作为生态系统工程师的植物通过对非生物条件和生物区系的影响而显著地改变环境,从而促进了原本在这些环境下无法存活的伴生物种的生长.然而,很多研究缺乏对互惠效应的了解,因为对植物-植物相互作用的研究通常只估计了对伴生物种的益处,而很少考虑一个营养水平如何直接和间接调节植物的反馈作用.我们对两个植物物种(Arenaria tetraquetra和Hormathophylla spinosa,他们或单独开花,或与伴生植物一起开花)进行了一项田间试验用以分解净效应并验证以下假设:授粉者介导的相互作用提供益处用来平衡植物产生促进作用的成本.我们发现,促进作用的净成本伴随授粉介导的益处而存在.尽管两种植物单株植物上的产花更少,但与单独开花相比,他们在与伴生植物一起开花时,每朵花上的授粉者多样性增强.尽管这两种植物单独开花时,单株种子产量较高,但坐果和结实的产量存在种间差异.四叶蕨属的植物(Arenaria tetraquetra),伴生植物对其坐果和结实有负面效应,而刺山蕨属的植物(Hormathophylla spinose),当其与伴生植物一起开花时,坐果更高,结实则不受影响.我们的研究表明,除了承受直接成本外,植物还可以从伴生物种的促进作用中受益,通过提高其对传粉者的可见性.因此,我们强调授粉的相互作用可以补偿物种促进作用的成本,这些促进作用依赖于作为生态系统工程师的植物.本研究阐明了植物-植物间直接相互作用的结果如何被包括第三方在内的间接相互作用来调节的.“,”Aims Ecosystem engineers substantially modify the environment via their impact on abiotic conditions and the biota,resulting in facilitation of associated species that would not otherwise grow.Yet,reciprocal effects are poorly understood as studies of plant-plant interactions usually estimate only benefits for associated species,while how another trophic level may mediate direct and indirect feedback effects for ecosystem engineers is hardly considered.Methods We ran a field experiment with two ecosystem engineers(Arenaria tetraquetra and Hormathophylla spinosa)blooming either alone or with associated plants to decompose net effects and to test the hypothesis that pollinator-mediated interactions provide benefits that balance costs of facilitation by ecosystem engineers.Important Findings We found that net costs of facilitation are accompanied by pollinator-mediated benefits.Despite ecosystem engineers producing fewer flowers per plant,they were visited by more and more diverse pollinators per flower when blooming with associated plants than when blooming alone.Although seed production per plant was higher when ecosystem engineers bloomed alone,fruit set and seed set varied between species.In one case(A.tetraquetra),fruit and seed sets were negatively affected by the presence of associated plants,whereas,in another case(H.spinosa),fruit set and seed set were higher and unaffected when ecosystem engineers bloomed with associated plants,respectively.Our findings suggest that besides experiencing direct costs,ecosystem engineers can also benefit from facilitating other species via increasing their own visibility to pollinators.Thus,we highlight that pollination interactions can compensate for costs of facilitation depending on ecosystem engineer species.This study illuminates how the outcome of direct plant-plant interactions might be mediated by indirect interactions including third players.