摘要
钾是农作物生长必需的大量营养元素之一,而我国钾肥资源短缺,因此通过生物转化将我国含量丰富的低品位含钾岩石生产成生物钾肥不但能拓宽我国钾肥生产资源,还可以缓解农业生产钾肥紧张的状况。微生物在含钾岩石生物转化过程中发挥重要作用,对含钾岩石生物转化过程中微生物的多样性进行研究,进而对其风化含钾岩石的机制进行探讨,有利于为进一步改进含钾岩石生物转化工艺提供理论依据。
为研究蚯蚓(Pheretima carnosa)对低品位含钾矿石风化的促进作用,用含钾矿粉喂养了蚯蚓,然后对喂养蚯蚓前后含钾矿粉水溶性和硝酸浸提液中钾、铝、铁、钙等元素进行了测定,结果表明：饲养过蚯蚓的钾矿粉与不加蚯蚓的对照样品相比,前者水溶性和硝酸浸提液中钾、铝、铁、钙等四种元素含量均明显高于后者,其中水溶性钾、铝、铁、钙等元素含量比后者分别高5.64%、20.25%、27.52%、17.21%,硝酸浸提液中钾、铝、铁、钙等元素含量比后者分别高6.59%、3.94%、4.36%、8.49%。上述结果说明蚯蚓能促进含钾矿粉的风化。为揭示与蚯蚓促进含钾矿粉风化相关的微生物群落结构,用16S rRNA基因克隆文库和限制性酶切片段长度多样性分析(ARDRA)技术研究了蚯蚓肠道内容物及喂养环境样品中细菌的多样性,研究结果表明：除含钾矿粉初始样品细菌的多样性很低外,喂养蚯蚓10d的含钾矿粉及在这些含钾矿粉喂养10d的蚯蚓肠道内容物、黄棕壤及黄棕壤中喂养10d的蚯蚓肠道内容物等四种样品中都具有很高的微生物多样性；蚯蚓肠道与蚯蚓生存环境虽共有一些相同种类的微生物,但各个样品细菌群落结构相差很大；γ-变形杆菌(Gammaproteobacteria)是蚯蚓肠道优势菌群,在蚯蚓肠道内对矿粉风化可能起重要作用；与含钾矿石风化相关的微生物菌落结构随环境变化而改变,含钾矿石的风化是由多种微生物共同作用的结果。
堆肥固态发酵是生物转化含钾岩石的有效技术。为获得堆肥固态发酵的有效菌种,我们对耐高温钾矿物溶解细菌进行了分离和筛选。从饲喂含钾矿粉的蚯蚓肠道内分离到了一株耐高温的钾矿物溶解菌EGT,16S rRNA基因序列和系统发育结果可以确定该菌为一种链霉菌(Streptomyces sp);采用固态发酵法用该链霉菌对以钾长石为主要成份的含钾矿粉进行发酵处理,ICP的检测结果表明用该链霉菌处理后的发酵物料水溶性Al、Fe和K的含量均明显高于接灭活菌未发酵和发酵14天的两个对照,其中比接灭活菌未发酵的对照分别高23.08%、123.19%和30.99%,比接接灭活菌发酵的对照分别高45.45%、52.48%和9.13%。上述结果说明该菌可以促进含钾矿物的风化。由于该链霉菌不但解钾,而且还耐高温,因此可以做为堆肥化处理含钾矿物制备钾肥的有效菌种。
为探讨含钾矿石的生物转化工艺及转化过程中的微生物多样性,我们采用先堆肥发酵再用蚯蚓处理的方法处理低品位含钾矿粉,然后用DGGE技术研究了该工艺过程中细菌群落结构的动态变化。堆肥有效钾含量的测定结果表明堆肥发酵和蚯蚓处理均能提高低品位含钾岩石粉的释钾量,其中经过堆肥发酵处理,有效钾含量提高了37.36%;经过蚯蚓处理,有效钾含量提高了9.77%；DGGE指纹图谱显示钾矿粉堆肥化处理过程中微生物群落结构变化很大。其中,堆肥固态发酵前期样品之间细菌群落结构差异较大,而堆肥固态发酵后期及加蚯蚓处理阶段样品之间细菌群落结构差异则相对较小；固态发酵前期堆肥中优势细菌种类变动较大,而后期优势细菌种类则比较稳定；在整个固态发酵阶段一直占据优势地位的细菌仅见条带12(可能属于交替单胞菌属Alteromonas)和14(壁厚菌门Firmicutes的一种细菌)所代表的细菌：加蚯蚓处理堆肥后,并未出现特殊的优势细菌。16SrRNA基因序列相似性和系统发育树聚类结果表明：DGGE指纹图上13个主要条带代表的细菌全部属于变形杆菌门(Proteobacteria)和壁厚菌门(Firmicutes),说明这两大类细菌在堆肥化处理过程中发挥主要作用。
含钾矿粉有机肥不仅含有丰富的活性钟,而且还有N、P等其他营养元素,将其作为底肥施用到农田后,不但会影响植物生长,而且也会影响植物根部内生细菌的组成和多样性。植物根部内生菌在帮助植物分解土壤矿物、摄取营养、抗病抑病及促进植物生长等方面发挥重要作用。苋菜(Amaranthus mangostanus L.)是一种富钾植物,其内生菌可能在协助苋菜风化土壤中的含钾矿物方面发挥重要作用。为探讨施含钾矿粉有机肥对苋菜内生细菌多样性的影响,我们用非培养的分子方法研究了在施含钾矿粉有机肥和不施肥苋菜根部内生细菌的群落结构。虽然两个文库挑选出的阳性克隆数差别不大,但不施肥苋菜根16S rRNA基因克隆文库内生细菌的OTUs数量和克隆数量远远大于施含钾矿粉有机肥苋菜根文库,说明不施肥苋菜根文库中代表内生细菌的克隆无论是种类和数量都远远大于施含钾矿粉有机肥的苋菜。与施含钾矿粉有机肥的苋菜根部内生细菌群落相比,γ-变形菌纲(Gammaproteobacteria)肠杆菌目(Enterobacteriales)和假单胞菌目(Pseudomonadales)的细菌在不施肥苋菜根部内生细菌群落占有绝对的优势,其中OTUs和克隆数量较大的主要有肠杆菌属(Enterobacter)、泛菌属(Pantoea)和假单胞属(Pseudomonas)。这些类群的细菌可能通过协助植物风化土壤矿物、固氮以及产生植物生长激素等途径在促进苋菜生长过程中扮演着重要角色。
上述研究不但解析了与蚯蚓促进含钾矿石风化相关的细菌群落结构,探明了堆肥固态发酵与加蚯蚓处理低品位含钾矿粉过程细菌群落的动态变化,还揭示了施用含钾矿粉有机肥后对苋菜生长及其根部内生菌多样性的影响,为进一步改进含钾岩石生物转化工艺,挖掘钾矿物溶解细菌资源,优化生物转化菌剂以及开发更高效的微生物钾肥奠定了理论基础。
Potassium is the third major macro nutrient for plant growth, but China is deficient in potassium fertilizer resources. China is fortunate to have rich deposit of potassium-bearing rock (PBR). If PBR could be converted into potassium fertilizer, it will provide enough potassium resource for China's agriculture. Microorganism play a key role during releasing potassium from PBR by bioconversion technology, so studying the bacterial diversities during bioconversion of PBR and understanding bio-weathering mechanism of PBR will be of benefit to improve bioconversion technology.
Feeding experiments were conducted to examine the accelerated degradation of potassium-bearing rock powders (PBRP) induced by earthworm. Results indicated some elements such as K, Al, Fe and Ca were released from the PBRP upon digestion by earthworm, and the contents of water-soluble K, Al, Fe and Ca in the PBRP weathered by earthworm respectively increased by5.64%,20.25%,27.52%,17.21%and its HNO3-extractable K Al, Fe and Ca respectively increased by6.59%,3.94%,4.36%and8.49%, thereby enhancing the degradation of PBRP. The microbial communities in earthworm's gut and the surrounding substrates including soil and PBRP were analyzed by ARDRA (Amplified Ribosomal DNA Restriction Analysis). The results indicated a higher bacterial diversity in the guts of the earthworms fed with PBRP and the PBRP fed to earthworm. The earthworms'gut and surrounding environment shared some bacteria, but bacterial communities showed very different in different stage. Some members from Gammaproteobacteria dominating in earthworm gut were likely to play an important role during degradation of mineral grains mediated by earthworm. The results also showed that the microbial communities relevant to mineral degradation varied with environmental conditions, but a large number of microorganisms played their role in the degradation process of PBRP.
Composting is an efficient technique to transform the potassium in PRRP into bioactive forms. To obtain high effective microorganisms which can solubilizing potassium from PBRP in compost, bacteria, tolerating high temperature, were screened. One mineral solubilizing bacterial strain designated EGT, which can be tolerant to high temperature, was isolated from the earthworm fed with PBRP. The sequence similarities and phylogenetic relationships based on16S rRNA gene showed that the strain belonged to the group of the genus Streptomyces. The strain was used as the agent releasing potassium from PBRP under solid state fermentation. Comparing to fresh substrate, the contents of water-soluble Al, Fe and K from the substrate inocubated by the strain respectively increased by23.08%、123.19%and30.99%. Comparing to the substrate inocubated by autoclaved inoculum, their contents respectively increased by45.45%,52.48%and9.13%. The increase of water-soluble elements indicated that the strain could weather PBRP. It may be as an effective inoculum of the compost which comprise PBRP and organic matter such as straw and cotton seed hull to obtain potassium fertilizer by composting.
The low-grade potassium-bearing rock powder (PBRP) was processed by composting to enhance the weathering of it for K-fertilizer's production. After compost was finished, the compost was treated again with earthworms'feeding to further increase its available potassium. The results revealed that the two processing could effectively accelerate the degradation of PBRP and increase the compost's available potassium. The dynamics of bacterial communities during composting and earthworm's feeding was investigated using DGGE technique. The profile of DGGE showed that the bacterial community of compost had a large difference in the earlier stage, but had a little difference in the latter stage. The dominant species of bacteria largely varied in the earlier stage of composting, but they were stable in the latter stage. Only band12and14, which belonged to Alteromonas and Firmicutes respectively, had been dominant during the whole course of composting. None of special dominant bacteria appeared after the compost was fed to earthworm. Selected bands were cut from the DGGE gel and then were sequenced after recycling and purifying. The similarities and phylogeny of the16S rRNA sequences indicated that the bacteria which the thirteen dominant bands represented were attributed only to Proteobacteria and Firmicutes. Therefore, the two phyla of bacteria may play an important role during composting of the material enriched by potassium-bearing rock powder.
The compost enriched by PBRP is abundant not only in bioactive potassium but also in nitrogen and phosphorus. Therefore, it would promote the growth of plants and affect its endophytic bacterial communities in root. The endophytic bacteria in plant root contributes to plant growth by dissolving soil mineral, providing nutrients for plant, protecting plant from disease and produing plant growth promoting molecules or substances. Amaranth (Amaranthus mangostanus L.) is a plant of rich potassium and might be effective in solubilizing mineral K of soil. Its endophytic bacteria in root may play an important role during amaranth releasing potassium from soil mineral. The compost enriched by PBRP was used as base fertilizer to amaranth which is a plant of rich potassium and might be effective in solubilizing mineral K of soil. The endophytic bacteria in amaranth root may play an important role during amaranth releasing potassium from soil mineral. The endophytic bacterial communities of amaranth's root fertilized by the compost and its control to which no fertilizer was applied were investigated through16S rRNA gene cloning library technology and and ARDRA. The number of clones in two libraries were similar, but the number of endophytic bacteria from library of the amaranth without fertilizer were much higher than those of the amaranth fertilized by the compost enriched by PBRP. The result showed that the former's endophytic bacteria were much richer than the latter's. Comparing to those of the amaranth fertilized by compost, some bacteria from Gammaproteobacteria were dominant in the root of the amaranth no fertilizer was applied. Among these bacteria of Gammaproteobacteria, main members were from the genera Enterobacter, Pantoea and Pseudomonas. These bacteria may play an important role in promoting amaranth's growth by releasing elements from soil, fixing nitrogen and producing growth hormones.
These studies not only revealed the bacterial communities associated with weathering of PBRP and the dynamics of bacterial communities in the compost enriched by PBRP during composting and feeding earthworms, but also showed the changes of amaranth's growth and its endophytic bacteria in root after providing the compost enriched by PBRP as a base fertilizer to it. The results will be useful in helping us improve the technology of bioconversion of PBRP, find new the resources of mineral potassium-solubilizing bacteria and produce more effective potassium bio-fertilizer for agriculture.
引文
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