平菇菌丝体对小麦生长发育抑制作用的形态学特征和代谢组学分析

摘要(中文译文)

随着栽培规模的持续扩大，产生了大量收获后的蘑菇废料（spent mushroom substrate，SMS）。这种有机废弃物若处置不当，将对土壤健康构成潜在威胁，包括污染和生态失衡。因此，严格的 safety assessment 对于支持 SMS 作为土壤改良剂的可持续且农艺可行的利用不可或缺。本研究对 SMS 进行了灭菌和非灭菌处理，并建立了整合菌丝体与小麦的受控共培养体系。该系统有助于在表型、细胞学和非靶向代谢组学（LC-MS）水平上全面评估残留菌丝体对小麦生长发育的影响。结果表明，非灭菌 SMS 直接田间施用严重损害了小麦表现，导致根系坏死并显著降低结实率。对比实验证实，是非灭菌 SMS——而非其灭菌对应物——产生了明显的植物毒性效应，显著抑制幼苗生长并引发萎蔫症状。为阐明菌丝体互作的时间动态，将小麦幼苗接种活菌丝体并共培养七天。在此条件下，对照组平均根长（10.82 cm）约为处理组的三倍。组织病理学分析揭示了从根尖起始并向基部延伸的渐进性感染模式；长时间暴露最终导致整个根系崩溃。扫描电镜进一步显示感染根表面有大量菌丝体定殖，并伴有特征性细胞损伤——包括严重的细胞壁皱缩和广泛的细胞死亡。LC-MS 分析鉴定出 1867 种注释化合物。比较分析揭示了次生代谢的显著失调，处理组中有 495 种代谢物上调、419 种代谢物下调。总体而言，这些发现提供了有力证据，表明未经处理的 SMS 直接施入土壤会带来切实的农艺风险。本研究为制定安全、循证的 SMS 增值利用与综合管理方案奠定了关键科学基础。

Abstract

With the continuous expansion of cultivation, substantial quantities of post-harvest spent mushroom substrate (SMS) are generated. Improper disposal of this organic waste poses potential threats to soil health, including contamination and ecological imbalance. Consequently, a rigorous safety assessment is indispensable to support the sustainable and agronomically viable utilization of SMS as a soil amendment. In this study, SMS was subjected to sterilized and non-sterilized treatments, and a controlled co-culture system integrating mycelium with wheat was established. This system facilitated a comprehensive evaluation of residual mycelium impacts on wheat growth and development at phenotypic, cytological, and non-targeted metabolomics (LC-MS) levels. Results demonstrated that direct field application of non-sterilized SMS severely compromised wheat performance, inducing root necrosis and significantly reducing grain set. Comparative experiments confirmed that non-sterilized SMS-not its sterilized counterpart-exerted pronounced phytotoxic effects, markedly inhibiting seedling growth and triggering wilting symptoms. To elucidate the temporal dynamics of mycelial interaction, wheat seedlings were inoculated with viable mycelium and co-cultured for seven days. Under these conditions, the mean root length of the control group (10.82 cm) was approximately threefold that of the treatment group. Histopathological analysis revealed a progressive infection pattern initiating at the root apex and extending basipetally; prolonged exposure ultimately caused complete root system collapse. Scanning electron microscopy further showed extensive mycelial colonization on infected root surfaces, accompanied by characteristic cellular damage-including severe cell wall wrinkling and widespread cell death. LC-MS profiling identified 1867 annotated compounds. Comparative analysis revealed significant dysregulation of secondary metabolism, with 495 metabolites upregulated and 419 metabolites downregulated in the treatment group. Collectively, these findings provide robust evidence that unprocessed SMS poses tangible agronomic risks upon direct soil application. This study establishes a critical scientific foundation for developing safe, evidence-based protocols for the valorization and integrated management of SMS.

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