Magnetotactic bacteria (MTB) are a diverse group of microorganisms that synthesize intracellular magnetic nanocrystals termed magnetosomes. In this study, a novel marine magnetotactic coccus, designated strain HHB-1, was magnetically enriched from intertidal sediments in Houhai Bay, southern China. Optical microscopy, electron microscopy and elemental analysis revealed that HHB-1 cells are relatively large coccoid-ovoid bacteria (3.9 ± 0.3 μm × 2.8 ± 0.2 μm) containing multiple chains of prismatic magnetite magnetosomes and prominent intracellular Ca/Mg-rich polyphosphate (Ca-Mg-polyP) granules. Whole-genome sequencing and phylogenomic analyses revealed that HHB-1 represents a novel and deeply branching lineage within the order Magnetococcales, exhibiting low average amino acid identity (57.3%-58.7%) with previously reported strains. The magnetosome gene cluster (MGC) of HHB-1 comprises a nearly complete set of mam (magnetosome membrane) genes with conserved gene order and structure, representing the first genomic and MGC characterization of a novel magnetococcus possessing multi-chain magnetosomes. These findings expand our understanding of the diversity, biomineralization strategies and evolutionary history of MTB in marine environments.

The intertidal sediments of Sanya Haitang Bay, a tropical coast, harbor abundant multicellular magnetotactic prokaryotes (MMPs). Using light and electron microscopy, micromanipulation sorting, and whole-genome amplification, we examined their diversity from morphological, phylogenetic, and ecological perspectives. Two types of MMPs were identified: ellipsoidal (eMMPs) and spherical (sMMPs). Their average abundance was 1.37 × 103 ind./dm3 in autumn and 0.27 × 103 ind./dm3 in spring, indicating strong seasonal variation. eMMPs averaged 9.74 × 8.15 µm, consisting of 80–100 cells arranged in layers, whereas sMMPs averaged 5.64 µm in diameter with 40–50 cells organized radially or spirally. Electron microscopy revealed bullet-shaped magnetosomes in both types: those in eMMPs averaged 90.1 × 34.0 nm, while those in sMMPs averaged 97.2 × 36.3 nm. Interestingly, Cu was homogenously detected in the magnetosomes of sMMPs. 16S rRNA gene analysis identified nine OTUs, including three potential new species in the Desulfobacteraceae family within Thermodesulfobacteriota phylum. Of these, two may represent a new genus, and one is affiliated with Candidatus Magnetananas. Global distribution analysis suggests that eMMPs prefer stable, nutrient-rich environments, whereas sMMPs occupy broader ecological niches. Together, these findings expand understanding of tropical MMP diversity and distribution, and the discovery of Cu-containing magnetosomes provides new insight into biomineralization mechanisms.

The Clarion-Clipperton Fracture Zone (CCFZ) in the eastern central Pacific Ocean is the world's largest area for potential deep-sea polymetallic nodule mining and is attracting increased scientific and commercial interest. Recent studies indicate that biogenic magnetite, generated intracellularly by magnetotactic bacteria (MTB), can carry a biogeochemical remanent magnetization in polymetallic nodules, although whether biogenic or physical-chemical processes are responsible for nodule formation remain poorly constrained. Here, we report a combination of magnetic, electron microscope and geochemical analyses on seafloor surface sediments from the eastern CCFZ to understand the spatial distribution of biogenic magnetite and possible relationships between MTB and polymetallic nodules. Experimental results indicate that sedimentary magnetic minerals from the northern and southern regions are dominated by detrital (eolian loess and volcanic material) and biogenic magnetic minerals (magnetosomes), respectively. Sediments from the intermediate region contain both detrital and biogenic magnetic minerals. Quantitative first-order reversal curve-principal component analysis indicates that biogenic magnetite has the highest concentration in the intermediate CCFZ region, coincident with the highest polymetallic nodule density. Combined with previous research, we speculate that MTB growth on the CCFZ seafloor is driven mainly by local redox conditions. Manganese nodule surfaces are rich in organic biofilms, which results in a relatively thick oxic-anoxic transition zone in high-abundance manganese nodule regions, which generates an optimal microenvironment for both MTB growth and magnetite biomineralization. This study provides new clues for understanding the ecological distribution of MTB and the biogeochemical remanent magnetization recorded by biogenic magnetite in deep-sea sediments. Polymetallic nodules are an important seafloor resource because they contain economically important minerals (e.g., Ni, Co, and Mn) that are not common on land. Hence, there is increased economic and scientific interest in understanding the formation mechanisms of these materials. Several studies indicate that biogenic magnetite crystals produced by magnetotactic bacteria (MTB) occur within polymetallic nodules. Testing the relationship between MTB abundance and polymetallic nodule distribution can help to better understand how these materials formed. Here, we study seafloor sediments from the Clarion-Clipperton Fracture Zone (CCFZ) and find that they mainly contain biogenic and detrital magnetic minerals. Detrital magnetic minerals come from volcanic chains and eolian dust, while biogenic magnetite forms within bacterial cells. We find that increased biogenic magnetite abundance coincides with higher polymetallic nodule occurrences in the CCFZ. This implies that MTB activity may play a significant role in the formation and growth of polymetallic nodules. Biogenic and detrital magnetic minerals are quantified in seafloor surface sediments from the Clarion-Clipperton Fracture Zone Biogenic magnetite concentrations and polymetallic nodule density are increased in the central Clarion-Clipperton Fracture Zone Polymetallic nodule growth likely produced reducing, organic-rich microenvironments suitable for magnetotactic bacteria

Magnetotactic bacteria (MTB) within the Nitrospirota phylum play important roles in biogeochemical cycles due to their outstanding ability to biomineralize large amounts of magnetite magnetosomes and intracellular sulfur globules. For several decades, Nitrospirota MTB were believed to only live in freshwater or low-salinity environments. While this group have recently been found in marine sediments, their physiological features and ecological roles have remained unclear. In this study, we combine electron microscopy with genomics to characterize a novel population of Nitrospirota MTB in a coral reef area of the South China Sea. Both phylogenetic and genomic analyses revealed it as representative of a novel genus, named as Candidatus Magnetocorallium paracelense XS-1. The cells of XS-1 are small and vibrioid-shaped, and have bundled chains of bullet-shaped magnetite magnetosomes, sulfur globules, and cytoplasmic vacuole-like structures. Genomic analysis revealed that XS-1 has the potential to respire sulfate and nitrate, and utilize the Wood-Ljungdahl pathway for carbon fixation. XS-1 has versatile metabolic traits that make it different from freshwater Nitrospirota MTB, including Pta-ackA pathway, anaerobic sulfite reduction, and thiosulfate disproportionation. XS-1 also encodes both the cbb3-type and the aa3-type cytochrome c oxidases, which may function as respiratory energy-transducing enzymes under high oxygen conditions and anaerobic or microaerophilic conditions, respectively. XS-1 has multiple copies of circadian related genes in response to variability in coral reef habitat. Our results implied that XS-1 has a remarkable plasticity to adapt the environment and can play a beneficial role in coral reef ecosystems.

Multicellular magnetotactic prokaryotes (MMPs) are a unique group of magnetotactic bacteria that are composed of 10–100 individual cells and show coordinated swimming along magnetic field lines. MMPs produce nanometer-sized magnetite (Fe3O4) and/or greigite (Fe3S4) crystals—termed magnetosomes. Two types of magnetosome gene cluster (MGC) that regulate biomineralization of magnetite and greigite have been found. Here, we describe a dominant spherical MMP (sMMP) species collected from the intertidal sediments of Jinsha Bay, in the South China Sea. The sMMPs were 4.78 ± 0.67 μm in diameter, comprised 14–40 cells helical symmetrically, and contained bullet-shaped magnetite and irregularly shaped greigite magnetosomes. Two sets of MGCs, one putatively related to magnetite biomineralization and the other to greigite biomineralization, were identified in the genome of the sMMP, and two sets of paralogous proteins (Mam and Mad) that may function separately and independently in magnetosome biomineralization were found. Phylogenetic analysis indicated that the sMMPs were affiliated with Deltaproteobacteria. This is the first direct report of two types of magnetosomes and two sets of MGCs being detected in the same sMMP. The study provides new insights into the mechanism of biomineralization of magnetosomes in MMPs, and the evolutionary origin of MGCs.

Magnetotactic bacteria (MTB) have been found in a wide variety of marine habitats, ranging from intertidal sediments to deep-sea seamounts. Deep-sea hydrothermal fields are rich in metal sulfides, which are suitable areas for the growth of MTB. However, MTB in hydrothermal fields have never been reported. Here, the presence of MTB in sediments from the Tangyin hydrothermal field was analyzed by 16S rRNA gene amplicon analysis, metagenomics, and transmission electron microscopy. Sequencing 16S rRNA gene yielded a total of 709 MTB sequences belonging to 20 OTUs, affiliated with Desulfobacterota, Alphaproteobacteria, and Nitrospirae. Three shapes of magnetofossil were identified by transmission electron microscopy: elongated-prismatic, bullet-shaped, and cuboctahedron. All of these structures were composed of Fe3O4. A total of 121 sequences were found to be homologous to the published MTB magnetosome-function-related genes, and relevant domains were identified. Further analysis revealed that diverse MTB are present in the Tangyin hydrothermal field, and that multicellular magnetotactic prokaryote (MMPs) might be the dominant MTB.

Endosomal sorting complex required transport (ESCRT) III proteins are essential for membrane remodeling and repair across all domains of life. Eukaryotic ESCRT-III and the cyanobacterial homologs PspA and Vipp1/Imm30 remodel membrane into vesicles, rings, filaments and tubular rods structures. Here our microscopy analysis showed that multicellular bacteria, referred to as magnetoglobules, possess multiple compartments including magnetosome organelles, polyphosphate granules, vesicles, rings, tubular rods, filaments and MVB-like structures. Therefore, membrane remodeling protein PspA might be required for the formation of these compartments, and contribute to the morphogenesis and evolution of multicellularity. To assess these hypotheses, we sequenced nine genomes of magnetoglobules and found a significant genome expansion compared to unicellular magnetotactic bacteria. Moreover, PspA was ubiquitous in magnetoglobules and formed a distinct clade on the tree of eubacterial and archaeal ESCRT-III. The phylogenetic feature suggested the evolution of magnetoglobules from a unicellular ancestor of deltaproteobacterium. Hetero-expression of ellipsoidal magnetoglobule pspA2 gene alone in Escherichia coli resulted in intracellular membrane aggregation. GFP fusion labeling revealed polar location of PspA2 in rod-shaped unicells and regular interval location in filamentous cells. Cryo-electron tomography analysis showed filament bundle, membrane sacculus, vesicles and MVB-like structure in the cells expressing PspA2. Moreover, electron-dense area with a similar distribution as GFP-PspA2 foci in filamentous cells changed the inward orientation of the septum, which might interfere with the cell division. Collectively, these results show the membrane remodeling function of magnetoglobule PspA proteins, which may contribute to morphogenesis and the evolution of multicellularity of magnetotactic bacteria.

Magnetotactic bacteria (MTB) are morphologically and phylogenetically diverse prokaryotes commonly able to produce magnetic nanocrystals within intracellular membrane-bound organelles (i.e.,magnetosomes) and to swim along geomagnetic field lines.We studied the diversity of MTB in the samples collected from a mangrove area in the Sanya River,Hainan,South China,using microscopic and microbial phylogenetic methods.Results of microanalysis and observation in microscopy and energy dispersive X-ray spectroscopy (EDXS) reveal a highly morphological diversity of MTB including unicellular cocci,vibrios,rod-shaped bacteria,and three morphotypes of multicellular magnetotactic prokaryotes (MMPs).In addition,analysis of the 16S rRNA gene showed that these MTB were clustered into 16 operational taxonomic units affiliated to the Alpha-,Delta-,and Gamma-proteobacteria classes within the Proteobacteria phylum.Meanwhile,by using the coupled fluorescence and transmission electron microscopy analysis,rod-shaped bacteria,vibrio,and cocci were phylogenetically and structurally identified at the single-cell level.This study demonstrated highly diverse MTB communities in the mangrove ecosystem and provide a new insight into the overall diversity of MTB.

Multicellular magnetotactic prokaryotes (MMPs) are a group of aggregates composed of 10-100 gram-negative cells synthesizing intracellular magnetic crystals. Two morphotypes of MMPs have been identified, including several species of globally distributed spherical mulberry-like MMPs (sMMPs), and ellipsoidal pineapple-like MMPs (eMMPs). We recently collected MMPs from the intertidal zone of Huiquan Bay, Qingdao. Optical microscopy showed that there were two types of MMPs in the area, including sMMPs and eMMPs. We observed the size of eMMPs was 9.25 ± 0.79 × 7.48 ± 0.79 μm (n = 24), and the average diameter of sMMPs was 5 ± 0.66 μm (n = 24). Transmission electron microscopy showed that these MMPs contained three sizes of bullet-shaped crystals in parallel chains or clusters. The length and width ratios of the sizes of these magnetosomes were 4.16±0.64, 3.07±0.29 and 2.51±0.36 (n=44). The 16S rRNA gene of micromanipulation-purified sMMPs and eMMPs were cloned and sequenced. Phylogenetic analysis based on the 16S rRNA gene sequence revealed that 565 sequences of MMPs belonged to 16 OTUs, affiliated with Deltaproteobacteria. Four OTUs displayed >3.48% sequence divergence and two OTUs displayed >7.26% sequence divergence with respect to previously reported MMPs. This result suggested that they represented six new species and two novel genera. These results indicated that the intertidal zone of Huiquan Bay has a high diversity of MMPs that bio-mineralize iron crystals and play an important role in iron cycling in such a complex environment. These observations provide a new perspective of the diversity of MMPs in general and expand knowledge of the occurrence of MMPs in the Huiquan Bay. Keywords: Intertidal zone, Diversity, Multicellular magnetotactic prokaryotes (MMPs), 16S rRNA gene, Magnetosomes

Unambiguous magnetic mineral identification in sediments is a prerequisite for reconstructing paleomagnetic and paleoenvironmental information from environmental magnetic parameters. We studied a deep-sea surface sediment sample from the Clarion Fracture Zone region, central Pacific Ocean, by combining magnetic measurements and scanning and transmission electron microscopic analyses. Eight titanomagnetite and magnetite particle types are recognized based on comprehensive documentation of crystal morphology, size, spatial arrangements, and compositions, which are indicative of their corresponding origins. Type-1 particles are detrital titanomagnetites with micron- and submicron sizes and irregular and angular shapes. Type-2 and -3 particles are well-defined octahedral titanomagnetites with submicron and nanometer sizes, respectively, which are likely related to local hydrothermal and volcanic activity. Type-4 particles are nanometer-sized titanomagnetites hosted within silicates, while type-5 particles are typical dendrite-like titanomagnetites that likely resulted from exsolution within host silicates. Type-6 particles are single domain magnetite magnetofossils related to local magnetotactic bacterial activity. Type-7 particles are superparamagnetic magnetite aggregates, while Type-8 particles are defect-rich single crystals composed of many small regions. Electron microscopy and supervised magnetic unmixing reveal that type-1 to -5 titanomagnetite and magnetite particles are the dominant magnetic minerals. In contrast, the magnetic contribution of magnetite magnetofossils appears to be small. Our work demonstrates that incorporating electron microscopic data removes much of the ambiguity associated with magnetic mineralogical interpretations in traditional rock magnetic measurements.

介绍了趋磁细菌的基本概念及研究状况,综述了海洋趋磁细菌的研究进展,着重总结了深海趋磁细菌的研究成果及现状,展望了深海大洋和海山生态环境中趋磁细菌的研究意义及前景,为海洋趋磁细菌研究提供借鉴与参考.

ABSTRACT Magnetotactic bacteria (MTB) are a group of microorganisms that have the ability to synthesize intracellular magnetic crystals (magnetosomes). They prefer microaerobic or anaerobic aquatic sediments. Thus, there is growing interest in their ecological roles in various habitats. In this study we found co-occurrence of a large rod-shaped deltaproteobacterial magnetotactic bacterium (tentatively named LR-1) in the sediment of a brackish lagoon with algal bloom. Electron microscopy observations showed that they were ovoid to slightly curved rods having a mean length of 6.3 ± 1.1 μm and a mean width of 4.1 ± 0.4 μm. Each cell had a single polar flagellum. They contained hundreds of bullet-shaped intracellular magnetite magnetosomes. Phylogenetic analysis revealed that they were most closely related to Desulfamplus magnetovallimortis strain BW-1, and belonged to the Deltaproteobacteria. Our findings indicate that LR-1 may be a new species of MTB. We propose that deltaproteobacterial MTB may play an important role in iron cycling and so may represent a reservoir of iron, and be an indicator species for monitoring algal blooms in such eutrophic ecosystems. These observations provide new clues to the cultivation of magnetotactic Deltaproteobacteria and the control of algal blooms, although further studies are needed.

Seamounts are subsurface mountains in the ocean. Examination of the abundance and distribution of Archaea in seamount ecosystems may provide a better understanding of their ecological functions. Most studies of marine archaeal assemblages in seamount area have focused on hydrothermal vents or ferromanganese crusts. We investigated the archaeal communities from a seamount of the Mariana Volcanic Arc, in the tropical western Pacific Ocean by using high-throughput sequencing. Thaumarchaeota was dominant in the sediments of all sample stations. Community diversity and species richness were greatest at stations near the top of the seamount, and lowest at the deepest station. One sample station on the steep southeast slope that faced the Yap-Mariana trench had a unique composition of Archaea. In summary, depth has an important influence on archaeal community structure, and the geographic properties and sediment characteristics may explain the unique distribution patterns of Archaea in this seamount. This study provides a foundation for future research on Archaea in seamounts.

荣成月湖是一个典型天然潟湖,潮间带沉积物中存在多种形态趋磁细菌.通过Roche 454高通量测序平台,对沉积物样品(B_S)和磁收样品(B_M)进行16S rRNA基因高通量测序,认识荣成潮间带沉积物的细菌群落结构,并了解趋磁细菌多样性及系统进化地位.研究结果表明沉积物样品中主要的细菌类群为δ-变形菌纲,占总细菌数的26.4％,其次是γ-变形菌纲和α-变形菌纲;而磁收样品中细菌多样性和种类明显降低,以α-变形菌纲占绝对优势,相对比例达72.6％.在磁收样品和沉积物样品分别发现了1612条和186条reads与趋磁细菌相关,分别占细菌总数的5.76％和0.85％,磁收样品中趋磁细菌数是沉积物样品的6.8倍.对两个样品中获得的趋磁细菌序列进行系统进化分析,发现这些序列多数属于变形菌门的 α-变形菌纲,以趋磁球菌占绝对优势,少数属于 δ-变形菌纲,与多细胞趋磁原核生物亲缘关系最近.海洋趋磁螺菌属仅在B_M样品中检出,趋磁弧菌属在B_M样品优势度高于B_S样品,而多细胞趋磁原核生物和趋磁螺菌属在B_S样品中优势度更高.通过分析样品间的差异OTUs,认为荣成潮间带沉积物中可能存在大量未知的趋磁细菌新类群.研究结果为下一步培养和开发趋磁细菌这一功能菌群,发现趋磁细菌新类群及趋磁细菌生态功能提供了基础资料.

A diversity of organisms can sense the geomagnetic field for the purpose of navigation. Magnetotactic bacteria are the most primitive magnetism-sensing organisms known thus far and represent an excellent model system for the study of the origin, evolution, and mechanism of microbial magnetoreception (or magnetotaxis). The present study is the first report focused on magnetosome gene cluster duplication in the Alphaproteobacteria , which suggests the important role of gene duplication in the evolution of magnetotaxis in the Alphaproteobacteria and perhaps the domain Bacteria . A novel scenario for the evolution of magnetotaxis in the Alphaproteobacteria is proposed and may provide new insights into evolution of magnetoreception of higher species.

While multicellular magnetotactic prokaryotes (MMPs) are ubiquitous in marine environments, the diversity of MMPs in sediments of coral reef ecosystems has rarely been reported. In this study, we made an investigation on the diversity and characteristics of MMPs in sediments at 11 stations in coral reef habitats of the Paracel Islands. The results showed that MMPs were present at nine stations, with spherical mulberry-like MMPs (s-MMPs) found at all stations and ellipsoidal pineapple-like MMPs (e-MMPs) found at seven stations. The maximum abundance of MMPs was 6 ind./cm3. Phylogenetic analysis revealed the presence of one e-MMP species and five s-MMP species including two species of a new genus. The results indicate that coral reef habitats of the Paracel Islands have a high diversity of MMPs that bio-mineralize multiple intracellular chains of iron crystals and play important role in iron cycling in such oligotrophic environment. These observations provide new perspective of the diversity of MMPs in general and expand knowledge of the occurrence of MMPs in coral reef habitats.

Magnetotactic bacteria (MTB) are cosmopolitan and ubiquitous in sediments of freshwater and marine habitats. Most studies of marine MTB have concerned intertidal zones, and there have been few reports of MTB from seafloor sediments. The objective of this study was to investigate the diversity and spatial distribution of MTB on the continent shelf of the Yellow Sea.

Magnetotactic bacteria are a group of Gram-negative bacteria that synthesize magnetic crystals, enabling them to navigate in relation to magnetic field lines. Morphologies of magnetotactic bacteria include spirillum, coccoid, rod, vibrio, and multicellular morphotypes. The coccid shape is generally the most abundant morphotype among magnetotactic bacteria. Here we describe a species of giant rod-shaped magnetotactic bacteria (designated QR-1) collected from sediment in the low tide zone of Huiquan Bay (Yellow Sea, China). This morphotype accounted for 90% of the magnetotactic bacteria collected, and the only taxonomic group which was detected in the sampling site. Microscopy analysis revealed that QR-1 cells averaged (6.71±1.03)×(1.54±0.20) μm in size, and contained in each cell 42–146 magnetosomes that are arranged in a bundle formed one to four chains along the long axis of the cell. The QR-1 cells displayed axial magnetotaxis with an average velocity of 70±28 μm/s. Transmission electron microscopy based analysis showed that QR-1 cells had two tufts of flagella at each end. Phylogenetic analysis of the 16S rRNA genes revealed that QR-1 together with three other rod-shaped uncultivated magnetotactic bacteria are clustered into a deep branch of Alphaproteobacteria .

There are two genetically distinct morphological types of multicellular magnetotactic prokaryotes (MMPs) in the intertidal zone of Lake Yuehu (China): ellipsoidal MMPs (eMMPs) and spherical MMPs (sMMPs). We studied the vertical distribution of both types of MMPs in the sediment at Lake Yuehu during 1 year. Both types of MMPs were observed at sediment depths ranging from 1 to 34 cm, depending on the seasons. The eMMPs distributed at depths of 2-34 cm during spring, 1-11 cm during summer, 2-21 cm during autumn and 9-32 cm during winter. The eMMP species Candidatus Magnetananas rongchenensis, with magnetite magnetosomes, dominated at all distribution depths. These results suggested that Ca. M. rongchenensis migrated vertically during four seasons. The vertical profiles of oxidation-reduction potential (ORP) in Lake Yuehu changed seasonally, and these changes coincided with the seasonal distribution of MMPs, suggesting that the ORP affected the vertical distribution of MMPs. In addition, high concentrations of ammonium and silicate were associated with low abundances of MMPs.