瘤胃氨同化通路及基因进化研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2024-0764

摘要/Abstract

摘要：

蛋白质饲料资源的短缺严重限制着我国畜牧业的持续稳定发展。反刍动物具有将非蛋白氮转化为氨基酸或蛋白质的独特能力，添加非蛋白氮是缓解反刍动物蛋白饲粮短缺的有效途径，但瘤胃氨同化是限制其利用效率的关键步骤。因此，如何提高反刍动物氮同化效率是缓解蛋白饲粮短缺的重要手段。氨同化指游离氨并入碳骨架形成含氮有机物的过程。瘤胃微生物可以将游离的氨同化为微生物蛋白质（microbial protein, MCP），为反刍动物提供49%-98%的可代谢蛋白质。瘤胃氨同化作用可通过4条通路进行：①谷氨酸脱氢酶（glutamate dehydrogenase, GDH）：氨与α-酮戊二酸在GDH的作用下发生胺化生成谷氨酸；②谷氨酰胺合成酶-谷氨酸合成酶（glutamine synthetase-glutamate synthase, GS-GOGAT）：氨和谷氨酸经GS催化后形成谷氨酰胺；③丙氨酸脱氢酶（alanine dehydrogenase, ADH）：氨和丙酮酸可经过ADH通路生成丙氨酸；④天冬酰胺合成酶（asparagine synthetase, AS）：天冬氨酸和氨可通过AS通路发生酰胺化生成天冬酰胺。本文深入探究了瘤胃微生物氨同化通路反应过程，综述了瘤胃4条氨同化通路过程及其关键微生物、酶和影响因素，并通过构建系统发育树解析了瘤胃细菌不同氨同化基因的进化关系，为反刍动物氨氮高效利用提供理论支持。

关键词: 反刍动物, 非蛋白氮, 氨同化通路, GDH通路, GS-GOGAT通路

Abstract:

The shortage of protein feed resources has severely restricted the sustained and stable development of animal husbandry in China. Ruminants possess the special ability to convert non-protein nitrogen into amino acids or proteins. Adding non-protein nitrogen is an effective approach to alleviate protein feed shortages in ruminants, but rumen ammonia assimilation is the key step limiting its utilization efficiency. Therefore, improving the nitrogen assimilation efficiency of ruminants is an important means of alleviating the shortage of protein feed. Ammonia assimilation refers to the process of free ammonia merging into the carbon skeleton to form nitrogen-containing organic compounds. Rumen microorganisms may assimilate free ammonia into microbial protein (MCP), providing 49%-98% of metabolizable protein for ruminants. Rumen ammonia assimilation can be carried out through four pathways: ① the glutamate dehydrogenase (GDH) pathway: ammonia and α-ketoglutarate undergo aminification catalyzed by GDH to produce glutamate; ② the glutamine synthetase-glutamate synthase (GS-GOGAT) pathway: ammonia and glutamate are catalyzed by GS to form glutamine;③ the alanine dehydrogenase (ADH) pathway: alanine can be produced by ammonia and pyruvate through the ADH pathway; ④ the asparagine synthetase (AS) pathway: aspartate and ammonia can form asparagine through the AS pathway. This article explores the process of ammonia assimilation pathway reaction in rumen microorganisms and reviews these four ammonia assimilation pathways of rumen microbes, their key enzymes, microorganisms involved, and influencing factors. By constructing phylogenetic trees to explain the evolutionary relationships of key genes, theoretical support for the utilization of ammonia nitrogen in ruminants is provided.

Key words: ruminants, non-protein nitrogen, ammonia assimilation pathway, GDH pathways, GS-GOGAT pathways

王彤, 肖汉杰, 谢昊炅, 张立申, 姚晓亮, 严慧, 纪守坤. 瘤胃氨同化通路及基因进化研究进展[J]. 生物技术通报, 2025, 41(2): 40-50.

WANG Tong, XIAO Han-jie, XIE Hao-jiong, ZHANG Li-shen, YAO Xiao-liang, YAN Hui, JI Shou-kun. Research Progress in Ammonia Assimilation and Genetic Evolution in Rumen[J]. Biotechnology Bulletin, 2025, 41(2): 40-50.

图/表 11

图1 GDH通路图

Fig. 1 Pathway of GDH

图2 GDH结构图A：GDH六聚体和亚基结构示意图，基于Butyrivibrio fibrisolvens全基因组数据（NCBI序列号：GCA_037113525.1），利用AlphaFold3平台构建；B：底物与辅酶结合后GDH构象变化示意图

Fig. 2 Structure of GDHA: Cartoon representation of hexameric and subunit GDH structure. The structure is built based on the whole genome of B. fibrisolvens using the AlphaFold3 platform (NCBI GenBank: GCA_037113525.1). B: Schematic diagram depicting conformational change in GDH structure upon substrate and coenzyme binding

图3 GS-GOGAT通路图

Fig. 3 Pathway of GS-GOGAT

图4 三类GS结构图来自PDB数据库，序列号分别为5ZLP、7v4i、8TGE

Fig. 4 Multimeric structures of the three types of GSThe structures are from the PDB database, with serial number 5ZLP, 7v4i, and 8TGE, respectively

图5 GOGAT结构图A：Fd-GOGAT结构示意图，结构源于PDB（1LLZ）；B：NADPH-GOGAT结构示意图，基于B. fibrisolvens（NCBI GenBank： GCA_037113525.1）全基因组数据，利用AlphaFold3平台预测

Fig. 5 Structure of GOGATA: Schematic diagram of Fd-GOGAT structure, from PDB (1 LLZ). B: Schematic diagram of NADPH-GOGAT structure. Structure based on B. fibrisolvens genome data (NCBI GenBank: GCA_037113525.1), predicted by AlphaFold3

图6 ADH通路图

Fig. 6 Pathway of ADH

图7 ADH结构图A：ADH六聚体结构图，不同颜色代表不同亚基；B：单个亚基结构示意图，其中球棒状模型代表NAD＋，红色六边形代表丙酮酸。图中所有结构均以B. fragilis全基因组数据为基础，利用AlphaFold3平台预测蛋白结构（NCBI序列号：GCA_016889925.1）

Fig. 7 Structure of ADHA: The structural diagram of ADH hexamer, with different colors representing different subunits. B: The structural diagram of ADH monomer, where the ball-and-stick model indicates NAD+ and the red hexagon indicates pyruvate. The structure in the figure is based on the whole genome data of B. fragilis, and protein structure prediction was conducted using the AlphaFold3 platform (NCBI GenBank: GCA_016889925.1)

图8 AS通路图

Fig. 8 Pathway of AS

图9 AS结构图A：AS-B单个亚基结构图；B：AS-B二聚体形式结构图；C：AS-A二聚体形式结构图。图中结构均基于B. fibrisolvens全基因组数据（NCBI GenBank：GCA_037113525.1），利用AlphaFold3平台构建

Fig. 9 Structure of ASA: The structural diagram of AS-B monomer. B: Structure diagram of AS-B dimer. C: Structure diagram of AS-A dimer. The structures are all built based on the whole genome data of B. fibrisolvens (NCBI GenBank: GCA_037113525.1) using the AlphaFold3 platform

图10 瘤胃细菌4条氨同化通路关系图

Fig. 10 Relationship diagram of four ammonia assimilation pathways in rumen bacteria

图11 16SrRNA和氨同化通路关键基因系统发育树A：31株瘤胃细菌的16S rRNA系统发育树；B：gdhA基因系统发育树（GDH通路）；C：glnA基因系统发育树（GS-GOGAT通路）；D：ald基因系统发育树（ADH通路）；E和F分别为asnA和asnB基因系统发育树（AS通路）。B-F中的虚线为进化树分支的基因分型位置。基于HMMER检索同源基因，使用MEGA 11 最大似然法构建进化树，bootstrap值设为100

Fig. 11 Phylogenetic trees of 16S rRNA and key genes in the ammonia assimilation pathwayA: 16S rRNA phylogenetic tree of 31 rumen bacterial strains. B: Phylogenetic tree of gdhA gene (GDH pathway). C: Phylogenetic trees of glnA gene (GS-GOGAT pathway). D: Phylogenetic tree of ald gene (ADH pathway); E and F is phylogenetic trees of asnA and asnB genes, respectively (AS pathway). The dashed lines in Fig. B-F indicate the location of genotyping. Homologous genes were retrieved using HMMER. Trees are constructed using MEGA 11 with maximum likelihood method. Bootstrap values are 100

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摘要

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