Project title: Genetic variation at range margins in monkeyflowers

Supervisors with affiliations: Alex Twyford, Richard Ennos

CASE partnership: no case partnership
Primary supervisor contact email: Alex.Twyford@ed.ac.uk
Project background - Species may experience exceptionally variable environments

across their range, with the most dramatic environmental differences seen between
populations at the extreme margins of their distribution. In widespread North
American species, major differences in climate occur over 30 degrees of latitude, and
as such these conditions are likely to select for locally adaptive variation that
increase survivorship at their home site. Despite an increasing interest in the
evolutionary implications of local adaptation, our knowledge of its genetic basis is
limited to relatively few study systems. This project aims to understand the genetic
basis underlying adaptive trait variation in the monkeyflower Mimulus guttatus. Using
new genomic approaches for genetically mapping trait differences among
populations, this project will investigate divergence between populations of M.
guttatus over their 4000km native range. By identifying the genomic regions involved
in adaptive trait divergence, this project aims to better understand the extent of
standing genetic variation in this exceptionally diverse model plant species.
Key research questions
a. Which genomic regions underlie local adaptation, as revealed by a genome-wide
association study in range-wide populations of M. guttatus?
b. How does variation for key traits relate to divergence in chromosome structure
between populations of M. guttatus at their range limits?
c. How do recombinant genotypes between northern and southern populations
survive in a novel environment?
Methodology
These questions will be addressed using a combination of new and pre-existing genomic
data, fieldwork in North America, and by using experimental growth trials. We have
previously genotyped 70 populations of M. guttatus for 40,000 SNP loci, and this data will
form the foundation of a genome-wide association study for local adaptation. This will be
supplemented with genotypes from new populations to be collected in the Pacific North
West. Additional growth trials will be performed at the Royal Botanic Garden Edinburgh,
where we are currently performing large-scale trails of experimental populations.
Training
A comprehensive training programme will be provided comprising both specialist scientific
training and generic transferable and professional skills. The student will be taught
experimental design for studies of quantitative genetics, how to generate next-generation
sequencing data, a variety of genetic analyses (particularly QTL mapping), and skills related
to experiments in plant evolutionary biology.
Requirements
No specific technical skills are required for this position - full training forms part of the PhD
process. Applicants should have a keen interest in evolutionary biology, with a desire to
perform both lab and field-based experiments, and a willingness to learn genetic analyses.
An interest in plant biology is beneficial but not essential. Similarly, prior basic statistical or
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bioinformatic skills will be considered an advantage. There will be opportunities to develop

the project to reflect the applicants interests and experience, which could include field work
in North America, and a population genetics component.
Further reading or any references referred to in the proposal
Wu et al. (2008) Mimulus is an emerging model system for the integration of ecological and
genomic studies. Heredity 100, 220-230.
Vergeer & Kunin (2013) Adaptation at range margins: common garden trials and the
performance of Arabidopsis lyrata across its nothwestern European range. New Phytologist
197: 989-1001.
Friedman & Willis (2013) Major QTLs for critical photoperiod and vernalization underlie
extensive variation in flowering in the Mimulus guttatus species complex. New
Phytologist 199: 571-583.
Project summary - This project seeks to understand the genetic basis of local adaptation in
the exceptionally widespread monkeyflower Mimulus guttatus, with an emphasis on
divergence between populations at the range margin.
This project fits within the NERC aim of understanding processes underlying terrestrial
biodiversity. This research will integrate quantitative genetic, genomic (with associated
bioinformatic) and ecological data to better understand local adaptation. This information will
be essential for understanding the resilience and responses of populations at the margins of
their distribution range under a projected scenario of climate change.