Characterization of Bacterial Magnetotactic Behaviors by Using a Magnetospectrophotometry Assay

Magnetotactic bacteria have the unique capacity of synthesizing intracellular single-domain magnetic particles called magnetosomes. The magnetosomes are usually organized in a chain that allows the bacteria to align and swim along geomagnetic field lines, a behavior called magnetotaxis. Two mechanisms of magnetotaxis have been described. Axial magnetotactic cells swim in both directions along magnetic field lines. In contrast, polar magnetotactic cells swim either parallel to the geomagnetic field lines toward the North Pole (north seeking) or antiparallel toward the South Pole (south seeking). In this study, we used a magnetospectrophotometry (MSP) assay to characterize both the axial magnetotaxis of “Magnetospirillum magneticum” strain AMB-1 and the polar magnetotaxis of magneto-ovoid strain MO-1. Two pairs of Helmholtz coils were mounted onto the cuvette holder of a common laboratory spectrophotometer to generate two mutually perpendicular homogeneous magnetic fields parallel or perpendicular to the light beam. The application of magnetic fields allowed measurements of the change in light scattering resulting from cell alignment in a magnetic field or in absorbance due to bacteria swimming across the light beam. Our results showed that MSP is a powerful tool for the determination of bacterial magnetism and the analysis of alignment and swimming of magnetotactic bacteria in magnetic fields. Moreover, this assay allowed us to characterize south-seeking derivatives and non-magnetosome-bearing strains obtained from north-seeking MO-1 cultures. Our results suggest that oxygen is a determinant factor that controls magnetotactic behavior.Magnetotactic bacteria are morphologically, metabolically, and phylogenetically diverse prokaryotes (1, 11). They synthesize unique intracellular organelles, the magnetosomes, which are single-domain magnetic crystals of the mineral magnetite or greigite enveloped by membranes. Magnetosomes are usually organized in a chain(s) within the cell and cause the cell to align along geomagnetic field lines while it swims. The highest numbers of magnetotactic bacteria are generally found at, or just below, the oxic-anoxic transition zone (OATZ) or redoxocline in aquatic habitats (1). Early studies showed that Northern Hemisphere magnetotactic bacteria swim preferentially northward in parallel with the geomagnetic field lines (north seeking [NS]) (2) and that those from the Southern Hemisphere swim preferentially antiparallel to the geomagnetic field lines to the magnetic South Pole (south seeking [SS]) (4). The geomagnetic field is inclined downward from horizontal in the Northern Hemisphere and upward in the Southern Hemisphere, with the inclination magnitude increasing from the equator to the poles. Therefore, magnetotaxis might guide cells in each hemisphere downward to less-oxygenated regions of aquatic habitats, where they would presumably stop swimming until conditions change (1). A recent study reported the coexistence of both NS and SS magnetotactic bacteria in the Northern Hemisphere, which conflicts with the prevalent model of the adaptive value of magnetotaxis (14).Under laboratory conditions, magnetotactic bacteria form microaerophilic bands of cells in oxygen-gradient medium. In fact, magnetotaxis and aerotaxis work together in these bacteria, and the behavior observed has been referred to as “magnetoaerotaxis.” Two different magnetoaerotactic mechanisms, termed polar and axial, are found in different bacterial species (6). The magnetotactic bacteria, principally the magnetotactic cocci, that swim persistently in one direction along the magnetic field (NS or SS) are polar magnetoaerotactic. Magnetotactic bacteria, especially the freshwater spirilla, that swim in either direction along the magnetic field lines with frequent, spontaneous reversals of swimming direction without turning around are axial magnetoaerotactic. For polar magnetotactic bacteria, the magnetic field provides an axis and a direction for motility, whereas for axial magnetotactic bacteria, the magnetic field provides only an axis of motility. The two mechanisms can best be seen in flattened capillary tubes containing suspensions of cells in reduced medium in a magnetic field oriented parallel to the capillary. An oxygen gradient forms along the tube, beginning at the ends of the capillary, with one oriented parallel and the other antiparallel to the magnetic field (1). Band formation by axial magnetoaerotactic cells, such as Magnetospirillum magnetotacticum cells, occurs at both ends of the capillary. Rotation of the magnetic field by 180° after the formation of the bands causes the cells in both bands to rotate 180°, but the bands remain intact. In contrast, band formation by polar magnetoaerotactic cells, such as the marine cocci, occurs only at the end of the capillary for which the magnetic field and the oxygen concentration gradient are oriented opposite to each other. Rotation of the magnetic field by 180° after the formation of the band causes the cells in the band to rotate 180° and swim away, resulting in the dispersal of the band (1). In this study, we developed a magnetospectrophotometry (MSP) assay that provides an alternative method for the quantitative and versatile characterization of the two magnetotactic mechanisms. Using this assay, we demonstrated the effect of artificial magnetic fields on the generation of homogeneous NS or SS magnetotactic bacterial populations.     

Magneto-aerotaxis in marine coccoid bacteria.

Magnetotactic cocci swim persistently along local magnetic field lines in a preferred direction that corresponds to downward migration along geomagnetic field lines. Recently, high cell concentrations of magnetotactic cocci have been found in the water columns of chemically stratified, marine and brackish habitats, and not always in the sediments, as would be expected for persistent, downward-migrating bacteria. Here we report that cells of a pure culture of a marine magnetotactic coccus, designated strain MC-1, formed microaerophilic bands in capillary tubes and used aerotaxis to migrate to a preferred oxygen concentration in an oxygen gradient. Cells were able to swim in either direction along the local magnetic field and used magnetotaxis in conjunction with aerotaxis, i.e., magnetically assisted aerotaxis, or magneto-aerotaxis, to more efficiently migrate to and maintain position at their preferred oxygen concentration. Cells of strain MC-1 had a novel, aerotactic sensory mechanism that appeared to function as a two-way switch, rather than the temporal sensory mechanism used by other bacteria, including Magnetospirillum megnetotacticum, in aerotaxis. The cells also exhibited a response to short-wavelength light (< or = 500 nm), which caused them to swim persistently parallel to the magnetic field during illumination.     

Single-step transfer of biosynthetic operons endows a non-magnetotactic Magnetospirillum strain from wetland with magnetosome biosynthesis

The magnetotactic lifestyle represents one of the most complex traits found in many bacteria from aquatic environments and depends on magnetic organelles, the magnetosomes. Genetic transfer of magnetosome biosynthesis operons to a non-magnetotactic bacterium has only been reported once so far, but it is unclear whether this may also occur in other recipients. Besides magnetotactic species from freshwater, the genus Magnetospirillum of the Alphaproteobacteria also comprises a number of strains lacking magnetosomes, which are abundant in diverse microbial communities. Their close phylogenetic interrelationships raise the question whether the non-magnetotactic magnetospirilla may have the potential to (re)gain a magnetotactic lifestyle upon acquisition of magnetosome gene clusters. Here, we studied the transfer of magnetosome gene operons into several non-magnetotactic environmental magnetospirilla. Single-step transfer of a compact vector harbouring >30 major magnetosome genes from M. gryphiswaldense induced magnetosome biosynthesis in a Magnetospirillum strain from a constructed wetland. However, the resulting magnetic cellular alignment was insufficient for efficient magnetotaxis under conditions mimicking the weak geomagnetic field. Our work provides insights into possible evolutionary scenarios and potential limitations for the dissemination of magnetotaxis by horizontal gene transfer and expands the range of foreign recipients that can be genetically magnetized.     

Acidthiobacillus ferrooxidans中磁小体的提取

At.f和趋磁细菌在生理特性和生长环境有一定的相似性,而且镜检发现At.f具有趋磁性,所以本文采用了趋磁细菌中磁小体的提取方法尝试提取At.f中的磁小体,用超声波破碎At.f后,以磁铁吸取其体内的磁性颗粒,经过检测,发现其体内确实存在含铁元素的磁性颗粒。提取粗样品经过电镜分析,证实其体内存在着少量由脂质包裹的磁小体。磁小体悬浮液经过蔗糖密度梯度离心纯化后,对其作透射电镜,可以清晰的看到磁小体。实验结果表明,At.f体内存在少量的磁小体,正是由于磁小体的存在,才使得At.f在外加磁场作用下发生磁生物效应。这是首次发现从酸性矿坑水分离的At.f具有趋磁性,并从中提取到了磁小体,可以利用At.f的趋磁性将其按照不同磁性进行分离,从而获得活性高的、对不同磁性矿物有特异性的高效浸矿菌种。     

磁泳分离细菌新方法的研究

从酸性矿坑水中富集培养分离到的嗜酸氧化亚铁硫杆菌（Acidithiobacillus ferrooxidans,A.ferrooxidans）[1-2] 菌同趋磁细菌具有一定的相似性。通过显微镜观察发现,部分浸矿细菌在外加磁场的作用下具有微弱的趋磁性,基于菌种的这种特性,设计了磁泳分离仪,对其在磁场作用下泳动（磁泳）进行分析,经磁泳后的近磁、远磁菌的生理特性有较大的差异。从用涂布平板法获得的近磁菌纯培养A. ferrooxidans菌体中,分离得到纳米磁性颗粒,能谱分析表明,其主要成分为Fe和O元素。实验结果证明,A. ferrooxidans具有微弱趋磁性,采用磁泳分离该类菌体内含有磁性颗粒的细菌是可行的,这一分离技术的进一步完善和改进将为传统的微生物菌种分离提供一种新型分离技术,也将大大促进趋磁细菌的研究,而且它与浸矿工艺的结合将大大促进我国生物冶金的研究步伐。     

Swimming behaviour of the multicellular magnetotactic prokaryote ‘Candidatus Magnetoglobus multicellularis’ under applied magnetic fields and ultraviolet light

Magnetotactic bacteria move by rotating their flagella and concomitantly are aligned to magnetic fields because they present magnetosomes, which are intracellular organelles composed by membrane-bound magnetic crystals. This results in magnetotaxis, which is swimming along magnetic field lines. Magnetotactic bacteria are morphologically diverse, including cocci, rods, spirilla and multicellular forms known as magnetotactic multicellular prokaryotes (MMPs). ‘Candidatus Magnetoglobus multicellularis’ is presently the best known MMP. Here we describe the helical trajectories performed by these microorganisms as they swim forward, as well as their response to UV light. We measured the radius of the trajectory, time period and translational velocity (velocity along the helix axis), which enabled the calculation of other trajectory parameters such as pitch, tangential velocity (velocity along the helix path), angular frequency, and theta angle (the angle between the helix path and the helix axis). The data revealed that ‘Ca. M. multicellularis’ swims along elongated helical trajectories with diameters approaching the diameter of the microorganism. In addition, we observed that ‘Ca. M. multicellularis’ responds to UV laser pulses by swimming backwards, returning to forward swimming several seconds after the UV laser pulse. UV light from a fluorescence microscope showed a similar effect. Thus, phototaxis is used in addition to magnetotaxis in this microorganism.     

Magnetite and magnetotaxis in microorganisms

Magnetotactic bacteria from freshwater and marine sediments orient and navigate along geomagnetic field lines. Their magnetotactic response is based on intracellular, single magnetic domains of ferrimagnetic magnetite, which impart a permanent magnetic dipole moment to the cell.     

Effect of light wavelength on motility and magnetic sensibility of the magnetotactic multicellular prokaryote ‘Candidatus Magnetoglobus multicellularis’

‘Candidatus Magnetoglobus multicellularis’ is a magnetotactic microorganism composed of several bacterial cells. Presently, it is the best known multicellular magnetotactic prokaryote (MMP). Recently, it has been observed that MMPs present a negative photoresponse to high intensity ultraviolet and violet-blue light. In this work, we studied the movement of ‘Candidatus Magnetoglobus multicellularis’ under low intensity light of different wavelengths, measuring the average velocity and the time to reorient its trajectory when the external magnetic field changes its direction (U-turn time). Our results show that the mean average velocity is higher for red light (628 nm) and lower for green light (517 nm) as compared to yellow (596 nm) and blue (469 nm) light, and the U-turn time decreased for green light illumination. The light wavelength velocity dependence can be understood as variation in flagella rotation speed, being increased by the red light and decreased by the green light relative to yellow and blue light. It is suggested that the dependence of the U-turn time on light wavelength can be considered a form of light-dependent magnetotaxis, because this time represents the magnetic sensibility of the magnetotactic microorganisms. The cellular and molecular mechanisms for this light-dependent velocity and magnetotaxis are unknown and deserve further studies to understand the biochemical interactions and the ecological roles of the different mechanisms of taxis in MMPs.     

Inactivation of the flagellin gene flaA in Magnetospirillum gryphiswaldense results in nonmagnetotactic mutants lacking flagellar filaments

Magnetotactic bacteria synthesize magnetosomes, which cause them to orient and migrate along magnetic field lines. The analysis of magnetotaxis and magnetosome biomineralization at the molecular level has been hindered by the unavailability of genetic methods, namely the lack of a means to introduce directed gene-specific mutations. Here we report a method for knockout mutagenesis by homologous recombination in Magnetospirillum gryphiswaldense. Multiple flagellin genes, which are unlinked in the genome, were identified in M. gryphiswaldense. The targeted disruption of the flagellin gene flaA was shown to eliminate flagella formation, motility, and magnetotaxis. The techniques described in this paper will make it possible to take full advantage of the forthcoming genome sequences of M. gryphiswaldense and other magnetotactic bacteria.     

Light effects on the multicellular magnetotactic prokaryote ‘<Emphasis Type="Italic">Candidatus</Emphasis> Magnetoglobus multicellularis’ are cancelled by radiofrequency fields: the involvement of radical pair mechanisms

‘Candidatus Magnetoglobus multicellularis’ is the most studied multicellular magnetotactic prokaryote. It presents a light-dependent photokinesis: green light decreases the translation velocity whereas red light increases it, in comparison to blue and white light. The present article shows that radio-frequency electromagnetic fields cancel the light effect on photokinesis. The frequency to cancel the light effect corresponds to the Zeeman resonance frequency (DC magnetic field of 4 Oe and radio-frequency of 11.5 MHz), indicating the involvement of a radical pair mechanism. An analysis of the orientation angle relative to the magnetic field direction shows that radio-frequency electromagnetic fields disturb the swimming orientation when the microorganisms are illuminated with red light. The analysis also shows that at low magnetic fields (1.6 Oe) the swimming orientation angles are well scattered around the magnetic field direction, showing that magnetotaxis is not efficiently in the swimming orientation to the geomagnetic field. The results do not support cryptochrome as being the responsible chromophore for the radical pair mechanism and perhaps two different chromophores are necessary to explain the radio-frequency effects.     

An MCP-Like Protein Interacts with the MamK Cytoskeleton and Is Involved in Magnetotaxis in Magnetospirillum magneticum AMB-1

Magnetotactic bacteria have the unique capacity of aligning and swimming along geomagnetic field lines, a behavior called magnetotaxis. Although this behavior has been observed for 40 years, little is known about its mechanism. Magnetotactic bacteria synthesize unique organelles, magnetosomes, which are magnetic crystals enveloped by membrane. They form chains with the help of the filamentous cytoskeletal protein MamK and impart a net magnetic-dipole moment to the bacterium. The current model proposes that magnetotaxis comprises passive magnetic orientation and active swimming due to flagellar rotation. We thought that magnetic sensing, via the widely used chemotaxis mechanism, might be actively involved in magnetotaxis. We found that the methyl-accepting chemotaxis protein Amb0994 of Magnetospirillum magneticum AMB-1 was capable of carrying out such a function. Amb0994 is encoded by a gene in the magnetosome island, in which genes essential for magnetosome biosynthesis and magnetotaxis are concentrated. Amb0994 lacks periplasmic sensing domain, which is generally involved in sensing stimuli from outside of cells. By constructing fusions with a derivative of yellow-fluorescent-protein, we showed that Amb0994 localizes to the cell poles, where methyl-accepting chemotaxis proteins are usually clustered. We then showed that Amb0994 specifically interacts, via its C-terminal domain, with MamK, using a bimolecular fluorescence complementation assay. Moreover, overproduction of Amb0994 slowed down the response of the bacterium to changes in the direction of the magnetic field. Most importantly, the C-terminal domain of Amb0994, which interacts with MamK, is responsible for this phenotype, suggesting that the interaction between Amb0994 and MamK plays a key role in magnetotaxis. These results lead to a novel explanation for magnetotaxis at the molecular level.     

Magneto-Chemotaxis in Sediment: First Insights

Magnetotactic bacteria (MTB) use passive alignment with the Earth magnetic field as a mean to increase their navigation efficiency in horizontally stratified environments through what is known as magneto-aerotaxis (M-A). Current M-A models have been derived from MTB observations in aqueous environments, where a >80% alignment with inclined magnetic field lines produces a one-dimensional search for optimal living conditions. However, the mean magnetic alignment of MTB in their most widespread living environment, i.e. sediment, has been recently found to be <1%, greatly reducing or even eliminating the magnetotactic advantage deduced for the case of MTB in water. In order to understand the role of magnetotaxis for MTB populations living in sediment, we performed first M-A observations with lake sediment microcosms. Microcosm experiments were based on different combinations of (1) MTB position with respect to their preferred living depth (i.e. above, at, and below), and (2) magnetic field configurations (i.e. correctly and incorrectly polarized vertical fields, horizontal fields, and zero fields). Results suggest that polar magnetotaxis is more complex than implied by previous experiments, and revealed unexpected differences between two types of MTB living in the same sediment. Our main findings are: (1) all investigated MTB benefit of a clear magnetotactic advantage when they need to migrate over macroscopic distances for reaching their optimal living depth, (2) magnetotaxis is not used by all MTB under stationary, undisturbed conditions, (3) some MTB can rely only on chemotaxis for macroscopic vertical displacements in sediment while other cannot, and (4) some MTB use a fixed polar M-A mechanisms, while other can switch their M-A polarity, performing what can be considered as a mixed polar-axial M-A. These observations demonstrate that sedimentary M-A is controlled by complex mechanical, chemical, and temporal factors that are poorly reproduced in aqueous environments.     

Formation of magnetosomes in magnetotactic bacteria

The ability of magnetotactic bacteria to orient and migrate along geomagnetic field lines is based on intracellular magnetic structures, the magnetosomes, which comprise nano-sized, membrane bound crystals of magnetic iron minerals. The formation of magnetosomes is achieved by a biological mechanism that controls the accumulation of iron and the biomineralization of magnetic crystals with a characteristic size and morphology within membrane vesicles. This paper focuses on the current knowledge about magnetotactic bacteria and will outline aspects of the physiology and molecular biology of magnetosome formation. The biotechnological potential of the biomineralization process is discussed.     

Genes and proteins involved in bacterial magnetic particle formation

Magnetic bacteria synthesize intracellular magnetosomes that impart a cellular swimming behaviour referred to as magnetotaxis. The magnetic structures aligned in chains are postulated to function as biological compass needles allowing the bacterium to migrate along redox gradients through the Earth's geomagnetic field lines. Despite the discovery of this unique group of microorganisms 28 years ago, the mechanisms of magnetic crystal biomineralization have yet to be fully elucidated. This review describes the current knowledge of the genes and proteins involved in magnetite formation in magnetic bacteria and the biotechnological applications of biomagnetites in the interdisciplinary fields of nanobiotechnology, medicine and environmental management.     

Inactivation of the Flagellin Gene flaA in Magnetospirillum gryphiswaldense Results in Nonmagnetotactic Mutants Lacking Flagellar Filaments

Magnetotactic bacteria synthesize magnetosomes, which cause them to orient and migrate along magnetic field lines. The analysis of magnetotaxis and magnetosome biomineralization at the molecular level has been hindered by the unavailability of genetic methods, namely the lack of a means to introduce directed gene-specific mutations. Here we report a method for knockout mutagenesis by homologous recombination in Magnetospirillum gryphiswaldense. Multiple flagellin genes, which are unlinked in the genome, were identified in M. gryphiswaldense. The targeted disruption of the flagellin gene flaA was shown to eliminate flagella formation, motility, and magnetotaxis. The techniques described in this paper will make it possible to take full advantage of the forthcoming genome sequences of M. gryphiswaldense and other magnetotactic bacteria.     

Magnetosome chain superstructure in uncultured magnetotactic bacteria

Magnetotactic bacteria produce magnetosomes, which are magnetic particles enveloped by biological membranes, in a highly controlled mineralization process. Magnetosomes are used to navigate in magnetic fields by a phenomenon called magnetotaxis. Two levels of organization and control are recognized in magnetosomes. First, magnetotactic bacteria create a spatially distinct environment within vesicles defined by their membranes. In the vesicles, the bacteria control the size, composition and purity of the mineral content of the magnetic particles. Unique crystal morphologies are produced in magnetosomes as a consequence of this bacterial control. Second, magnetotactic bacteria organize the magnetosomes in chains within the cell body. It has been shown in a particular case that the chains are positioned within the cell body in specific locations defined by filamentous cytoskeleton elements. Here, we describe an additional level of organization of the magnetosome chains in uncultured magnetotactic cocci found in marine and freshwater sediments. Electron microscopy analysis of the magnetosome chains using a goniometer showed that the magnetic crystals in both types of bacteria are not oriented at random along the crystal chain. Instead, the magnetosomes have specific orientations relative to the other magnetosomes in the chain. Each crystal is rotated either 60°, 180° or 300° relative to their neighbors along the chain axis, causing the overlapping of the (1?1?1) and [Formula in text] capping faces of neighboring crystals. We suggest that genetic determinants that are not present or active in bacteria with magnetosomes randomly rotated within a chain must be present in bacteria that organize magnetosomes so precisely. This particular organization may also be used as an indicative biosignature of magnetosomes in the study of magnetofossils in the cases where this symmetry is observed.     

Quantifying the magnetic advantage in magnetotaxis

Magnetotactic bacteria are characterized by the production of magnetosomes, nanoscale particles of lipid bilayer encapsulated magnetite, that act to orient the bacteria in magnetic fields. These magnetosomes allow magneto-aerotaxis, which is the motion of the bacteria along a magnetic field and toward preferred concentrations of oxygen. Magneto-aerotaxis has been shown to direct the motion of these bacteria downward toward sediments and microaerobic environments favorable for growth. Herein, we compare the magneto-aerotaxis of wild-type, magnetic Magnetospirillum magneticum AMB-1 with a nonmagnetic mutant we have engineered. Using an applied magnetic field and an advancing oxygen gradient, we have quantified the magnetic advantage in magneto-aerotaxis as a more rapid migration to preferred oxygen levels. Magnetic, wild-type cells swimming in an applied magnetic field more quickly migrate away from the advancing oxygen than either wild-type cells in a zero field or the nonmagnetic cells in any field. We find that the responses of the magnetic and mutant strains are well described by a relatively simple analytical model, an analysis of which indicates that the key benefit of magnetotaxis is an enhancement of a bacterium's ability to detect oxygen, not an increase in its average speed moving away from high oxygen concentrations.     

Deep-Etching Electron Microscopy of Cells of <Emphasis Type="Italic">Magnetospirillum magnetotacticum</Emphasis>: Evidence for Filamentous Structures Connecting the Magnetosome Chain to the Cell Surface

Magnetospirillum magnetotacticum are magnetotactic bacteria that form a single chain of magnetite magnetosomes within its cytoplasm. Here, we studied the ultrastructure of M. magnetotacticum by freeze-fracture and deep-etching to understand the spatial correlation between the magnetosome chain and the cell envelope and its possible implications for magnetotaxis. Magnetosomes were found mainly near the cell envelope, forming chains that were closely associated with the granular cytoplasmic material. The membrane surrounding the magnetosomes could be visualized in deep-etching preparations. Thin connections between magnetosome chains and the cell envelope were observed in deep-etching images. These results strengthen the hypothesis for the existence of structures that transfer the torque from the magnetosome chains to the whole cell during the orientation of magnetotactic bacteria to a magnetic field lines.     

Comparative genome analysis of four magnetotactic bacteria reveals a complex set of group-specific genes implicated in magnetosome biomineralization and function

Magnetotactic bacteria (MTB) are a heterogeneous group of aquatic prokaryotes with a unique intracellular organelle, the magnetosome, which orients the cell along magnetic field lines. Magnetotaxis is a complex phenotype, which depends on the coordinate synthesis of magnetosomes and the ability to swim and orient along the direction caused by the interaction with the Earth's magnetic field. Although a number of putative magnetotaxis genes were recently identified within a conserved genomic magnetosome island (MAI) of several MTB, their functions have remained mostly unknown, and it was speculated that additional genes located outside the MAI might be involved in magnetosome formation and magnetotaxis. In order to identify genes specifically associated with the magnetotactic phenotype, we conducted comparisons between four sequenced magnetotactic Alphaproteobacteria including the nearly complete genome of Magnetospirillum gryphiswaldense strain MSR-1, the complete genome of Magnetospirillum magneticum strain AMB-1, the complete genome of the magnetic coccus MC-1, and the comparative-ready preliminary genome assembly of Magnetospirillum magnetotacticum strain MS-1 against an in-house database comprising 426 complete bacterial and archaeal genome sequences. A magnetobacterial core genome of about 891 genes was found shared by all four MTB. In addition to a set of approximately 152 genus-specific genes shared by the three Magnetospirillum strains, we identified 28 genes as group specific, i.e., which occur in all four analyzed MTB but exhibit no (MTB-specific genes) or only remote (MTB-related genes) similarity to any genes from nonmagnetotactic organisms and which besides various novel genes include nearly all mam and mms genes previously shown to control magnetosome formation. The MTB-specific and MTB-related genes to a large extent display synteny, partially encode previously unrecognized magnetosome membrane proteins, and are either located within (18 genes) or outside (10 genes) the MAI of M. gryphiswaldense. These genes, which represent less than 1% of the 4,268 open reading frames of the MSR-1 genome, as yet are mostly of unknown functions but are likely to be specifically involved in magnetotaxis and, thus, represent prime targets for future experimental analysis.     

趋磁细菌运动特性的研究进展

细菌的运动性是影响其生存及致病的一个关键条件,同时也为合成和开发仿生运动体、微型机器人等提供了有效的模型。趋磁细菌具有胞内磁小体从而能够感知磁场的变化,进而影响其运动行为。目前,这种外部磁场与生物体的远程响应模式已在环境、医疗、材料等领域有广泛应用。因此,聚焦于趋磁细菌的运动特性,综述了趋磁细菌运动行为的表征、运动机理以及应用等方面的最新研究进展,并对该领域的发展和面临的挑战进行了展望。