<p>Small, evergreen, and omnipresent, the bryophytes—comprising the liverworts, hornworts, and mosses—receive little attention, even with more than 19,000 species distributed across the globe (Brinda and Atwood, <span>2024</span>). They colonize almost any habitat and can find a place to settle between and on rocks, on other plants, on soil, on walls, on cars, and elsewhere. Having no vascular system and lacking roots makes them the ideal colonizers on any substrate. But they are small. They do not flower. They are difficult to identify. And yet they fulfill crucial ecosystem functions (Eldridge et al., <span>2023</span>), are used frequently in biotechnology (e.g., Horn et al., <span>2021</span>), and serve as models in physiological and genetic studies (e.g., Beaulieu et al., <span>2025</span>). They are not “lower plants” that have led to the evolution of “higher plants.” Rather, both vascular plants and bryophytes are derived from a complex ancestral land plant (Harris et al., <span>2022</span>). Recent viewpoints have synthesized the important roles that bryophytes play across ecosystems, calling for renewed attention to and inclusion of bryophytes in empirical and theoretical research (Deilmann et al., <span>2024</span>; Rousk and Villarreal, <span>2025</span>). One key feature of bryophytes is their ubiquitous associations with microorganisms, including N<sub>2</sub>-fixing prokaryotes (diazotrophs) that can supply ecosystems with readily available nitrogen (N). This association was first described in 1909—in half a sentence—by plant ecologist Eugene Warming in lecture notes at the University of Copenhagen. The field has progressed tremendously since then, and many of the abiotic controls of this key ecosystem function (i.e., N<sub>2</sub> fixation) have been identified in the past decade. It is now time to look ahead.</p><p>All plants associate with microorganisms, and one key role that microorganisms play is the fixation of atmospheric N<sub>2</sub>, converting inert N<sub>2</sub> into plant-available N. One enigmatic group that performs this ecosystem function is the Cyanobacteria, whose members often associate with vascular and non-vascular plants. Although liverwort-cyanobacteria and hornwort-cyanobacteria associations have been studied intensively, moss-cyanobacteria interactions remain comparatively understudied—even though all mosses are colonized by N<sub>2</sub>-fixing cyanobacteria. However, the degree of colonization varies widely among moss species, leading to large differences in N<sub>2</sub> fixation rates. Nevertheless, in unpolluted ecosystems, such as arctic tundra and boreal or tropical cloud forests, moss-cyanobacteria associations can contribute half of total ecosystem N input (Permin et al., <span>2022</span>). Nitrogen availability and humidity are the key drivers of cyanobacterial N<sub>2</sub> fixation associated with mosses, independent of habitat (Alvarenga and Rousk, <span>2022</span>). These abiotic controls h
作为一个整体,(1)苔藓很容易取样、运输和储存;(二)在环境条件不同的生境中发现的;(3)它们在数量、身份和位置方面与蓝藻的共生关系存在很大差异;(4)共生是可塑的(即它们可以随着环境条件的变化而变化)。因此,这些关联可以帮助我们回答诸如植物和重氮营养体参与什么类型的相互作用以及共生关系如何起源等问题。除了作为研究共生相互作用进化的模型系统外,这些关联也可以用来测试生命策略的进化理论。苔藓-蓝藻的关联是普遍超越北极,北方和热带栖息地。在温带草原(Calabria et al., 2020)和地中海橡树林等森林生态系统中发现了它们。在地中海生境中,苔藓相关重氮营养体的固氮作用比热带云雾森林高10倍,比温带森林高3个数量级(图2A)。温带地区的低N2固定率可能是由于大气沉降的相对高的背景N输入抑制了N2固定活性(Wang et al., 2022),地中海生态系统也可能是这种情况。尽管如此,地中海森林中与苔藓相关的氮固定高于其他被调查的生态系统,特别是在降雨事件之后。这暗示除氮沉降外还有其他控制因素。地中海苔藓在一年中的大部分时间里都是干燥的,因此其固氮活性可以忽略不计。然而,在降雨事件之后,苔藓的N2固定率可以迅速复苏(图2B)。这在其他生态系统中并不适用,这可能是生物适应干旱和短活动窗口的不同生命策略的结果。r/K选择理论是在20世纪70年代提出的(MacArthur和Wilson, 1967),其中r选择策略(快速而短暂的生命,通常在不稳定的环境中)与K选择策略(物种的承载能力,通常在稳定的环境中发现)形成对比。这个概念起源于生殖策略科学,已被应用于其他领域,包括保护生物学和植物传播,因为它是关于在不同环境中最大限度地适应。也许在零星降雨的地中海栖息地中,苔藓定植的蓝藻是r选择的策略,而在气候稳定的热带生态系统中,蓝藻是k选择的策略。这些动态使苔藓-蓝藻共生成为一个强大的、可访问的系统,用于将宏观生态学和进化概念应用于与植物宿主相关的微生物群落。苔藓-蓝藻关联代表了一个有前途的和未充分利用的模型系统,用于探索共生相互作用和微生物生活史策略的进化。苔藓物种和相关蓝藻在定植模式和对变化环境条件的响应方面的差异暗示了苔藓群体内不同的进化轨迹。此外,苔藓叶可以作为研究微生物与微生物相互作用和微观尺度上固定N2命运的运动场,因为蓝藻被大量其他微生物包围,这些微生物可能在苔藓宿主接触到固定N2之前就吸收了固定N2。通过整合生理学、生态学和进化的观点,苔藓-蓝藻系统的研究可以加深我们对植物-微生物相互作用的认识,并为我们对陆地生命共同进化的理解提供更广泛的见解。因此,这些系统在理论框架和实证研究中都值得更多的关注。
{"title":"Moss-cyanobacteria associations: A model for studying symbiotic interactions and evolutionary strategies","authors":"Kathrin Rousk","doi":"10.1002/ajb2.70086","DOIUrl":"10.1002/ajb2.70086","url":null,"abstract":"<p>Small, evergreen, and omnipresent, the bryophytes—comprising the liverworts, hornworts, and mosses—receive little attention, even with more than 19,000 species distributed across the globe (Brinda and Atwood, <span>2024</span>). They colonize almost any habitat and can find a place to settle between and on rocks, on other plants, on soil, on walls, on cars, and elsewhere. Having no vascular system and lacking roots makes them the ideal colonizers on any substrate. But they are small. They do not flower. They are difficult to identify. And yet they fulfill crucial ecosystem functions (Eldridge et al., <span>2023</span>), are used frequently in biotechnology (e.g., Horn et al., <span>2021</span>), and serve as models in physiological and genetic studies (e.g., Beaulieu et al., <span>2025</span>). They are not “lower plants” that have led to the evolution of “higher plants.” Rather, both vascular plants and bryophytes are derived from a complex ancestral land plant (Harris et al., <span>2022</span>). Recent viewpoints have synthesized the important roles that bryophytes play across ecosystems, calling for renewed attention to and inclusion of bryophytes in empirical and theoretical research (Deilmann et al., <span>2024</span>; Rousk and Villarreal, <span>2025</span>). One key feature of bryophytes is their ubiquitous associations with microorganisms, including N<sub>2</sub>-fixing prokaryotes (diazotrophs) that can supply ecosystems with readily available nitrogen (N). This association was first described in 1909—in half a sentence—by plant ecologist Eugene Warming in lecture notes at the University of Copenhagen. The field has progressed tremendously since then, and many of the abiotic controls of this key ecosystem function (i.e., N<sub>2</sub> fixation) have been identified in the past decade. It is now time to look ahead.</p><p>All plants associate with microorganisms, and one key role that microorganisms play is the fixation of atmospheric N<sub>2</sub>, converting inert N<sub>2</sub> into plant-available N. One enigmatic group that performs this ecosystem function is the Cyanobacteria, whose members often associate with vascular and non-vascular plants. Although liverwort-cyanobacteria and hornwort-cyanobacteria associations have been studied intensively, moss-cyanobacteria interactions remain comparatively understudied—even though all mosses are colonized by N<sub>2</sub>-fixing cyanobacteria. However, the degree of colonization varies widely among moss species, leading to large differences in N<sub>2</sub> fixation rates. Nevertheless, in unpolluted ecosystems, such as arctic tundra and boreal or tropical cloud forests, moss-cyanobacteria associations can contribute half of total ecosystem N input (Permin et al., <span>2022</span>). Nitrogen availability and humidity are the key drivers of cyanobacterial N<sub>2</sub> fixation associated with mosses, independent of habitat (Alvarenga and Rousk, <span>2022</span>). These abiotic controls h","PeriodicalId":7691,"journal":{"name":"American Journal of Botany","volume":"112 8","pages":""},"PeriodicalIF":2.7,"publicationDate":"2025-08-12","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://bsapubs.onlinelibrary.wiley.com/doi/epdf/10.1002/ajb2.70086","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144820373","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":2,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}