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磷(P)是植物生长和发育所必需的营养物质,在植物细胞分裂、生物合成等生理过程扮演着重要角色。研究显示,增加土壤磷有效性对森林生态系统生产力的提升具有显著影响[1-5],而采用NaHCO3、酸性氟化铵(NH4F)等单一提取剂评估土壤磷有效性的方法无法完全反映土壤中不同形态磷间的复杂转化关系[6]。恰当的土壤磷分级方法能较好地理解土壤磷素的转化过程,Chang等[7]将土壤无机磷分为钙结合态磷(Ca-P)、铝结合态磷(Al-P)、铁结合态磷(Fe-P)和闭蓄态磷(Occluded-P)4个组分,适用于石灰土壤;Hedley等[8]将土壤磷分为树脂交换态磷、土壤微生物磷、NaOH溶性磷和残留磷等7个组分,兼顾了土壤无机磷和有机磷,适用于酸性土壤[9-10]。这些方法虽能评估土壤磷有效性程度,却无法充分反应根际过程带来的贡献,如有机酸、质子和胞外酶的分泌降解作用[11]。DeLuca等[12]从生物学利用的难易程度研发出评价生物有效性磷的分级方法(BBP法),该方法主要依据植物和微生物分泌有机酸、酸性磷酸酶等机制进行分类,将土壤磷素分为:(1)CaCl2提取的可溶性磷(CaCl2-P),直接可被根际截留或扩散的磷酸根离子;(2)柠檬酸提取的磷(Citrate-P),可被有机酸活化释放的无机磷;(3)酶提取的磷(Enzyme-P),被植物和微生物分泌的磷酸酶和植酸酶矿化的有机磷;(4)盐酸提取的无机磷(HCl-P),可被H+活化的最大潜力磷库。
毛竹(Phyllostachys edulis (Carrière) J. Houz.)是一种典型的克隆植物,主要分布于我国南方丘陵红壤地区,具有生长速度快、木材质量优、观赏价值高等特点,是我国南方重要的经济树种[13]。近年来,随着毛竹人工林面积扩大及间伐期缩减,竹林资源质量下降,地力衰退,已严重影响了竹林的经营发展。研究显示,磷已成为毛竹林生产力提高的主要限制因子之一[14],而过度施用化学磷肥导致竹林土壤形成储量可观的潜在磷库,故如何活化竹林土壤潜在磷库,缓解竹林土壤有效磷短缺是竹林可持续经营发展亟需解决的问题。解磷微生物作为土壤有益微生物种类之一,可通过释放有机酸、质子等方式将土壤潜在磷库中的难溶性磷酸盐转化为可溶性磷,促进植物生长。以往部分研究表明,添加生物有机肥可提高土壤相关酶活性,从而提升土壤有效养分[15],而解磷微生物在调控土壤磷有效性含量中是否存在土壤酶活性介导作用仍有待证实,且关于解磷微生物对毛竹林土壤磷组分有何影响还尚未明确,解磷微生物如何调控土壤不同磷组分进而影响毛竹生长的报道甚少。
课题组前期从毛竹根际土壤中获得1株高效解磷真菌金黄蓝状菌(Talaromyces aurantiacus (J.H. Mill., Giddens & A.A.Foster))JXBR04,具有较好的解无机磷和有机磷能力[16]。本实验基于生物有效性的磷分级方法,通过温室盆栽试验开展该菌株对毛竹根际土壤不同磷组分含量的影响及促生作用,明确该菌株对毛竹根际土壤磷有效性的调控作用,有望阐明解磷真菌提升竹林土壤有效磷的生物途径及对相关促生机理做出诠释。
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表1表明:施用解磷菌株JXBR04制剂能显著提高毛竹叶片生物量和净光合速率,增幅分别为51.74%[16]、51.74%。
表 1 施用菌株JXBR04制剂对毛竹生长量和净光合速率的影响
Table 1. Effect of added strain JXBR04 on biomass and net photosynthetic rate of moso bamboo seedlings.
观测指标
Observed variable对照
Control施用菌剂
Adding strain生物量 Biomass/g[16]* 3.73 ± 0.21 b 5.66 ± 0.13 a 净光合速率
Net photosynthetic rate/(μmol·m−2·s−1)2.72 ± 0.41 b 3.78 ± 1.86 a 注:同行不同字母表示处理间差异显著p < 0.05),下同。*生物量数据来源于文献[16]。
Notes: Different letters in the same row indicated a significant difference between treatments at the 0.05 level, The same as below. *The date of biomass was derived from reference [16]. -
表2表明:施用菌株JXBR04制剂180 d后改变了毛竹根际土壤的化学性质,其中,显著提高了土壤中矿质氮、有效磷和有机质的含量,与对照相比分别增加了13.3%、78.3%、12.5%;而施用菌株JXBR04制剂短期内对土壤pH值并无显著影响。
表 2 施用菌株JXBR04制剂对毛竹土壤化学性质的影响
Table 2. Effect of added strain JXBR04 on soil chemical properties of moso bamboo seedlings.
化学性质
Chemical property对照
Control施用菌剂
Adding strain矿质氮
Mineral nitrogen/(mg·kg−1)3.15 ± 0.04 b 3.57 ± 0.08 a 有效磷
Available phosphorus/(mg·kg−1)3.27 ± 0.43 b 5.83 ± 0.16 a 全氮
Total nitrogen/(g·kg−1)0.47 ± 0.12 a 0.58 ± 0.12 a 全磷
Total phosphorus/(g·kg−1)0.26 ± 0.05 a 0.24 ± 0.04 a 有机质
Organic matter/(g·kg−1)6.82 ± 0.62 b 7.67 ± 0.97 a pH值 6.40 ± 0.15 a 5.98 ± 0.04 a -
图1表明:毛竹根际土壤各磷组分含量存在显著差异,其中,盐酸磷(HCl-P)含量最高(21.18 mg·kg−1),柠檬酸磷含量次之(3.40 mg·kg−1),氯化钙磷(CaCl2-P)含量随后(0.33 mg·kg−1),酶磷含量最低(0.30 mg·kg−1)。施用菌株JXBR04制剂极显著提高了毛竹根际土壤HCl-P和CaCl2-P的含量,分别增加了46.0%和51.6%,而对土壤柠檬酸磷和酶磷含量并无显著影响。
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图2表明:施用解磷菌剂影响了毛竹根际土壤酶活性,其中,极显著增强了土壤脲酶和过氧化氢酶的活性,与对照相比分别增加了25.9%和101.0%,而酸性磷酸酶和碱性磷酸酶含量与对照相比差异不显著。
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Pearson相关性分析结果(表3)显示:土壤磷组分与土壤化学性质和生长指标相关性不尽一致,其中,土壤有效磷、毛竹生物量与CaCl2-P、HCl-P均呈极显著正相关,土壤pH与CaCl2-P、HCl-P均呈极显著负相关;土壤有效磷与土壤柠檬酸磷、酶磷不显著相关,土壤酶磷与毛竹生物量不显著相关;土壤全磷与4个磷组分均不显著相关。
表 3 土壤磷组分与各测定指标之间的相关性
Table 3. Relationships between various measurement indexes and phosphorus fraction
观测指标
Observed variable氯化钙磷
CaCl2−P柠檬酸磷
Citrate-P酶磷
Enzyme-P盐酸磷
HCl-PpH值 −0.967** −0.886* −0.604 −0.937** 生物量 Total biomasss 0.970** 0.850* 0.680 0.970** 全磷 Total phosphorus −0.615 −0.684 −0.563 −0.414 有机质 Organic matter 0.874* 0.783 0.818* 0.843* 矿质氮 Mineral nitrogen 0.830* 0.804 0.921** 0.739 有效磷 Available phosphorus 0.930** 0.780 0.770 0.930** 过氧化氢酶 Catalase −0.920 0.897 0.840 −0.980 酸性磷酸酶 Acid phosphatase −0.988 0.762 0.683 −0.901 碱性磷酸酶 Alkaline phosphatase −0.723 −0.052 −0.167 −0.202
金黄蓝状菌对毛竹土壤磷组分及苗木生物量的影响
Influence of Talaromyces aurantiacus on the Soil Phosphorus Fraction and Biomass of Moso Bamboo Seedling
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摘要:
目的 为探究解磷真菌金黄蓝状菌(Talaromyces aurantiacus)JXBR04在毛竹促生作用中对土壤生物有效性磷组分的贡献。 方法 采用温室盆栽法研究该菌株对毛竹根际土壤生物有效磷组分含量、土壤有效磷、矿质氮含量及相关土壤酶活性的影响,并分析该菌株对毛竹的促生效果。 结果 与对照(CK)相比,施用菌株JXBR04制剂显著提高了毛竹根际土壤中盐酸磷和氯化钙磷组分含量,显著增强了土壤脲酶和过氧化氢酶活性,而对酶磷、柠檬酸磷组分含量和土壤酸性磷酸酶、碱性磷酸酶活性并无影响,且土壤有效磷和矿质氮含量分别提高了78.3%和13.3%;土壤盐酸磷、氯化钙磷组分含量和土壤有效磷含量、毛竹生物量均呈极显著正相关,表明施用菌株JXBR04制剂显著促进了毛竹的生物量。 结论 金黄蓝状菌JXBR04主要通过提高土壤氯化钙磷和盐酸磷组分含量来增强土壤有效磷供给,进而促进毛竹对土壤有效磷的吸收利用和生物量增长,且该过程并无土壤磷酸酶介导。本研究可为竹林施用磷肥高效利用和管理提供理论依据。 Abstract:Objective To explore the contribution of the phosphorus-solubilizing fungi Talaromyces aurantiacus JXBR04 to soil phosphorus bioavailability in growth promotion of Phyllostachys edulis. Method Greenhouse potting experiment was used to study the effect of the strain on the content of phosphorus bioavailability in the rhizosphere soil of Ph. edulis. The contents of available phosphorus, mineral nitrogen the activities of related soil enzymes in the soil were analyzed, and the growth-promoting effect on Ph. edulis by inoculating on JXBR04 was evaluated. Results Compared with the control, the application of JXBR04 significantly increased the content of HCl-P and CaCl2-P fraction, increased the activities of soil urease and catalase in the rhizosphere soil of Ph. edulis, while the content of enzyme-P, citric acid-P fraction, the activities of soil acid phosphatase and alkaline phosphatase were not affected. The content of soil available phosphorus and mineral nitrogen increased by 78.3% and 13.3%, respectively. The content of HCl-P and CaCl2-P in soil was significantly positively correlated with the content of available phosphorus in soil and the biomass of Ph. edulis, indicating that the application of JXBR04 significantly promoted the biomass of Ph. edulis. Conclusion The increase of available phosphorus in soil is driven mainly by the content of CaCl2-P and HCl-P fraction though inoculating on strain JXBR04, which can promote Ph. edulis to absorb and utilize the available phosphorus to increase biomass. This regulated process is not mediated by soil phosphatase. -
Key words:
- Phyllostachys edulis
- / Talaromyces aurantiacus
- / phosphorus fraction
- / biomass
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表 1 施用菌株JXBR04制剂对毛竹生长量和净光合速率的影响
Table 1. Effect of added strain JXBR04 on biomass and net photosynthetic rate of moso bamboo seedlings.
观测指标
Observed variable对照
Control施用菌剂
Adding strain生物量 Biomass/g[16]* 3.73 ± 0.21 b 5.66 ± 0.13 a 净光合速率
Net photosynthetic rate/(μmol·m−2·s−1)2.72 ± 0.41 b 3.78 ± 1.86 a 注:同行不同字母表示处理间差异显著p < 0.05),下同。*生物量数据来源于文献[16]。
Notes: Different letters in the same row indicated a significant difference between treatments at the 0.05 level, The same as below. *The date of biomass was derived from reference [16].表 2 施用菌株JXBR04制剂对毛竹土壤化学性质的影响
Table 2. Effect of added strain JXBR04 on soil chemical properties of moso bamboo seedlings.
化学性质
Chemical property对照
Control施用菌剂
Adding strain矿质氮
Mineral nitrogen/(mg·kg−1)3.15 ± 0.04 b 3.57 ± 0.08 a 有效磷
Available phosphorus/(mg·kg−1)3.27 ± 0.43 b 5.83 ± 0.16 a 全氮
Total nitrogen/(g·kg−1)0.47 ± 0.12 a 0.58 ± 0.12 a 全磷
Total phosphorus/(g·kg−1)0.26 ± 0.05 a 0.24 ± 0.04 a 有机质
Organic matter/(g·kg−1)6.82 ± 0.62 b 7.67 ± 0.97 a pH值 6.40 ± 0.15 a 5.98 ± 0.04 a 表 3 土壤磷组分与各测定指标之间的相关性
Table 3. Relationships between various measurement indexes and phosphorus fraction
观测指标
Observed variable氯化钙磷
CaCl2−P柠檬酸磷
Citrate-P酶磷
Enzyme-P盐酸磷
HCl-PpH值 −0.967** −0.886* −0.604 −0.937** 生物量 Total biomasss 0.970** 0.850* 0.680 0.970** 全磷 Total phosphorus −0.615 −0.684 −0.563 −0.414 有机质 Organic matter 0.874* 0.783 0.818* 0.843* 矿质氮 Mineral nitrogen 0.830* 0.804 0.921** 0.739 有效磷 Available phosphorus 0.930** 0.780 0.770 0.930** 过氧化氢酶 Catalase −0.920 0.897 0.840 −0.980 酸性磷酸酶 Acid phosphatase −0.988 0.762 0.683 −0.901 碱性磷酸酶 Alkaline phosphatase −0.723 −0.052 −0.167 −0.202 -
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