硅酸盐矿物与微生物相互作用对于矿物风化、土壤形成、元素生物地球化学循环和长时间尺度大气CO2的固定有重要的作用，细菌对硅酸盐矿物的风化效应与机制研究已有不少报道。然而有关不同硅酸盐矿物与细菌相互作用机制的研究还比较薄弱。本研究进一步开展了硅酸盐矿物与矿物风化细菌相互作用机制的研究，研究结果对矿物与微生物相互作用机制的深入认识有重要的意义。

本文以黑云母和钾长石为供试矿物，以本课题组保藏的一株高效矿物风化细菌Pseudomonas azotoformans ZY-1作为供试菌株，开展了不同硅酸盐矿物和不同粒径大小矿物对菌株ZY-1生长代谢和矿物风化相关基因表达的影响；基于代谢组学技术分析了黑云母和钾长石对菌株ZY-1分泌代谢产物的影响；通过RT-qPCR和基因敲除技术验证了菌株ZY-1中与矿物风化相关的基因功能。本文主要结果如下：

（1）硅酸盐矿物对菌株ZY-1生长代谢及矿物风化相关基因表达的影响

BHm培养基中20 h时添加黑云母和钾长石发酵液中细胞数量比不添加矿物增加了26%-71%，添加黑云母的发酵液中细胞数量比添加钾长石的增加了35%。添加钾长石的发酵液pH值与不添加矿物无显著差异，添加黑云母的发酵液pH值显著高于添加钾长石。添加黑云母和钾长石后，菌株ZY-1葡萄糖脱氢酶基因（gcd）相比与无矿物上调了1.53-3.43倍，黏附素基因（adh）相比与无矿物上调了1.29-1.50倍。添加黑云母后菌株ZY-1葡萄糖脱氢酶基因（gcd）相比与添加钾长石上调了2.24倍，黏附素基因（adh）下调了1.16倍。

（2）菌株ZY-1风化黑云母和钾长石的机制研究——基于代谢组学分析

菌株发酵液中共鉴定到799种代谢物质，主要包括脂类和类脂分子、有机酸及其衍生物、有机杂环化合物、有机氧化物、苯环型化合物等；添加黑云母后，显著改变的代谢途径包括辅因子的生物合成、色氨酸代谢、嘌呤代谢、谷胱甘肽代谢等；添加钾长石后，显著改变的代谢途径包括精氨酸生物合成、赖氨酸生物合成、戊糖和葡萄糖醛酸的相互转化、脂肪酸生物合成等；黑云母组与钾长石组相比，显著改变的代谢途径包括嘌呤代谢、酪氨酸代谢、辅因子的生物合成、苯丙氨酸代谢等；前二十丰度的代谢物较多与葡萄糖代谢产物有关，如D-葡萄糖、D-葡萄糖酸、2-酮-D-葡萄糖酸、D-葡萄糖庚糖；添加黑云母后，N-乙酰丙氨酸、苯丙氨酰脯氨酸、γ-谷氨酰半胱氨酸等代谢物质发生了上调；添加钾长石后，S-亚硝基谷胱甘肽、天冬氨酰-色氨酸、N-癸酰甘氨酸等物质发生了上调；添加黑云母与添加钾长石相比，N-乙酰-L-蛋氨酸、半乳糖酸、5-羟基吲哚乙酰甘氨酸等物质发生了上调；添加黑云母与添加钾长石这两组比较中，共有的上调代谢物是N-乙酰-L-蛋氨酸、半乳糖酸、D-2-羟基戊二酸等物质；上述三组比较中同时上调的代谢物质为5-羟基吲哚乙酰甘氨酸；KEGG代谢通路和基因组查找，共找到不同比较中上调的与矿物风化相关的8种代谢物、9种代谢途径、14种相关的酶和基因。

（3）代谢组学中矿物风化相关基因功能验证

RT-qPCR分析表明，添加黑云母后，菌株ZY-1 argD（编码乙酰鸟氨酸氨基转移酶）、cysE（编码L-丝氨酸O-乙酰转移酶）、ilvE（编码支链氨基酸氨基转移酶）基因分别上调了1.30倍、2.30倍、2.85倍；添加钾长石后，菌株ZY-1 argD、phhA-PAH （编码苯丙氨酸4-单加氧酶）、orf00874_1（编码醛脱氢酶）、cysE基因分别上调了4.35倍、1.44倍、1.22倍、1.45倍，黑云母与钾长石组比较中，基因ilvE、cysE分别上调了4.32倍、1.59倍；利用同源重组技术构建突变株ZY-1ΔargD、ZY-1ΔphhA-PAH、ZY-1Δorf00874_1、ZY-1ΔcysE、ZY-1ΔilvE；菌株风化黑云母效果显示，突变株ZY-1ΔargD、ZY-1ΔphhA-PAH、ZY-1Δorf00874_1与野生株相比，发酵液中铁浓度降低了19%-32%，铝浓度降低了17%-27%；菌株风化钾长石效果显示，突变株ZY-1ΔargD、ZY-1ΔphhA-PAH、ZY-1ΔilvE与野生株相比，发酵液中铁浓度降低了31%-42%，铝浓度降低了17%-28%。

本文主要结论：（1）不同硅酸盐矿物和不同粒径大小的矿物对菌株ZY-1生长代谢、风化矿物及基因表达都有不同的影响；（2）添加黑云母和钾长石改变了菌株ZY-1与矿物风化相关的代谢通路和代谢产物；（3）乙酰鸟氨酸氨基转移酶基因argD和苯丙氨酸4-单加氧酶基因phhA-PAH对菌株ZY-1风化黑云母和钾长石有着重要影响，醛脱氢酶基因orf00874_1和支链氨基酸氨基转移酶基因ilvE分别对菌株ZY-1风化黑云母和钾长石有着重要影响。

The interaction between silicate minerals and microorganisms plays an important role in mineral weathering, soil formation, element biogeochemistry cycle and long-term atmospheric CO2 fixation. There have been many reports on the weathering effect and mechanism of bacteria on silicate minerals. However, research on the interaction mechanism between different silicate minerals and bacteria is still relatively weak. This study further investigated the interaction mechanism between silicate minerals and mineral weathering bacteria, and the research results are of great significance for a deeper understanding of the interaction mechanism between mineral and microorganisms.

This article uses biotite and potassium feldspar as the test minerals, and a highly efficient mineral weathering bacterium Pseudomonas azotoformans ZY-1 as the test strain, which is preserved by our research group. The effects of different silicate minerals and minerals with different particle sizes on the growth metabolism and mineral weathering related gene expression of strain ZY-1 were studied; Based on metabolomics techniques, the effects of biotite and potassium feldspar on the metabolites of strain ZY-1 were analyzed; The gene function related to mineral weathering in strain ZY-1 was verified through RT-qPCR and gene knockout techniques. The main results of this article are as follows:

(1) Effects of silicate minerals on the growth metabolism and weathering related genes of strain ZY-1

At 20 hours in BHm culture medium, the number of cells in the fermentation broth with biotite and potassium feldspar added increased by 26%-71% compared to without minerals. The number of cells in the fermentation broth with biotite added increased by 35% compared to that with potassium feldspar added. The pH value of the fermentation broth with and without the addition of potassium feldspar showed no significant difference, while the pH value of the fermentation broth with the addition of biotite was significantly higher than that with the addition of potassium feldspar. After adding biotite and potassium feldspar, the glucose dehydrogenase gene (gcd) of strain ZY-1 was up-regulated 1.53-3.43 times compared to no minerals, and the adhesin gene (adh) was up-regulated 1.29-1.50 times compared to no minerals. After adding biotite, the glucose dehydrogenase gene (gcd) of strain ZY-1 was up-regulated by 2.24 times compared to adding potassium feldspar, and the adhesin gene (adh) was down-regulated by 1.16 times.

(2) Mechanism study on the weathering of biotite and potassium feldspar by strain ZY-1 based on metabolomics analysis

A total of 799 metabolic substances were identified in the fermentation broth of the strain, mainly including lipids and lipid molecules, organic acids and their derivatives, organic heterocyclic compounds, organic oxides, benzene ring compounds, etc; After adding biotite, the metabolic pathways significantly changed include cofactor biosynthesis, tryptophan metabolism, purine metabolism, glutathione metabolism, etc; After the addition of potassium feldspar, the significantly altered metabolic pathways include arginine biosynthesis, lysine biosynthesis, mutual conversion of pentose and glucuronic acid, and fatty acid biosynthesis; Compared with potassium feldspar group, biotite group has significantly changed metabolic pathways including purine metabolism, tyrosine metabolism, cofactor biosynthesis, phenylalanine metabolism, etc; The first twenty abundances of metabolites are mostly related to glucose metabolites, such as D-glucose, D-gluconic acid, 2-keto-D-gluconic acid, and D-glucoheptose; After adding biotite, some metabolic substances are up-regulated such as N-acetylalanine, phenylalanylproline, γ-glutamylcysteine; After adding potassium feldspar, some metabolic substances are up-regulated such as S-nitrosoglutathione, aspartyl-tryptophan, N-decanoylglycine; Compared with potassium feldspar, after adding biotite, N-acetyl-L-methionine, galactose acid, 5-hydroxyindole acetylglycine and other substances were up-regulated; In the comparison between biotite and potassium feldspar, the common up-regulated metabolites are N-acetyl-L-methionine, galactose acid, D-2-hydroxyglutaric acid, etc; The metabolic substance up-regulated simultaneously in the above three groups of comparisons is 5-hydroxyindole acetylglycine; KEGG metabolic pathway and genome search, a total of 8 up-regulated metabolites related to mineral weathering, 9 metabolic pathways, 14 related enzymes and genes were found in different comparisons.

(3) Functional verification of genes related to mineral weathering in metabonomics

RT-qPCR analysis showed that the genes of ZY-1 argD (encoding acetylornithine aminotransferase), cysE (encoding L-serine O-acetyltransferase), and ilvE (encoding branched chain amino acid aminotransferase) were up-regulated by 1.30, 2.30, and 2.85 times respectively after biotite addition; After adding potassium feldspar, strains ZY-1 argD, phhA-PAH (encoding phenylalanine 4-monooxygenase), orf00874_1 (encoding aldehyde dehydrogenase) and cysE genes were up-regulated 4.35, 1.44, 1.22, and 1.45 times respectively. In comparison with biotite and potassium feldspar groups, genes ilvE and cysE were up-regulated 4.32 and 1.59 times respectively; Mutant ZY-1ΔargD, ZY-1ΔphhA-PAH, ZY-1Δorf00874_1, ZY-1ΔcysE, ZY-1ΔilvE were structured using homologous recombination technology; The weathered biotite effect of the strain showed that the mutant ZY-1ΔargD, ZY-1ΔphhA-PAH, ZY-1Δorf00874_1 of iron concentration in the fermentation broth decreased by 19%-32%, and the aluminum concentration decreased by 17%-27% compared to wild strains; The weathered potassium feldspar effect of the strain showed that the mutant ZY-1ΔargD, ZY-1ΔphhA-PAH, ZY-1ΔilvE of iron concentration in the fermentation broth decreased by 31%-42%, and the aluminum concentration decreased by 17%-28% compared to wild strains.

The main conclusions of this article are as follows: (1) Different silicate minerals and minerals with different particle sizes have different effects on the growth metabolism, weathered minerals, and gene expression of strain ZY-1; (2) The addition of biotite and potassium feldspar altered the metabolic pathways and metabolic substances related to strain ZY-1; (3) The acetylornithine aminotransferase gene argD and phenylalanine 4-monooxygenase gene phhA-PAH have important effects on the weathering biotite and potassium feldspar of strain ZY-1, and the aldehyde dehydrogenase gene orf00874_1 and the branched chain amino acid aminotransferase gene ilvE have significant effects on the weathering biotite and potassium feldspar of strain ZY-1, respectively.

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