浓香型大曲后熟过程中的真菌群落演替及其组装规律

doi:10.13560/j.cnki.biotech.bull.1985.2025-1459

摘要/Abstract

摘要：

目的解析不同等级浓香型大曲在后熟过程中（0-90 d）的发酵性能变化、真菌群落构成及其动态演替规律，揭示其真菌群落的组装机制。方法基于高通量测序技术跟踪不同等级（优级、一级和二级）大曲在后熟期间的真菌群落动态变化，探究大曲真菌群落与生化指标之间的相关性，利用中性模型与零模型分析大曲后熟期间的真菌群落组装规律。结果在后熟期间，大曲的酯化力、糖化力、发酵力和纤维素酶活性显著升高（P<0.05），且优级曲显著高于一级和二级曲。嗜热子囊菌（44.18%-54.51%）、嗜热真菌（37.88%-49.42%）、曲霉（1.69%-6.31%）既是3个等级大曲共有的优势菌属，也是区分大曲等级和后熟阶段的关键差异类群。进一步发现，大曲生化指标与真菌菌群之间具有显著相关性。其中，酯化力与毕赤酵母属（Pichia）呈显著正相关，与丝衣霉属（Byssochlamys）、威克汉姆酵母属呈极显著负相关。糖化力、发酵力与嗜热真菌属呈显著正相关，与丝衣霉属呈显著负相关。纤维素酶、蛋白酶（酸性和碱性）与假丝酵母属（Candida）呈显著正相关，与威克汉姆酵母属呈显著负相关。中性模型和零模型分析表明，不同等级大曲在后熟期间的真菌群落动态演替主要受随机性过程影响，且主要由漂变和扩散限制导致菌群结构发生随机变化。结论优级大曲的真菌群落结构更趋稳定，且生化指标相对更优越，尤其后熟3个月后的大曲特性更趋理想水平，可以投入酿酒生产。

关键词: 浓香大曲, 等级差异, 后熟过程, 生化指标, 真菌群落特征, 演替规律

Abstract:

Objective This study investigates the changes in fermentation performance, fungal community composition, and dynamic succession patterns of different grades of Nongxiang Daqu during the post-ripening period (0–90 days), aiming to elucidate the assembly mechanisms of the fungal community. Method We used high-throughput sequencing technology to track the dynamic succession of fungal communities in different grades of Daqu (premium, first-grade, and second-grade) during post-ripening, and to explore the correlations between these communities and biochemical indicators. We also applied neutral and null models to analyze the assembly patterns of the fungal communities during the post-ripening process. Result During post-ripening, the esterification capacity, saccharification capacity, fermentation capacity, and cellulase activity of Daqu increased significantly (P<0.05), with premium-grade Daqu exhibiting significantly higher values than first-grade and second-grade Daqu. Thermoascus (44.18%–54.51%), Thermomyces (37.88%–49.42%), and Aspergillus (1.69%–6.31%) were not only the dominant fungal genera shared by three Daqu grades but also served as key differential taxa that differentiated between Daqu grades and post-ripening stages. Moreover, we observed significant correlations between biochemical indicators of Daqu and fungal communities. Specifically, esterification capacity showed a significant positive correlation with Pichia and a highly significant negative correlation with Byssochlamys and Wickerhamomyces. Saccharification and fermentation capacities were significantly positively correlated with Thermomyces and significantly negatively correlated with Byssochlamys. Cellulase, acidic protease, and alkaline protease activities were significantly positively correlated with Candida and significantly negatively correlated with Wickerhamomyces. Neutral and null model analyses reveal that stochastic processes predominantly affect the dynamic succession of fungal communities in different grades of Daqu during post-ripening, with drift and dispersal limitation serving as the primary drivers of stochastic changes in community assembly. Conclusion The fungal community structure of premium Daqu is more stable, and its biochemical indicators are superior. Particularly after three months of post-ripening, the characteristics of Daqu approach an ideal state, making it suitable for liquor production.

Key words: Nongxiang Daqu, grade difference, post-ripening process, biochemical indicators, fungal community characteristics, succession pattern

张如敏, 张博, 林杰豪, 乔晶, 赵洪源, 郭晓鹏, 任海伟. 浓香型大曲后熟过程中的真菌群落演替及其组装规律[J]. 生物技术通报, 2026, 42(5): 203-212.

ZHANG Ru-min, ZHANG Bo, LIN Jie-hao, QIAO Jing, ZHAO Hong-yuan, GUO Xiao-peng, REN Hai-wei. Fungal Community Succession and Assembly Mechanisms during the Post-ripening Process of Nongxiang Daqu[J]. Biotechnology Bulletin, 2026, 42(5): 203-212.

图/表 7

图1 不同等级大曲后熟过程中的生化指标P、F和S分别代表大曲等级为优级、一级和二级；0、45、90分别代表后熟0 d、后熟45 d、后熟90 d。不同大写字母表示同一等级不同后熟期大曲差异显著（P＜0.05）；不同小写字母表示同一后熟期不同等级大曲差异显著（P＜0.05），下同

Fig. 1 Biochemical indicators of different grades of Daqu during the post-ripening processP, F, and S refer to premium, first, and second grades of Daqu, respectively. 0, 45, and 90 refer to post-ripening for 0 d, 45 d, and 90 d, respectively. Different capital letters indicate significant differences (P<0.05) between Daqu of the same grade at different post-ripening stages, while different lowercase letters indicate significant differences (P<0.05) between Daqu of different grades at the same post-ripening stage, the same below

图2 不同等级大曲后熟过程中真菌群落的α多样性分析

Fig. 2 Analysis of fungal community alpha diversity during the post-ripening process of different grades of Daqu

图3 不同等级大曲真菌OTU水平组成相似性（A）和主成分分析（B）

Fig. 3 Similarity of fungal OTU composition (A) and principal component analysis (B) in different grades of Daqu

图4 不同等级大曲后熟过程中真菌门水平（A）和属水平（B）物种组成情况

Fig. 4 Species composition of different grades of Daqu during the post-ripening process at phylum(A) and genus (B)

图5 不同等级大曲后熟过程中真菌属水平LDA柱形图

Fig. 5 Bar chart of LDA at the fungi genus level for different grades of Daqu in the post-ripening process

图6 大曲优势真菌属与生化指标（A）及与优势细菌属之间（B）的相关性分析相关性均采用Pearson方法计算，*P<0.05，**P<0.01。图B中细菌丰度数据来源于课题组已发表成果［15］

Fig. 6 Correlation of dominant fungal genera with biochemical indicators(A) and dominant bacterial genera(B) in DaquCorrelations were calculated with Pearson’s method, * P<0.05, and ** P<0.01. The bacterial abundance data in Figure B were derived from the published results of our research group [15]

图7 不同等级和后熟期大曲的真菌中性模型（A-F）和零模型（G、H）A：0 d；B：45 d；C：90 d；D：优级曲；E：一级曲；F：二级曲。蓝色实线表示中性模型的最佳拟合，虚线表示模型预测的95%置信区间。Rsqr值代表中性群落模型的整体拟合优度，Rsqr值越高表明微生物群落受随机性过程的影响越大，受确定性过程的影响越小

Fig. 7 Neutral models (A-F) and null models (G, H) of fungi in Daqu of different grades and ripening periodsA: 0 d; B: 45 d; C: 90 d; D: premium grade of Daqu; E: first grade of Daqu; F: second grade of Daqu. The solid blue line indicates the best fit of the neutral model, and the dashed lines represent the 95% confidence interval of the model prediction. The Rsqr value represents the overall goodness-of-fit of the neutral community model. A higher Rsqr value indicates a greater influence of stochastic processes and a lesser influence of deterministic processes on the microbial community

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