没有毒害作用[1]。酸性条件下(pH<5.5),难溶性的铝会逐渐解离转变为有毒的离子态铝离子(Al3+),Al(OH)2+和Al(OH)2+,酸雨的沉降加上长期施用生理酸性化肥可进一步增进土壤的酸化过程,土壤中活性铝的增加会对植物生长造成极为不利的影响。目前,已经对玉米Zea mays,小麦Triticum aestivum,高粱Sorghum bicolor,马尾松Pinus massoniana,杉木Cunninghamia lanceolata,桉树Eucalyptus,柚木Tectona grandis等农林作物开展了有关铝毒害及耐铝毒机制的研究[2]。铝对植物的危害主要是通过抑制植物根系的生长,进而影响根系对水分和养分的吸收,最终影响植物的生长和发育[3]。不同植物或同种植物不同基因型对铝毒的耐性存在着一定的差异[4]。铝会对植物细胞产生氧化损伤,而这些损伤可以得到抗氧化酶的保护[5]。植物器官在铝胁迫下受到伤害时往往会发生膜脂过氧化作用,丙二醛(MDA)是膜脂过氧化过程的最终分解产物,其含量可以反映植物遭受酸铝伤害的程度。植物体内的过氧化物酶(POD),过氧化氢酶(CAT),超氧化物歧化酶(SOD),多酚氧化酶(PPO),抗坏血酸过氧化物酶(APX)和谷胱甘肽还原酶(GR)等是重要的抗氧化酶,在清除金属等诱发产生的氧自由基和过氧化物、抑制膜脂过氧化、保护细胞免遭伤害等方面起着重要作用。在许多作物抗性机制研究中,这类抗氧化酶活性的变化已广泛作为指示植物抵御逆境伤害的指标[6]。目前,关于铝处理下抗氧化系统的变化多集中在农作物及经济作物方面,而有关林木方面的研究相对较少。桉树具有丰产、优质、适应性强、用途广泛等特点。目前,中国桉树人工林面积已达300万hm2,广西是其主要栽培区之一[7]。桉树人工林生产力的维持与养分供应有很大关系,持续施肥可加剧土壤酸化,使土壤活性铝不断增多[8]。西班牙桉树林土壤中铝含量及桉树根系中铝的积累受土壤酸度影响,且存在潜在的铝毒危险性,赤桉E. camaldulensis,蓝桉E. globulus,巨桉E. grandis,邓恩桉E. dunnii等实生苗在铝胁迫下分泌苹果酸、柠檬酸和草酸[9],低浓度铝离子甚至会促进桉树生长[10]。笔者曾对4个速生桉优良无性系(巨尾桉9号E. grandis × E. urophylla No.9,巨尾桉12号E. grandis × E. urophylla,尾叶桉4号E. urophylla,韦赤桉3号E. wetarensis × E. Camaldulensis No.3等的耐铝性进行了研究,结果为巨尾桉9号具有较强的耐铝性,而尾叶桉4号的耐铝性较差,且耐铝桉树无性系根部铝的积累量较低[11-12]。在此基础上,本研究进一步探讨了不同耐铝型桉树无性系在根、叶器官中对铝的抗性生理响应差异,为阐明桉树耐铝机制提供参考依据。
Al stress with lipid peroxidation and antioxidant enzyme activities in eucalyptus roots and leaves
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摘要: 为了阐明不同桉树无性系对铝的抗逆性生理响应机制,采用室内水培法,以广西区2个桉树无性系耐铝型巨尾桉9号Eucalyptus grandis×E.urophylla No.9(记为G9)和铝敏感型尾叶桉4号E.urophylla No.4(记为G4)为研究对象,在4.4 mmol·L-1铝离子浓度下处理24 h(以pH 4.0,0.5 mmol·L-1氯化钙为对照),从根系及叶片内细胞膜透性(CMP),丙二醛(MDA)质量摩尔浓度及过氧化氢酶(CAT),多酚氧化酶(PPO),超氧化物歧化酶(SOD),抗坏血酸过氧化物酶(APX)和谷胱甘肽还原酶(GR)等抗氧化酶活性方面探讨了铝胁迫下供试苗木的耐铝机制。采用SPSS 21.0软件分别对根和叶中G9和G4的各测定指标进行单因素方差分析和Duncan多重比较。根细胞膜透性、丙二醛质量摩尔浓度均显著(P<0.05)高于叶,铝对2个桉树无性系苗木的毒害主要表现在根部。在4.4 mmol·L-1铝离子处理下,G4根相对电导率与丙二醛质量摩尔浓度最高,分别为48.8%,11.5 μmol·g-1,而G9根内过氧化氢酶、抗坏血酸过氧化物酶、谷胱甘肽还原酶活性极显著(P<0.01)大于G4,且G9根过氧化氢酶活性比相应对照增加145.0%,G4根过氧化氢酶活性比相应对照仅增加43.0%,过氧化氢酶在G9根中对铝毒害的缓解起到了更为重要的作用。G9清除活性氧能力较G4强,表明耐铝型桉树对铝毒害具有较好的适应能力。图7参28Abstract: Eucaplytus is the main timber tree species in south China with enrichment of aluminum (Al) in soil, but the physiological mechanisms of Al tolerance in eucalyptus trees is not well understood. To clarify the physiological response mechanism of Al resistance of eucalyptus, seedlings of the two eucalyptus genotypes (Eucalyptus grandis×Eucalyptus urophylla No.9, Al-resistant type, designated G9; E. urophylla No.4, Al-sensitive type, designated G4) were grown for 24 hours in 0.5 mmol·L-1 CaCl2 solutions (pH 4.0)containing 0 and 4.4 mmol·L-1 Al respectively. The indexes of plant stress resistance were measured by cell membrane permeability (CMP), malondialdehyde content (MDA), catalase activities (CAT), polyphenol oxidase (PPO), superoxide dismutase (SOD), ascorbate peroxidase (APX), and glutathione reductase (GR) in the roots and leaves. The significance of data was analyzed with one-way Anova and Duncan multiple comparison by SPSS 21.0 system. The contents of CMP and MDA in the roots were significant higher (P<0.05) than those in the leaves indicating that Al toxicity mainly happened in the roots of the two eucalyptus genotypes. With 4.4 mmol·L-1 of Al stress in the roots of G4, the highest relative electrical conductivity was 48.8% and MDA content was 11.5 μmol·g-1. CAT, APX, and GR activities in roots of G9 were extremely significant higher (P<0.01) than G4. CAT play an important role in the detoxification of reactive oxygen species in Al-tolerant eucalyptus clone, because it rose by 145% in the roots of G9 in response to Al treatment, only rose by 43% in G4. Conclusively, the Al-resistant eucalyptus genotype G9 was adapted to Al toxicity with active physiological characteristics to remove reactive oxygenresistant.[Ch, 7 fig. 28 ref.]
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https://zlxb.zafu.edu.cn/article/doi/10.11833/j.issn.2095-0756.2016.06.012