四川核桃主产区根际解磷细菌研究
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摘要
核桃具有重要的经济价值和生态作用,在我国林业建设和生态环境建设中具有十分重要的意义,特别是对于提高四川省山区农业经济水平具有积极的作用。但是,核桃植株树体高大,生长旺盛,养分消耗高,特别是对磷素需求较大,易造成土壤肥力下降。因此,施肥尤其是磷肥的施用成为核桃生产必不可少的技术措施。然而,随着化肥使用量的增加,其负面影响日益凸显。因此,探寻新的肥源,尤其是生物肥源以替代或部分替代化肥的研究倍受关注。目前,关于解磷菌剂在核桃上的应用尚未见报道。
本研究以四川省10个核桃主产区10年生核桃根际土壤为材料,进行了高效解磷细菌的分离,筛选,并通过形态特征、生理生化特性和16S rDNA鉴定其种属,优化了高效解磷菌株的基本培养条件,同时将高效解磷菌剂回接至美国山核桃(Carya illinoensis)1年生实生苗,观测其定殖规律,并探讨了解磷菌剂对核桃植株生长、养分吸收、土壤养分、土壤微生物区系及土壤酶活性的影响,获得以下主要研究结果：
1、利用选择性培养基,对四川10个核桃主产区核桃根际解磷细菌进行了分离、筛选,并分析比较不同产区解磷细菌的数量、类群以及影响因素。研究发现：(1)10个核桃主产区根际解磷细菌数量在3.71×105～35.21×105cfu/g干土之间；(2)海拔高度对四川核桃主产区根际土壤解磷细菌数量没有明显的影响。不同土壤类型中,褐土对根际解磷细菌的数量影响最大,依次为黄壤、棕壤。根际解磷细菌数量及种群结构与土壤有机质含量和碱解氮含量呈显著正相关,相关系数分别为0.639和0.724(P<0.05),与土壤速效磷含量呈显著的负相关,相关系数为-0.896(P<0.05)；(3)根据菌落形态、生理生化特征初步鉴定其类群分别属于9个属,其中以芽胞杆菌属(Bacillus)、假单胞菌属(Pseudomonas)和节杆菌属(Arthrobacter)为优势种群；(4)各核桃产区所处的生态环境不同,根际解磷细菌的种群多样性也存在一定的差异。解磷细菌多样性指数以汉源县最高,广元朝天区最低。丰富度指数以汉源县最高,兴汶县和广元朝天区丰富度指数最低。种群均匀度指数以广元朝天区最高,南江县最低。种群优势度指数以兴汶县最高,而以汉源县最低。
2、分别以溶磷圈法、液体振荡培养试验测定了分离后经多次纯化仍保持一定溶磷活性的37个菌株的解磷能力。结果表明：(1)参试菌株在Pikovskaya培养液中培养7 d后,解磷量为81.09 mg/L-233.35 mg/L,培养结束时pH值与解磷量呈显著负相关(r=-0.698)；(2)采用16S rDNA序列分析对菌株进行了鉴定,确定其中15株解磷菌株属于假单胞菌属(Pseudomonas),9株属于芽胞杆菌属(Bacillus),3株属于寡养单胞菌属(Stenotrophomonas),2株属于不动杆菌属(Acinetobacter),1株属于贪铜菌属(Cupriavidus),1株属于土壤杆菌属(Agrobacterium),1株属于节杆菌属(Arthrobacter),1株属于泛菌属(Pantoea),1株属于红球菌属(Rhodococcus)。
3、液体振荡培养法结合正交设计优化了高效解磷细菌W25、W9、W12的最适培养条件。结果表明：3株解磷细菌在不同碳源、氮源、初始pH值、接种量、装液量条件下,其生长量及解Ca3(PO4)2能力不同。菌株长势和解磷能力之间具有一定的正相关性。(1)解磷细菌W25的最佳培养基配方为：碳源(葡萄糖)20g/L,氮源(NH4C1) 1.0 g/L,初始pH值为7.5,接种量为4%,装液量为75 mL,其影响因素依次为碳源>氮源>装液量>初始pH值>接种量。经优化培养后的解磷活性为204.56 mg/L,较普通解磷培养基中的解磷活性高16.28%；(2)解磷细菌W9的最佳配方为：碳源(蔗糖)30g/L,氮源(NH4C1) 1.0 g/L,初始pH值为7.0,接种量为6%,装液量为100 mL。其影响因素依次为碳源>初始pH值>氮源>接种量>装液量。经优化培养后的解磷活性为192.25mg/L,较普通解磷培养基中的解磷活性高12.43%；(3)解磷细菌W12的最佳培养基配方为：碳源(葡萄糖)30g/L氮源(NH4NO3) 1.0 g/L,初始pH值为6.0,接种量为7%,装液量为75 mL。其影响因素依次为氮源>装液量>碳源>接种量>初始pH值。经优化培养后的解磷活性为192.12 mg/L,较普通解磷培养基中的解磷活性高18.44%。
4、在对解磷细菌W25,W9,W12进行耐药性标记、并得到稳定的链霉素抗性突变体菌株W25',W9'和W12'的基础上,通过盆栽试验,采用选择性培养基分离,研究了3株标记菌株在核桃根际的定殖动态。结果表明：(1)3株菌株均能成功地在核桃根际定殖；(2)两种土壤处理方式下,3株标记菌株的定殖动态变化规律相似。W25'在灭菌土和未灭菌中均为接种20 d后细菌数量达到最高,分别为1.28x108cfu/g根土、188x108cfu/g根土,接种后60 d细菌数量均趋向稳定；W9'在灭菌土和未灭菌土条件下,接种10 d后,根际土壤细菌数量均达到最高值,分别为2.02x107cfu/g根土和2.36x107cfu/g根土,之后逐渐下降,到60 d时趋向稳定；W12'在灭菌土和未灭菌土条件下,接种后20 d在根际的定殖水平最高,分别为1.94×107cfu/g根土和2.09×107cfu/g根土,之后逐渐下降,接种后60 d趋向稳定,菌株数量分别为3.38×103cfu/g根土和3.40×103cfu/g根土；(3)3株标记菌株在未灭菌土壤中的定殖能力高于灭菌土壤。
5、采用盆栽试验系统研究了解磷细菌对美国山核桃1年生实生苗生长发育、光合特性、土壤养分、土壤微生物区系及酶活性的影响,结果表明：(1)接种4种解磷菌剂均可不同程度地促进美国山核桃实生苗的株高、生物量、净光合速率、植株全氮含量和全磷含量；(2)接种解磷菌剂后,根际土壤速效磷、碱解氮含量也有不同程度的提高；(3)接种4种解磷菌剂均能增加根际土壤中细菌和放线菌的数量,减少真菌数量,提高土壤磷酸酶和脲酶活性。核桃植株根际细菌数量以新梢生长期最多,放线菌数量为新梢生长期>落叶期>展叶期,真菌数量为新梢生长期<落叶期<展叶期。根际土壤磷酸酶和脲酶活性均为新梢生长期>落叶期>展叶期；(4)解磷菌剂在土壤有效磷含量较低的情况下,能更好的发挥其溶磷作用及促生效果；(5)在接种解磷菌剂、磷水平处理相同的情况下,在未灭菌土壤里生长的核桃植株,其株高、生物量、净光合速率、植株全磷、全氮含量、土壤速效养分含量等各项指标均高于灭菌土壤；(6)综合分析表明,解磷菌剂能改善核桃植株根际土壤微生态环境,增强土壤酶活性,而土壤酶活性的提高又促进了土壤养分的有效化,加强了植株对N、P营养的吸收,从而促进核桃植株的生长发育。
Walnut plays an important role in agriculture economy and ecological environment of mountainous areas in Sichuan province, because of economic nuts and water and soil conservation. However, walnut plants often grow vigorously into 30-40 feet in height, and assimilate and consume lots of soil nutrients, especially for phosphorus. So to fertilize, including phosphorus is absolutely necessary to get high economic effect of walnut production. Whereas, the more closely the negative effects face to us, the more chemical fertilizer used in field, such as less unproducible phosphate and harder environmental pollution. Biological fertilizer is one of effective ways to absolve this puzzledom. Up to now, no reports have been found for phosphorus sollubilizing bacteria (PSB) in walnut field management.
In this study, rhizosphere soil of 10-year-old walnut plants were sampled separately in the ten important production areas in Sichuan province. The soil microorganisms were isolated and the PSBs with high ability of phosphorus solubilizing were screened out from all the rhizosphere soil samples. These PSBs were identified effectively according to their biophysical and biochemical characteristics and 16S rDNA sequencing. The culture conditions were optimized through orthogonal experiments for the three PSBs. At last, this three PSBs with high ability of phosphorus sollubilizing were inoculate into one-year-old seedlings of pecan (Carya illinoensis) to observe the inoculation effects on such as plant growth, soil nutrients, microorganism and activities of soil enzymes.
The main results according to this study are as follows:
1. By using selective culture media, the phosphate solubilizing bacteria (PSB) in the walnut rhizosphere were isolated and identified with their community structure and genus studied. The results showed that:(1) The number of PSB that formed clear zones was among 3.71×105～35.21×105cfu/g dry soil (DS) across all the sites. (2) The altitude could not influence the PSB number. The maximum PSB number was recorded in cinnamon soil followed by yellow soil. The distribution of PSB was influenced by soil organic matter, soil available phosphorus, the content of alkali-hydrolysable nitrogen. The PSB number were found to be significantly positive correlated with organic matter or alkali-hydrolysable nitrogen of rhizosphere soil(r=0.639, r=0.724), but were significantly negative correlated to available phosphor content of soil(r=-0.896). (There were a strong negative correlation between the amount of PSB and available phosphorus content of soil. (r=-0.896). (3) The PSB isolated from all the rhizosphere soil were belonged to nine genera according to the colony morphogenesis, biochemical characterization, and Bacillus, Psedumonas and Arthrobacter were the dominant ones. (4) The diversity of PSB population could vary with different ecological environments. The diversity index of PSB was highest in Hanyuan site compared with the lowest index in Guangyuan site. Richness index changed as similar to the diversity index among all the sampling sites. The evenness index was highest in Guangyuan site and was the lowest in Nanjiang site. As for domination index, it was the highest in Xingwen site, whereas the lowest in Hanyuan site.
2. Phosphate solubilizing activities of 37 strains of PSBs were tested on solid tricalcium and liquid phosphate medium. The soluble-P concentration ranged between 81.09 mg/L and 233.35 mg/L with variations among different strains. A strong negative correlation (r=-0.698) between pH and soluble P concentration was observed. Identification of 37 PSBs were carried out by 16S rDNA sequencing. Fifteen strains belonged to genus Pseudomonas, nine to genus Bacillus, three to genus Stenotrophomonas, two to genus Acinetobacter, one to genus Cupriavidus, one to genus Agrobacterium, one to genus Arthrobacter, one to genus Pantoe, one to genus Rhodococcus.
3. The optimal culture media and phosphate solubilizing conditions of PSB W25, W9 and W12 were studied separately. We found that their growth and phosphate dissolving ability of these three bacteria strains were different under conditions of different carbon sources, nitrogen sources, initial pH, inoculums and liquid volumes. The growth ability of bacteria strains was positively correlated with their phosphorus dissolving abilities. The results of orthogonal experiments showed that:(1) for W25, the optimal carbon and nitrogen source were glucose at the concentration of 20.0 g/L, NH4C1 at the concentration of 1.0 g/L with pH7.5, inoculums of 4%, liquid volume of 75 mL. The importance of the four factors are ranked in decreasing order:carbon source> nitrogen source> liquid volume> the initial pH> inoculums. The phosphate solubilizing capacity of it was up to 204.56 mg/L under the optimized conditions of cultivation, being 16.28% higher than CK; (2) While for W9, the optimal carbon and nitrogen source were sucrose at the concentration of 30.0 g/L and NH4C1 at 1.0 g/L with pH7.0, inoculums of 6%, liquid volume of 100 mL. The importance of the four factors are ranked in decreasing order:carbon source> the initial pH> nitrogen source> inoculum> liquid volume. The phosphate solubilizing capacity of this bacteria strain was 192.25 mg/L under the optimized conditions of media, being 12.43% higher than CK; (3) For W12, the optimal carbon and nitrogen source were glucose at the concentration of 30.0 g/L, NH4NO3 at the concentration of 1.0 g/L with pH6.0, inoculums of 7%, liquid volume of 75 mL; The importance of the four factors are ranked in decreasing order: nitrogen source>liquid volume> carbon source> inoculum> the initial pH. The phosphate solubilizing activity of W12 was 192.12 mg/L under the optimized conditions of cultivation, being 18.44% higher than CK.
4. The PSB strains W25, W9, W12 were marked with streptomycin, and the stable streptomycin resistance strains W25', W9'and W12'were obtained. Their colonization dynamics in walnut rhizosphere were studied under pot experiment condition by counting with selective medium. The results of potted experiment showed that:(1) the three strains could successfully colonize in the rhizosphere of walnut seedlings. (2) Both in sterilized and unsterilized soil, the colonization dynamics was similar for all the three bacteria strains. The highest colonization level of W25'and W12'were reached on 20 days after inoculation,1.28x108 cfu/g DS and 1.88x108 cfu/g DS,1.94x107 cfu/g DS and 2.09×107 cfu/g DS, respectively. But the survival number of W9'reached the peak value after inoculation of 10 days,2.02×107 cfu/g DS and 2.36×107 cfu/g DS under conditions of sterilized and unsterilized soil, respectively. And then, the population of the three strains decreased gradually and tended to be stable on 60 days after inoculation; (3) Colonization level in unsterilized soil was higher than sterilized soil.
5. The effects of PSB strains on growth, photosynthesis characteristics, nutrient uptake and rhizosphere properties of walnut under pot experiment were studied. The results show that: (1) The plant height, biomass, net photosynthesis rate (Pn), total content of nitrogen and phosphorus of walnut seedlings were higher after inoculation with the PSB strains; (2) Application of the three PSB strains increased the content of available phosphorus and alkali-hydrolyzable nitrogen in the rhizosphere of walnut; (3) Amounts of bacteria and actinomyces of rhizosphere soil increased, but the amount of soil fungi decreased to some extent after inoculation with strains W25, W9 and W12. At the same time, activities of phosphatase and urease were improved effectively. The higher phosphorus solubilizing and growth promoting effects were observed for all the three PSBs in the low content of available phosphorus of potted soil. The maximum number of population of rhizosphere bacteria and actinomyces were measured at the shoot growth stage followed by defoliation and leaf expansion stage. Nevertheless, the amounts of rhizosphere fungi changed in opposite direction. For phosphatase and urease activities, both of them reached its maximum at the shoot growth stage followed by defoliation and leaf expansion stage of walnut seedlings; (4) The PSB strain could act with higher phosphate-solubilizing ability and growth promoting effect of plant under conditions of low available P in soil; (5) At the same phosphorus level of potted soil and bacteria strain, some important indexes including plant height, biomass, Pn value, total phosphorus and nitrogen content of plant, available nutrient of soil under conditions of unsterilized soil were higher than that under conditions of sterilized soil. (6) In conclusion, PSB strains could improve micro-ecological environment in rhizosphere soil of walnut plant, increase the enzyme activities which would promote effective absorption of N and P nutrition for walnut seedlings.
本研究以四川省10个核桃主产区10年生核桃根际土壤为材料,进行了高效解磷细菌的分离,筛选,并通过形态特征、生理生化特性和16S rDNA鉴定其种属,优化了高效解磷菌株的基本培养条件,同时将高效解磷菌剂回接至美国山核桃(Carya illinoensis)1年生实生苗,观测其定殖规律,并探讨了解磷菌剂对核桃植株生长、养分吸收、土壤养分、土壤微生物区系及土壤酶活性的影响,获得以下主要研究结果：
1、利用选择性培养基,对四川10个核桃主产区核桃根际解磷细菌进行了分离、筛选,并分析比较不同产区解磷细菌的数量、类群以及影响因素。研究发现：(1)10个核桃主产区根际解磷细菌数量在3.71×105～35.21×105cfu/g干土之间；(2)海拔高度对四川核桃主产区根际土壤解磷细菌数量没有明显的影响。不同土壤类型中,褐土对根际解磷细菌的数量影响最大,依次为黄壤、棕壤。根际解磷细菌数量及种群结构与土壤有机质含量和碱解氮含量呈显著正相关,相关系数分别为0.639和0.724(P<0.05),与土壤速效磷含量呈显著的负相关,相关系数为-0.896(P<0.05)；(3)根据菌落形态、生理生化特征初步鉴定其类群分别属于9个属,其中以芽胞杆菌属(Bacillus)、假单胞菌属(Pseudomonas)和节杆菌属(Arthrobacter)为优势种群；(4)各核桃产区所处的生态环境不同,根际解磷细菌的种群多样性也存在一定的差异。解磷细菌多样性指数以汉源县最高,广元朝天区最低。丰富度指数以汉源县最高,兴汶县和广元朝天区丰富度指数最低。种群均匀度指数以广元朝天区最高,南江县最低。种群优势度指数以兴汶县最高,而以汉源县最低。
2、分别以溶磷圈法、液体振荡培养试验测定了分离后经多次纯化仍保持一定溶磷活性的37个菌株的解磷能力。结果表明：(1)参试菌株在Pikovskaya培养液中培养7 d后,解磷量为81.09 mg/L-233.35 mg/L,培养结束时pH值与解磷量呈显著负相关(r=-0.698)；(2)采用16S rDNA序列分析对菌株进行了鉴定,确定其中15株解磷菌株属于假单胞菌属(Pseudomonas),9株属于芽胞杆菌属(Bacillus),3株属于寡养单胞菌属(Stenotrophomonas),2株属于不动杆菌属(Acinetobacter),1株属于贪铜菌属(Cupriavidus),1株属于土壤杆菌属(Agrobacterium),1株属于节杆菌属(Arthrobacter),1株属于泛菌属(Pantoea),1株属于红球菌属(Rhodococcus)。
3、液体振荡培养法结合正交设计优化了高效解磷细菌W25、W9、W12的最适培养条件。结果表明：3株解磷细菌在不同碳源、氮源、初始pH值、接种量、装液量条件下,其生长量及解Ca3(PO4)2能力不同。菌株长势和解磷能力之间具有一定的正相关性。(1)解磷细菌W25的最佳培养基配方为：碳源(葡萄糖)20g/L,氮源(NH4C1) 1.0 g/L,初始pH值为7.5,接种量为4%,装液量为75 mL,其影响因素依次为碳源>氮源>装液量>初始pH值>接种量。经优化培养后的解磷活性为204.56 mg/L,较普通解磷培养基中的解磷活性高16.28%；(2)解磷细菌W9的最佳配方为：碳源(蔗糖)30g/L,氮源(NH4C1) 1.0 g/L,初始pH值为7.0,接种量为6%,装液量为100 mL。其影响因素依次为碳源>初始pH值>氮源>接种量>装液量。经优化培养后的解磷活性为192.25mg/L,较普通解磷培养基中的解磷活性高12.43%；(3)解磷细菌W12的最佳培养基配方为：碳源(葡萄糖)30g/L氮源(NH4NO3) 1.0 g/L,初始pH值为6.0,接种量为7%,装液量为75 mL。其影响因素依次为氮源>装液量>碳源>接种量>初始pH值。经优化培养后的解磷活性为192.12 mg/L,较普通解磷培养基中的解磷活性高18.44%。
4、在对解磷细菌W25,W9,W12进行耐药性标记、并得到稳定的链霉素抗性突变体菌株W25',W9'和W12'的基础上,通过盆栽试验,采用选择性培养基分离,研究了3株标记菌株在核桃根际的定殖动态。结果表明：(1)3株菌株均能成功地在核桃根际定殖；(2)两种土壤处理方式下,3株标记菌株的定殖动态变化规律相似。W25'在灭菌土和未灭菌中均为接种20 d后细菌数量达到最高,分别为1.28x108cfu/g根土、188x108cfu/g根土,接种后60 d细菌数量均趋向稳定；W9'在灭菌土和未灭菌土条件下,接种10 d后,根际土壤细菌数量均达到最高值,分别为2.02x107cfu/g根土和2.36x107cfu/g根土,之后逐渐下降,到60 d时趋向稳定；W12'在灭菌土和未灭菌土条件下,接种后20 d在根际的定殖水平最高,分别为1.94×107cfu/g根土和2.09×107cfu/g根土,之后逐渐下降,接种后60 d趋向稳定,菌株数量分别为3.38×103cfu/g根土和3.40×103cfu/g根土；(3)3株标记菌株在未灭菌土壤中的定殖能力高于灭菌土壤。
5、采用盆栽试验系统研究了解磷细菌对美国山核桃1年生实生苗生长发育、光合特性、土壤养分、土壤微生物区系及酶活性的影响,结果表明：(1)接种4种解磷菌剂均可不同程度地促进美国山核桃实生苗的株高、生物量、净光合速率、植株全氮含量和全磷含量；(2)接种解磷菌剂后,根际土壤速效磷、碱解氮含量也有不同程度的提高；(3)接种4种解磷菌剂均能增加根际土壤中细菌和放线菌的数量,减少真菌数量,提高土壤磷酸酶和脲酶活性。核桃植株根际细菌数量以新梢生长期最多,放线菌数量为新梢生长期>落叶期>展叶期,真菌数量为新梢生长期<落叶期<展叶期。根际土壤磷酸酶和脲酶活性均为新梢生长期>落叶期>展叶期；(4)解磷菌剂在土壤有效磷含量较低的情况下,能更好的发挥其溶磷作用及促生效果；(5)在接种解磷菌剂、磷水平处理相同的情况下,在未灭菌土壤里生长的核桃植株,其株高、生物量、净光合速率、植株全磷、全氮含量、土壤速效养分含量等各项指标均高于灭菌土壤；(6)综合分析表明,解磷菌剂能改善核桃植株根际土壤微生态环境,增强土壤酶活性,而土壤酶活性的提高又促进了土壤养分的有效化,加强了植株对N、P营养的吸收,从而促进核桃植株的生长发育。
Walnut plays an important role in agriculture economy and ecological environment of mountainous areas in Sichuan province, because of economic nuts and water and soil conservation. However, walnut plants often grow vigorously into 30-40 feet in height, and assimilate and consume lots of soil nutrients, especially for phosphorus. So to fertilize, including phosphorus is absolutely necessary to get high economic effect of walnut production. Whereas, the more closely the negative effects face to us, the more chemical fertilizer used in field, such as less unproducible phosphate and harder environmental pollution. Biological fertilizer is one of effective ways to absolve this puzzledom. Up to now, no reports have been found for phosphorus sollubilizing bacteria (PSB) in walnut field management.
In this study, rhizosphere soil of 10-year-old walnut plants were sampled separately in the ten important production areas in Sichuan province. The soil microorganisms were isolated and the PSBs with high ability of phosphorus solubilizing were screened out from all the rhizosphere soil samples. These PSBs were identified effectively according to their biophysical and biochemical characteristics and 16S rDNA sequencing. The culture conditions were optimized through orthogonal experiments for the three PSBs. At last, this three PSBs with high ability of phosphorus sollubilizing were inoculate into one-year-old seedlings of pecan (Carya illinoensis) to observe the inoculation effects on such as plant growth, soil nutrients, microorganism and activities of soil enzymes.
The main results according to this study are as follows:
1. By using selective culture media, the phosphate solubilizing bacteria (PSB) in the walnut rhizosphere were isolated and identified with their community structure and genus studied. The results showed that:(1) The number of PSB that formed clear zones was among 3.71×105～35.21×105cfu/g dry soil (DS) across all the sites. (2) The altitude could not influence the PSB number. The maximum PSB number was recorded in cinnamon soil followed by yellow soil. The distribution of PSB was influenced by soil organic matter, soil available phosphorus, the content of alkali-hydrolysable nitrogen. The PSB number were found to be significantly positive correlated with organic matter or alkali-hydrolysable nitrogen of rhizosphere soil(r=0.639, r=0.724), but were significantly negative correlated to available phosphor content of soil(r=-0.896). (There were a strong negative correlation between the amount of PSB and available phosphorus content of soil. (r=-0.896). (3) The PSB isolated from all the rhizosphere soil were belonged to nine genera according to the colony morphogenesis, biochemical characterization, and Bacillus, Psedumonas and Arthrobacter were the dominant ones. (4) The diversity of PSB population could vary with different ecological environments. The diversity index of PSB was highest in Hanyuan site compared with the lowest index in Guangyuan site. Richness index changed as similar to the diversity index among all the sampling sites. The evenness index was highest in Guangyuan site and was the lowest in Nanjiang site. As for domination index, it was the highest in Xingwen site, whereas the lowest in Hanyuan site.
2. Phosphate solubilizing activities of 37 strains of PSBs were tested on solid tricalcium and liquid phosphate medium. The soluble-P concentration ranged between 81.09 mg/L and 233.35 mg/L with variations among different strains. A strong negative correlation (r=-0.698) between pH and soluble P concentration was observed. Identification of 37 PSBs were carried out by 16S rDNA sequencing. Fifteen strains belonged to genus Pseudomonas, nine to genus Bacillus, three to genus Stenotrophomonas, two to genus Acinetobacter, one to genus Cupriavidus, one to genus Agrobacterium, one to genus Arthrobacter, one to genus Pantoe, one to genus Rhodococcus.
3. The optimal culture media and phosphate solubilizing conditions of PSB W25, W9 and W12 were studied separately. We found that their growth and phosphate dissolving ability of these three bacteria strains were different under conditions of different carbon sources, nitrogen sources, initial pH, inoculums and liquid volumes. The growth ability of bacteria strains was positively correlated with their phosphorus dissolving abilities. The results of orthogonal experiments showed that:(1) for W25, the optimal carbon and nitrogen source were glucose at the concentration of 20.0 g/L, NH4C1 at the concentration of 1.0 g/L with pH7.5, inoculums of 4%, liquid volume of 75 mL. The importance of the four factors are ranked in decreasing order:carbon source> nitrogen source> liquid volume> the initial pH> inoculums. The phosphate solubilizing capacity of it was up to 204.56 mg/L under the optimized conditions of cultivation, being 16.28% higher than CK; (2) While for W9, the optimal carbon and nitrogen source were sucrose at the concentration of 30.0 g/L and NH4C1 at 1.0 g/L with pH7.0, inoculums of 6%, liquid volume of 100 mL. The importance of the four factors are ranked in decreasing order:carbon source> the initial pH> nitrogen source> inoculum> liquid volume. The phosphate solubilizing capacity of this bacteria strain was 192.25 mg/L under the optimized conditions of media, being 12.43% higher than CK; (3) For W12, the optimal carbon and nitrogen source were glucose at the concentration of 30.0 g/L, NH4NO3 at the concentration of 1.0 g/L with pH6.0, inoculums of 7%, liquid volume of 75 mL; The importance of the four factors are ranked in decreasing order: nitrogen source>liquid volume> carbon source> inoculum> the initial pH. The phosphate solubilizing activity of W12 was 192.12 mg/L under the optimized conditions of cultivation, being 18.44% higher than CK.
4. The PSB strains W25, W9, W12 were marked with streptomycin, and the stable streptomycin resistance strains W25', W9'and W12'were obtained. Their colonization dynamics in walnut rhizosphere were studied under pot experiment condition by counting with selective medium. The results of potted experiment showed that:(1) the three strains could successfully colonize in the rhizosphere of walnut seedlings. (2) Both in sterilized and unsterilized soil, the colonization dynamics was similar for all the three bacteria strains. The highest colonization level of W25'and W12'were reached on 20 days after inoculation,1.28x108 cfu/g DS and 1.88x108 cfu/g DS,1.94x107 cfu/g DS and 2.09×107 cfu/g DS, respectively. But the survival number of W9'reached the peak value after inoculation of 10 days,2.02×107 cfu/g DS and 2.36×107 cfu/g DS under conditions of sterilized and unsterilized soil, respectively. And then, the population of the three strains decreased gradually and tended to be stable on 60 days after inoculation; (3) Colonization level in unsterilized soil was higher than sterilized soil.
5. The effects of PSB strains on growth, photosynthesis characteristics, nutrient uptake and rhizosphere properties of walnut under pot experiment were studied. The results show that: (1) The plant height, biomass, net photosynthesis rate (Pn), total content of nitrogen and phosphorus of walnut seedlings were higher after inoculation with the PSB strains; (2) Application of the three PSB strains increased the content of available phosphorus and alkali-hydrolyzable nitrogen in the rhizosphere of walnut; (3) Amounts of bacteria and actinomyces of rhizosphere soil increased, but the amount of soil fungi decreased to some extent after inoculation with strains W25, W9 and W12. At the same time, activities of phosphatase and urease were improved effectively. The higher phosphorus solubilizing and growth promoting effects were observed for all the three PSBs in the low content of available phosphorus of potted soil. The maximum number of population of rhizosphere bacteria and actinomyces were measured at the shoot growth stage followed by defoliation and leaf expansion stage. Nevertheless, the amounts of rhizosphere fungi changed in opposite direction. For phosphatase and urease activities, both of them reached its maximum at the shoot growth stage followed by defoliation and leaf expansion stage of walnut seedlings; (4) The PSB strain could act with higher phosphate-solubilizing ability and growth promoting effect of plant under conditions of low available P in soil; (5) At the same phosphorus level of potted soil and bacteria strain, some important indexes including plant height, biomass, Pn value, total phosphorus and nitrogen content of plant, available nutrient of soil under conditions of unsterilized soil were higher than that under conditions of sterilized soil. (6) In conclusion, PSB strains could improve micro-ecological environment in rhizosphere soil of walnut plant, increase the enzyme activities which would promote effective absorption of N and P nutrition for walnut seedlings.
引文
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