溶磷微生物分离、应用及其相关基因的克隆与功能鉴定
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摘要
从适应土壤定殖的溶磷微生物中筛选到一株适应生产的溶磷真菌,并进行了其土壤溶磷特性和溶磷机理初步研究,选取溶磷效果较好的菌株进行了溶磷相关基因的克隆、表达和功能鉴定,取得以下主要结果:
在磷酸三钙为唯一磷源的无机盐培养基上,以透明圈为指示,通过初筛和复筛,获得效果稳定的细菌23株,分属6个属11个种;真菌16株,其中有9株草酸青霉(Penicillium oxalicum),2株变幻青霉(Penicillium variabile),1株刺孢青霉(Penicillium aculeatum)、1株绿色木霉(Trichoderma viride)、2株黑曲霉(Aspergillus niger)、1株构巢曲霉(Aspergillus nidulans)。
菌株对不同难溶磷(磷酸三钙、磷酸锌、磷酸铝和磷酸铁)的溶解能力实验表明:细菌对不同种类难溶磷溶解能力差异显著,真菌的溶磷效果显著高于细菌,16株真菌显著降低PDB培养液的pH值,最低达到1.43,固体培养基上培养5 d能将各种难溶磷全部溶解。
通过16株真菌在不同土壤中的定殖能力和释放难溶磷的效果比较,筛选土壤中定殖和溶磷效果好的菌株。土培实验结果表明:真菌在土壤中具有较强的定殖能力;不同土壤中真菌的定殖能力差异显著,同一种的真菌在土壤中的定殖能力也差异显著。真菌释放土壤难溶磷的能力与其土壤定殖的能力显著相关,菌株定殖能力越强,释放土壤难溶磷效果越好。其中,真菌Z33、Zh、ZC、Zh3、ZI1、Z15、ZQ3和Z30在土壤中定殖效果较好,28 d时,菌株数量能达到起始值的10~48倍,有效磷含量增加44%~160%。
通过微生物与作物、土壤的匹配性实验,筛选适应玉米生产的溶磷真菌,并开展了溶磷菌对土壤微生物多样性影响分析。溶磷微生物与作物的匹配性实验表明:16株真菌在平板上对玉米根系分泌物的适应性差异显著, Z15+、Zh、ZQ3、ZI1、ZJ1和Z30能够利用玉米的根系分泌物快速生长,并且能溶解平板中的磷酸三钙。选取这6株真菌进行土培实验,结果也证明Z15~+、ZQ3、ZI1、Zh和Z30都能以玉米根系分泌物为营养进行繁殖,第21天时,菌株数量达到起始的9.70~48.03倍,土壤有效磷增加了62.08%~147.55%。在盆栽玉米的土壤中,ZI1和Zh定殖的速度最快,菌剂ZI1和Zh处理后,玉米干重和湿重显著增加,在大田试验中,菌株Zh连续2年都具有较高的增产作用,而菌株ZI1只在2010年的试验中具有较好的增产作用。因此本文获得适应大田玉米生产的溶磷真菌为构巢曲霉Zh。溶磷菌剂对土壤微生物的结构产生显著的影响,总体趋势为细菌、真菌、放线菌的数量比对照有所增加,土壤中微生物多样性变化与土壤中细菌数量变化成正相关,土壤中细菌数量越多,土壤微生物多样性越高。
选取具有较好增产效果的构巢曲霉Zh和草酸青霉ZI1,对其产酸的种类,不同温度条件下土壤溶液pH值的变化、菌株定殖能力以及土壤中不同级别难溶磷的转化规律进行研究,探讨了溶磷真菌在土壤中溶磷机理和溶磷特性。利用离子色谱对菌株产酸种类进行分析,结果表明:构巢曲霉Zh和草酸青霉ZI1在PDB溶液中能够大量产生乳酸和草酸,培养26 h时,溶液中乳酸和草酸的含量分别达到了372.5μg/mL和593.9μg/mL。菌株Zh和ZI1在15、20和30℃土培条件下,都有很强的定殖能力,并能使土壤溶液的pH值显著降低,土壤溶液pH值变化与菌株在土壤中定殖的规律一致。不同的温度下,菌株Zh和ZI1对土壤中不同级别的难溶磷的转化规律总体趋势一致,土壤中Ca_2-P、Ca_8-P、Al-P、Fe-P和Ca_(10)-P形态的无机磷含量都发生了变化,菌株Zh转化土壤中Ca_8-P量较高,而菌株ZI1对Al-P、Fe-P转化效率较高。菌株Zh和ZI1溶磷机理为菌株在土壤繁殖过程中分泌有机酸与铁、铝、钙等离子螯合,从而使难溶磷转化为有效磷。
构建黑曲霉Zh3和草酸青霉ZI1的cDNA文库,在磷酸三钙为唯一磷源的无机盐培养基上以透明圈为指示,筛选溶磷克隆子,结果从菌株Zh3和ZI1的cDNA文库中分别筛选到携带溶磷相关基因的克隆子60个和72个,这些克隆子在第二代转接后功能稳定,确认这些克隆子所携带的基因具有溶磷的功能。黑曲霉Zh3文库中筛选的克隆子所携带基因序列全长为1407bp,与Aspergillus niger CBS 513.88和Aspergillus niger contig An07c0220的同源性达到98%,与其他菌的同源性低于80%,但目前还没报到该基因在溶磷功能方面的研究,将该基因命名为pag T。草酸青霉ZI1文库中筛选的克隆子所携带基因序列全长为1160bp,与Penicillium chrysogenum Wisconsin 54-1255和Aspergillus niger CBS 513.88的同源性只有68%,与其他菌的同源性更低,表明该基因为一个新的基因,将该基因命名为ppg T。
分析溶磷相关基因ppq T和paq T的开放阅读框,构建重组表达载体,获得转化子,并分析转化子产生有机酸种类和溶液pH值的变化。草酸青霉ZI1中溶磷相关基因ppg T开放阅读框长702 bp,蛋白长233个氨基酸,该蛋白与Penicillium chrysogenum Wisconsin 54-1255的Pc21g23580同源性最高,为59%,目前还没有该类蛋白的功能研究报道,确定本文获得的蛋白PPG-T为新的蛋白。黑曲霉Zh3中溶磷相关基因pag T开放阅读框长819 bp,蛋白长333个氨基酸,该蛋白与Aspergillus niger CBS 513.88的bax Inhibitor family protein同源性达到100%,但该蛋白还没有溶磷功能的报道,属于该蛋白的新功能。将ppq T和paq T的开放阅读框连接到pET28和pGEX表达载体中,构建了4株转化子,产生的乙酸和甲酸将难溶磷无机盐培养液的pH值降低到3.6左右,在LB培养中,4株转化子不能将溶液的pH值降低,表明基因的表达需要难溶磷的诱导。ppq T基因在E. coli. DH5α中的溶磷机理是将葡萄糖氧化生成乙酸,溶解无机磷;而paq T基因在E. coli. DH5α中的溶磷机理是将葡萄糖氧化生成乙酸和甲酸,溶解无机磷。
Phosphate solubilizing microorganisms which can be suitable for colonization were isolated from soils. And One isolated wae screened, which can be suitable for corn production. Its colonization and phosphate-dissolving characteristics in soil were conducted. Cloning and identification of P-dissolving gene were studied in this article. The main conclusions were as follows:
Twenty-three bacteria strains with high performance of phosphate dissolving were screened. Sixteen fungi were isolated, including 9 strains of Penicillium oxalicum, 2 strain of Aspergillus niger, 2 strains of Penicillium variabile, 1 strain of Penicillium aculeatum, 1 strain of Aspergillus nidulans ,1 strain of Trichoderma viride.
The P-dissolving capbility of strains on different insoluble phosphorus(Ca_3(PO_4)_2, Zn_3(PO_4)_2, FePO_4, AlPO_4) were conducted. The P-dissolving ability of fungi were better than bacteria. 16 strains reduced the pH value from 7.0 to lower 2.0 in 18 h.They could dissolve all phosphate in solid culture.
Colonization and P-dissolving characteristics in soil results showed fungi presented high colonization ability in soil. However, colonization ability of fungi in different soils was significant differences. The P-dissolving ability of isolated were significantly correlated with their colonization. 6 strains of Z33、Zh、ZC、Zh3、ZI1 and Z30 presented high colonization ability in soil. The number of 6 strains increased 9.83-48 times than initial number in 28 days and the content of available phosphorous was increased 44%~160% than initial content.
The plate experiment and soil experiment of the matching between strains and crop in laboratory indicated that the 6 strains of Z15+、Zh、ZQ3、ZI1、ZJ1 and Z30 could propagate in plates and soil useing the root secretion of corn as sole carbon source and dissolve insoluble phosphorus. Six strains of Z33、Zh、ZC、Zh3、ZI1 and Z30 presented high colonization ability in soil, the number of 6 strains increased 9.60-48.03 times and the content of available phosphorous was increased 62.08%-147.55% in 21 days than initial content. The pot experiment inoculating with 5 strains was conducted in greenhouse by planting corn. The result demonstrated that the content of available phosphorous was increased remarkably with strains ZI1 and Zh treatment. The dry matter yield of corn significantly increased than CK, Then strains ZI1 and Zh were used in field experiment as microbial inoculums. The yield of corn treated with strain Zh increased remarkably in two years. However, the yield of corn treated with strain ZI1 presented yield increased in one year. The application of P-solubilizing microorganisms had a significant impact on microbial community structure . The numbers of bacteria, fungi, actinomycetes were increased than CK. There were positive correlation between the biodiversity and the number of bacteria.
Aspergillus nidulans Zh and Penicillium oxalicum ZI1 secrete lactic acid and acetic acid, the content of acid at 49th day was 372.5μg/mLand 593.9μg/mL , respectively. The strains ZI1 and Zh could colonizate in soil at 15、20、30℃and remarkably decreased the pH of soil. There were negative correlation between pH value decreased of soil and the number of fungi. The strain ZI1 and Zh promoted the transformation of insoluble phosphate which was the same law in different temperature. The content of Ca_2-P、Ca_8-P、Al-P、Fe-P and Ca_(10)-P were changed with ZI1 and Zh treatment. The transformation efficiency of Ca8-P of the strain ZI1was higher, but transformation efficiency of Al-P and Fe-P of the strain Zh was higher in soil. The mechanism of P-dissolving of the strain ZI1 and Zh was that they secrete organic acids in the progress of propagation chelating Ca~(2+), Fe~(3+), Al~(3+) for dissolving phosphate.
Sixty clone were screened from cDNA library of Aspergillus niger Zh3 according to the transparent zone of P-dissolving in plate. And 72 clone were screened from cDNA library of Penicillium oxalicum ZI1. The function of these clone were stable at the second generation. The gene from A.niger Zh3 was named pag T, sequence analysis showed that it was 1407bp in length, and shared 98% identity at the nucleotide level to Aspergillus niger contig An07c0220, and shared less than 80% identity to other isolated. The gene from Penicillium oxalicum ZI1 was named ppg T, Sequence analysis showed that it was 1160bp in length, and shared only 68% identity at the nucleotide level to Penicillium chrysogenum Wisconsin 54-1255, it showed ppg T was a novel gene.
Sequence analysis showed that the reading box of gene ppg T has 702bp, and its deduced 233 amino acid, this protein shares 59% homology with the Pc21g23580 in Penicillium chrysogenum Wisconsin 54-1255—the highest homology. And there was no report about the P-dissolving function of this protein, so the function of this protein PPG– T is a novel one. Sequence analysis showed that the reading box of gene pag T in Zh3 has 819bp, and its deduced 333 amino acids, this protein shared 100% homology with the bax Inhibitor family protein in Aspergillus niger CBS 513.88, but there is no report about the P-dissolving function of this protein. So it’s a novel function of the protein.Connecting ppqT and paqT to expression vectors pET28 and pGEX ,we constructed four engineering strains, which mainly produce acetic acid and formic acid.They could reduce the pH of the inorganic undissolved phosphorus medium to 3.6 or so .The 4 engineering strains was inducted by undissolved phosphorus, because they couldn’t reduce the pH value of medium without undissolved phosphorus. The P-dissolving mechanism of ppq T in E. coli. DH5αis the oxidation of glucose into acetic acid to dissolve phosphorus ; The phosphorus-dissolving mechanism of ppq T in E. coli. DH5αis the oxidation of glucose into acetic acid and formic acid to dissolve phosphorus.
在磷酸三钙为唯一磷源的无机盐培养基上,以透明圈为指示,通过初筛和复筛,获得效果稳定的细菌23株,分属6个属11个种;真菌16株,其中有9株草酸青霉(Penicillium oxalicum),2株变幻青霉(Penicillium variabile),1株刺孢青霉(Penicillium aculeatum)、1株绿色木霉(Trichoderma viride)、2株黑曲霉(Aspergillus niger)、1株构巢曲霉(Aspergillus nidulans)。
菌株对不同难溶磷(磷酸三钙、磷酸锌、磷酸铝和磷酸铁)的溶解能力实验表明:细菌对不同种类难溶磷溶解能力差异显著,真菌的溶磷效果显著高于细菌,16株真菌显著降低PDB培养液的pH值,最低达到1.43,固体培养基上培养5 d能将各种难溶磷全部溶解。
通过16株真菌在不同土壤中的定殖能力和释放难溶磷的效果比较,筛选土壤中定殖和溶磷效果好的菌株。土培实验结果表明:真菌在土壤中具有较强的定殖能力;不同土壤中真菌的定殖能力差异显著,同一种的真菌在土壤中的定殖能力也差异显著。真菌释放土壤难溶磷的能力与其土壤定殖的能力显著相关,菌株定殖能力越强,释放土壤难溶磷效果越好。其中,真菌Z33、Zh、ZC、Zh3、ZI1、Z15、ZQ3和Z30在土壤中定殖效果较好,28 d时,菌株数量能达到起始值的10~48倍,有效磷含量增加44%~160%。
通过微生物与作物、土壤的匹配性实验,筛选适应玉米生产的溶磷真菌,并开展了溶磷菌对土壤微生物多样性影响分析。溶磷微生物与作物的匹配性实验表明:16株真菌在平板上对玉米根系分泌物的适应性差异显著, Z15+、Zh、ZQ3、ZI1、ZJ1和Z30能够利用玉米的根系分泌物快速生长,并且能溶解平板中的磷酸三钙。选取这6株真菌进行土培实验,结果也证明Z15~+、ZQ3、ZI1、Zh和Z30都能以玉米根系分泌物为营养进行繁殖,第21天时,菌株数量达到起始的9.70~48.03倍,土壤有效磷增加了62.08%~147.55%。在盆栽玉米的土壤中,ZI1和Zh定殖的速度最快,菌剂ZI1和Zh处理后,玉米干重和湿重显著增加,在大田试验中,菌株Zh连续2年都具有较高的增产作用,而菌株ZI1只在2010年的试验中具有较好的增产作用。因此本文获得适应大田玉米生产的溶磷真菌为构巢曲霉Zh。溶磷菌剂对土壤微生物的结构产生显著的影响,总体趋势为细菌、真菌、放线菌的数量比对照有所增加,土壤中微生物多样性变化与土壤中细菌数量变化成正相关,土壤中细菌数量越多,土壤微生物多样性越高。
选取具有较好增产效果的构巢曲霉Zh和草酸青霉ZI1,对其产酸的种类,不同温度条件下土壤溶液pH值的变化、菌株定殖能力以及土壤中不同级别难溶磷的转化规律进行研究,探讨了溶磷真菌在土壤中溶磷机理和溶磷特性。利用离子色谱对菌株产酸种类进行分析,结果表明:构巢曲霉Zh和草酸青霉ZI1在PDB溶液中能够大量产生乳酸和草酸,培养26 h时,溶液中乳酸和草酸的含量分别达到了372.5μg/mL和593.9μg/mL。菌株Zh和ZI1在15、20和30℃土培条件下,都有很强的定殖能力,并能使土壤溶液的pH值显著降低,土壤溶液pH值变化与菌株在土壤中定殖的规律一致。不同的温度下,菌株Zh和ZI1对土壤中不同级别的难溶磷的转化规律总体趋势一致,土壤中Ca_2-P、Ca_8-P、Al-P、Fe-P和Ca_(10)-P形态的无机磷含量都发生了变化,菌株Zh转化土壤中Ca_8-P量较高,而菌株ZI1对Al-P、Fe-P转化效率较高。菌株Zh和ZI1溶磷机理为菌株在土壤繁殖过程中分泌有机酸与铁、铝、钙等离子螯合,从而使难溶磷转化为有效磷。
构建黑曲霉Zh3和草酸青霉ZI1的cDNA文库,在磷酸三钙为唯一磷源的无机盐培养基上以透明圈为指示,筛选溶磷克隆子,结果从菌株Zh3和ZI1的cDNA文库中分别筛选到携带溶磷相关基因的克隆子60个和72个,这些克隆子在第二代转接后功能稳定,确认这些克隆子所携带的基因具有溶磷的功能。黑曲霉Zh3文库中筛选的克隆子所携带基因序列全长为1407bp,与Aspergillus niger CBS 513.88和Aspergillus niger contig An07c0220的同源性达到98%,与其他菌的同源性低于80%,但目前还没报到该基因在溶磷功能方面的研究,将该基因命名为pag T。草酸青霉ZI1文库中筛选的克隆子所携带基因序列全长为1160bp,与Penicillium chrysogenum Wisconsin 54-1255和Aspergillus niger CBS 513.88的同源性只有68%,与其他菌的同源性更低,表明该基因为一个新的基因,将该基因命名为ppg T。
分析溶磷相关基因ppq T和paq T的开放阅读框,构建重组表达载体,获得转化子,并分析转化子产生有机酸种类和溶液pH值的变化。草酸青霉ZI1中溶磷相关基因ppg T开放阅读框长702 bp,蛋白长233个氨基酸,该蛋白与Penicillium chrysogenum Wisconsin 54-1255的Pc21g23580同源性最高,为59%,目前还没有该类蛋白的功能研究报道,确定本文获得的蛋白PPG-T为新的蛋白。黑曲霉Zh3中溶磷相关基因pag T开放阅读框长819 bp,蛋白长333个氨基酸,该蛋白与Aspergillus niger CBS 513.88的bax Inhibitor family protein同源性达到100%,但该蛋白还没有溶磷功能的报道,属于该蛋白的新功能。将ppq T和paq T的开放阅读框连接到pET28和pGEX表达载体中,构建了4株转化子,产生的乙酸和甲酸将难溶磷无机盐培养液的pH值降低到3.6左右,在LB培养中,4株转化子不能将溶液的pH值降低,表明基因的表达需要难溶磷的诱导。ppq T基因在E. coli. DH5α中的溶磷机理是将葡萄糖氧化生成乙酸,溶解无机磷;而paq T基因在E. coli. DH5α中的溶磷机理是将葡萄糖氧化生成乙酸和甲酸,溶解无机磷。
Phosphate solubilizing microorganisms which can be suitable for colonization were isolated from soils. And One isolated wae screened, which can be suitable for corn production. Its colonization and phosphate-dissolving characteristics in soil were conducted. Cloning and identification of P-dissolving gene were studied in this article. The main conclusions were as follows:
Twenty-three bacteria strains with high performance of phosphate dissolving were screened. Sixteen fungi were isolated, including 9 strains of Penicillium oxalicum, 2 strain of Aspergillus niger, 2 strains of Penicillium variabile, 1 strain of Penicillium aculeatum, 1 strain of Aspergillus nidulans ,1 strain of Trichoderma viride.
The P-dissolving capbility of strains on different insoluble phosphorus(Ca_3(PO_4)_2, Zn_3(PO_4)_2, FePO_4, AlPO_4) were conducted. The P-dissolving ability of fungi were better than bacteria. 16 strains reduced the pH value from 7.0 to lower 2.0 in 18 h.They could dissolve all phosphate in solid culture.
Colonization and P-dissolving characteristics in soil results showed fungi presented high colonization ability in soil. However, colonization ability of fungi in different soils was significant differences. The P-dissolving ability of isolated were significantly correlated with their colonization. 6 strains of Z33、Zh、ZC、Zh3、ZI1 and Z30 presented high colonization ability in soil. The number of 6 strains increased 9.83-48 times than initial number in 28 days and the content of available phosphorous was increased 44%~160% than initial content.
The plate experiment and soil experiment of the matching between strains and crop in laboratory indicated that the 6 strains of Z15+、Zh、ZQ3、ZI1、ZJ1 and Z30 could propagate in plates and soil useing the root secretion of corn as sole carbon source and dissolve insoluble phosphorus. Six strains of Z33、Zh、ZC、Zh3、ZI1 and Z30 presented high colonization ability in soil, the number of 6 strains increased 9.60-48.03 times and the content of available phosphorous was increased 62.08%-147.55% in 21 days than initial content. The pot experiment inoculating with 5 strains was conducted in greenhouse by planting corn. The result demonstrated that the content of available phosphorous was increased remarkably with strains ZI1 and Zh treatment. The dry matter yield of corn significantly increased than CK, Then strains ZI1 and Zh were used in field experiment as microbial inoculums. The yield of corn treated with strain Zh increased remarkably in two years. However, the yield of corn treated with strain ZI1 presented yield increased in one year. The application of P-solubilizing microorganisms had a significant impact on microbial community structure . The numbers of bacteria, fungi, actinomycetes were increased than CK. There were positive correlation between the biodiversity and the number of bacteria.
Aspergillus nidulans Zh and Penicillium oxalicum ZI1 secrete lactic acid and acetic acid, the content of acid at 49th day was 372.5μg/mLand 593.9μg/mL , respectively. The strains ZI1 and Zh could colonizate in soil at 15、20、30℃and remarkably decreased the pH of soil. There were negative correlation between pH value decreased of soil and the number of fungi. The strain ZI1 and Zh promoted the transformation of insoluble phosphate which was the same law in different temperature. The content of Ca_2-P、Ca_8-P、Al-P、Fe-P and Ca_(10)-P were changed with ZI1 and Zh treatment. The transformation efficiency of Ca8-P of the strain ZI1was higher, but transformation efficiency of Al-P and Fe-P of the strain Zh was higher in soil. The mechanism of P-dissolving of the strain ZI1 and Zh was that they secrete organic acids in the progress of propagation chelating Ca~(2+), Fe~(3+), Al~(3+) for dissolving phosphate.
Sixty clone were screened from cDNA library of Aspergillus niger Zh3 according to the transparent zone of P-dissolving in plate. And 72 clone were screened from cDNA library of Penicillium oxalicum ZI1. The function of these clone were stable at the second generation. The gene from A.niger Zh3 was named pag T, sequence analysis showed that it was 1407bp in length, and shared 98% identity at the nucleotide level to Aspergillus niger contig An07c0220, and shared less than 80% identity to other isolated. The gene from Penicillium oxalicum ZI1 was named ppg T, Sequence analysis showed that it was 1160bp in length, and shared only 68% identity at the nucleotide level to Penicillium chrysogenum Wisconsin 54-1255, it showed ppg T was a novel gene.
Sequence analysis showed that the reading box of gene ppg T has 702bp, and its deduced 233 amino acid, this protein shares 59% homology with the Pc21g23580 in Penicillium chrysogenum Wisconsin 54-1255—the highest homology. And there was no report about the P-dissolving function of this protein, so the function of this protein PPG– T is a novel one. Sequence analysis showed that the reading box of gene pag T in Zh3 has 819bp, and its deduced 333 amino acids, this protein shared 100% homology with the bax Inhibitor family protein in Aspergillus niger CBS 513.88, but there is no report about the P-dissolving function of this protein. So it’s a novel function of the protein.Connecting ppqT and paqT to expression vectors pET28 and pGEX ,we constructed four engineering strains, which mainly produce acetic acid and formic acid.They could reduce the pH of the inorganic undissolved phosphorus medium to 3.6 or so .The 4 engineering strains was inducted by undissolved phosphorus, because they couldn’t reduce the pH value of medium without undissolved phosphorus. The P-dissolving mechanism of ppq T in E. coli. DH5αis the oxidation of glucose into acetic acid to dissolve phosphorus ; The phosphorus-dissolving mechanism of ppq T in E. coli. DH5αis the oxidation of glucose into acetic acid and formic acid to dissolve phosphorus.
引文
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