Towards designing synthetic molecular motors: in situ visualization of the progressive evolution of molecular gearing by bacteria
Lead Research Organisation:
Imperial College London
Department Name: Life Sciences
Abstract
We are entering a new era in which we will make our own tiny 'nanoscale' machinery. We hope that this machinery will produce food, fuel, and clean water, help fight disease, accurately read and write information to DNA, and perform emission-free mechanical tasks. Recent successes have come from using machines that have evolved inside microbes in making chemicals using living cells and storing information in DNA, but we haven't done much work to make our own "molecular motors" that perform physical tasks inside cells. This is mainly because we've been limited by our techniques: we haven't been able to see the machines that have naturally evolved inside cells over the billenia, so we've found it difficult to understand how they work or evolved - and how to do it ourselves (instead we've had to break open cells and purify the machines to study them, a process that often badly affects the machinery that we're trying to study). Thankfully recently we've developed ways to to image machines in 3-D inside cells by freezing the bacteria and putting them in an electron microscope.
I recently trained to use this new technology. I've used it to study probably the most captivating example of molecular motors, a motor that spins a spiral-shaped propellor (the "flagellum") to push bacteria in good directions. But the flagellar motor is more than just a fascinating example of the wonders of life on earth, and studying it promises many useful things to us. One use would be to work out how to harness its emission-less power. A single motor from E. coli (a very well studied bacterium), if it were the man-sized, would be as powerful as an airplane turboprop engine!
It has been known for some time that different motors are stronger or weaker than the E. coli motor, but we don't know why because we haven't been able to see them. This all changed when we used the new technique for imaging whole cells to look at a wide range of bacteria to see their motors. A bacterium called Campylobacter jejuni, or just "Campy", struck us as interesting: not only was it known to swim through very sticky fluids better than other bacteria, but the part that generates rotation is bigger than other bacteria, which we think makes a stronger motor that helps Campy swim. What makes this particularly interesting is that Campy has evolved this adaptation. If we could understand how it did this, we might try to copy it to modify motors to our own specifications. Then we might control bacteria to ferry cargoes around, selectively seek and destroy cancers cells, push miniature rotors, or mix fluids together.
I aim to collect data to fully understand how the Campy motor evolved. To do this I'll image Campy, and simultaneously alter the DNA to change the appearance of the motor. In this way we'll be able to work out where parts of the motor are. At the same time I'll image a bacterium that looks like it's halfway between Campy and E. coli to see if this "half-way" cousin tells us anything about the evolutionary pathway that Campy had to follow. Finally we'll take each part of the motors from all of these bacteria and use their DNA sequences to work out their ancestry. This will help us to see where the Campy motor recruited the additional parts that it uses to be more powerful.
As well as these aims I'm planning on developing methods to work out which parts of the motor go where. To do this I'm going to develop computer programs to design better ways to 'tag' components of the motor with extra bits. These extra bits will be easily visible in the 3-D structures that we collect, enabling us to directly visualize where the tagged components are. Indeed, if this works out it will open the next chapter: instead of using these tags to understand the motor, we'll use them as modifications that we can improve by directing evolution. Maybe someday we'll have a motor that we can gear as we wish.
I recently trained to use this new technology. I've used it to study probably the most captivating example of molecular motors, a motor that spins a spiral-shaped propellor (the "flagellum") to push bacteria in good directions. But the flagellar motor is more than just a fascinating example of the wonders of life on earth, and studying it promises many useful things to us. One use would be to work out how to harness its emission-less power. A single motor from E. coli (a very well studied bacterium), if it were the man-sized, would be as powerful as an airplane turboprop engine!
It has been known for some time that different motors are stronger or weaker than the E. coli motor, but we don't know why because we haven't been able to see them. This all changed when we used the new technique for imaging whole cells to look at a wide range of bacteria to see their motors. A bacterium called Campylobacter jejuni, or just "Campy", struck us as interesting: not only was it known to swim through very sticky fluids better than other bacteria, but the part that generates rotation is bigger than other bacteria, which we think makes a stronger motor that helps Campy swim. What makes this particularly interesting is that Campy has evolved this adaptation. If we could understand how it did this, we might try to copy it to modify motors to our own specifications. Then we might control bacteria to ferry cargoes around, selectively seek and destroy cancers cells, push miniature rotors, or mix fluids together.
I aim to collect data to fully understand how the Campy motor evolved. To do this I'll image Campy, and simultaneously alter the DNA to change the appearance of the motor. In this way we'll be able to work out where parts of the motor are. At the same time I'll image a bacterium that looks like it's halfway between Campy and E. coli to see if this "half-way" cousin tells us anything about the evolutionary pathway that Campy had to follow. Finally we'll take each part of the motors from all of these bacteria and use their DNA sequences to work out their ancestry. This will help us to see where the Campy motor recruited the additional parts that it uses to be more powerful.
As well as these aims I'm planning on developing methods to work out which parts of the motor go where. To do this I'm going to develop computer programs to design better ways to 'tag' components of the motor with extra bits. These extra bits will be easily visible in the 3-D structures that we collect, enabling us to directly visualize where the tagged components are. Indeed, if this works out it will open the next chapter: instead of using these tags to understand the motor, we'll use them as modifications that we can improve by directing evolution. Maybe someday we'll have a motor that we can gear as we wish.
Technical Summary
Designed nanomachinery holds promise to transport nanocargoes, seek cancer cells, power microscale mechanics, and mix microfluidics. Yet we have been restricted to using pre-existing machinery instead of adapting or designing our own because we lack techniques to visualize and understand function and evolution.
The bacterial flagellar motor is a model nanomachine. It is tantalising to note that different motors produce different torques and if we could understand their evolutionary fine-tuning we might capitalize on this knowledge to fine-tune our own through directed evolution. We recently identified three motors that may represent descendants of an evolutionary path to a high-torque motor. We used electron cryo-tomography to image motors in situ in Escherichia coli, Vibrio fischeri, and Campylobacter jejuni. C. jejuni can push through very viscous fluids using a high-torque motor. Strikingly, the diameter of the "C-ring" responsible for torque transmission is wider in C. jejuni. C. jejuni also has a large extra motor-associated structure, one protein of which is also found in V. fischeri. To understand its function we obtained results locating component proteins, and saw that the torque-generation stator protein MotB is spaced at a wider radius than E. coli, in correspondence with the wider C-ring.
We hypothesize that C. jejuni evolved from a Vibrio-esque motor that enabled evolution of a wider C-ring to generate higher torque. It would be of high impact to understand this evolution to mimic it to evolve our own machinery. I propose to:
1. Determine locations and numbers of novel components of the C. jejuni disk complex using gene deletions, truncations, and fusion of designed peptide tags.
2. Perform a similar analysis of the "missing link", V. fischeri.
3. Contextualize results and identify other intermediary bacteria by calculating phylogenies for each protein family, infer their order of recruitment, and what conditions predisposed them to recruitment.
The bacterial flagellar motor is a model nanomachine. It is tantalising to note that different motors produce different torques and if we could understand their evolutionary fine-tuning we might capitalize on this knowledge to fine-tune our own through directed evolution. We recently identified three motors that may represent descendants of an evolutionary path to a high-torque motor. We used electron cryo-tomography to image motors in situ in Escherichia coli, Vibrio fischeri, and Campylobacter jejuni. C. jejuni can push through very viscous fluids using a high-torque motor. Strikingly, the diameter of the "C-ring" responsible for torque transmission is wider in C. jejuni. C. jejuni also has a large extra motor-associated structure, one protein of which is also found in V. fischeri. To understand its function we obtained results locating component proteins, and saw that the torque-generation stator protein MotB is spaced at a wider radius than E. coli, in correspondence with the wider C-ring.
We hypothesize that C. jejuni evolved from a Vibrio-esque motor that enabled evolution of a wider C-ring to generate higher torque. It would be of high impact to understand this evolution to mimic it to evolve our own machinery. I propose to:
1. Determine locations and numbers of novel components of the C. jejuni disk complex using gene deletions, truncations, and fusion of designed peptide tags.
2. Perform a similar analysis of the "missing link", V. fischeri.
3. Contextualize results and identify other intermediary bacteria by calculating phylogenies for each protein family, infer their order of recruitment, and what conditions predisposed them to recruitment.
Planned Impact
BBSRC Strategic Priorities. This project has relevance to multiple strategic priorities. Campylobacter is found in two-thirds of commercially available chicken, causing 460 000 cases of food poisoning, 22 000 hospital admissions, and 110 deaths per year. Campylobacter requires a working flagellar motor for virulence, making this work directly relevant to BBSRC Research Priority "Food Security" ("Healthy and Safe food"). This work also meets the BBSRC "Energy" priority: although harnessing nanomachinery is still underdeveloped, this project could lead to specifically designed motors for future application. Finally, this research meets all sub-priorities of the "Exploiting new ways of working" priority. It is "Data driven biology" (electron cryo-tomography produces terabytes of 3-D images on whole cells, and our tagging methodology mines PDB data to identify ideal tags for visualizing protein location). "Synthetic biology" is at the core of this proposal and the ultimate construction of a toolkit of parts to build motors to our own specification via directed evolution. Electron cryo-tomography is inherently a "Systems approach to the biosciences", imaging whole cells without stains or fixatives, and macromolecular machinery being imaged in situ instead of the standard reductionist approach of purifiying components. Finally, this work is "Technology development for the biosciences" to develop techniques for in situ dissection by method development in generation of minimally-perturbatory tags.
The following people beyond my academic peers may also benefit:
Industry. Results from this research may lead to commercial ventures to develop novel antibiotics that specifically target the Campylobacter or Vibrio flagellar motor. This benefit would be realized through Imperial Innovations, the Imperial College London technology commercialization company. In addition, novel methods to structurally adapt macromolecular machinery may stem from this work, and in this eventuality a patent will be sought through Imperial Innovations.
Basic life scientists. The synthetic biology, evolutionary biology, biophysics, microbiology, C. jejuni and Vibrio communities will learn about the structure, function, and mechanisms and evolution of molecular machinery. This information will be communicated by myself and the unnamed postdoctoral research assistant through peer-reviewed publications, and invited and informal talks at conferences and research institutes.
The public. I am promoting my work to the public in a number of ways. I will exhibit at the yearly Imperial Festival (I already presented my work at a stall in the 2013 Festival, which attracted 10 000 visitors over a weekend). I will also apply for future Royal Society Summer Science Exhibition to present my work. I am also engaging with artist colleagues to disseminate concepts on biological self-assembly and self-organization. For example I am developing plans for a sculpture series with Los Angeles-based artist Michael Parker (http://michaelparker.org) to illustrate the concept of biological self-assembly (we plan to submit an application to the Wellcome Trust Small Arts Awards program), and in spring 2013 delivered a talk on self-organisation and it's relation to bacterial multiprotein machinery at another artist's gallery opening in Cologne, Joel Kyack (http://ghebaly.com/artists/joel-kyack). I anticipate similar collaborations in the future. Despite being international, all work will prominently feature the Imperial and BBSRC brand identities. All of these projects will also focus my communication abilities with a non-specialist audience.
Students. As with non-scientists, students are excited to learn about molecular machinery, particularly when the methods used for study incorporate new, exciting methods such as electron cryo-tomography. I will incorporate this research in my lectures to inspire undergraduates. PhD students who I will mentor will also benefit.
The following people beyond my academic peers may also benefit:
Industry. Results from this research may lead to commercial ventures to develop novel antibiotics that specifically target the Campylobacter or Vibrio flagellar motor. This benefit would be realized through Imperial Innovations, the Imperial College London technology commercialization company. In addition, novel methods to structurally adapt macromolecular machinery may stem from this work, and in this eventuality a patent will be sought through Imperial Innovations.
Basic life scientists. The synthetic biology, evolutionary biology, biophysics, microbiology, C. jejuni and Vibrio communities will learn about the structure, function, and mechanisms and evolution of molecular machinery. This information will be communicated by myself and the unnamed postdoctoral research assistant through peer-reviewed publications, and invited and informal talks at conferences and research institutes.
The public. I am promoting my work to the public in a number of ways. I will exhibit at the yearly Imperial Festival (I already presented my work at a stall in the 2013 Festival, which attracted 10 000 visitors over a weekend). I will also apply for future Royal Society Summer Science Exhibition to present my work. I am also engaging with artist colleagues to disseminate concepts on biological self-assembly and self-organization. For example I am developing plans for a sculpture series with Los Angeles-based artist Michael Parker (http://michaelparker.org) to illustrate the concept of biological self-assembly (we plan to submit an application to the Wellcome Trust Small Arts Awards program), and in spring 2013 delivered a talk on self-organisation and it's relation to bacterial multiprotein machinery at another artist's gallery opening in Cologne, Joel Kyack (http://ghebaly.com/artists/joel-kyack). I anticipate similar collaborations in the future. Despite being international, all work will prominently feature the Imperial and BBSRC brand identities. All of these projects will also focus my communication abilities with a non-specialist audience.
Students. As with non-scientists, students are excited to learn about molecular machinery, particularly when the methods used for study incorporate new, exciting methods such as electron cryo-tomography. I will incorporate this research in my lectures to inspire undergraduates. PhD students who I will mentor will also benefit.
Organisations
- Imperial College London (Lead Research Organisation)
- Francis Crick Institute (Collaboration)
- University of Hawaii (Collaboration)
- The University of Texas at San Antonio (Collaboration)
- The Francis Crick Institute (Project Partner)
- University of Texas SW Medical Centre (Project Partner)
- University of Wisconsin–Madison (Project Partner)
People |
ORCID iD |
Morgan Beeby (Principal Investigator) |
Publications
Beeby M
(2020)
Propulsive nanomachines: the convergent evolution of archaella, flagella and cilia.
in FEMS microbiology reviews
Beeby M
(2016)
Diverse high-torque bacterial flagellar motors assemble wider stator rings using a conserved protein scaffold.
in Proceedings of the National Academy of Sciences of the United States of America
Beeby M
(2015)
Motility in the epsilon-proteobacteria.
in Current opinion in microbiology
Chaban B
(2015)
The flagellum in bacterial pathogens: For motility and a whole lot more.
in Seminars in cell & developmental biology
Chaban B
(2018)
Evolution of higher torque in Campylobacter-type bacterial flagellar motors.
in Scientific reports
Ferreira JL
(2021)
The "Jack-of-all-Trades" Flagellum From Salmonella and E. coli Was Horizontally Acquired From an Ancestral ß-Proteobacterium.
in Frontiers in microbiology
Henderson LD
(2018)
High-Throughput Electron Cryo-tomography of Protein Complexes and Their Assembly.
in Methods in molecular biology (Clifton, N.J.)
Oikonomou C
(2016)
Imaging Bacterial Molecules, Structures and Cells
Rossmann FM
(2020)
In situ structure of the Caulobacter crescentus flagellar motor and visualization of binding of a CheY-homolog.
in Molecular microbiology
Rossmann FM
(2018)
Insights into the evolution of bacterial flagellar motors from high-throughput in situ electron cryotomography and subtomogram averaging.
in Acta crystallographica. Section D, Structural biology
Description | The proposal for this grant award was concerned with how different bacteria have evolved different swimming abilities. We thought we could rationalize this in terms of the architecture of the molecular motors with which they swim. We speculated that a wider rotor component enabled higher torque. Significantly, work under this grant has enabled unprecedentedly large amounts of data collection via the engineering of a unique roboticized device for automated electron microscope maintenance. This major innovation has enabled us to make considerable biological insights, most strikingly the identification of the rotor components of the bacterial motors that we studied, considerably supporting our earlier speculations. This has now been published in the prestigious international journal PNAS, setting the stage for further discoveries. Subsequently we published another paper that addressed the third aim of this proposal, that of the evolutionary pathway to higher torque, this time published in the journal Scientific Reports. |
Exploitation Route | This provides fundamental insights into the mechanisms of both molecular motor function and molecular evolution. This is of major impact to the field. Additionally, this may provide the basis for us to engineer or modify our own molecular motors int he future. Furthermore technological developments with automated data collection are widely appealing to electron microscopists internationally and I regularly received solicitations for advice to build similar devices in other universities. I am currently preparing a publication describing the steps needed to do this. This work has lead to award of an MRC grant on another aspect of Campylobacter motility. |
Sectors | Agriculture Food and Drink Energy Environment |
Description | Publications from this work have been widely publicized by the media, promoting understanding by a lay audience of molecular evolution. |
First Year Of Impact | 2017 |
Sector | Other |
Impact Types | Cultural |
Description | Darwin rwinDa: rewinding and rerunning evolution to study innovation in action |
Amount | $465,000 (USD) |
Funding ID | RGP0028/2021 |
Organisation | Human Frontier Science Program (HFSP) |
Sector | Charity/Non Profit |
Country | France |
Start | 07/2021 |
End | 07/2024 |
Description | MRC |
Amount | £523,977 (GBP) |
Organisation | Medical Research Council (MRC) |
Sector | Public |
Country | United Kingdom |
Start | 05/2017 |
End | 05/2020 |
Description | Wellcome Trust Summer Studentship |
Amount | £2,000 (GBP) |
Organisation | Wellcome Trust |
Department | Wellcome Trust Vacation Scholarship |
Sector | Charity/Non Profit |
Country | United Kingdom |
Start | 05/2016 |
End | 09/2016 |
Description | Dave Hendrixson |
Organisation | University of Texas |
Country | United States |
Sector | Academic/University |
PI Contribution | Electron cryo-tomography |
Collaborator Contribution | Campylobacter genetics |
Impact | Paper accepted at PNAS (2016) |
Start Year | 2010 |
Description | Ned Ruby |
Organisation | University of Hawaii |
Country | United States |
Sector | Academic/University |
PI Contribution | Electron cryo-tomography |
Collaborator Contribution | Vibrio fischeri genetics |
Impact | Paper accepted at PNAS (2016) |
Start Year | 2010 |
Description | Willie Tayor |
Organisation | Francis Crick Institute |
Country | United Kingdom |
Sector | Academic/University |
PI Contribution | Electron cryo-tomography and theoretical discussions |
Collaborator Contribution | Protein structure prediction bioinformatics |
Impact | Sir Francis Crick Institute PhD studentship awarded. |
Start Year | 2013 |
Description | "Electron cryo-tomographic studies of the evolution of a bacterial molecular machine", VIB Training/Imaging@VIB VIII: Single Molecule Approaches in Imaging, VIB, Ghent, Belgium |
Form Of Engagement Activity | A formal working group, expert panel or dialogue |
Part Of Official Scheme? | No |
Geographic Reach | International |
Primary Audience | Professional Practitioners |
Results and Impact | "Electron cryo-tomographic studies of the evolution of a bacterial molecular machine", VIB Training/Imaging@VIB VIII: Single Molecule Approaches in Imaging, VIB, Ghent, Belgium |
Year(s) Of Engagement Activity | 2018 |
Description | "Evolution of a molecular motor", BacNet17, Sant Feliu de Guixols, Spain |
Form Of Engagement Activity | A formal working group, expert panel or dialogue |
Part Of Official Scheme? | No |
Geographic Reach | International |
Primary Audience | Professional Practitioners |
Results and Impact | "Evolution of a molecular motor", BacNet17, Sant Feliu de Guixols, Spain |
Year(s) Of Engagement Activity | 2017 |
Description | "Studying the evolution of macromolecular machines using high-throughput electron cryo-tomography" talk, CCP-EM Spring Symposium, Diamond Light Source Campus, UK |
Form Of Engagement Activity | A formal working group, expert panel or dialogue |
Part Of Official Scheme? | No |
Geographic Reach | International |
Primary Audience | Professional Practitioners |
Results and Impact | "Studying the evolution of macromolecular machines using high-throughput electron cryo-tomography" talk, CCP-EM Spring Symposium, Diamond Light Source Campus, UK |
Year(s) Of Engagement Activity | 2017 |
Description | "Studying the evolution of macromolecular machines using high-throughput electron cryo-tomography", MMC2017, Manchester, UK |
Form Of Engagement Activity | A formal working group, expert panel or dialogue |
Part Of Official Scheme? | No |
Geographic Reach | National |
Primary Audience | Professional Practitioners |
Results and Impact | "Studying the evolution of macromolecular machines using high-throughput electron cryo-tomography", talk, MMC2017, Manchester, UK |
Year(s) Of Engagement Activity | 2017 |
Description | Bacterial Cell Surfaces talk |
Form Of Engagement Activity | Participation in an activity, workshop or similar |
Part Of Official Scheme? | No |
Geographic Reach | International |
Primary Audience | Study participants or study members |
Results and Impact | Presentation at a conference. |
Year(s) Of Engagement Activity | 2016 |
Description | Bacterial Locomotion and Signal Transduction (BLAST) conference presentation |
Form Of Engagement Activity | Participation in an activity, workshop or similar |
Part Of Official Scheme? | No |
Geographic Reach | International |
Primary Audience | Study participants or study members |
Results and Impact | Presentation at a conference. |
Year(s) Of Engagement Activity | 2017 |
Description | Chair and speaker, "Molecular Evolution in Bacteria: insights into the diversification of life" session, ASM Microbe session chair, Atlanta, Georgia, USA |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | International |
Primary Audience | Postgraduate students |
Results and Impact | Chair and speaker, "Molecular Evolution in Bacteria: insights into the diversification of life" session, ASM Microbe session chair, Atlanta, Georgia, USA |
Year(s) Of Engagement Activity | 2018 |
Description | Press coverage of PNAS paper |
Form Of Engagement Activity | A press release, press conference or response to a media enquiry/interview |
Part Of Official Scheme? | No |
Geographic Reach | International |
Primary Audience | Public/other audiences |
Results and Impact | Press release on 2016 Beeby et al PNAS paper that was picked up and covered by New Scientist, Popular Science, Nanowerk.com, mentalfloss.com, "Science et Vie"(French popular science magazine). Also press release was listed as a "top ten" news item for Imperial College. |
Year(s) Of Engagement Activity | 2016 |
Description | School visit and talk (North London Collegiate School) |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Schools |
Results and Impact | Talk to sixth-form school students about my research. |
Year(s) Of Engagement Activity | 2015 |
Description | Science festival exhibit: "Science Uncovered" at the Natural History Museum |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Public/other audiences |
Results and Impact | Stall in the Natural History Museum in South Kensington aimed at communicating the lab's research project to the general public, generating much discussion |
Year(s) Of Engagement Activity | 2016 |
Description | Science festival exhibit: Imperial Festival, Imperial College London |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Regional |
Primary Audience | Public/other audiences |
Results and Impact | Stall at the Imperial College Festival intended to communicate the lab's research to a wider audience, triggering much discussion with visitors. |
Year(s) Of Engagement Activity | 2013,2014,2015,2016 |
URL | http://www.imperial.ac.uk/festival/ |
Description | Sensory Transduction in Microorganisms conference (California) |
Form Of Engagement Activity | Participation in an activity, workshop or similar |
Part Of Official Scheme? | No |
Geographic Reach | International |
Primary Audience | Study participants or study members |
Results and Impact | Conference with scientific peers |
Year(s) Of Engagement Activity | 2016 |
Description | Talk at 'Nerd Nite' London |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Public/other audiences |
Results and Impact | Presentation of my work and general aspects of evolution to a general audience. |
Year(s) Of Engagement Activity | 2016 |
Description | Talk, "Electron cryo-microscopy reveals the mechanisms and evolution of bacterial flagellar motors", University of Bristol |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Electron cryo-microscopy reveals the mechanisms and evolution of bacterial flagellar motors", University of Bristol |
Year(s) Of Engagement Activity | 2019 |
Description | Talk, "Evolution of bacterial flagellar motors", 2019 IPBS Students Symposium, Toulouse, France |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Evolution of bacterial flagellar motors", 2019 IPBS Students Symposium, Toulouse, France |
Year(s) Of Engagement Activity | 2019 |
Description | Talk, "Evolution of bacterial flagellar motors", 6th Molecular Microbiology Meeting, Newcastle, UK |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | International |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Evolution of bacterial flagellar motors", 6th Molecular Microbiology Meeting, Newcastle, UK |
Year(s) Of Engagement Activity | 2019 |
Description | Talk, "Evolution of bacterial flagellar motors", Bioscience Seminar, University of Exeter |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Evolution of bacterial flagellar motors", Bioscience Seminar, University of Exeter |
Year(s) Of Engagement Activity | 2019 |
Description | Talk, "Evolution of bacterial flagellar motors", CNRS Marseille, France |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Regional |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Evolution of bacterial flagellar motors", CNRS Marseille, France |
Year(s) Of Engagement Activity | 2019 |
Description | Talk, "Evolution of bacterial flagellar motors", Humboldt University of Berlin, Germany |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Evolution of bacterial flagellar motors", Humboldt University of Berlin, Germany |
Year(s) Of Engagement Activity | 2019 |
Description | Talk, "Evolution of bacterial flagellar motors", TU Delft, Netherlands |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Evolution of bacterial flagellar motors", TU Delft, Netherlands |
Year(s) Of Engagement Activity | 2019 |
Description | Talk, "Evolution of bacterial flagellar motors", University of Copenhagen, Denmark |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Evolution of bacterial flagellar motors", University of Copenhagen, Denmark |
Year(s) Of Engagement Activity | 2019 |
Description | Talk, "Evolution of bacterial flagellar motors", University of Tübingen |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Evolution of bacterial flagellar motors", University of Tübingen |
Year(s) Of Engagement Activity | 2019 |
Description | Talk, "Evolution of high torque bacterial flagellar motors", Biozentrum Basel, Switzerland |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Local |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Evolution of high torque bacterial flagellar motors", Biozentrum Basel, Switzerland |
Year(s) Of Engagement Activity | 2018 |
Description | Talk, "Ratcheting complexity in evolution: Structural studies of the bacterial flagellum as case study", |
Form Of Engagement Activity | A talk or presentation |
Part Of Official Scheme? | No |
Geographic Reach | Regional |
Primary Audience | Postgraduate students |
Results and Impact | Talk, "Ratcheting complexity in evolution: Structural studies of the bacterial flagellum as case study", Munich, Germany |
Year(s) Of Engagement Activity | 2020 |