（1）植物在生长过程中常面临生物胁迫（如病害）和非生物胁迫（如高温）。根际微生物群对植物抵抗胁迫至关重要，而植物通过根系分泌物主动招募有益微生物。但植物如何通过微生物同时抵抗多种胁迫的机制尚不明确。以药用植物三七为模型，研究其在36℃热胁迫下的反应。热胁迫虽对植物造成损伤，但也能增强后续植物的耐热性和抗病性。热胁迫后，三七根系分泌大量核苷酸（如嘌呤和嘧啶）。这些核苷酸特异性招募两种有益微生物：伯克霍尔德氏菌（Burkholderia sp.）真菌 Saitozyma podzolica这些微生物抑制根腐病病原菌，并帮助植物抵抗高温。

（2）外源添加核苷酸，在自然土壤中添加这些核苷酸，同样富集了这两种有益微生物，说明核苷酸是招募的关键信号。跨物种验证：在辣椒和番茄中重复实验，发现：核苷酸 + 有益微生物联合使用，能协同增强植物的耐热性。

Nucleotides enriched under heat stress recruit beneficial rhizomicrobes to protect plants from heat and root-rot stresses 在热胁迫下富集的核苷酸招募有益的根茎微生物来保护植物免受热和根腐胁迫

摘要

（1）Plants thrive under biotic and abiotic stresses with the help of rhizomicrobiota. Root exudates play a pivotal role in recruiting beneficial microbes that assist plants in surviving environmental challenges, but the mechanisms of plant-microbiome interactions to resist multiple stresses remain elusive. 植物在生物和非生物胁迫下，借助于根状微生物群茁壮成长。根分泌物在招募有益微生物以帮助植物在环境挑战中生存方面发挥着关键作用，但植物-微生物相互作用以抵抗多种胁迫的机制仍然难以捉摸。 

（2）We investigated how heat stress alters the rhizomicrobiomes of Panax notoginseng and how these heat stress-regulated microbes confer enhanced heat tolerance and disease resistance. 我们研究了热胁迫如何改变三七的根茎微生物群，以及这些热胁迫调节的微生物如何增强耐热性和抗病性。 

（3）We revealed that heat stress at 36 °C caused thermal damage to plants while enhancing heat tolerance and disease resistance for the survival of subsequent plants. 我们揭示了36℃的热胁迫对植物造成热损伤，同时增强了随后植物存活的耐热性和抗病性。

（4）Specifically, the beneficial microbes Burkholderia sp. and Saitozyma podzolica were recruited by the heat-stressed P. notoginseng and were confirmed to be responsible for resisting multiple stresses. Heat stress-induced plant roots secrete nucleotides such as purines and pyrimidines to promote the proliferation of these two beneficial microbes rather than root-rot pathogens. 特别是有益微生物伯克霍尔德氏菌。和Saitozyma podzolica被热胁迫的三七所招募，并被证实负责抵抗多种胁迫。热胁迫诱导的植物根分泌核苷酸，如嘌呤和嘧啶，以促进这两种有益微生物的增殖，而不是根腐病菌。

（5）The exogenous application of these nucleotides to natural soil also resulted in the enrichment of the same beneficial microbes. Cross-species validation experiments in Capsicum annuum (pepper) and Solanum lycopersicum (tomato) further demonstrated that co-application of nucleotides with beneficial microbes synergistically enhanced heat tolerance. 这些核苷酸对天然土壤的外源应用也导致了相同有益微生物的富集。在辣椒(pepper)和番茄(Solanum lycopersicum，番茄)中的跨物种验证实验进一步证明了核苷酸与有益微生物的共同应用协同增强了耐热性。

（6）Our findings highlight a plant strategy for thriving under multiple adversities and propose a potential pathway by leveraging nucleotide-mediated recruitment of beneficial microbes for enhancing plant resilience against multiple stresses. 我们的发现强调了植物在多重逆境下茁壮成长的策略，并提出了一种通过利用核苷酸介导的有益微生物募集来增强植物对多重胁迫的适应性的潜在途径

（7）Result

（8）Heat stress enhances the survival and heat tolerance of subsequent P. notoginseng plants 热胁迫提高了后续三七植株的存活率和耐热性

（9）Enrichment of Burkholderia sp. and Saitozyma podzolica enhances heat tolerance and disease resistance in subsequent P. notoginseng 伯克霍尔德氏菌的富集。并且Saitozyma podzolica增强了后续三七的耐热性和抗病性

（10） Heat stress enhances root nucleotide secretion 热胁迫增强根核苷酸分泌

（11） Nucleotides promote the growth of beneficial microbes B36 and SP 核苷酸促进有益微生物B36和SP的生长

（12） Beneficial microbes and nucleotides synergistically enhance heat tolerance and growth in pepper and tomato 有益微生物和核苷酸协同增强辣椒和番茄的耐热性和生长

The conceptual figure illustrates the strategy for thriving in heat stress to combat multifarious stresses. Heat stress was found to enhance nucleotide metabolism, leading to the enrichment of purine and pyrimidine metabolites in root exudates. Nucleotides significantly promoted the growth of beneficial microbes Burkholderia sp. (B36) and Saitozyma podzolica (SP). Two beneficial microbes exhibited antagonistic effects against root rot pathogens and enhanced the heat tolerance of P. notoginseng. Consequently, the survival of subsequent plants of P. notoginseng under multiple stresses was notably improved. 热胁迫（逆境）→ 植物代谢响应（分泌核苷酸）→ 特异性招募有益根际微生物 → 微生物提供双重保护（抗病+耐热）→ 形成良性土壤遗产，提高后续植物在多重胁迫下的生存能力。

but the mechanisms of plant-microbiome interactions to resist multiple stresses remain elusive