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                <forename type="first">Zhen-Yu</forename>
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            <funder>national Natural Science Foundation of China (42177194), Yangtze Delta RegionHealthy Agriculture Institute (HAI-YFWW-202403-01), the Joint Funds of the Zhejiang Provincial Natural ScienceFoundation of China (LLSSZ24C030001), and One Health Interdisciplinary Research Project, Ningbo University(HY202406).FIG. 6. Conceptual framework for future drilosphere research. (a) micro-interfacial processes: spatial-temporal P dynamicsresolved via molecular-scale imaging of speciation, microbial communities, and enzyme activities. (b) Biogeographical linkages:earthworm diversity (epigeic, anecic, and endogeic) and distribution patterns correlated with soil P cycling across ecosystems. (c)agricultural synergies: earthworm-crop interactions and management practices (e.g., organic amendments) to optimize P-useefficiency and soil health. (d) Climate-driven dynamics: impacts of temperature, precipitation, and extreme events onearthworm-mediated P transformations and ecosystem resilience.</funder>
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                <title xml:lang="en">Phosphorus dynamics in the drilosphere: Unraveling earthworm-mediated soil biogeochemistry</title>
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                <term xml:lang="en">soil hotspots</term>
                <term xml:lang="en">spatial heterogeneity</term>
                <term xml:lang="en">Esther Alvarez-Ayuso</term>
                <term xml:lang="en">phosphorus biogeochemistry</term>
                <term xml:lang="en">nutrient cycling</term>
                <term xml:lang="en">earthworm ecosystem engineering</term>
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              <p>The drilosphere, encompassing soil zones influenced by earthworm activities, represents a crucial biogeochemical hotspot for phosphorus (P) cycling and soil health. While earthworms are recognized for their positive impacts on soil P dynamics through feeding, burrowing, and casting activities, comprehensive understanding of P cycling mechanisms within the drilosphere and their relative importance across different soil environments remains limited. This review synthesized recent advances through four interconnected domains. First, we examined drilosphere formation and characteristics, revealing how earthworm ecosystem engineering creates distinct biogeochemical zones through burrow construction, mucus secretion, and cast deposition. Second, we analyzed P transformation mechanisms within the drilosphere, documenting how burrowing and casting pathways mobilize P through competitive adsorption, enzymatic mineralization, and microbial regulation. Third, we characterized the drilosphere's contribution to terrestrial P cycling, demonstrating distinctive outward P diffusion patterns compared to rhizosphere inward movement and examining ecosystem-scale impacts. Fourth, we discussed emerging imaging techniques that offer unprecedented opportunities to visualize drilosphere P dynamics, though systematic applications remain limited. Our analysis identified critical research frontiers in micro-interfacial characterizations, biogeographical patterns, climate change impacts, and agricultural applications, providing a framework for advancing sustainable soil management strategies that leverage earthworm-mediated P mobilization.</p>
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