摘要
深层土壤储存了超过一半的陆地有机碳，但调控地下环境中碳持久性的病毒机制仍不清楚。研究人员通过对四类农业生态系统开展0---700厘米土壤剖面调查，并综合全球地下土壤宏基因组，建立了深层土壤病毒资源数据库，包含50,949个病毒操作分类单元和3,902个辅助代谢基因。病毒丰富度随深度显著增加，同时伴随由碳可利用性下降驱动的溶原性转变。深土病毒组携带参与聚合物降解及细胞内裂解的辅助代谢基因和溶菌酶。病毒内溶素显著促进微生物残体碳积累，进而促进矿物结合态有机碳增加。对全球138个土壤剖面的综合分析进一步揭示，病毒丰度对土壤有机碳具有正向影响。这些发现建立了一个解释框架：病毒通过辅助代谢基因促进宿主碳获取，并通过内溶素活性促进残体形成，从而强化深层土壤的微生物碳泵。
关键词： 深层土壤；病毒组；辅助代谢基因；微生物残体；矿物结合态有机碳。
- Wu, X.-J., Han, W.-X., Liang, C., et al. (2026). Soil viruses as key regulators of the microbial carbon pump in deep soils. Nature Ecology & Evolution. https://doi.org/10.1038/s41559-026-03207-x
- 期刊：Nature Ecology & Evolution（2025年 IF=17.1；2026版JCR）
- 在线发表时间：2026年10月21日
研究总结
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path,#mermaid-svg-NtpIFYQn54yiAtky .section-root circle,#mermaid-svg-NtpIFYQn54yiAtky .section-root polygon{fill:hsl(240, 100%, 46.2745098039%);}#mermaid-svg-NtpIFYQn54yiAtky .section-root text{fill:#ffffff;}#mermaid-svg-NtpIFYQn54yiAtky .section-root span{color:#ffffff;}#mermaid-svg-NtpIFYQn54yiAtky .section-2 span{color:#ffffff;}#mermaid-svg-NtpIFYQn54yiAtky .icon-container{height:100%;display:flex;justify-content:center;align-items:center;}#mermaid-svg-NtpIFYQn54yiAtky .edge{fill:none;}#mermaid-svg-NtpIFYQn54yiAtky .mindmap-node-label{dy:1em;alignment-baseline:middle;text-anchor:middle;dominant-baseline:middle;text-align:center;}#mermaid-svg-NtpIFYQn54yiAtky :root{--mermaid-font-family:"trebuchet ms",verdana,arial,sans-serif;} 深土病毒与碳稳定化
科学问题
活跃分解者与长期储碳并存
研究设计
四类农田与七米剖面
微宇宙干预与全球数据综合
核心结果
病毒丰富度与溶原性随深度增加
残体积累连接矿物结合态碳
证据解释
基因潜力与群落干预相互支持
特定内溶素作用仍需直接验证
背景介绍
深土碳的持久性不能仅用"微生物休眠、分解缓慢"解释，因为深层仍存在具有复杂有机物降解潜力的活跃群落。微生物碳泵（microbial carbon pump，MCP）强调微生物将有机物转化为生物量，继而形成残体；残体与矿物结合，又可能进入相对稳定的碳库。本文检验病毒是否同时参与碳获取与残体生成，把病毒感染过程纳入土壤碳转化框架。
重要结果
七米剖面揭示碳利用策略的深度分化
作者在桃园、麦玉轮作、棉田和菜地的18个土层采集216份样品。随深度增加，有机碳及其可利用性下降，植物细胞壁和易利用糖降解基因减少，而几丁质降解潜力略有增加。深度对碳获取基因组成的解释度为14.0%，高于农业生态系统类型的4.2%，支持深层群落更依赖微生物来源碳的解释；这些指标首先代表遗传潜力，而非直接测得的底物通量。
图1｜四类农业生态系统0---700厘米剖面中216份宏基因组的微生物碳利用。a，桃、麦玉轮作、棉花及蔬菜生态系统，分别覆盖18个土层。b，依据剖面中的Al₂O₃、CaO、Fe₂O₃、K₂O、MgO、Na₂O和SiO₂组成计算化学蚀变指数CIA，n＝36个独立样品。c---e，216份样品中植物细胞壁降解基因------纤维素与木聚糖，c；易利用糖降解基因------岩藻糖与甘露糖，d；微生物细胞壁降解基因------几丁质，e的相对丰度，含3个生物学重复。TPM为每百万转录本的标准化指标，此处用于基因相对丰度。红色虚线标示表土0---30厘米的界限。数据为均值±标准误，阴影为95%置信区间；采用双侧线性回归检验。b---e的精确P值依次为：CIA，8.004×10⁻⁶；纤维素，0.00076；木聚糖，2.2×10⁻¹⁶；甘露糖，3.254×10⁻¹⁶；岩藻糖，2.2×10⁻¹⁶；几丁质，0.00091。
深土病毒组富集溶原性策略与碳转化相关功能
数据库汇集约5.1万个病毒操作分类单元（vOTUs），病毒丰富度和预测溶原性比例随深度增加，并与碳资源下降相联系。作者鉴定出3,902个辅助代谢基因（auxiliary metabolic genes，AMGs），其中707个具有碳水化合物活性酶相关注释；内溶素基因在较深土层相对丰富。原文摘要报告50,949个vOTUs，方法及图2a标为50,947，存在两条记录的内部差异，不影响此处对整体趋势的解读。
图2｜深层土壤病毒组的多样性、生活史与AMGs。a，0---30、30---100和100---700厘米土层的vOTU分类组成。b，0---700厘米剖面中50,949个vOTUs的生活史------溶原型与裂解型------及预测宿主；数据为均值±标准误，插图显示216份样品中溶原性病毒比例的变化，含3个生物学重复。c、d，同一批样品的vOTU Chao1多样性指数，以及糖苷水解酶GH和辅助活性酶AA相关AMG数量；阴影为95%置信区间，灰点表示全球综合数据中的样品。e，剖面病毒中鉴定的GH与AA类碳水化合物活性酶AMGs。f，216份样品中病毒内溶素编码基因的相对丰度，阴影为95%置信区间。相关性采用双侧Spearman检验；b、c、f的精确P值分别为1.3×10⁻⁶、2.2×10⁻¹⁶和0.02166。
前病毒为宿主碳获取提供基因组层面的线索
453个非冗余宏基因组组装基因组（MAGs）中，263个检测到前病毒。前病毒携带的碳水化合物活性酶和内溶素基因数量与深度相关，连接了病毒功能注释与具体宿主背景。代表性深土MAG的代谢重建显示宿主与前病毒具有互补的功能组成，但这种基因共存并不直接证明每个候选基因均在体内发挥相应代谢作用。
图3｜病毒促进深土细菌宿主碳获取的基因组证据。a，依据453个合格MAG的保守核糖体蛋白重建的最大似然树，平均完整度91%、污染度3%；星号表示检测到前病毒。b，病毒编码GH相关AMGs和内溶素，可能促进深土宿主碳获取的示意图。c，前病毒携带的GH23、GH108、GH24、GH19------肽聚糖降解；GH105------葡萄糖醛酸苷酶；GH28------果胶降解相关基因。插图为前病毒中相关酶及内溶素基因数量与0---700厘米深度的Spearman相关性；双侧检验中两星号表示P＜0.01，三星号表示P＜0.001，阴影为95%置信区间。对应精确P值为0.00032和0.0012。d，来自650---700厘米的代表性MAG SZB54_bin.5及其前病毒的代谢重建，完整度95.4%、污染度0.8%。
活性病毒处理增加残体碳，并与矿物结合态碳相关
微宇宙实验设置缓冲液对照、失活病毒和活性病毒处理。活性病毒处理相对失活处理使细菌残体碳增加11.3%，真菌残体碳增加3.3%；深土残体与矿物结合态有机碳（MAOC）的关联强于表土。路径模型支持"内溶素相关指标---残体---碳组分"的联系。实验操纵的是病毒群落活性，未单独操纵某一种内溶素。 因而群落处理效应、基因富集和路径关联共同支持机制假说，但不等于已完成单酶因果验证。
图4｜病毒内溶素与残体形成介导的微生物碳泵。a、b，微宇宙获得的10,613个vOTUs的生活史及预测宿主；c，其AMG组成；d，科水平分类。e，总微生物残体与颗粒态有机碳POC、MAOC含量的双侧Spearman相关，n＝72个独立样品，阴影为95%置信区间。f，偏最小二乘路径模型PLS-PM，展示病毒内溶素和碳循环基因与残体碳------以氨基糖表征------POC、MAOC及CO₂排放的联系，n＝72；一星号P＜0.05，三星号P＜0.001，拟合优度0.63，各响应变量框旁标示R²。病毒至残体、残体至POC、残体至MAOC、基因多样性至残体及至CO₂排放路径的精确P值分别为0.0373、0.0127、0.013、4.88×10⁻⁵和1.19×10⁻⁷。g，缓冲液对照CK、失活病毒DV与活性病毒AV处理的细菌和真菌残体碳。h、i，0---30及500---700厘米土壤整个微生物组中GH13葡萄糖苷酶和GH120木糖苷酶基因。j、k，各处理和土层中携带GH23、GH108内溶素基因的病毒相对丰度。g---k均为72个独立样品；处理间采用双侧t检验并进行Benjamini--Hochberg假发现率校正，数据为均值±标准误。点形状代表不同农业生态系统，字母不同表示处理间差异显著；h---k精确P值见原文补充表1。
全球综合数据支持关联，但并非所有深度趋势均重复
全球138个剖面的1,169份扩增子和503份宏基因组数据，支持植物来源聚合物和易利用糖降解基因随深度减少。病毒相对丰度与有机碳密度正相关，路径模型同时纳入地理及土壤理化变量。值得区分的是，全球肽聚糖降解基因的深度趋势并不显著，不能将局部几丁质降解结果概括为所有微生物细胞壁降解功能均随深度上升。
图5｜全球深土剖面的碳利用与病毒组驱动因素。a，1,169份扩增子与503份宏基因组的采样位置，深度分别覆盖0---1,800与0---500厘米；点大小为样品数，颜色代表农田、森林、草地、冻土和沉积物等生态系统。扩增子数据的优势细菌谱系变化见补充材料。b，503份独立宏基因组中植物细胞壁------纤维素和木质素、易利用糖------岩藻糖和半乳糖，以及微生物细胞壁------肽聚糖降解基因相对丰度；双侧Spearman检验精确P值依次为0.00076、0.00014、0.00039、0.000094和0.1668。c，PLS-PM评估环境因素对SOC密度的直接和间接效应，n＝503；纳入经纬度、土深、容重与粉粒含量、阳离子交换量CEC与总氮TN、病毒群落及SOC密度，拟合优度0.662，三星号P＜0.001。d，各路径的标准化直接、间接及总效应；星号表示总效应显著，P＜0.05，精确P值见补充表2。OCD为有机碳密度。
微生物周转与矿物稳定化可共同增强
作者提出双路径框架：病毒相关代谢潜力参与宿主碳获取，裂解活动则增加残体供应，残体随后可能被矿物稳定化。因此，更快的微生物碳周转不必然等于更多碳矿化损失；净效应取决于残体生成、再利用和矿物保护之间的平衡。这一框架仍需跨生态系统与长期碳通量观测检验。
图6｜病毒加速全球深层土壤微生物碳周转的概念模型。病毒通过两条路径参与深土碳循环：一是GH相关AMGs促进宿主碳获取；二是病毒内溶素促进微生物残体形成及MAOC积累。MCP为微生物碳泵，MnCP为矿物碳泵。
方法学参考
研究将剖面宏基因组、MAG与前病毒关联、功能注释、微宇宙处理和全球数据综合相结合。生物信息分析中，病毒识别、序列质量、宿主预测与AMG甄别分别回答不同问题，不能用单一注释结果替代完整证据链。特别是作者将直接参与细胞裂解的溶菌酶与辅助代谢基因分开处理；TPM在宏基因组中的数值也应解释为相对基因丰度，而不是转录活性。
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宏基因组组装与质量控制
病毒和宿主基因组识别
功能注释与深度关联分析
微宇宙处理与残体碳测定
综合解释碳周转与稳定化
总结
研究将深土病毒从群落组成描述推进到碳周转过程解释，支持其与微生物残体及矿物结合态碳的联系。农田干预和全球关联提供互补证据；特定酶的独立作用、自然生态系统中的效应大小及长期净固碳结果，仍需进一步测定。
参考文献
Wu, X.-J., Han, W.-X., Liang, C., et al. (2026). Soil viruses as key regulators of the microbial carbon pump in deep soils. Nature Ecology & Evolution. https://doi.org/10.1038/s41559-026-03207-x