生防芽孢杆菌对作物耐盐性的影响及其抗盐机理的研究
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摘要
芽孢杆菌(Bacillus spp.)是土壤和植物微生态的优势微生物种群之一,具有很强的抗逆能力和抑菌防病增产作用,且其中的许多菌株能够产生植酸酶。目前已发现,在磷限量条件下施加微生物来源的植酸酶能够促进植物生长,但关于土壤生防微生物胞外植酸酶对植物耐盐性的影响尚未见报道。
本文从作物根际土壤及本实验室已有的生防细菌中筛选出高产植酸酶且对小麦纹枯病菌及其它多种植物病原真菌具有拮抗作用的生防芽孢杆菌,并对其产生的胞外植酸酶在增强植物耐盐性中的作用进行了研究,具体结果如下:
1.通过盆栽试验研究了在土壤中直接施用高产植酸酶的广谱拮抗生防芽孢杆菌(Bacillus subtilis)T2对作物耐盐性的影响。盆栽试验结果显示:无盐胁迫及180 mmol·L~(-1)、360 mmol·L~(-1)、540 mmol·L~(-1) NaCl胁迫条件下,土壤施加产胞外植酸酶芽孢杆菌菌悬液可促进小麦幼苗的生长,小麦株高、鲜重、可溶性糖含量、脯氨酸含量、过氧化物酶(POD)活性和过氧化氢酶(CAT)活性均得以不同程度的增高;无盐胁迫及180 mmol·L~(-1) NaCl胁迫条件下小麦叶片叶绿素含量增高;180 mmol·L~(-1)、360 mmol·L~(-1)、540 mmol·L~(-1) NaCl胁迫下植株超氧化物歧化酶(SOD)活性升高,MDA含量降低。表明土壤中施加具有胞外植酸酶活力的生防芽孢杆菌可增强作物对一定浓度范围内盐胁迫的耐受能力。
2.为进一步明确芽孢杆菌胞外植酸酶在其增强作物耐盐性过程中的作用和地位,本研究进行了胞外植酸酶突变菌株的诱变选育。以T2为出发菌株,经微波(Microwave)和硫酸二乙酯(DES)诱变,得到遗传特性稳定的植酸酶正负突变株。酶活力测定结果显示:与出发菌株相比,培养72 h的正突变株M15胞外植酸酶活力提高31.7%,负突变株M3植酸酶活力降低93.3%。
3.利用水培试验研究了出发菌株与负突变株对作物耐盐性的不同影响。采用有效磷限量、含NaCl 150 mmol L~(-1)的植物生长水培液对小麦进行盐胁迫,在植酸存在的条件下,水培液中施加T2发酵液或植酸酶均可促进盐胁迫下小麦幼苗的生长,小麦株高、鲜重、叶片叶绿素含量和根系活力分别提高7.7%、2.5%,39.4%、19.3%,12.2%、2.1%和178.3%、173.9%。而植株丙二醛(MDA)含量分别降低85.1%和70.2%。失去植酸酶活力的负突变株M3发酵液则不具有提高小麦株高鲜重和叶绿素含量的作用,对根系活力的提高和对MDA含量的降低程度也均低于T2发酵液处理。上述结果表明生防芽孢杆菌胞外植酸酶活力在其增强小麦耐盐性过程中起了重要作用。
4.将突变株在平板上连续传10代,考察其遗传稳定性。结果显示:突变株产酶能力无明显变化,表明获得的突变株具有良好的遗传稳定性。这对于该菌株在各种生态环境中长期稳定地发挥作用具有重要意义,并为将来高效菌肥的进一步开发奠定了基础。
Bacillus species, which are both resistant to adverse environment and usually against bacterial and fungal pathogens, are among the dominant microorganisms in the soil and plant microecological systems. Bacillus spp. is one of the important sources of phytase applied in industry. However, little is known about the role of phytase in the alleviation of the effect of salt stress to plant produced by Bacillus strains.
Therefore, in order to evaluate whether the phytase secreted by Bacillus spp. can enhance the saline tolerance of plant, Bacillus strains, which were potential biocontrol agent against a broad spectrum of plant pathogenic fungi with the ability to produce extracellular phytase, were isolated. The objectives of this study were:
(1) to reveal whether Bacillus strains can enhance the saline tolerance of plant in edaphic culturing experiment,
(2) to obtain phytase mutants by means of DES and Microwave treatments,
(3) to further investigate the activity of increasing resistance of wheat to salt stress of mutants and the wild type,
(4) to identify the genetic stability of phytase mutants.
The main conclusions of this paper are as follows:
1. To reveal whether Bacillus strains T2 with phytase activity can alleviate the salt stress effect of NaCl to wheat seedlings in soil, physiological and biochemical activities of wheat under NaCl stress were studied by edaphic culture. The results showed that Bacillus subtilis had positive effect on wheat growth under free salt stress and salt stress. Compared to free Bacillus subtilis treatment, Bacillus subtilis accelerated the increase of plant height, plant fresh weight, total leaf chlorophyll content, soluble sugar content, praline content and enhanced the activities of protective enzymes, such as SOD, POD and CAT, but decreased the content of MDA under free NaCl stress, 180 mmol·L~(-1) NaCl, 360 mmol·L~(-1) NaCl and 540 mmol·L~(-1) NaCl stress. Significant increases in total leaf chlorophyll content were observed under free NaCl stress and 180 mmol·L~(-1) NaCl stress. Simultaneously, increase in SOD activities and decrease in content of MDA were observed under 180 mmol·L~(-1) NaCl, 360 mmol·L~(-1) NaCl and 540 mmol·L~(-1) NaCl stress. These may provide strong evidence that B. subtilis with phytase activity is effective for the increasing resistance of wheat to salt stress in soil.
2. Strain Bacillus subtilis T2 was a potential biocontrol agent against a broad-spectrum plant pathogenic fungi with the ability to produce extracellular phytase.To improve its saline stress alleviating capacity, mutagenetic program was undertaken for the construction of extracellular phytase derivates. Two phytase mutants with genetic stability were obtained by means of DES and Microwave treatments. It was revealed that extracellular phytase activity of the positive mutant M15 was elevated by 31.7% and the negative mutant M3 decreased by 93.3% after 72 h cultivation compared to the wild type T2.
3. To further evaluate whether the phytase secreted by T2 can enhance the saline tolerance of plant, hydroponic culturing experiment was undertaken. A phytase minus mutant M3 with genetic stability was obtained by means of DES treatment. The results of biochemical experiments revealed that the culture filtrates of wild-type strain T2 with phytase activity and purified phytase stimulated growth of wheat seedlings under phosphate limitation in the presence of 150 mmol·L~(-1) NaCl and phytate. Significant increases in plant height, fresh weight, total leaf chlorophyll content and TTC reducing power and decrease in content of MDA were observed in response to addition of phytase and culture filtrates of T2 to culture liquids. However, culture filtrates obtained from phytase-negative mutant strain M3 did not stimulate wheat seedlings growth under salt stress. These may provide strong evidence that phytase activity of B. subtilis is important for the increasing resistance of wheat to salt stress under phosphate limitation.
4. It is important for the mutants with genetic stability to function normally in various environments. Two phytase mutants were taken to inherit from generation to generation. The result showed that the capacity of enzyme production and the salt stress alleviation effect did not change significantly compared to the wild type. So, they can be applied into practice stably.
本文从作物根际土壤及本实验室已有的生防细菌中筛选出高产植酸酶且对小麦纹枯病菌及其它多种植物病原真菌具有拮抗作用的生防芽孢杆菌,并对其产生的胞外植酸酶在增强植物耐盐性中的作用进行了研究,具体结果如下:
1.通过盆栽试验研究了在土壤中直接施用高产植酸酶的广谱拮抗生防芽孢杆菌(Bacillus subtilis)T2对作物耐盐性的影响。盆栽试验结果显示:无盐胁迫及180 mmol·L~(-1)、360 mmol·L~(-1)、540 mmol·L~(-1) NaCl胁迫条件下,土壤施加产胞外植酸酶芽孢杆菌菌悬液可促进小麦幼苗的生长,小麦株高、鲜重、可溶性糖含量、脯氨酸含量、过氧化物酶(POD)活性和过氧化氢酶(CAT)活性均得以不同程度的增高;无盐胁迫及180 mmol·L~(-1) NaCl胁迫条件下小麦叶片叶绿素含量增高;180 mmol·L~(-1)、360 mmol·L~(-1)、540 mmol·L~(-1) NaCl胁迫下植株超氧化物歧化酶(SOD)活性升高,MDA含量降低。表明土壤中施加具有胞外植酸酶活力的生防芽孢杆菌可增强作物对一定浓度范围内盐胁迫的耐受能力。
2.为进一步明确芽孢杆菌胞外植酸酶在其增强作物耐盐性过程中的作用和地位,本研究进行了胞外植酸酶突变菌株的诱变选育。以T2为出发菌株,经微波(Microwave)和硫酸二乙酯(DES)诱变,得到遗传特性稳定的植酸酶正负突变株。酶活力测定结果显示:与出发菌株相比,培养72 h的正突变株M15胞外植酸酶活力提高31.7%,负突变株M3植酸酶活力降低93.3%。
3.利用水培试验研究了出发菌株与负突变株对作物耐盐性的不同影响。采用有效磷限量、含NaCl 150 mmol L~(-1)的植物生长水培液对小麦进行盐胁迫,在植酸存在的条件下,水培液中施加T2发酵液或植酸酶均可促进盐胁迫下小麦幼苗的生长,小麦株高、鲜重、叶片叶绿素含量和根系活力分别提高7.7%、2.5%,39.4%、19.3%,12.2%、2.1%和178.3%、173.9%。而植株丙二醛(MDA)含量分别降低85.1%和70.2%。失去植酸酶活力的负突变株M3发酵液则不具有提高小麦株高鲜重和叶绿素含量的作用,对根系活力的提高和对MDA含量的降低程度也均低于T2发酵液处理。上述结果表明生防芽孢杆菌胞外植酸酶活力在其增强小麦耐盐性过程中起了重要作用。
4.将突变株在平板上连续传10代,考察其遗传稳定性。结果显示:突变株产酶能力无明显变化,表明获得的突变株具有良好的遗传稳定性。这对于该菌株在各种生态环境中长期稳定地发挥作用具有重要意义,并为将来高效菌肥的进一步开发奠定了基础。
Bacillus species, which are both resistant to adverse environment and usually against bacterial and fungal pathogens, are among the dominant microorganisms in the soil and plant microecological systems. Bacillus spp. is one of the important sources of phytase applied in industry. However, little is known about the role of phytase in the alleviation of the effect of salt stress to plant produced by Bacillus strains.
Therefore, in order to evaluate whether the phytase secreted by Bacillus spp. can enhance the saline tolerance of plant, Bacillus strains, which were potential biocontrol agent against a broad spectrum of plant pathogenic fungi with the ability to produce extracellular phytase, were isolated. The objectives of this study were:
(1) to reveal whether Bacillus strains can enhance the saline tolerance of plant in edaphic culturing experiment,
(2) to obtain phytase mutants by means of DES and Microwave treatments,
(3) to further investigate the activity of increasing resistance of wheat to salt stress of mutants and the wild type,
(4) to identify the genetic stability of phytase mutants.
The main conclusions of this paper are as follows:
1. To reveal whether Bacillus strains T2 with phytase activity can alleviate the salt stress effect of NaCl to wheat seedlings in soil, physiological and biochemical activities of wheat under NaCl stress were studied by edaphic culture. The results showed that Bacillus subtilis had positive effect on wheat growth under free salt stress and salt stress. Compared to free Bacillus subtilis treatment, Bacillus subtilis accelerated the increase of plant height, plant fresh weight, total leaf chlorophyll content, soluble sugar content, praline content and enhanced the activities of protective enzymes, such as SOD, POD and CAT, but decreased the content of MDA under free NaCl stress, 180 mmol·L~(-1) NaCl, 360 mmol·L~(-1) NaCl and 540 mmol·L~(-1) NaCl stress. Significant increases in total leaf chlorophyll content were observed under free NaCl stress and 180 mmol·L~(-1) NaCl stress. Simultaneously, increase in SOD activities and decrease in content of MDA were observed under 180 mmol·L~(-1) NaCl, 360 mmol·L~(-1) NaCl and 540 mmol·L~(-1) NaCl stress. These may provide strong evidence that B. subtilis with phytase activity is effective for the increasing resistance of wheat to salt stress in soil.
2. Strain Bacillus subtilis T2 was a potential biocontrol agent against a broad-spectrum plant pathogenic fungi with the ability to produce extracellular phytase.To improve its saline stress alleviating capacity, mutagenetic program was undertaken for the construction of extracellular phytase derivates. Two phytase mutants with genetic stability were obtained by means of DES and Microwave treatments. It was revealed that extracellular phytase activity of the positive mutant M15 was elevated by 31.7% and the negative mutant M3 decreased by 93.3% after 72 h cultivation compared to the wild type T2.
3. To further evaluate whether the phytase secreted by T2 can enhance the saline tolerance of plant, hydroponic culturing experiment was undertaken. A phytase minus mutant M3 with genetic stability was obtained by means of DES treatment. The results of biochemical experiments revealed that the culture filtrates of wild-type strain T2 with phytase activity and purified phytase stimulated growth of wheat seedlings under phosphate limitation in the presence of 150 mmol·L~(-1) NaCl and phytate. Significant increases in plant height, fresh weight, total leaf chlorophyll content and TTC reducing power and decrease in content of MDA were observed in response to addition of phytase and culture filtrates of T2 to culture liquids. However, culture filtrates obtained from phytase-negative mutant strain M3 did not stimulate wheat seedlings growth under salt stress. These may provide strong evidence that phytase activity of B. subtilis is important for the increasing resistance of wheat to salt stress under phosphate limitation.
4. It is important for the mutants with genetic stability to function normally in various environments. Two phytase mutants were taken to inherit from generation to generation. The result showed that the capacity of enzyme production and the salt stress alleviation effect did not change significantly compared to the wild type. So, they can be applied into practice stably.
引文
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