AbstractBiofilm formation by Shewanella oneidensis has been extensively studied under oxic conditions; however, relatively little is known about biofilm formation under anoxic conditions and how biofilm architecture and composition can positively influence current generation in bioelectrochemical systems.

摘要在有氧条件下，已经广泛研究了Shewanella oneidensis的生物膜形成；然而，关于缺氧条件下生物膜的形成以及生物膜的结构和组成如何对生物电化学系统中的电流产生产生积极影响，人们知之甚少。

In this study, we utilized a recently developed microfluidic biofilm analysis setup with automated 3D imaging to investigate the effects of extracellular electron acceptors and synthetic modifications to the extracellular polymeric matrix on biofilm formation. Our results with the wild type strain demonstrate robust biofilm formation even under anoxic conditions when fumarate is used as the electron acceptor.

在这项研究中，我们利用最近开发的具有自动3D成像的微流体生物膜分析装置来研究细胞外电子受体和细胞外聚合物基质的合成修饰对生物膜形成的影响。我们对野生型菌株的研究结果表明，当富马酸盐用作电子受体时，即使在缺氧条件下也能形成强大的生物膜。

However, this pattern shifts when a graphite electrode is employed as the electron acceptor, resulting in biofilm formation falling below the detection limit of the optical coherence tomography imaging system. To manipulate biofilm formation, we aimed to express BpfG with a single amino acid substitution in the catalytic center (C116S) and to overexpress bpfA.

然而，当石墨电极用作电子受体时，这种模式发生变化，导致生物膜形成低于光学相干断层扫描成像系统的检测限。为了操纵生物膜的形成，我们旨在在催化中心（C116S）中用单个氨基酸取代表达BpfG并过表达bpfA。

Our analyses indicate that, under oxic conditions, overarching mechanisms predominantly influence biofilm development, rather than the specific mutations we investigated. Under anoxic conditions, the bpfG mutation led to a quantitative increase in biofilm formation, but both strains exhibited significant qualitative changes in biofilm architecture compared to the controls.

我们的分析表明，在有氧条件下，总体机制主要影响生物膜的发育，而不是我们研究的特定突变。在缺氧条件下，bpfG突变导致生物膜形成的定量增加，但与对照相比，两种菌株的生物膜结构均表现出显着的质变。

When an anode was used as the sole electron acceptor, both the bpfA and bpfG mutations positively impacted mean current density, yielding a 1.8-fold increase for each mutation..

当阳极用作唯一的电子受体时，bpfA和bpfG突变均对平均电流密度产生积极影响，每个突变产生1.8倍的增加。。

IntroductionShewanella oneidensis MR1 is the best understood model organism regarding dissimilatory metal reduction and extracellular electron transfer onto carbon electrodes in bioelectrochemical systems (BES)1. However, strains of S. oneidensis generally exhibit lower rates of dissimilatory iron reduction and lower current densities compared to the other prominent model organism, Geobacter sulfurreducens2.

引言Shewanella oneidensis MR1是生物电化学系统（BES）1中关于异化金属还原和细胞外电子转移到碳电极上的最佳理解模式生物。然而，与其他突出的模式生物Geobacter sulfurreducens2相比，S.oneidensis菌株通常表现出较低的异化铁还原率和较低的电流密度。

The prevailing view is that G. sulfurreducens has evolved as a specialist for anaerobic respiration with insoluble electron acceptors, while S. oneidensis is as a respiration generalist, which is corroborated by its ability of respiring with the widest range of electron acceptors (including oxygen) known to date.

普遍的观点是，G.sulfurreducens已经发展成为具有不溶性电子受体的无氧呼吸专家，而S.oneidensis是呼吸通才，这一点得到了迄今为止已知的最广泛的电子受体（包括氧气）呼吸能力的证实。

A comparison of the biochemical machinery involved in extracellular electron transfer reveals that the overall strategies for transporting electrons to the cell surface are quite similar in both S. oneidensis and G. sulfurreducens3,4,5. Both organisms have developed an electron conduit through the outer membrane, consisting of one c-type cytochrome on the periplasmic side and one or two c-type cytochromes on the extracellular side of the outer membrane.

对涉及细胞外电子转移的生化机制的比较表明，在S.oneidensis和G.sulfurreducens3,4,5中，将电子传输到细胞表面的总体策略非常相似。两种生物都形成了一个穿过外膜的电子导管，该电子导管由周质侧的一个c型细胞色素和外膜胞外侧的一个或两个c型细胞色素组成。

The primary difference in their electron transfer strategies lies in the extension of the electron transfer chain beyond the cell surface in G. sulfurreducens6,7,8,9. While the mechanisms for electron transfer from the cytoplasmic membrane through the periplasm and the outer membrane are similar, G.

它们的电子转移策略的主要区别在于电子转移链在G.sulfurreducens6,7,8,9中延伸到细胞表面之外。虽然电子从细胞质膜通过周质和外膜转移的机制相似，但G。

sulfurreducens has developed the ability to conductively connect cells with one another and with electrodes using nanowires8,10,11,12,13,14. This capability has led to the hypothesis that the formation of multilayered, thick biofilms under anoxic conditions on solid electron acceptors is directly related to the o.

sulfurreducens已经开发出使用纳米线8,10,11,12,13,14将细胞彼此导电连接并与电极导电连接的能力。这种能力导致了这样的假设，即在缺氧条件下在固体电子受体上形成多层厚生物膜与o直接相关。

(1)

(1)

In order to automate data acquisition and processing as far as possible and to counteract the imperfect repeatability in x-y-positioning of the gantry robot, the OCT images were larger (as seen from above) than the actual cultivation channel. Consequently, parts of the image had to be cropped to be excluded from the calculations.

为了尽可能自动化数据采集和处理，并抵消龙门机器人x-y定位的不完全重复性，OCT图像比实际的栽培通道更大（从上面可以看出）。因此，必须对图像的部分进行裁剪才能从计算中排除。

To automate this step, only 50% of the respective cultivation channel was analyzed. Thus, the lateral walls were not included in the calculation of biovolume, biofilm height, surface coverage and porosity. Only the height maps show the entire channel.Preparation of cells for single-cell force spectroscopyThe S.

为了自动化这一步骤，仅分析了各自培养通道的50%。因此，侧壁不包括在生物体积，生物膜高度，表面覆盖率和孔隙率的计算中。只有高度贴图显示整个通道。单细胞力谱分析细胞的制备。

oneidensis variants were cultured as follows: One bacterial colony of each strain was added to 5 mL of LB medium (LB-Medium (Lennox), Carl Roth, Karlsruhe, Germany) and incubated overnight at 30 °C and 150 rpm for 16 h. To ensure that only bacteria from the exponential phase are used, another 40 µL of the overnight solution were cultivated in 4 mL LB-Medium for 2.5 h at 30 °C and 150 rpm, resulting in an optical density of ~ 0.05 at 600 nm.

如下培养oneidensis变体：将每种菌株的一个细菌菌落加入5 mL LB培养基（LB培养基（Lennox），Carl Roth，Karlsruhe，Germany）中，并在30°C和150 rpm下孵育过夜16小时。为了确保仅使用指数期的细菌，将另外40µL过夜溶液在4 mL LB培养基中于30°C和150 rpm培养2.5小时，在600 nm处的光密度约为0.05。

Afterwards the bacterial solution was washed three times with phosphate-buffered saline (PBS, pH 7.3) (Tablets, Sigma-Aldrich Merck KGaA, Darmstadt, Germany) and centrifuged for 3 min at 17,000 g.Substrate preparationFor the adhesion measurements, hydrophobized silicon wafers were used as test surfaces.

然后，将细菌溶液用磷酸盐缓冲盐水（PBS，pH 7.3）（片剂，Sigma-Aldrich-Merck KGaA，达姆施塔特，德国）洗涤三次，并以17000 g离心3分钟。基板制备为了进行附着力测量，使用疏水化硅片作为测试表面。

To fabricate these surfaces, the wafers were cleaned and coated with a monolayer of OTS (octadecyltrichlorosilane) (CAS 112-04-9, abcr GmbH, Karlsruhe, Germany), following a protocol by Lessel et al.36. Before the surfaces were used in the experiments, they were initially cleaned in ethanol (99,8% VWR International, Radnor, PA, USA) for a period of five minutes, followed by another five.

。在将表面用于实验之前，首先将它们在乙醇（99.8%VWR International，Radnor，PA，USA）中清洗五分钟，然后再清洗五分钟。

Data availability

数据可用性

The data sets generated and analyzed during this study are shown in the manuscript or can be obtained from the corresponding author on request.

在这项研究中生成和分析的数据集显示在手稿中，或者可以根据要求从通讯作者那里获得。

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Download referencesAcknowledgementsThis work was financially supported from the projects “Continuous Bioproduction Using a Tailored Biocatalyst for Electrode Assisted Fermentation” (grant no: 031B0847C) of the Federal Ministry of Education and Research (BMBF) and “Synthetic engineering of conductive biofilm development in the γ-proteobacterium Shewanella oneidensis“ of the German Research Foundation (DFG).

下载参考文献致谢这项工作得到了联邦教育和研究部（BMBF）的项目“使用定制的生物催化剂进行电极辅助发酵的连续生物生产”（批准号：031B0847C）和德国研究基金会（DFG）的“γ-变形杆菌Shewanella oneidensis中导电生物膜开发的合成工程”的资助。

The authors H.H., H.H. and K.J. thank the German Research Foundation (DFG) for funding within the Collaborative Research Center SFB 1027. This research was also supported by DFG large instrument funding under grant number INST 256/542-1 FUGG (project number 449375068). K.J. would like to thank the German Federal Ministry of Education and Research (BMBF) for their support of the Max Planck School Matter to Life in collaboration with the Max Planck society.FundingOpen Access funding enabled and organized by Projekt DEAL.Author informationAuthors and AffiliationsInstitute of Technical Microbiology, University of Technology Hamburg, 21073, Hamburg, GermanyEdina Marlen Klein, René Wurst, Simon Schuldt & Johannes GescherExperimental Physics, Center for Biophysics, Saarland University, 66123, Saarbrücken, GermanyHannah Heintz, Hendrik Hähl & Karin JacobsMax Planck School Matter to Life, 69120, Heidelberg, GermanyKarin JacobsAuthorsEdina Marlen KleinView author publicationsYou can also search for this author in.

作者H.H.，H.H.和K.J.感谢德国研究基金会（DFG）在合作研究中心SFB 1027内提供资金。这项研究也得到了DFG大型仪器基金的支持，资助号为INST 256/542-1 FUGG（项目号449375068）。K、 J.要感谢德国联邦教育和研究部（BMBF）与马克斯·普朗克学会合作，支持马克斯·普朗克学校的生命问题。。作者信息作者和附属机构汉堡理工大学技术微生物学研究所，21073，汉堡，GermanyEdina Marlen Klein，RenéWurst，Simon Schuldt＆Johannes Gescher Saarland大学生物物理中心实验物理，66123，Saarbrücken，GermanyHannah Heintz，Hendrik Hähl＆Karin JacobsMax Planck School Matter to Life，69120，Heidelberg，GermanyKarin JacobsAuthorsEdina Marlen KleinView作者出版物您也可以在中搜索这位作者。

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PubMed Google ScholarContributionsE.K.: Writing – original draft, Writing – review & editing, Supervision, Visualization, Validation, Methodology, Investigation, Formal analysis, Data curation, Conceptualization. Ha.H.: Formal analysis, Investigation, Methodology, Visualization, Writing – review & editing.

PubMed谷歌学术贡献。K、 ：写作-原稿，写作-评论和编辑，监督，可视化，验证，方法论，调查，正式分析，数据管理，概念化。哈。H、 ：形式分析、调查、方法论、可视化、写作-评论和编辑。

R.W.: Writing – review & editing, Validation, Supervision, Investigation. S.S.: Investigation. He.H.: Formal analysis, Methodology, Supervision, Visualization, Writing – review & editing. K.J.: Conceptualization, Data Curation, Funding acquisition, Project administration, Resources, Supervision, Writing – review & editing.

R、 W.：写作-审查和编辑，验证，监督，调查。S、 美国：调查。他。H、 ：形式分析，方法论，监督，可视化，写作-评论和编辑。K、 J.：概念化，数据管理，资金获取，项目管理，资源，监督，写作-审查和编辑。

J.G.: Writing – original draft, Writing – review & editing, Validation, Supervision, Resources, Project administration, Methodology, Funding acquisition, Conceptualization.Corresponding authorCorrespondence to.

J、 。对应作者对应。

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Reprints and permissionsAbout this articleCite this articleKlein, E.M., Heintz, H., Wurst, R. et al. Comparative analysis of the influence of BpfA and BpfG on biofilm development and current density in Shewanella oneidensis under oxic, fumarate- and anode-respiring conditions.

转载和许可本文引用本文Klein，E.M.，Heintz，H.，Wurst，R。等人在有氧，富马酸盐和阳极呼吸条件下比较分析BpfA和BpfG对Shewanella oneidensis生物膜发育和电流密度的影响。

Sci Rep 14, 23174 (2024). https://doi.org/10.1038/s41598-024-73474-wDownload citationReceived: 18 July 2024Accepted: 17 September 2024Published: 05 October 2024DOI: https://doi.org/10.1038/s41598-024-73474-wShare this articleAnyone you share the following link with will be able to read this content:Get shareable linkSorry, a shareable link is not currently available for this article.Copy to clipboard.

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