Understanding major transitions in individuality

Lead Research Organisation: University of St Andrews
Department Name: Biology

Abstract

Major transitions in individuality occur when a group of separate organisms evolves into a higher-level organism in its own right. For example, the ancestors of the social insects used to live a solitary existence, but today they form highly elaborate, cooperative societies that can be considered "superorganisms" in their own right. Current theory describing major transitions in individuality highlights the importance of clonal reproduction and systems of policing in maintaining the integrity of the superorganism. However, the social insects appear to have evolved a higher level of individuality on the basis of strict lifetime monogamy, which has abolished conflicts of interest within the colony. I will develop new theory to incorporate this monogamy hypothesis into the theory of major transitions in individuality, and I will investigate how the quirky genetics of some species can lead to other forms of conflict arising within their societies.

Planned Impact

The successful media coverage of my previous work on social evolution has demonstrated that the general public has a profound interest in this topic, so they stand to benefit from the research in this respect. In general, the public are interested in evolution, because we are all interested in our origins. Social evolution is of particular interest, because it illuminates our social origins, and the reasons for love and strife within families. The British public also take a special interest in natural history, including the alien biology of the insect world.

Of more applied interest, my research may eventually inform agricultural practice and policy. The social insects are of enormous economic significance: for example, the honeybee is responsible for much pollination of crop plants. Understanding the factors that maintain the integrity of the colony may help protect bee colonies from collapse due to pressure from social parasites. My work on arbuscular mycorrhizal fungi and their mutualism with plant roots may lead to the development of new cultivation techniques to maximize crop productivity. Although I will not be directly developing new agricultural methodology in this research, my collaborative partnership with Dr Kiers -- who does interact closely with farmers -- will ensure that such insights are carried through to novel agricultural technology.

Publications

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Faria GS (2017) Sexual selection modulates genetic conflicts and patterns of genomic imprinting. in Evolution; international journal of organic evolution

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Farrell EJ (2015) Intragenomic Conflict over Dispersal. in The American naturalist

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Gardner A (2015) More on the genetical theory of multilevel selection in Journal of Evolutionary Biology

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Gardner A (2015) Group selection versus group adaptation. in Nature

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Gardner A (2023) The rarer-sex effect. in Philosophical transactions of the Royal Society of London. Series B, Biological sciences

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Gardner A (2015) Hamilton's Rule. in The American naturalist

 
Description 1. I have developed a theory of multi-level selection for application to class-structured populations. Published in Journal of Evolutionary Biology.

2. I have developed a general model of the evolution of genome elimination (including paternal genome elimination, an important form of male haploidy) under inbreeding, XY/XO/ZW/ZO sex determination, zygotic/maternal/paternal effects, autosomal or sex-chromosomal inheritance, and imprinted versus unimprinted gene expression. This work has explained why, across 1000s of animal species, genome expression typically occurs under male heterogamety, under inbreeding and in the form of elimination of the paternal genome. Published in Ecology Letters.

3. To facilitate my investigation of the interplay between kin selection and male haploidy, I have proven the crucial mathematical result that, collectively, juvenile females have twice the reproductive value of juvenile males in male haploid populations, even though, collectively, the total reproductive value of all females need not be twice that of all males, in the context of age-structure. Published in Journal of Theoretical Biology.

4. I have developed theory for genomic imprinting conflict to explain the observed imprinting of gene Grb10 in mouse pups and their mothers' mammary tissue, showing that this pattern of gene expression may be driven by a conflict of interest between paternal-origin genes versus maternal-origin genes in both pups and their mothers. Published in PLOS Biology.

5. I have provided formal mathematical support for key features of the analogy of 'biological markets' as applied to plant-mycorrhizal interactions and demonstrated its theoretical connections with evolutionary game theory. Published in Evolution.

6. I have reviewed and synthesised research on the topic of the major transitions in individuality. Published in Proceedings of the National Academy of Sciences, USA.

7. I have developed theory showing that two celebrated sex-ratio invariant results, concerning constancy in the number of males with respect to variation in dispersal rate or variance in fecundity, break down when both sources of variation are occurring simultaneously. Published in Proceedings B.

8. I have further developed the theory of multi-level selection in class-structured populations, with reference to the Price-Hamilton versus contextual-analysis approaches to multi-level selection. Published in Journal of Evolutionary Biology.

9. I have developed theory on individual-level and group-level adaptation in social spiders. Published in Nature.

10. I have developed theory on intragenomic conflict with regard to whether or not the individual should engage in dispersal behaviour, from which I have derived testable predictions concerning patterns of genomic imprinting of genes underpinning dispersal phenotypes. Published in American Naturalist.

11. I have developed theory showing that mutualistic partners may be favoured to harm each other in order to stabilise the mutualism, with particular application to plant-mycorrhizal interactions. Published in Nature Communications.

12. I have developed theory investigating the consequences of kin selection on the evolution of sexual-conflict behaviours. Published in Journal of Evolutionary Biology.

13. I have developed theory demonstrating an invariant result for the evolution of dispersal, whereby each mother produces the same number of non-dispersing offspring irrespective of variation in the total number of offspring produced by each mother. Published in Journal of Evolutionary Biology.

14. I have developed theory demonstrating that a 'loner' strategy can stabilise bacterial cooperation and ecological diversity, and have confirmed the predictions of this theory using evolutionary experiments with the opportunities human pathogen Pseudomonas aeruginosa. Published in Proceedings B.

15. I have developed theory demonstrating the potential existence of intragenomic conflict with regard to soldiering behaviour in polyembryonic parasitoid wasps, deriving testable predictions concerning patterns of genomic imprinting that may allow discrimination between competing hypotheses as to these soldiers' primary function. To be published in American Naturalist.

16. I have synthesised theory and recent recent research findings concerning the bizarre female biased sex ratios of quasi social parasitoid wasps, pointing out directions for future research. To be published in a special issue of Entomologia Experimentalis et Applicata.

17. I have developed theory showing that sex-biased dispersal can drive intragenomic conflict and genomic imprinting in relation to sexual conflict phenotypes. To be published in Evolution.

18. I have developed theory showing that sex-biased dispersal can drive intrafamily and intragenomic conflict in relation to human warfare, yielding predictions for corresponding patterns of genomic imprinting and associated human behavioural disorders. To be published in Proceedings of the Royal Society of London B.

19. I have developed theory showing the interplay between sex allocation and the evolution of sib-altruism, pointing to a key role for the ecology of sex in driving patterns of sociality in arthropods. Published in Ecology Letters.

20. I have reviewed misconceptions on the application of the theory of biological markets to plant-fungus symbioses. Published in Nature Plants.
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21. I have developed and analysed models of viral "germline" investment in the context of chronic viral infections. Published in Trends in Microbiology.

22. I have developed a formal and fully comprehensive theory of intragenomic conflict. Published in Nature Ecology & Evolution.

23. I have discussed the rationale and formal underpinnings of optimality / adaptationist approaches in evolutionary biology. Published in Interface Focus.

24. I have developed and analysed models of intragroup and intragenomic conflict in relation to anti-predator chemical defence in insects, yielding predictions that may facilitate gene discovery in relation to agriculturally important products. Published in Ecology and Evolution.

25. I have developed theory showing how the effect of infraspecific relatedness on the evolution of public goods cooperation is diminished in the context of multi species communities. Published in Evolution.

26. I have developed theory showing that male-only warfare may evolve even when there are no underlying sex differences in ecology, owing to positive feedbacks that are able to magnify even chance differences that arise between the sexes. Published in Proceedings B.

27. I have developed theory on human brain evolution suggesting that ecological challenges rather than social challenges better explain why humans have evolved large brains. Published in Nature.

28. I have provided an overview and synthesis of the theory of group selection. Published in Oxford Bibliographies in Evolutionary Biology.

29. I have developed theory showing the formal link between R. A. Fisher's "sexy son effect" in the sexual selection literature, on the one hand, and W. D. Hamilton's "greenbeard effect" in the kin selection literature, on the other hand. Published in Evolution Letters.

30. I have developed theory on the application of kin selection to social interaction among affinal kin. Published in Biology Letters.

31. I have developed theory showing that female monogamy is a robust promoter of altruistic sterility in insect societies. Published in Royal Society Open Science.

32. I have reviewed and synthesised the evolutionary literature on the link between dispersal and genomic imprinting. Published in Current Opinion in Behavioural Science.

33. I have developed theory on the social evolutionary drivers of sleep, with a focus on sex differences, intragenomic conflicts and sleep disorders / pathologies. Published in Proceedings B.

34. I have developed theory on the evolution of haplodiploidy, determining under which sex determination mechanisms a haploid offspring developing from an unfertilised egg is expected to be male, and showing that inbreeding does not impede the evolutionary invasion of male haploidy as had previously been thought. Published in Evolution Letters.

35. I have developed theory on sexual selection, investigating the role for interspecific reproductive interference in structured populations. Published in Evolution.

36. I have developed theory on the evolution of human language, showing that language genes are expected to be embroiled in intragenomic conflicts and to evolve genomic imprinting, with patterns of imprint yielding information as to the selection pressures that have shaped language. Published in BioEssays.

37. I have provided an account of the use of the agent concept in evolutionary biology, highlighting its scientific utility. Published in Metascience.

38. I have developed theory showing that resource heterogeneity promotes the evolution of cooperation and provided experimental confirmation using bacteria. Published in Evolution Letters.

39. I have shown that the demography of human warfare is expected to modulate the evolution of social behaviours in a sex-specific way. Published in Evolutionary Human Sciences.

40. I have shown that a gene's-eye view of sexual antagonism resolves recent apparently contradictory observations and enables novel predictions concerning the effects of dosage compensation, inbreeding and age structure on sex-specific evolution. Published in Proceedings B.

41. I have developed theory investigating how females should adjust their sex allocation in the context of local mate competition in response to variation in relatedness and co-foundress number. Published in Journal of Evolutionary Biology.

42. I have explained the usefulness of Price's equation, clarifying its role in the definition rather than the prediction of evolutionary forces. Published in Phil Trans B.

43. I have developed theory on the role for kin discrimination and budding dispersal to modulate the degree to which males harm females in pursuit of enhanced mating success. Published in Nature Ecology & Evolution.

44. I have developed theory pinpointing the conditions under which kin discrimination is expected to promote as opposed to inhibit the evolution of cooperation, linking this with Jensen's inequality. Published in Biology Letters.

45. I have used genetic analysis to provide evidence in support of the hypothesis that the cottony cushion scale insect Icerya purchasi has an androdioecious mating system. Published in Evolution.

46. I have developed theory showing how different portions of the genome may come into conflict with respect to harming behaviour that males enact towards females. Published in Proceedings B.

47. I have developed theory showing how haplodiploidy and related systems of inheritance modulate the evolution of sexual antagonism. Published in Evolution.

48. I have explained the role of reproductive value as a modulator of the evolution of altruism, with application to diverse taxa including microbes, insects and birds. Published in Trends in Ecology and Evolution.
Exploitation Route Several of the contributions made so far have been methodological, and will enable me to conduct the more applied parts of the research proposal. They will, however, be useful for other researchers pursuing other research topics. Moreover, even for these primarily methodological contributions, I have developed novel theoretical predictions that empirical researchers may now go on to test as part of their own research programmes.
Sectors Agriculture, Food and Drink,Communities and Social Services/Policy,Environment,Healthcare,Government, Democracy and Justice,Pharmaceuticals and Medical Biotechnology

URL http://synergy.st-andrews.ac.uk/gardner/
 
Description Consolidator Grant
Amount € 1,968,688 (EUR)
Funding ID 771387 
Organisation European Research Council (ERC) 
Sector Public
Country Belgium
Start 05/2018 
End 04/2023
 
Description John Templeton Foundation
Amount £130,779 (GBP)
Organisation The John Templeton Foundation 
Sector Academic/University
Country United States
Start 06/2017 
End 05/2019
 
Description Marie Sklodowska-Curie Fellowship
Amount £130,303 (GBP)
Organisation European Commission 
Sector Public
Country European Union (EU)
Start 10/2016 
End 09/2018
 
Description Newton Fellowship
Amount £92,550 (GBP)
Organisation The Royal Society 
Sector Charity/Non Profit
Country United Kingdom
Start 01/2017 
End 12/2018
 
Title Complete Numerical Solutions For "Inference Of Ecological And Social Drivers Of Human Brain-Size Evolution" By Mauricio González-Forero And Andy Gardner 
Description This zip file contains the complete numerical solutions across the parameter sweep over the P parameters for the six cases considered. /1RatioForm/ -> solutions for power competence.
/2DiffForm/ -> solutions for exponential competence. /1RatioForm/1BenchmarkFromSimpleInitialGuess -> solution for the step 1 of initialization (section 5 of the SI).
/1RatioForm/2Benchmark/ -> solution for the step 2 of the initialization (section 5 of the SI).
/1RatioForm/3AdditiveCoop/ -> solutions across the P parameter combinations for additive cooperation.
/1RatioForm/4MultCoop/ -> solutions across the P parameter combinations for multiplicative cooperation.
/1RatioForm/5SubMultCoop/ -> solutions across the P parameter combinations for submultiplicative cooperation. The contents of /2DiffForm/ are analogous. The P vector is written in these files in the order (etas,etac,etaC,etag), where
etas -> P1
etac -> P3
etaC -> P2
etag -> P4 The files 1runNotesACMC.pdf and 2runNotesSC.pdf contain the tree structure of the parameter sweep, specifying which parameter combination was used as the resident and which combinations converged to an uninvadable strategy (those with a checkmark). The file 3runNotesMaternalCareOptimization.pdf contains the 10 parameter combinations that yielded the best adult fit, which then were subject to variation in the parameter phi to find the combination that yielded the best ontogenetic fit. The file 4runNotesDuplicates.pdf gives the parameter combinations that were not run because they are equivalent to other parameter combinations. Running [T,N1,run,Tshort,N1short,runShort]=etaCombinations in Matlab and typing run.seed{i}.parallel{:} gives the "next" parameter combinations from parameter combination i (where i is a number 1,2,...) for PC-AC, EC-MC, PC-SC, and EC-SC. The meaning of "next" is explained in step 4 of the parameter sweep (section 5 of the SI). Typing runShort.seed{i}.parallel{:} gives the "next" parameter combinations from parameter combination i for PC-MC and EC-AC. The terminal folders contain the solutions and have the following files:
brainNashDeep.m -> the master file launching the iteration of best responses.
brainMainDeep.m -> the file launching one iteration solving the optimal control problem to find best response.
brainMainTestRunDeep.m -> runs a test to check if there are infeasibility warnings.
brainContinuous.m -> specifies the dynamic constraints.
brainEndpoint.m -> specifies the terminal constraints.
parameters.m -> specifies the parameter values and rescales them to rescale units as specified in section 5 of the SI.
getSolution.m -> extracts solution.
plots.m -> plots solutions over best response iterations.
brainPlot.m -> plots solution of a given best response iteration.
guessDeep.mat -> initial guess and resident used.
solutionNashDeep.mat -> solutions over best response iterations.
solutionDeep.mat -> solution of last best response iteration. 
Type Of Material Database/Collection of data 
Year Produced 2018 
Provided To Others? Yes  
 
Title Data from: Co-foundress confinement elicits kinship effects in a naturally sub-social parasitoid 
Description Kinship among interacting individuals is often associated with sociality and also with sex ratio effects. Parasitoids in the bethylid genus Goniozus are sub-social, with single foundress females exhibiting post-ovipositional maternal care via short-term aggressive host and brood defence against conspecific females. Due to local mate competition (LMC) and broods normally being produced by a single foundress, sex ratios are female biased. Contests between adult females are, however, not normally fatal and aggression is reduced when competing females are kin, raising the possibility of multi-foundress reproduction on some hosts. Here we screen for further life-history effects of kinship by varying the numbers and relatedness of foundresses confined together with a host resource and also by varying the size of host. We confined groups of 1 to 8 Goniozus nephantidis females together with a host for 5+ days. Multi-foundress groups were either all siblings or all non-siblings. Our chief expectations included that that competition for resources would be more intense among larger foundress groups but diminished by both larger host size and closer foundress relatedness, affecting both foundress mortality and reproductive output. From classical LMC theory, we expected that offspring group sex ratios would be less female biased when there were more foundresses and from extended LMC theory we expected that sex ratios would be more female biased when foundresses were close kin. We found that confinement led to the death of some females (11% overall) but only when host resources were most limiting. Mortality of foundresses was less common when foundresses were siblings. Developmental mortality among offspring was considerably higher in multi-foundress clutches but was unaffected by foundress relatedness. Groups of sibling foundresses collectively produced similar numbers of offspring to non-sibling groups. There was little advantage for individual females to reproduce in multi-foundress groups: single foundresses suppressed even the largest hosts presented and had the highest per capita production of adult offspring. Despite single-foundress reproduction being the norm, G. nephantidis females in multi-foundress groups appear to attune sex allocation according to both foundress number and foundress relatedness: broods produced by sibling foundresses had sex ratios similar to broods produced by single foundresses (ca. 11% males) whereas the sex ratios of broods produced by non-sibling females were approximately 20% higher and broadly increased with foundress number. We conclude that relatedness and host size may combine to reduce selection against communal reproduction on hosts and that, unlike other studied parasitoids, G. nephantidis sex ratios conform to predictions of both classical and extended LMC theory. 
Type Of Material Database/Collection of data 
Year Produced 2020 
Provided To Others? Yes  
URL http://datadryad.org/stash/dataset/doi:10.5061/dryad.905qftth8
 
Title Data from: Monogamy promotes altruistic sterility in insect societies 
Description  
Type Of Material Database/Collection of data 
Year Produced 2018 
Provided To Others? Yes  
 
Title Data from: Presence of a loner strain maintains cooperation and diversity in well-mixed bacterial communities 
Description Cooperation and diversity abound in nature despite cooperators risking exploitation from defectors and superior competitors displacing weaker ones. Understanding the persistence of cooperation and diversity is therefore a major problem for evolutionary ecology, especially in the context of well-mixed populations, where the potential for exploitation and displacement is greatest. Here, we demonstrate that a 'loner effect', described by economic game theorists, can maintain cooperation and diversity in real-world biological settings. We use mathematical models of public-good-producing bacteria to show that the presence of a loner strain, which produces an independent but relatively inefficient good, can lead to rock-paper-scissor dynamics, whereby cooperators outcompete loners, defectors outcompete cooperators and loners outcompete defectors. These model predictions are supported by our observations of evolutionary dynamics in well-mixed experimental communities of the bacterium Pseudomonas aeruginosa. We find that the coexistence of cooperators and defectors that produce and exploit, respectively, the iron-scavenging siderophore pyoverdine, is stabilized by the presence of loners with an independent iron-uptake mechanism. Our results establish the loner effect as a simple and general driver of cooperation and diversity in environments that would otherwise favour defection and the erosion of diversity. 
Type Of Material Database/Collection of data 
Year Produced 2016 
Provided To Others? Yes  
URL http://datadryad.org/stash/dataset/doi:10.5061/dryad.k5n4k
 
Title Data from: Sexual selection in complex communities: integrating interspecific reproductive interference in structured populations 
Description  
Type Of Material Database/Collection of data 
Year Produced 2019 
Provided To Others? Yes  
 
Title Population viscosity promotes altruism under density-dependent dispersal (Simulation data) 
Description This dataset contains data generated by individual-based simulations described in the paper 'Population viscosity promotes altruism under density-dependent dispersal' by Jasmeen Kanwal and Andy Gardner. Twenty-five separate simulations were run: five different values of patch size (n = 1, 2, 3, 5, 10) each with five different values of cost of dispersal (c = 0, 0.2, 0.4, 0.6, 0.8). The dataset consists of a compressed folder which must be unzipped to extract 50 CSV files (which can then be opened with any text editor). Each simulation generated two CSV files. The files are named using the following formats. n_[value]_c_[value]_[date & time generated].csv - One line per generation, each with six values: population MEDIAN values for gene 1 (dispersal intercept 1), gene 2 (dispersal intercept 2), and gene 3 (level of altruism), and population MEAN values for genes 1, 2, and 3. n_[value]_c_[value]_[date & time generated]_disp.csv - Detailed dispersal data for the last 1000 generations of a simulation run. One line per patch, each with three values: generation number, number of surviving offspring in patch, number of these that disperse. The python code used to generate the data, and the Jupyter notebook used to analyse it, are both freely available at https://github.com/jkanwal/viscosity-promotes-altruism. 
Type Of Material Database/Collection of data 
Year Produced 2021 
Provided To Others? Yes  
URL https://risweb.st-andrews.ac.uk/portal/en/datasets/population-viscosity-promotes-altruism-under-dens...
 
Description Alberto Micheletti & Graeme Ruxton 
Organisation University of St Andrews
Department Earth & Environmental Sciences
Country United Kingdom 
Sector Academic/University 
PI Contribution PhD supervision, project formulation, mathematical modelling, paper writing.
Collaborator Contribution PhD supervision, project formulation, mathematical modelling, paper writing.
Impact Micheletti AJC, Ruxton GD & Gardner A (in press) Intrafamily and intragenomic conflicts in human warfare. Proceedings of the Royal Society of London Series B - Biological Sciences.
Start Year 2015
 
Description Angus Buckling and colleagues 
Organisation University of Exeter
Country United Kingdom 
Sector Academic/University 
PI Contribution Project formulation, development and analysis of mathematical models, paper writing.
Collaborator Contribution Project formulation, experimental research, statistical analysis, paper writing.
Impact Stilwell P, O'Brien S, Hesse E, Lowe C, Gardner A & Buckling A (2020) Resource heterogeneity and the evolution of public-goods cooperation. Evolution Letters doi: 10.1002/ev13.158
Start Year 2019
 
Description Antonio Rodrigues 
Organisation University of Cambridge
Country United Kingdom 
Sector Academic/University 
PI Contribution Co-authored papers
Collaborator Contribution Co-authored papers
Impact Rodrigues AMM & Gardner A (2016) The constant philopater hypothesis: a new life history invariant for dispersal evolution. Journal of Evolutionary Biology 29, 153-166. Rodrigues AMM & Gardner A (2015) Simultaneous failure of two sex-allocation invariants: implications for sex-ratio variation within and between populations. Proceedings of the Royal Society of London Series B - Biological Sciences 282, 20150570.
Start Year 2014
 
Description Erica Kiers, Stuart West, Gregory Wyatt, Heike Bücking, Gijsbert Werner 
Organisation Free University of Amsterdam
Country Netherlands 
Sector Academic/University 
PI Contribution paper writing
Collaborator Contribution paper writing
Impact Kiers ET, West SA, Wyatt GAK, Gardner A, Bücking H & Werner G (2016) Misconceptions on the application of biological market theory to the mycorrhizal symbiosis. Nature Plants 2, 16063.
Start Year 2016
 
Description Francisco Ubeda 
Organisation Royal Holloway, University of London
Country United Kingdom 
Sector Academic/University 
PI Contribution Mathematical modelling & paper writing
Collaborator Contribution Mathematical modelling & paper writing
Impact Úbeda F & Gardner A (in press) Mother and offspring in conflict: why not? PLoS Biology.
Start Year 2014
 
Description Ian Hardy and colleagues 
Organisation University of Nottingham
Country United Kingdom 
Sector Academic/University 
PI Contribution Co-authored paper
Collaborator Contribution Co-authored paper
Impact Kapranas A, Hardy ICW, Tang X, Gardner A & Li B (in press) Sex ratios, virginity and local resource enhancement in a quasisocial parasitoid. Entomologia Experimentalis et Applicata.
Start Year 2015
 
Description Laura Ross 
Organisation University of Edinburgh
Country United Kingdom 
Sector Academic/University 
PI Contribution Mathematical modelling & paper writing
Collaborator Contribution Paper writing
Impact Gardner A & Ross L (in press) Mating ecology explains patterns of genome elimination. Ecology Letters.
Start Year 2014
 
Description Lythgoe and colleagues 
Organisation University of Oxford
Country United Kingdom 
Sector Academic/University 
PI Contribution Developing and analysing mathematical models of viral "germline" optimal investment in the context of chronic viral infections.
Collaborator Contribution Provided framing of these mathematical results in their biological context.
Impact Lythgoe KA, Gardner A, Pybus OG & Grove J (2017) Short-sighted virus evolution and a germline hypothesis for chronic viral infections. Trends in Microbiology 25, 336-348.
Start Year 2016
 
Description Pau Carazo and colleagues 
Organisation University of Valencia
Country Spain 
Sector Academic/University 
PI Contribution Project formulation, development and analysis of mathematical models, paper writing.
Collaborator Contribution Project formulation, interpretation of results, paper writing.
Impact Faria GS, Gardner A & Carazo P (2020) Kin discrimination and demography modulate patterns of sexual conflict. Nature Ecology & Evolution 4, 1141-1148.
Start Year 2019
 
Description Pau Carazo and colleagues 
Organisation University of Valencia
Country Spain 
Sector Academic/University 
PI Contribution Project formulation, development and analysis of mathematical models, paper writing.
Collaborator Contribution Project formulation, interpretation of results, paper writing.
Impact Faria GS, Gardner A & Carazo P (2020) Kin discrimination and demography modulate patterns of sexual conflict. Nature Ecology & Evolution 4, 1141-1148.
Start Year 2019
 
Description Petri Rautiala 
Organisation University of Jyvaskyla
Country Finland 
Sector Academic/University 
PI Contribution Co-authored paper
Collaborator Contribution Co-authored paper
Impact Rautiala P & Gardner A (in press) Intragenomic conflict over soldier allocation in polyembryonic parasitoid wasps. American Naturalist.
Start Year 2015
 
Description Rolf Kuemmerli & colleagues 
Organisation University of Zurich
Country Switzerland 
Sector Academic/University 
PI Contribution Co-authored paper
Collaborator Contribution Co-authored paper
Impact Inglis RF, Biernaskie JM, Gardner A & Kümmerli R (2016) Presence of a loner strain maintains cooperation and diversity in well-mixed bacterial communities. Proceedings of the Royal Society of London Series B - Biological Sciences 283, 20152682.
Start Year 2014
 
Description Silvia Paracchini 
Organisation University of St Andrews
Country United Kingdom 
Sector Academic/University 
PI Contribution Co-supervision of PhD project (60%) on genomic imprinting analysis.
Collaborator Contribution Co-supervision of PhD project (30%) on genomic imprinting analysis.
Impact None at this time.
Start Year 2017
 
Description Susana Varela 
Organisation University of Lisbon
Country Portugal 
Sector Academic/University 
PI Contribution Co-supervision of PhD student
Collaborator Contribution Co-supervision of PhD student
Impact Faria GS, Varela SAM & Gardner A (2015) Sex-biased dispersal, kin selection and the evolution of sexual conflict. Journal of Evolutionary Biology 28, 1901-1910.
Start Year 2015
 
Title Computer Code For "Inference Of Ecological And Social Drivers Of Human Brain-Size Evolution" 
Description This software performs the numerical solutions and produces the parameter-sweep figures for the paper "Inference of ecological and social drivers of human brain-size evolution", by Mauricio González-Forero and Andy Gardner. 
Type Of Technology Software 
Year Produced 2018 
 
Description YAS-MSP Pairing Scheme 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach National
Primary Audience Policymakers/politicians
Results and Impact I developed the Royal Society of Edinburgh Young Academy of Scotland's YAS-MSP Pairing Scheme, which sees five YAS members paired up with five Members of the Scottish Parliament for reciprocal shadowing at the Scottish Parliament, during the MSP's constituency work and at the YAS member's institution (i.e. university). The scheme also involves a day of group activities for ~20 YAS members at the Scottish Parliament, including a tour, briefing from the Scottish Parliament Information Centre (SPICe) and sitting in on Committees. The 2019 round of the scheme was intended as a pilot, to assess its viability and usefulness. It was judged a success, and so will now continue in perpetuity.

I also took part in the pairing during the first round, shadowing Andy Wightman MSP at the Scottish Parliament and hosting a visit from him at the University of St Andrews.
Year(s) Of Engagement Activity 2019
URL https://www.youngacademyofscotland.org.uk/our-work/smarter/msp-pairing/