Control of plastid biogenesis by the ubiquitin-proteasome system

Lead Research Organisation: University of Oxford
Department Name: Plant Sciences

Abstract

Chloroplasts and mitochondria are normal components of many cells - they are sub-cellular structures called organelles. Interestingly, these two organelles evolved from bacteria that were engulfed by other cells over a billion years ago, and in many ways they still resemble free-living bacteria. Chloroplasts are found in plant cells, contain the green pigment chlorophyll, and are responsible for the reactions of photosynthesis (the process that captures sunlight energy and uses it to power the activities of the cell). Since photosynthesis is the only significant mechanism of energy-input into the living world, chloroplasts are of inestimable importance, not just to plants but to all life on Earth. Actually, chloroplasts belong to a wider family of related organelles called plastids. Other members of the family are the highly-pigmented chromoplasts in ripe fruits, and etioplasts in dark-grown plants.
Although plastids do contain DNA (a relic from their evolutionary past as free-living photosynthetic bacteria), and so can make some of their own proteins, most of the proteins needed to form a functional plastid are encoded on DNA in the cell nucleus; these proteins are made outside of the plastid in the cellular matrix known as the cytosol. As plastids are each surrounded by a double membrane, or envelope, that is impervious to the passive movement of proteins, this presents a significant problem. To overcome the problem, plastids evolved a sophisticated protein import apparatus, which uses energy (in the form of ATP) to drive the import of proteins from the cytosol, across the envelope, to the plastid interior. This import apparatus comprises two molecular machines: one in the outer envelope membrane called TOC (an abbreviation of "Translocon at the outer envelope membrane of chloroplasts"), and another in the inner envelope membrane called TIC. Each machine is made up of several different proteins which cooperate to ensure the efficiency of import.
We work on a model plant called Arabidopsis that has many advantages for research, such as an availability of numerous mutants (each one with a mutation in a specific gene). One such mutant plant, ppi1, has a defect in a TOC gene such that plastid protein import does not work efficiently. Several years ago, we identified another mutation called sp1 (this stands for "suppressor of ppi1") that counteracts the negative effects of ppi1. The gene disrupted by sp1 (the SP1 gene) encodes a type of regulatory protein called a "ubiquitin E3 ligase". These work by labelling-up unwanted proteins and targeting them for removal. Because this control mechanism was not previously known to operate in plastids, this discovery was an important breakthrough in biology. The SP1 E3 ligase carefully controls the composition of the TOC machinery so that the right proteins are always imported (this is normally good, but in the abnormal ppi1 background it is apparently a hindrance). Such control is very important when plastids need to convert from one form to another; e.g. when dark-germinated plants emerge into the light, etioplasts must change into chloroplasts so that photosynthesis can begin.
In this project we will investigate whether SP1 is important for the conversion of chloroplasts into chromoplasts in tomato fruit. If it is, then our work may have commercial, agricultural importance by enabling the manipulation of fruit ripening in crops (e.g. tomato, bell pepper, citrus). We will also study in much greater detail how SP1 and related proteins control plastid development. For example, our work may elucidate how plants respond to stresses like salinity and drought, which are major limits on crop yield across the world. Photosynthetic performance (and thus the energy available to plants for growth) is strongly affected by stress, and we suspect that SP1 is involved in this process. Thus, knowledge gained from our work may enable improved adaptation of crops to adverse environmental conditions.

Technical Summary

Arabidopsis SP1 encodes a RING-type E3 ligase of the chloroplast outer membrane. It targets plastid protein import machinery components (i.e. TOC proteins) for ubiquitination and degradation. SP1 regulates which proteins are imported, and this in turn controls the plastid's proteome, development and functions. Thus, the ubiquitin-proteasome system (UPS) directly regulates plastids. Here we will elucidate the roles of SP1 and UPS in plastid biogenesis, using tomato and Arabidopsis.
SP1 is important for developmental transitions in which plastids convert from one form to another (e.g. de-etiolation). A commercially important example is fruit ripening, when chloroplasts change into chromoplasts; we will study SP1's role in this by altering its expression in transgenic tomato plants. This may eventually enable the manipulation of fruit ripening in crops.
E3 ligases are typically regulated by post-translational modification (PTM). We found that SP1 is developmentally controlled, that it can be ubiquitinated, that it is likely phosphorylated, and that some of its targets are potentially sumoylated. We will study these PTMs to elucidate how SP1 is regulated to optimize plastid protein import. We will also study SP1's role in stress responses, which may involve reorganizing the plastid proteome to optimize photosynthetic activity against photo-oxidative damage.
As chaperones aid extraction of UPS targets from ER and mitochondrial membranes, we propose a similar role for them in plastids. We will test for plastid localization of such chaperones, and assess the effects of their inactivation in planta on TOC protein levels and polyubiquitination, and plastid protein import.
Finally, we will seek new factors involved in the UPS-control of plastid proteins. We will study Arabidopsis SP1 homologues by assessing the effects of gene inactivation in planta, and by using TAP tagging to identify their targets and regulators. We will also use informatics to seek novel factors.

Planned Impact

Academic impact will be large due to the work's interdisciplinarity as detailed in the Academic Beneficiaries section. This will manifest itself in several ways: 1) The work will contribute to scientific advancement, providing new knowledge with relevance in several overlapping fields and disciplines. 2) The work will stimulate international collaboration via the link with Prof Grimm of Humboldt University and via the involvement of Dr Ling as RA. Dr Ling maintains strong links with the prestigious Shanghai institute where he completed his PhD, and may eventually return to China as an independent research leader (whereupon we would expect to maintain collaborative links). 3) The work will contribute to the health of UK plant science due to publicity surrounding the project, the interactions it will foster, and by our hosting of visitors from schools as this will generate enthusiasm for plant biology. 4) The work will deliver highly-trained individuals who will also contribute to the health of UK plant science. Training will result not only from the direct involvement of the research staff but also from Prof Jarvis' supervision of PhD and MSc students who will work on projects related to the proposed work and have daily interaction with the staff.

Our work may have agricultural, commercial and societal impact. It may enable the manipulation of chromoplast development during fruit ripening in crops like tomato, bell pepper and citrus, which are rich in carotenoid pigments of dietary importance. By altering SP1 expression, we might accelerate or inhibit fruit ripening or otherwise alter fruit properties. Thus a preliminary patent application has been filed. We will begin to explore these possibilities using tomato as a model. We also plan to apply for Pathfinder and Follow-On Funding to further develop these commercial ideas. In the longer term our work may facilitate improved adaptation of crops to the environment, as preliminary data suggest that SP1 acts in stress responses, perhaps through effects on photosynthetic performance. Stresses like salinity and drought are major limits on crop yield across the world, which may become more prominent in the future due to climate change.
Unforeseen benefits may arise in the future due to the fundamental importance of the area in which the project is focused: chloroplasts are the site of photosynthesis and synthesize a diversity of products (e.g. starch, amino acids, fatty acids), many of which are vital in mammalian diets. Plastids offer many possibilities for industrial exploitation. Raw materials for biofuel production are derived largely via the actions of plastids. Chloroplasts dominate plant cells enabling foreign proteins to be expressed at very high levels in plastids; greater understanding of plastid biology may facilitate their use as bioreactors. New knowledge on plastids may even have medical applications, as apicomplexan parasites (e.g. malaria) contain a relict plastid. Moreover, our work on SP1 may elucidate related processes in mitochondrial biogenesis; significantly, mitochondria and the ubiquitin proteasome system are both implicated in ageing and neurodegenerative diseases.

The general public and schools will benefit as we will engage with them in various ways. We will develop a schools engagement activity on Chloroplast Biology as part of a two-day event entitled Dynamic DNA organized by GENIE, a Centre for Excellence in Teaching and Learning at the University. Through the University's Botanic Garden (which receives 40k visitors annually) we will contribute to a publicly-circulated newsletter, deliver a public lecture on project-related topics, prepare a display board for placement at the gardens, and contribute to well-attended educational activities for local schools. Finally, we will continue to accept visitors into our lab via different schemes (e.g. sixth-formers funded by the Nuffield Foundation) and engage the media via the University Press Office.

Publications

10 25 50
 
Description Proteins are a diverse group of molecules with primary responsibility for the myriad functions that constitute life. There are thousands of different proteins in any given cell, and the presence/absence or abundance of each one is an important parameter in the governance of cellular function; this is controlled by the balance of protein synthesis and degradation. A critically important mechanism for protein degradation is mediated by the ubiquitin proteasome system (UPS), which was previously thought to act only in central parts of the cell. Our work shows that the UPS also acts on subcellular compartments in plant cells called chloroplasts, which are responsible for the light-driven reactions of photosynthesis that power almost all life on Earth. Identification of this previously-unsuspected link between the UPS and chloroplasts constitutes a major breakthrough in biology, and may enable the manipulation of chloroplast functions in crops.

Using a forward-genetic screen in Arabidopsis, we identified a ubiquitin E3 ligase, termed SP1, embedded in the chloroplast outer envelope membrane. We found that SP1 targets components of the chloroplast protein import machinery (the so-called TOC apparatus) for degradation by the UPS, and in doing so regulates chloroplast biogenesis. As most of the ~3,000 different proteins in chloroplasts are imported via the TOC machinery, its regulation represents an efficient mechanism by which the cell may control the organelle's proteome and functions. The identification of semi-autonomous, endosymbiotic chloroplasts as new, previously-unsuspected targets of UPS activity further extends the known field of influence of this remarkably pervasive and ubiquitous eukaryotic regulatory network.
Exploitation Route Our results suggest that SP1 is important for the developmental transitions that plastids undergo during different phases of plant growth (for example, the differentiation of chloroplasts from etioplasts following the emergence of germinating seedlings into the light, leading to the establishment of photoautotrophic growth). Commercially important plastid developmental transitions (chloroplast to chromoplast) occur during fruit ripening in crops such as tomato, red bell pepper, and citrus. We are exploring the possibility that SP1 may be utilized to manipulate this and other aspects of crop plant development. This technology is covered by a patent application, and is being promoted commercially by PBL, Norwich.
Sectors Agriculture, Food and Drink,Energy

 
Description This work led to the realization that multiple, client-specific protein import pathways operate in plastids, and that these pathways are enabled by the existence of a diversity of client-specific protein import pathways, and by the operation of a regulator of the import machinery called SP1. We believe that manipulation of these import pathways will find beneficial applications in crops, by enabling manipulation of plastid functions and plastid development in a variety of different ways. This technology is covered by a patent application and is being promoted by PBL, Norwich.
First Year Of Impact 2012
Sector Agriculture, Food and Drink
Impact Types Societal

 
Description Application of the plastidic E3 ligase SP1 in crop improvement, using tomato and rice as models
Amount £152,584 (GBP)
Funding ID BB/R005591/1 
Organisation Biotechnology and Biological Sciences Research Council (BBSRC) 
Sector Public
Country United Kingdom
Start 03/2018 
End 03/2019
 
Description BBSRC Follow-On Funding Pathfinder: "Manipulation of the chloroplast-associated protein degradation pathway (CHLORAD) - applications in plant breeding and biotechnology" (Jan - Jul 2019)
Amount £10,897 (GBP)
Funding ID BB/S013873/1 
Organisation Biotechnology and Biological Sciences Research Council (BBSRC) 
Sector Public
Country United Kingdom
Start 01/2019 
End 07/2019
 
Description BBSRC Responsive Mode
Amount £615,440 (GBP)
Funding ID BB/N006372/1 
Organisation Biotechnology and Biological Sciences Research Council (BBSRC) 
Sector Public
Country United Kingdom
Start 04/2016 
End 03/2019
 
Description BBSRC iCASE Studentship
Amount £95,042 (GBP)
Funding ID BB/M015165/1 
Organisation Biotechnology and Biological Sciences Research Council (BBSRC) 
Sector Public
Country United Kingdom
Start 10/2015 
End 09/2019
 
Description Chloroplast-Associated Degradation (CHLORAD): Molecular definition of a ubiquitin-dependent system for plastid protein removal in plants
Amount £537,125 (GBP)
Funding ID BB/R009333/1 
Organisation Biotechnology and Biological Sciences Research Council (BBSRC) 
Sector Public
Country United Kingdom
Start 04/2018 
End 04/2021
 
Description Elucidating the role of SP2 and the SP1-SP2 machinery in chloroplast protein degradation
Amount £498,394 (GBP)
Funding ID BB/R016984/1 
Organisation Biotechnology and Biological Sciences Research Council (BBSRC) 
Sector Public
Country United Kingdom
Start 10/2018 
End 09/2021
 
Title A method to analyse chloroplast protein import using chloroplasts isolated from stressed plants 
Description We developed a method to analyse chloroplast protein import in vitro using chloroplasts isolated from stressed plants. The method enables the assessment of stress treatments on chloroplast protein import. 
Type Of Material Technology assay or reagent 
Year Produced 2015 
Provided To Others? Yes  
Impact These resources supported two publications (see below), and a successful BBSRC grant application (BB/N006372/1). Ling, Q. and Jarvis, P. (2015) Regulation of chloroplast protein import by the ubiquitin E3 ligase SP1 is important for stress tolerance in plants. Curr. Biol. 25: 2527-2534. Ling, Q. and Jarvis, P. (2016) Analysis of Protein Import into Chloroplasts Isolated from Stressed Plants. J. Vis. Exp. (117), e54717, doi:10.3791/54717. 
 
Title Vectors and resulting transgenic plants (Arabidopsis) that over/underexpress SP1 or its homologues, or mutant or tagged (e.g. TAP) forms thereof 
Description Vectors and resulting transgenic plants (Arabidopsis) that over/underexpress SP1 or its homologues, or mutant or tagged (e.g. TAP) forms thereof 
Type Of Material Biological samples 
Provided To Others? No  
Impact Furtherance of our BBSRC-funded research 
 
Title Vectors and resulting transgenic plants (Arabidopsis) that over/underexpress a cytosolic Hsp100-type chaperone 
Description Vectors and resulting transgenic plants (Arabidopsis) that over/underexpress a cytosolic Hsp100-type chaperone. This chaperone is of interest because we hypothesize that it acts on the SP1 pathway for chloroplast protein degradation. 
Type Of Material Biological samples 
Provided To Others? No  
Impact Furtherance of our BBSRC-funded research 
 
Title Vectors and resulting transgenic plants (brassica, AG DH1012) that overexpress or are silenced for SP1 
Description Vectors and resulting transgenic plants (brassica, AG DH1012) that overexpress or are silenced for SP1. The transgenic brassica plants were generated at BRACT, John Innes Centre, Norwich, with support of Plant Biosciences Ltd. (PBL). 
Type Of Material Biological samples 
Provided To Others? No  
Impact The plants are being used to provide proof-of-concept for the application of SP1 in crop improvement. The analysis is still in progress. 
 
Title Vectors and resulting transgenic plants (rice, Kitaake) that overexpress or are silenced for SP1 
Description Vectors and resulting transgenic plants (rice, Kitaake) that overexpress or are silenced for SP1. 
Type Of Material Biological samples 
Provided To Others? No  
Impact The plants are being used to provide proof-of-concept for the application of SP1 in crop improvement. The analysis is still in progress. 
 
Title Vectors and resulting transgenic plants (tomato, Ailsa Craig) that overexpress or are silenced for SP1 and its homologue SPL2 
Description Vectors and resulting transgenic plants (tomato, Ailsa Craig) that overexpress or are silenced for SP1 and its homologue SPL2 
Type Of Material Biological samples 
Provided To Others? No  
Impact Furtherance of our BBSRC-funded research. This work is covered by a PCT stage patent application, and by a licensing agreement in place with PBL, Norwich. 
 
Title Vectors and resulting transgenic plants (wheat, Fielder) that overexpress or are silenced for SP1 
Description Vectors and resulting transgenic plants (wheat, Fielder) that overexpress or are silenced for SP1. The transgenic wheat plants were generated at NIAB, Cambridge, as part of the BBSRC-funded Community Resource for Wheat Transformation. 
Type Of Material Biological samples 
Year Produced 2013 
Provided To Others? Yes  
Impact Furtherance of our BBSRC-funded research. This work is covered by a PCT stage patent application, and by a licensing agreement in place with PBL, Norwich. 
 
Title Vectors encoding site-directed mutant forms of the SP1 protein, such that it lacks ubiquitination sites, for bacterial and plant expression, and resulting transgenic plants (Arabidopsis) 
Description Vectors encoding site-directed mutant forms of the SP1 protein, such that it lacks ubiquitination sites, for bacterial and plant expression, and resulting transgenic plants (Arabidopsis) 
Type Of Material Biological samples 
Provided To Others? No  
Impact Furtherance of our BBSRC-funded research 
 
Description Dr Gail Preston, Department of Plant Sciences, University of Oxford 
Organisation University of Oxford
Department Department of Experimental Psychology
Country United Kingdom 
Sector Academic/University 
PI Contribution We are collaborating on the role of SP1 in biotic stress responses in Arabidopsis and brassica, via a PhD studentship
Collaborator Contribution Expertise in plant pathology
Impact The project is on-going.
Start Year 2015
 
Description Dr Masanori Izumi 
Organisation Tohoku University
Department Graduate School of Life Sciences
Country Japan 
Sector Academic/University 
PI Contribution We are collaborating on the role of ubiquitination in chloroplast degradation by autophagy. Dr Izumi visited the group in Oxford for four months in 2015 on a Japanese government fellowship.
Collaborator Contribution Expertise in autophagy.
Impact The work is on-going.
Start Year 2015
 
Description NIAB, Cambridge 
Organisation National Institute of Agronomy and Botany (NIAB)
Country United Kingdom 
Sector Charity/Non Profit 
PI Contribution Generation of constructs, and analysis of transgenic wheat plants
Collaborator Contribution Transformation of wheat to generate transgenic plants with altered expression of the SP1 protein
Impact Transgenic plants with altered expression of the SP1 protein, and improved understanding of SP1 function and possible applications in crops
Start Year 2013
 
Description Prof. Matthew Terry, University of Southampton 
Organisation University of Southampton
Country United Kingdom 
Sector Academic/University 
PI Contribution We are providing expertise in the area of chloroplast protein import, as well as seeds of relevant mutant genotypes. We will be conducting analyses of the levels of components of the protein import machinery in a new mutant identified by our collaborators that displays defective plastid signalling.
Collaborator Contribution Our collaborators are supplying the new plastid signalling mutant to us, and are conducting detailed analyses on the mutant seed lines that we are providing.
Impact This collaboration is on-going and so has not yet generated any outputs.
Start Year 2016
 
Description Prof. Ralf Bock, Max-Planck Institute of Molecular Plant Physiology, Potsdam, Germany 
Organisation Max Planck Society
Department Max Planck Institute of Molecular Plant Physiology
Country Germany 
Sector Charity/Non Profit 
PI Contribution We are providing expertise in the area of chloroplast protein import in order to understand the role of a plastid signalling mutant. We hosted a visiting postdoctoral researcher from Germany in order that skills in this area may be transferred to our collaborators.
Collaborator Contribution Our collaborators provided the initial observations and the mutant genotypes of interest, and are completing the analyses in readiness for publication.
Impact The collaboration is on-going and so has not yet generated any outputs.
Start Year 2015
 
Title ATSP1, AN E3 UBIQUITIN LIGASE, AND ITS USE 
Description The invention relates to plants with improved phenotypes and related methods. These improved phenotypes are conferred by altering the expression of the SP1 gene which is involved in plastid development or altering the activity of the SP1 protein. 
IP Reference 16/643507 
Protection Patent application published
Year Protection Granted 2015
Licensed Yes
Impact A BBSRC Follow-on Funding Pathfinder grant was awarded in 2013 (BB/FOF/PF/15/12), which related to earlier patent filings of the same technology. A subsequent BBSRC Follow-on Funding Standard grant was applied for in 2017 (BB/R005591/1; application pending).
 
Title CONTROL OF PLASTID ASSOCIATED PROTEIN DEGRADATION 
Description The patent application covers the possibility of manipulating CHLORAD to modify diverse aspects of chloroplast function, enabling novel crop improvement strategies; for example, improving the tolerance of crop plants to abiotic stress. 
IP Reference GB1815206.6 
Protection Patent application published
Year Protection Granted 2018
Licensed No
Impact Too early for impact to be assessed.
 
Title CONTROL OF PLASTID ASSOCIATED PROTEIN DEGRADATION I 
Description The patent application covers the possibility of manipulating SP2 and CHLORAD to modify diverse aspects of chloroplast function, enabling novel crop improvement strategies; for example, improving the tolerance of crop plants to abiotic stress. 
IP Reference GB1803833.1 
Protection Patent application published
Year Protection Granted 2018
Licensed No
Impact Too early for impact to be assessed.
 
Title CONTROL OF PLASTID ASSOCIATED PROTEIN DEGRADATION II 
Description The patent application covers the possibility of manipulating PUX10 (CDC48) and CHLORAD to modify diverse aspects of chloroplast function, enabling novel crop improvement strategies; for example, improving the tolerance of crop plants to abiotic stress. 
IP Reference GB1803834.9 
Protection Patent application published
Year Protection Granted 2018
Licensed No
Impact Too early for impact to be assessed.
 
Title Transgenic Plants. AtSP1, An E3 Ubiquitin Ligase, and Its Use 
Description The invention relates to plants with improved phenotypes and related methods. These improved phenotypes are conferred by altering the expression of the SP1 gene which is involved in plastid development or altering the activity of the SP1 protein. 
IP Reference WO2014037735 
Protection Patent application published
Year Protection Granted 2014
Licensed Yes
Impact The technology is currently in development.
 
Description Gordon Research Conference on Mitochondria and Chloroplasts, Barga, Italy (July 6-11, 2014) 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? Yes
Type Of Presentation keynote/invited speaker
Geographic Reach International
Primary Audience Other audiences
Results and Impact Presentation delivered at conference.

Promoted awareness and outcomes of my BBSRC-funded work, and made contact with researchers in related fields leading possible future collaboration.
Year(s) Of Engagement Activity 2014
URL http://www.grc.org/programs.aspx?year=2014&program=mitochon
 
Description Plant Biology Europe FESPB/EPSO Congress, Dublin, Ireland (June 22-26, 2014) 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? Yes
Type Of Presentation keynote/invited speaker
Geographic Reach International
Primary Audience Other audiences
Results and Impact Presentation as keynote speaker delivered at congress.

Promoted awareness and outcomes of my BBSRC-funded work, and made contact with researchers in related fields leading possible future collaboration.
Year(s) Of Engagement Activity 2014
URL http://europlantbiology.org/
 
Description Article in the BBSRC Business magazine, Winter 2016 
Form Of Engagement Activity A magazine, newsletter or online publication
Part Of Official Scheme? No
Geographic Reach National
Primary Audience Industry/Business
Results and Impact This article highlighted our recent latest results, published in Current Biology, showing that the regulation of protein import into chloroplasts by the SP1 ubiquitin E3 ligase is an important component of the way in which plants tolerate environmental stress.
Year(s) Of Engagement Activity 2016
URL http://www.bbsrc.ac.uk/news/business-magazine/2016/winter-2016/
 
Description BBSRC press release associated with Ling and Jarvis 2015 Current Biology paper on SP1 
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 Media (as a channel to the public)
Results and Impact Much media interest was generated as a result of this press release, which focused on the role of chloroplast protein import regulation by SP1 in plant responses to abiotic stress, published in October 2015 (Current Biology 25: 2527-34).

The press release was the headline story on the front page of the BBSRC website on publication.
Year(s) Of Engagement Activity 2015
URL http://www.bbsrc.ac.uk/news/food-security/2015/150918-pr-plant-discovery-help-develop-stress-resista...
 
Description FEBS Workshop, Plant Organellar Signaling - From Algae to Higher Plants, 2011, Primošten, Croatia 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? Yes
Geographic Reach International
Primary Audience Other audiences
Results and Impact Participant; poster presentation International networking; inspiration for research; esteem

no actual impacts realised to date
Year(s) Of Engagement Activity 2011
 
Description Grenoble Alliance for Integrated Structural Cell Biology (GRAL) Workshop, Autrans, France (April 7-8, 2014) 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? Yes
Type Of Presentation keynote/invited speaker
Geographic Reach International
Primary Audience Other audiences
Results and Impact Keynote presentation delivered at scientific workshop.

Promoted awareness and outcomes of my BBSRC-funded work, and made contact with researchers in related fields leading possible future collaboration.
Year(s) Of Engagement Activity 2014
URL http://www.labex-gral.fr/events
 
Description Hosting A-level student on Nuffield Research Placement (Adam Bills, King Edward VII College, Coalville) 
Form Of Engagement Activity Participation in an open day or visit at my research institution
Part Of Official Scheme? Yes
Geographic Reach Local
Primary Audience Schools
Results and Impact The student, Adam Bills from King Edward VII College, Coalville, participated in scientific research for a one month period, working alongside Dr Jocelyn Bedard in the laboratory.

The student developed better understanding of scientific research and biology during the course of his placement, and this will be likely to positively inform his future career choices.
Year(s) Of Engagement Activity 2013
URL http://www.nuffieldfoundation.org/nuffield-research-placements
 
Description International Symposium on the Regulation of Photosynthetic Function, Guilin, China (August 16-20, 2014) 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? Yes
Type Of Presentation keynote/invited speaker
Geographic Reach International
Primary Audience Other audiences
Results and Impact Presentation delivered at congress.

Promoted awareness and outcomes of my BBSRC-funded work, and made contact with researchers in related fields leading possible future collaboration.
Year(s) Of Engagement Activity 2014
URL http://rpf2014.csp.escience.cn/dct/page/1
 
Description Interview for Australian Radio, ABC Rural 
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 Media (as a channel to the public)
Results and Impact An interview with Australian ABC Rural radio was conducted in order to promote our latest results, published in Current Biology, which showed that the regulation of protein import into chloroplasts by the SP1 ubiquitin E3 ligase is an important component of the way in which plants tolerate environmental stress.
Year(s) Of Engagement Activity 2015
URL http://www.abc.net.au/news/2015-10-05/plant-stress-discovery/6823334
 
Description Interview for The Guardian 
Form Of Engagement Activity A press release, press conference or response to a media enquiry/interview
Part Of Official Scheme? No
Geographic Reach National
Primary Audience Media (as a channel to the public)
Results and Impact An interview with The Guardian was conducted in order to promote our latest results, published in Current Biology, which showed that the regulation of protein import into chloroplasts by the SP1 ubiquitin E3 ligase is an important component of the way in which plants tolerate environmental stress.
Year(s) Of Engagement Activity 2015
URL http://www.theguardian.com/environment/world-on-a-plate/2015/oct/07/scientists-find-gene-that-makes-...
 
Description Invited speaker at GRC Protein Transport Across Cell Membranes Meeting entitled "Protein Transport Across Cell Membranes: Mechanism, Structure, and Regulation" (Texas, USA, 2016) 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact I was an invited speaker at this prestigious international meeting, which took place during 6-11 March 2016, at Hotel Galvez, Galveston, TX, USA.
Year(s) Of Engagement Activity 2016
URL https://www.grc.org/protein-transport-across-cell-membranes-conference/2016/
 
Description Keynote Speaker at Society of Experimental Biology (SEB) Annual Meeting (Gothenburg, Sweden, 2017) 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact I was a keynote speaker at this prestigious international meeting, which took place during 3-6 July 2017, at Gothenburg, Sweden.
Year(s) Of Engagement Activity 2017
URL http://www.sebiology.org/events/event/seb-gothenburg
 
Description Plenary Speaker and Session Chair at the 27th International Conference on Arabidopsis Research (ICAR) (Gyeong Ju, Korea, 2016) 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact I was a plenary speaker and session at the is prestigious international meeting, which took place during 29 June-3 July 2016, at Gyeong Ju, South Korea.
Year(s) Of Engagement Activity 2016
URL http://www.arabidopsisresearch.org/images/ICAR/ICAR2016_programofICAR2016_160418.pdf
 
Description Talk given to A-level students as part of Study Day at Oxford's Museum of Natural History 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach Local
Primary Audience Schools
Results and Impact This talk was part of an engagement event (a Study Day) organized by Oxford's Museum of Natural History and Botanic Garden. It was a special day for A-level biologists and it focused on cells; my contribution was a presentation covering the evolution of the plant cell with a particular focus on chloroplast evolution. Approximately 300 students attended from a mixture of local schools. All students were between 16 and 18 years old and had chosen to study biology.
Year(s) Of Engagement Activity 2014
URL http://www.oum.ox.ac.uk/educate/index.htm
 
Description University press release associated with Ling and Jarvis 2015 Current Biology paper on SP1 
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 Media (as a channel to the public)
Results and Impact Much media interest was generated as a result of this press release, which focused on the role of chloroplast protein import regulation by SP1 in plant responses to abiotic stress, published in October 2015 (Current Biology 25: 2527-34).
Year(s) Of Engagement Activity 2015
URL http://www.ox.ac.uk/news/2015-09-17-photosynthesis-gene-could-help-crops-grow-adverse-conditions