食品工业科技

差异比较组名称	差异代谢物数目	下调代谢物数目	上调代谢物数目
DR45 vs. DR40	1071	314	757
DR50 vs. DR40	1219	226	993
DR50 vs. DR45	1077	373	704
DR55 vs. DR40	1210	315	895
DR55 vs. DR45	629	192	437
DR55 vs. DR50	577	328	249

差异比较组名称	差异代谢物数目	下调代谢物数目	上调代谢物数目
DR45 vs. DR40	1071	314	757
DR50 vs. DR40	1219	226	993
DR50 vs. DR45	1077	373	704
DR55 vs. DR40	1210	315	895
DR55 vs. DR45	629	192	437
DR55 vs. DR50	577	328	249

不同烘干温度的黔产红托竹荪非靶向代谢组学研究

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郭银萍 , 黄晓润 , 龚光禄 , 刘宏宇 , 黄万兵 , 卢颖颖 , 刘倾城 , 杨通静 ^*

食品工业科技 | 分析检测 2026,47(10): 322-334

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食品工业科技 |分析检测 2026 , 47 (10) : 322 -334

不同烘干温度的黔产红托竹荪非靶向代谢组学研究

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郭银萍（1995−），女，硕士，研究实习员，研究方向：食用菌栽培与加工，E-mail：1797415307@qq.com

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郭银萍（1995−），女，硕士，研究实习员，研究方向：食用菌栽培与加工，E-mail：1797415307@qq.com

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郭银萍, 黄晓润, 龚光禄, 刘宏宇, 黄万兵, 卢颖颖, 刘倾城, 杨通静^*

作者信息

贵州省农业科学院农作物品种资源研究所，贵州省农业微生物全省重点实验室，贵州省食用菌育种重点实验室，贵州贵阳 550006

通讯作者:

杨通静（1984−），男，硕士，副研究员，研究方向：食用菌栽培，E-mail：736932018@qq.com。

作者简介:

郭银萍（1995−），女，硕士，研究实习员，研究方向：食用菌栽培与加工，E-mail：1797415307@qq.com

Non-targeted Metabolomics Study of Dictyophora rubrovalvata from Guizhou at Different Drying Temperatures

Yinping GUO, Xiaorun HUANG, Guanglu GONG, Hongyu LIU, Wanbing HUANG, Yingying LU, Qingcheng LIU, Tongjing YANG^*

Affiliations

Institute of Crop Variety Resources, Guizhou Academy of Agricultural Sciences, Guizhou Provincial Key Laboratoryof Agricultural Microbiology, Guizhou Provincial Key Laboratory of Edible Fungi Breeding, Guiyang 550006, China

出版时间: 2026-05-15 doi: 10.13386/j.issn1002-0306.2025040259

文章导航

摘要

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红托竹荪（Dictyophora rubrovalvata，DR）作为贵州特色珍稀食用菌，其干品加工中干燥温度对功能成分保留的影响是产业升级的关键问题之一。本研究以40、45、50、55 ℃空气源热泵干燥至恒重的黔产红托竹荪（分别标记为DR40、DR45、DR50、DR55）为试材，采用液相色谱-串联质谱（Liquid Chromatography-Mass Spectrometry，LC-MS）非靶向代谢组学技术，结合主成分分析（Principal Components Analysis，PCA）、正交偏最小二乘判别分析（Orthogonal Partial Least Squares Discriminant Analysis，OPLS-DA）、KEGG通路富集及差异丰度得分（DA Score）分析，系统解析不同干燥温度下代谢物动态变化及代谢网络响应。结果表明，共检测到4470种代谢物，主要分布于脂质、脂肪酰类、甾体等21个大类（占比88.46%）；40~45 ℃干燥时代谢组稳定性高，组间差异代谢物仅1071个（DR45 vs. DR40），且上调（757个）与下调（314个）代谢物数量差异较小；当温度升至50 ℃时，差异代谢物增至1219个（DR50 vs. DR40），上调代谢物（993个）显著多于下调（226个），反映代谢系统通过激活保护通路维持稳态；50~55 ℃干燥时，差异代谢物骤降至577个（DR55 vs. DR50），且下调代谢物（328个）反超上调（249个），提示高温突破阈值后代谢系统受损。KEGG富集分析显示，高温（≥50 ℃）显著影响脂质、黄酮及抗氧化相关代谢通路，代谢网络发生重构。研究确认红托竹荪干燥存在约50 ℃的温度敏感阈值，40~45 ℃干燥能较好保留原始代谢特征。本研究为红托竹荪工业化干燥工艺温控优化提供了理论依据，建议生产中控制干燥温度低于50 ℃以减少功能成分流失。

关键词

红托竹荪 / 干燥温度 / 非靶向代谢组学 / 代谢通路 / 代谢物

Abstract

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As a rare and valuable edible fungus native to Guizhou Province, challenges in industrial upgrading due to the impact of drying temperatures on the retention of functional components during dry product processing were faced by Dictyophora rubrovalvata (DR). In this study, the investigation on air-source heat pump-dried DR samples at 40, 45, 50, and 55 ℃ (labeled as DR40, DR45, DR50, and DR55, respectively) was conducted using liquid chromatography-mass spectrometry (LC-MS)-based non-targeted metabolomics. By using principal component analysis (PCA), orthogonal partial least squares discriminant analysis (OPLS-DA), KEGG pathway enrichment analysis, and differential abundance (DA) score analysis, dynamic metabolic changes and network responses under varying drying temperatures were systematically characterized. The detection of a total of 4470 metabolites was conducted, primarily distributed across 21 major classes (88.46% of total), including lipids, fatty acyls, and steroids. At 40~45 ℃, metabolic stability was high, with only 1071 differential metabolites (DR45 vs. DR40), in which a small disparity between upregulated (757) and downregulated (314) metabolites was shown. At 50 ℃, differential metabolites increased to 1219 (DR50 vs. DR40), with significantly more upregulated (993) than downregulated (226) metabolites, indicating metabolic system activation to maintain homeostasis. At 50~55 ℃, differential metabolites sharply decreased to 577 (DR55 vs. DR50), with downregulated metabolites (328) surpassing upregulated ones (249), so metabolic system damage beyond the temperature threshold was presented. From KEGG enrichment analysis, high temperatures (≥50 ℃) were shown to significantly affect lipid, flavonoid, and antioxidant-related pathways, so metabolic network reconstruction was caused. Based on the study, a temperature-sensitive threshold of approximately 50 ℃ for DR drying was confirmed, with 40~45 ℃ preserving original metabolic characteristics, thus offering a theoretical basis for optimizing industrial drying processes, recommending temperatures below 50 ℃ to minimize functional component loss.

Key words

Dictyophora rubrovalvata / drying temperature / untargeted metabolomics / metabolic pathways / metabolites

引用本文

郭银萍, 黄晓润, 龚光禄, 刘宏宇, 黄万兵, 卢颖颖, 刘倾城, 杨通静. 不同烘干温度的黔产红托竹荪非靶向代谢组学研究. 食品工业科技, 2026 , 47 (10) : 322 -334 . DOI: 10.13386/j.issn1002-0306.2025040259

Yinping GUO, Xiaorun HUANG, Guanglu GONG, Hongyu LIU, Wanbing HUANG, Yingying LU, Qingcheng LIU, Tongjing YANG. Non-targeted Metabolomics Study of Dictyophora rubrovalvata from Guizhou at Different Drying Temperatures[J]. Science and Technology of Food Industry, 2026 , 47 (10) : 322 -334 . DOI: 10.13386/j.issn1002-0306.2025040259

正文

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红托竹荪（Dictyophora rubrovalvata，DR）隶属鬼笔科竹荪属真菌，是一种极具价值的珍稀食用菌^[1]。其主要生长于我国西南地区，尤其以贵州为代表产区，菌丝体发育过程对竹阔混交林腐殖质基质具有严格生态适应性^[2]。化学分析表明，红托竹荪富含蛋白质、多糖、氨基酸、维生素以及矿物质等营养成分^[3]。其中，多糖成分表现出显著的抗氧化、抗肿瘤、抗疲劳以及免疫调节等生物活性^[4]，醇提物还表现出降血脂、降血糖等多方面的保健潜力^[5]。2025年贵州省将红托竹荪纳入全省“3+5”特色产业布局，推动其产业化发展。然而，由于其鲜品水分活度高，采后易腐败^[6]，干制成为主要加工方式。干燥过程中温度参数对功能成分保留具有显著影响，高温易导致热敏成分降解，影响复水性、口感及风味品质，成为制约产业升级的关键因素^[7]。

目前，红托竹荪干制工艺多采用热风、真空红外、空气源热泵等技术^[8]，相关研究主要集中在多糖含量、色差等理化指标评价^[9]，缺乏对干燥过程代谢物动态变化的系统解析。为深入解析红托竹荪在热加工过程中的整体物质基础响应，特别是识别和量化未知或未预期的关键代谢物变化，非靶向代谢组学（Untargeted metabolomics）作为一种高通量、无偏向性的小分子代谢物分析技术，已广泛应用于食药用菌加工过程研究，能够揭示热加工过程中代谢物的降解与转化规律，为优化干制工艺提供理论依据。已有研究表明，不同干燥方式对香菇、银耳等食用菌的风味物质及活性成分具有显著影响，但针对红托竹荪的代谢组学研究较少，亟需开展系统性探索^[10]。

本研究以空气源热泵干燥技术为基础，设置不同温度处理红托竹荪样品，采用非靶向代谢组学方法对其代谢物进行全面分析，旨在揭示干燥温度对红托竹荪代谢的影响及关键功能成分的变化。研究结果有望为红托竹荪干制工艺优化提供科学依据，推动其营养品质提升与产业高质量发展。

1　材料与方法

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1.1　材料与仪器

红托竹荪　来自贵州省农业科学院农作物品种资源研究所。本研究以40、45、50、55 ℃四个温度干燥至恒定干重（含水率9.5%±0.6%）的红托竹荪（编号及外观如图1所示）为试材；甲醇、乙腈　色谱纯，德国Merck公司；甲酸　色谱纯，上海阿拉丁生化科技股份有限公司。

TripleTOF 6600+质谱仪　美国SCIEX公司；LC-30A超高效液相色谱仪　日本Shimadzu公司；Biomek i5自动化工作站　美国Beckman Coulter有限公司；MU-G02-0448恒温金属混匀仪　杭州米欧仪器有限公司；CentriVap离心浓缩仪　美国LABCONCO公司；MM 400球磨仪　德国Retsch公司。

1.2　实验方法

1.2.1　红托竹荪样品制备

红托竹荪采摘后，立即剔除菌托和菌盖，随后，样品送入烘干工序直至达到恒重。烘干结束后，样品置于−80 ℃下保存。分析前经63 h真空冷冻干燥处理，采用液氮预脆化辅助粉碎技术进行前处理。将冻干样品置于MM 400球磨仪中进行粉碎处理（频率30 Hz，持续时间90 s），获得粒径≤150 μm的均质粉末^[11]。精确称取50.0±0.3 mg样本粉末，按固液比1:24（m/v）加入−20 ℃预冷的70%甲醇-水提取液，提取过程采用间歇式涡旋-静置模式，按30 min间隔进行6次涡旋振荡，单次持续时间30 s，振幅设定为8 mm，全程维持4 ℃低温环境。离心分离参数设定为离心力13400×g，时间3 min。上清液经0.22 μm聚偏氟乙烯（PVDF）滤膜过滤后，转移至预冷的2 mL进样瓶，立即置于−80 ℃超低温保存箱中，保存时限不超过48 h，待UPLC-MS/MS分析^[12]。

1.2.2　红托竹荪干燥温度代谢组分组对比

本研究选取16个样品，分为4组进行代谢研究，每组4个平行。烘干温度40、45、50、55 ℃，分别标记为DR40、DR45、DR50、DR55。两两进行代谢组对比，分别为DR45 vs. DR40、DR50 vs. DR45、DR50 vs. DR40、DR55 vs. DR40、DR55 vs. DR45、DR55 vs. DR50，以系统解析温度梯度对代谢物的影响。

1.2.3　色谱质谱采集条件

1.2.3.1　内标溶液制备

精密称取1.00 mg标准品溶于1 mL 70%甲醇溶液（v/v），配制1000 μg/mL母液；取250 μL母液用70%甲醇定容至1 mL，获得250 μg/mL内标工作液。

1.2.3.2　色谱质谱采集条件与内标溶液制备

液相色谱条件参考文献[13]。色谱柱：Waters ACQUITY UPLC HSS T3 Column 1.8 µm，2.1 mm×100 mm，流动相A：超纯水（0.1%的甲酸）；流动相B：乙腈（0.1%的甲酸），柱温：40 ℃；流速：0.40 mL/min；进样量：4 µL。

色谱分离采用梯度洗脱程序：0~5.0 min，流动相A从95%线性降至35%，B相从5%升至65%；5.0~6.0 min，A相从35%骤降至1%，B相升至99%；6.0~7.5 min维持等度洗脱（A：1%，B：99%）；7.6 min立即切换至初始比例（A：95%，B：5%），并平衡至10.0 min。

1.2.3.3　质谱条件

质谱分析采用电喷雾电离（ESI）正负离子双模式采集，参数如下^[14]：采集时间10 min；离子源温度550 ℃；喷雾气压50 psi，辅助加热气压60 psi，气帘气压35 psi；离子化电压±5000 V（正模式+5000 V，负模式−4000 V）；去簇电压±60 V；MS1碰撞能量±10 V，MS2碰撞能量±30 V；碰撞能量步长15 V。

1.2.4　差异代谢物KEGG富集分析

KEGG通路富集分析基于差异代谢物筛选结果展开，其核心参数包含两个维度指标：富集度指数（Rich Factor）表征为特定代谢通路内差异代谢物数量与其注释代谢物总量的比值^[15]，该参数定量表征通路富集强度，数值增加反映显著性水平提升^[16]；统计显著性通过超几何分布检验确定^[17]，其概率计算模型如公式所示：

$\rm P=1-\sum \nolimits_{\mathrm{i}=0}^{\mathrm{m}-1}\frac{\binom{\mathrm{M}}{\mathrm{i}}\binom{\mathrm{N}-\mathrm{M}}{\mathrm{n}-\mathrm{i}}}{\binom{\mathrm{N}}{\mathrm{n}}} $

(1)

式中，N代表全部代谢物中具有KEGG注释的代谢物数量，n代表N中差异代谢物的数量，M代表N中某KEGG通路的代谢物数量，m代表M中某KEGG通路的差异代谢物数量。

1.2.5　差异代谢物代谢通路整体变化分析

差异丰度得分（Differential Abundance Score，DA Score）是一种基于通路的代谢变化分析方法，差异丰度得分可以捕捉到某一途径中所有代谢物的总体变化，其计算公式为：

$\begin{split} & {\text{差异丰度得分}=}\\& {\dfrac{\text{该通路上调差异代谢物个数}-\text{该通路下调差异代谢物个数}}{\text{注释到该通路的所有代谢物个数}} }\end{split}$

(2)

1.3　数据处理

原始文件经ProteoWizard软件转换为mzML标准化格式，采用XCMS算法执行特征峰检测（peak picking）及保留时间漂移校正及信号峰对齐（peak alignment）。基于质量控制系统，剔除各组样本中缺失值比例超过50%的特征峰，并通过K-最近邻插补法（KNN imputation）填补数据空白，继而应用支持向量回归（SVR）模型优化峰强度定量准确性。经上述质控优化的特征峰矩阵经由武汉迈维代谢生物科技股份有限公司的多维注释系统（集成自建数据库、公共资源库、预测数据库及metDNA算法）完成代谢物结构鉴定。采用Excel、Proteowizard、R软件（www.r-project.org/）、SIMCA 14.1及其omics skin组件、Orgin 2022、KEGG（http://www.kegg.com/）、MBRole 2.0通路分析平台处理数据并制图。

2　结果与分析

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2.1　红托竹荪非靶向代谢组学分析

本研究共得到4470种代谢物，并对其分类，结果见图2。由图2可知，正离子模式下检测到2549个，负离子模式下检测到1921个。其中，脂质（A）、脂肪酰类（B）、甾体（C）、杂环化合物（D）、有机酸（E）、萜类（F）、生物碱（G）、色胺、胆碱、色素（H）、鞣质（I）、鞘脂类（J）、木脂素和香豆素（L）、醌类（M）、黄酮（N）、核苷酸及其衍生物（O）、甘油脂类（P）、甘油磷脂类（Q）、酚酸类（R）、醇、胺类（S）、苯及其衍生物（T）、氨基酸及其衍生物（U）占比为88.46%，其他类（K）占比为11.54%。而K类中酮类化合物为174个，占到K类33.72%，内酯类化合物为151个，占到K类29.26%，糖类为102个，占到K类19.77%。

2.2　红托竹荪样本质控分析

质量控制（QC）样本由等比例混合样本提取物构建，用于评估实验流程的稳定性。质谱分析序列中，每10个实验样本插入1个QC样本以监控系统波动^[18]。代谢物提取重现性评价通过多批次QC样本的总离子流图（TIC）重叠度分析实现，该方法可同步验证前处理流程与仪器检测的批间一致性^[19]。

4组温度梯度下红托竹荪代谢物的正负离子模式下的TIC图如图3A和图3B所示。总离子流图（TIC）分析（图3A~图3B）显示，各组样本的保留时间漂移量及峰强度变异系数（RSD≤30%）均符合代谢组学质控标准，表明检测体系稳定性良好^[20]。

QC样本的皮尔逊相关矩阵分析（图3C~图3D）显示，样本间代谢物丰度呈高度线性相关（P<0.001）^[21]。变异系数（CV）评估表明，QC样本中CV<0.3的代谢物占比超过75%（图3E~图3F），进一步验证了数据的可靠性^[22]。

2.3　主成分分析（PCA）

采用主成分分析（Principal component analysis，PCA）对含质控样本（QC）的实验数据集进行系统评估^[23]。分析结果显示，不同处理组间存在显著分离趋势（见图4A得分图），同时组内样本呈现紧密聚集特征。该结果证实了生物学重复设置的可靠性，为后续代谢差异分析奠定方法学基础。

由图4A可知，PC1的贡献率为28.21%，PC2的贡献率为22.89%。DR40、DR45、DR50、DR55、QC之间相互分离，组内之间又相互靠近聚集，只有少量数据重叠，且范围较小。说明组间具有差距，整个分析方法，包括预处理方法和仪器分析方法，都是稳定可靠的^[24]。

2.4　正交偏最小二乘法判别分析（OPLS-DA）

主成分分析（PCA）虽在高维数据处理中展现出独特优势，但其对弱相关变量的敏感性缺陷在代谢组学研究中日益凸显。偏最小二乘判别分析（PLS-DA）通过构建分类变量与代谢轮廓的定量关联模型，有效弥补这一局限^[25]。正交偏最小二乘判别分析（OPLS-DA）进一步采用正交投影技术，将数据变异解耦为预测分量（与响应变量相关）和正交分量（非特异性干扰），通过系统性剥离噪声信号，显著提高差异代谢物鉴别可靠性^[26]。

图5C和图5F显示，图5C中组间距离（横坐标）较大，且主成分解释率较高（PC1>50%），表明40 ℃与55 ℃干燥条件下代谢组差异最显著。图5F中组间距离较小，但正交主成分（纵坐标）的组内差异较低，表明50 ℃与55 ℃的代谢组差异可能集中在少数关键代谢物上。图5A和图5D中组间差异较小，解释率偏低，提示相邻温度区间的代谢组差异较小，可能由微量代谢物波动引起。

在所有验证图（图5A₁~图5F₁）中，原始模型的R²Y和Q²值显著高于随机排列模型（橙色和紫色），尤其是Q²值（Q²>0.5），表明模型具有较高的预测能力且未过拟合。图5C₁中R²Y和Q²值最高（R²Y>0.9，Q²>0.8），进一步支持40 ℃与55 ℃干燥条件下代谢组差异最显著。图5C₂中的S-plot右上角和左下角分布大量红色点（VIP>1），提示该组存在显著差异的代谢物（如氨基酸、糖苷类、脂质衍生物），可能与高温引起的酶失活或氧化反应增强相关。图5F₂中的红色点集中在右上方，可能涉及热稳定性较差的次生代谢物（如多酚、萜类）。图5A₂中绿色点（VIP≤1）占主导，表明差异代谢物较少，反映温和干燥条件下（40~45 ℃）代谢组的稳定性较高。红托竹荪代谢组对干燥温度的敏感性呈梯度变化，55 ℃干燥显著改变代谢物谱，而40~50 ℃区间的影响较小。高温（≥50 ℃）可能通过促进美拉德反应、脂质氧化或破坏热敏感代谢物（如抗氧化成分）导致功能性成分流失^[27]；而低温（40~45 ℃）则有助于保留代谢物的原始组成。

2.5　差异代谢物筛选

通过融合单变量与多变量统计方法，构建了多维分析框架，以数据特征为基础实现精准的代谢物解析^[28]。在OPLS-DA模型（生物学重复≥4）中，变量重要性投影（VIP）作为主要筛选指标，有效识别了组间差异代谢物。该方法结合单变量显著性（P值）与定量变化（FC值），构建了多层次筛选体系。根据多维筛选标准（VIP>1，FC≥2和FC≤0.5），成功构建了红托竹荪四个温度下的差异代谢物特征谱（表1），筛选过程严格遵循代谢组学标准化分析流程。表1显示，差异代谢物数目随温度变化呈现“先增后减”趋势，40到50 ℃时总数递增（DR50 vs. DR40达1219个峰值），且上调代谢物（704~993）显著多于下调（226~373），反映红托竹荪通过激活保护通路维持代谢稳态；50到55 ℃时总数骤降至577个，且下调代谢物（328）反超上调（249），表明高温突破阈值A Score负值通路与酶活抑制的关联系统受损，功能成分进入降解主导阶段^[29]。提示代谢组在此温度区间发生了拐点。文献表明，多种与品质和风味密切相关的酶（如多酚氧化酶PPO、过氧化物酶POD）在50 ℃左右即开始出现显著的热失活与构象改变，导致下游代谢流向受阻或重新分配，并伴随细胞膜脂质重排和蛋白质去折叠等过程。CHENG等^[30]在蘑菇的研究中发现，PPO在50 ℃下的D值（使活性降低90%所需时间）在10~20 min之间，且随温度升高呈指数下降趋势，表明50 ℃已进入快速失活区间。WANG等^[31]分析显示，多种植物源酶（含PPO/POD）在45~55 ℃时开始出现α-螺旋含量减少、疏水性残基暴露的跃变，刺激蛋白质部分去折叠和聚集。表明50 ℃左右为温度敏感阈值，40~45 ℃为最佳干燥区间。

2.6　差异代谢物火山图

差异代谢物的可视化分析通过火山图（Volcano Plot）实现，横轴表示两组样本间代谢物的定量变化（log₂ fold change），纵轴则反映统计显著性（-lg P-value）^[32]。该方法通过设定筛选阈值（|log₂FC|>1且P<0.05）有效地定位显著差异代谢物。本文采用这一可视化方法，展示了各比较组代谢物的整体分布特征及其显著性差异，具体数据详见图6。

图6显示，在40到50 ℃之间，上调的代谢物数量显著增加，这表明红托竹荪在这一温度区间内可能激活了热胁迫和抗氧化防御机制，这与XIE等^[33]关于藜麦在中温胁迫下的响应非常相似。而当温度从50 ℃升至55 ℃时，下调的代谢物数量明显增加，说明过高温度可能导致关键酶活性下降或代谢物降解，从而抑制了某些生物合成途径。HASANUZZAMAN等^[34]指出，适度温度升高能激活细胞防御反应，而过高温度则会导致抗氧化系统失衡，使得热敏性代谢产物降解加剧。这个转变支持了50 ℃左右作为红托竹荪温度敏感阈值的结论。

2.7　差异代谢物聚类热图

聚类热图是一种常用的数据可视化工具，通过颜色编码和层次聚类将大量代谢物数据直观地展示出来。其中行向量代表差异代谢物，列向量映射样本分组，色阶梯度反映标准化后的代谢物丰度在各组间的波动。图7展示了该多维数据的可视化结果，直观揭示了温度梯度对红托竹荪代谢网络的调控特征。

图7展示了不同烘干温度（40、45、50和55 ℃）下红托竹荪次级代谢产物的相对丰度分布。各子图（图7a~图7f）采用标准化后的颜色梯度反映代谢物的表达变化。总体而言，40 ℃与45 ℃样品的代谢分布较为相似，而在温度升至50 ℃及55 ℃时，部分代谢物的表达由中性区域向极端红或绿跃变，表明在50 ℃左右可能存在温度敏感阈值，使代谢网络发生急剧的转变。ZANDALINAS等^[35]揭示了高温胁迫下植物内关键代谢通路的调控变化，发现当温度超过适宜范围时，细胞防御机制被抑制，部分代谢物显著下调，证明了代谢网络存在温度“拐点效应”。这种“拐点效应”提示红托竹荪的代谢调控呈现非线性特征，在低温时，细胞可能维持保护性响应，而当温度超过50 ℃后，其代谢网络迅速转向受损状态，主要体现在黄酮类及脂质代谢通路等关键途径上。ZENG等^[36]发现，高温条件下天然产物中部分代谢途径会发生突变性调控；同时，MICHAEL等^[27]亦讨论了高温对细胞内代谢途径作用的非线性效应。

2.8　差异代谢物KEGG富集分析

P-value越接近于0，表示富集越显著。图8中点的大小代表富集到相应通路上的差异显著代谢物个数。选择P-value排名前20个通路由小到大进行展示。

从图8的KEGG富集分析结果来看，各子图（图8a~图8f）展示了不同温度比较组中，红托竹荪差异代谢物参与各代谢通路的调控情况。图中采用颜色和数据点大小分别反映富集的统计显著性和参与通路的差异代谢物数量^[37]。图8a显示，温差仅为5 ℃时，大部分数据点呈现较浅色且体积较小，表明在40 ℃到45 ℃之间，红托竹荪的基础代谢状态较为平稳，细胞内的代谢网络变化较为温和。相反，在图8b和图8c的比较中，随着温度由40 ℃升至50 ℃或55 ℃，明显出现了深红色数据点及较大体积，这意味着在较大温差下，多条代谢通路中观察到的差异代谢物数量显著增加，从而诱发了更为剧烈的代谢调控。这种变化可能反映出高温对细胞能量代谢、应激响应等基础过程的调控作用。进一步地，在相邻温度组比较如图8d与图8e中，虽然温差仍为5 ℃，但部分通路的数据点依然显示出较高的显著性和较多的计数，提示红托竹荪对微小温度变化也具有敏感的代谢调控响应。尤其在图8f，数据点普遍呈现出深红并且体积较大，这表明从50 ℃升至55 ℃时，细胞内某些关键通路中的差异代谢物激增，可能存在明显的代谢阈值效应^[38]。这些不同温差组的比较结果直观地反映了烘干温度逐步升高所引起的细胞内代谢网络的重构。BITA等^[39]讨论了植物应对高温胁迫的分子和生理机制，并提到温度超过一定阈值后，细胞保护体系可能被破坏，导致代谢网络重构。在低温区间内代谢较为平衡，而温度达到一定阈值后，则激活更多的差异代谢通路，进而可能对红托竹荪的品质和功能产生影响^[40]。在富集结果（图8）中，高温（≥50 ℃）相比低温主要富集于多种与抗氧化、防御及能量代谢相关的通路，包括苯丙氨酸代谢、黄酮生物合成、氨基酸生物合成、ABC转运蛋白等。结合这些通路中的差异代谢物类别，可以说明功能成分流失原因。黄酮类及苯丙素衍生物在DR55 vs. DR50组中显著下调，这些分子是主要的自由基清除剂，其减少直接导致抗氧化能力下降，并伴随色泽和风味品质劣变。能量代谢通路中富集于糖酵解/糖异生及三羧酸循环通路中的关键酶底物含量波动，显示细胞在高温胁迫下能量供应被重新分配，可能牺牲二次代谢的积累以维持基本生命活动。由此可见，高温（≥50 ℃）烘干不仅在数量上减少了功能性二次代谢产物，也通过抑制核心抗氧化与能量代谢通路，削弱了红托竹荪的生物活性和品质。

2.9　差异代谢物代谢通路整体变化分析

选择P-value排名前20个通路由小到大进行展示。图9显示，不同温度对比组的多个代谢通路呈显著变化（图9a~图9f）。在图9a，由于仅相差5 ℃，大部分通路的DA Score分布接近于0，数据点普遍较小且颜色较淡，这表明在温度由40 ℃提升至45 ℃时，上调和下调代谢物的数量大致平衡，各通路响应温度变化的幅度不足以导致系统性失衡。相较之下，在图9c中，多数通路的DA Score显著偏离零值，不仅有明显正向（上调优势）的通路，也有呈现负向（下调优势）的通路，其数据点尺寸较大且呈现深红色，显示出极高的统计显著性（P值极低），这说明温度从40 ℃升至55 ℃时，红托竹荪内部分关键代谢通路受到强烈调控，多条通路内同时存在较多差异代谢物。图9f中虽然温差仅为5 ℃，大部分代谢通路的DA Score集中在负值区间，这反映出在从50 ℃上升到55 ℃时，红托竹荪中参与这些通路的差异代谢物主要处于下调状态。在这一温度区间内，高温应激倾向于抑制关键代谢途径的活性，可能是由于热诱导酶的失活、热敏性代谢物的降解或整体代谢平衡向降解代谢的转变^[41]。KRASENSKY等^[42]讨论了环境胁迫下（包括高温）植物细胞内代谢重编程的机制，指出温度超过一定阈值后，抗氧化和保护性代谢途径失衡，导致关键代谢物下降。总体来看，图9f提示当温度从50 ℃提升到55 ℃时，红托竹荪的代谢网络中大多数通路经历了显著的下调，这种现象可能表明存在一个温度“阈值效应”，即在超过某一温度后，细胞代谢调控主要转向抑制状态，从而降低了与生物合成或功能性成分积累相关的活性^[43]。50~55 ℃区间各主要代谢通路的DA Score均呈负值，表明在该温度段下，红托竹荪的多条功能性代谢通路受到整体抑制。如抗氧化相关通路（类黄酮降解、苯丙氨酸代谢）均表现出明显的负向DA Score，表明这些通路内的关键功能性代谢物整体下调。结合火山图和热图中黄酮类（N类）、酚酸类（R类）等次生代谢物在DR55样本丰度普遍低于DR50，可推断高温（>50 ℃）处理会导致抗氧化黄酮和酚酸类化合物的流失，从而降低干品的抗氧化能力与品质特性。

通过对比不同温差的组间（图9），可以发现，低温组内各通路变化温和，而温度超过50 ℃后，部分通路迅速展现出上调或下调优势，显示温度的微小变化在高温区间内引发了显著的代谢适应性调控。这些发现与已有文献中关于温度影响食用菌代谢调控的结论相似，ABU等^[44]揭示了温度胁迫下真菌内多个代谢与应激相关通路的调控变化，部分结果表明高温下细胞内代谢活动趋于抑制；同时，DING等^[45]探讨了番茄对高温胁迫的响应，结果表明适中温度时特定抗热代谢物上调，而温度进一步升高后一些关键防御代谢通路转向抑制，表现出明显的温度敏感性，为热胁迫下代谢重构提供了数据支撑。

3　结论

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本研究采用液相色谱-质谱联用的非靶向代谢组学方法，系统分析了40、45、50与55 ℃四种干燥温度下黔产红托竹荪的代谢物动态变化。共鉴定出4470种代谢物，分布于21个大类，其中脂质、脂肪酰类、甾体等基础代谢成分占比88.46%。多重统计（PCA、OPLS-DA）、KEGG通路富集及差异丰度得分（DA Score）表明，40~45 ℃区间代谢组整体差异有限，1071个代谢物出现微幅波动，它们主要集中于基础代谢通路且变化幅度有限，关键功能通路及功能性成分水平未见显著波动，核心功能性成分（黄酮、酚酸、谷胱甘肽等）并未在KEGG富集中显著下调或失活。随着温度升至50 ℃，差异代谢物数量达峰值，代谢系统通过激活能量与氨基酸代谢通路维持稳态；超过50 ℃后（50→55 ℃），差异代谢物骤降并主要集中于抗氧化和次生代谢物通路，DA Score明显为负，反映功能性成分出现拐点式流失。

基于上述结果，红托竹荪干燥存在约50 ℃的温度敏感阈值。40~45 ℃的温和干燥能够良好保留原始代谢特征与功能性成分，而50 ℃以上温度则诱导抗氧化黄酮、酚酸及保护性分子的降解。建议工业化生产中将空气源热泵干燥温度严格控制在50 ℃以下，以最大程度减少功能成分流失，提升产品营养品质与市场竞争力。未来研究应扩大样本规模，并结合湿度、风速等多因素协同效应，进一步完善干燥工艺优化方案。

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2026年第47卷第10期

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doi: 10.13386/j.issn1002-0306.2025040259

接收时间：2025-04-20
首发时间：2026-10-24
出版时间：2026-05-15

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收稿日期：2025-04-20

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贵州省农业科学院农作物品种资源研究所，贵州省农业微生物全省重点实验室，贵州省食用菌育种重点实验室，贵州贵阳 550006

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杨通静（1984−），男，硕士，副研究员，研究方向：食用菌栽培，E-mail：736932018@qq.com。

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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