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EASTBIO 400 Million Years of Sugar Transport in Plants: unearthing the origin, evolution and genetic toolkit of the phloem

School of Biological Sciences

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Dr S Hetherington , Prof Justin Goodrich No more applications being accepted Competition Funded PhD Project (Students Worldwide)

About the Project

Plants require an internal conducting network to transport food and water around their bodies. This conducting network is termed vasculature and consists of two tissues; water conducting xylem and food conducting phloem. The acquisition of these tissues during plant evolution was key for the origin of trees and crops from tiny moss-like ancestors. Despite the importance of the phloem for transporting sugars as well as a diversity of other compounds and signalling molecules throughout plants we know almost nothing about its evolution or how it responds to climate change. The aim for this project is to study the evolution of the phloem over its 400 million year history and to answer key evolution questions such as: (i) when did the phloem originate, (ii) how has its structure, function and genetic toolkit evolved over the past 400 million years, (iii) how has phloem evolution been driven by climate change?
To tackle these questions requires a truly interdisciplinary approach at the interface between the Life and Earth Sciences. The Molecular Palaeobotany and Evolution Group (, based in the Institute of Molecular Plants Sciences at the University of Edinburgh, provides the ideal place to investigate the origin and evolution of the phloem. Using the extensive living plant collections at the Royal Botanic Gardens Edinburgh you will make predictions about how the structure of the phloem has changed through evolutionary time. Next, you will use the fossil record to test these predictions [1]. Using new 3D imaging techniques [2] of fossils in the collections at the National Museums of Scotland, including fossils from the world famous Rhynie chert deposit [2], the key features that define phloem will be identified. Finally, using molecular biological approaches and comparative genomics and transcriptomics in the Institute of Molecular Plant Sciences, the evolution of the genetic toolkit of the phloem will be uncovered [3]. Taking these complementary interdisciplinary approaches you will shed new light on phloem evolution.
This project will provide an extensive training in evolutionary techniques spanning both living and fossil plants as well as molecular and omics approaches and is suitable for a candidate from either a Life or Earth Sciences background.

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How to Apply:
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Funding Notes

This 4 year PhD project is part of a competition funded by EASTBIO BBSRC Doctoral Training Partnership This opportunity is open to UK and International students and provides funding to cover stipend and UK level tuition fees. The fee difference will be covered by the University of Edinburgh for successful international applicants. Please refer to UKRI website ( and Annex B of the UKRI Training Grant Terms and Conditions ( for full eligibility criteria.


[1] Hetherington AJ, Berry CM, Dolan L. 2020. Multiple origins of dichotomous and lateral branching during root evolution. Nature Plants. 6: 454-459. DOI:
[2] Hetherington AJ, Dolan L. 2018. Stepwise and independent origins of roots among land plants. Nature. 561: 235–238. DOI:
[3] Hetherington AJ, Emms DM, Kelly S, Dolan L. 2019. Gene expression data support the hypothesis that Isoetes rootlets are true roots and not modified leaves. bioRxiv. DOI:
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