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A drought stress-induced MYB transcription factor regulates pavement cell shape in leaves of European aspen (Populus tremula)

Sijia Liu, Siamsa M. Doyle, Kathryn M. Robinson, Zahra Rahneshan, Nathaniel R. Street, Stéphanie Robert

Posted on: 20 February 2026 , updated on: 24 February 2026

Preprint posted on 16 January 2026

Leaf pavement cells look deceptively simple, yet their shapes encode developmental history and environmental response. A MYB transcription factor contributes to fine-tuning this process in aspen.

Selected by Jeny Jose

What are pavement cells?

Pavement cells are the most abundant cells in the leaf epidermis. Although they do not perform photosynthesis, they play a crucial role in protecting the leaf and maintaining its
structural integrity. Their characteristic wavy, interlocking “jigsaw puzzle” shape (Figure 1) has fascinated plant biologists for decades.

But, why do these cells adopt such complex geometries?

Figure 1 from Trozzi et al., 2023 (preprint; bioRxiv). Figure made available under a CC-BY NC-ND 4.0 International license.

 

One influential hypothesis, supported by computational modelling, proposes that lobed pavement cell shapes help to relieve mechanical stress generated during leaf expansion. As plant cell walls are rigid and constrain growth direction, changes in cell wall composition or mechanical properties directly influence where and how lobes form.
Understanding how pavement cell shape is regulated, therefore, provides broader insight into tissue morphogenesis.

A distinct developmental programme in European aspen

Most of what we know about pavement cell development comes from Arabidopsis thaliana. In Arabidopsis leaves, pavement cells on both the adaxial and abaxial surfaces largely originate from meristemoid cells within the stomatal lineage. These cells undergo asymmetric divisions before expanding and interlocking to form complex lobed shapes. In contrast, Liu and colleagues reveal that European aspen (Populus tremula) follows a distinct developmental program. Aspen stomata are restricted to the abaxial surface, and pavement cells on both sides of the leaf appear to arise mainly through symmetric divisions of pre-existing interdigitated cells. These initially straight-walled sister cells later acquire undulating borders as they expand. This divergence highlights that pavement cell morphogenesis is not universally conserved and may be regulated by species-specific mechanisms. Consistent with this idea, pavement cell morphology shows high variability and weak phylogenetic signal across eudicots.

Natural genetic variation in pavement cell shape in Populus tremula

To uncover the genetic basis of this variation, the authors quantified pavement cell shape and size across a natural population of P. tremula genotypes (the SwAsp collection).
Although overall shape complexity was relatively modest, significant natural variation was observed. A genome-wide association study identified a single SNP significantly associated with increased branch count, located in the first intron of a MYB family transcription factor, MYB305a.

MYB305a emerges as a regulator of pavement cell shape complexity

MYB305a is closely related to Arabidopsis MYB71 and MYB79, ABA-responsive transcription factors implicated in stress responses. To test its function, the authors generated MYB305a native promoter-GFP reporter lines, CRISPR/Cas9 deletion lines, and over-expression lines in hybrid aspen clone T89. Both loss- and gain-of-function lines showed reduced pavement cell shape complexity, with increased circularity and fewer lobes, without significant changes in cell area. These results indicate that precise, fine-tuned levels of MYB305a are required for proper pavement cell shape acquisition, independently of overall cell growth. GFP reporter analysis revealed that MYB305a expression is low and patchy in young leaves, with only subsets of pavement cells expressing the gene, but becomes uniformly high in older leaves, supporting a dosage-sensitive model linking developmental stage to shape acquisition (Figure 2).

Figure 2 MYB305a promoter expression increases in pavement cells as they grow and acquire their complex shape. The figure was reused with express permission from the authors

Drought stress modulates MYB305a activity and cell shape outcomes

Given that many MYB transcription factors are implicated in drought and ABA responses, Liu and team next asked whether MYB305a contributes to drought-induced changes in pavement cell shape. Drought treatment reduced shoot height in all genotypes, indicating a robust stress response irrespective of MYB305a genotype. In wild-type trees, drought also led to a simplification of pavement cell shape, with increased circularity and reduced branch count and lobe number. Strikingly, this drought-induced remodelling of pavement cell shape was partially uncoupled in MYB305a mutants and over-expressors. While the mutants showed resistance to the reduction in branch count and lobe number, the over-expressors showed enhanced sensitivity in terms of reduced lobe number and branching, suggesting that MYB305a modulates pavement cell responses to water deficit. Drought strongly induced MYB305a promoter activity in young leaves, while older leaves exhibited high baseline expression, reinforcing the idea that finely tuned MYB305a levels regulate pavement cell shape according to developmental stage and environmental conditions.

How the SwAsp population data strengthen the story

To place these findings in an ecological context, the authors returned to the natural SwAsp population. Clustering of genotypes based on pavement cell shape traits separated trees with more and less complex cell shapes, and these clusters differed in their climatic origins. Genotypes with more complex pavement cell shapes tended to originate from sites with lower annual precipitation, as well as more northern and eastern locations. Together, these results suggest that pavement cell shape complexity in European aspen reflects both short-term developmental plasticity in response to drought and longer-term evolutionary adaptation to water availability, with MYB305a acting as one component of the underlying regulatory network.

What I particularly liked about this study is that while, it provides a step towards unravelling the molecular mechanisms controlling leaf pavement cell shape in aspen, it also integrates natural population data with functional genetics. Rather than relying solely on artificial growth conditions, the authors connect a cellular phenotype to native habitats and environmental history, offering a compelling framework for understanding how plant cell shapes evolve and respond to stress in real-world settings.

Glossary:

Abaxial side: The lower surface of a leaf, oriented away from the apical meristem during its formation.
Adaxial side: The upper surface of a leaf, oriented towards the apical meristem during its formation.
Stomatal lineage: The series of cell divisions and cell fate transitions that lead from a protodermal cell to a mature stoma

Tags: aspen, cell shape, drought response, pavement cells

doi: https://doi.org/10.1242/prelights.42848

Read preprint (No Ratings Yet)

Response from the authors

Siamsa M. Doyle and Stéphanie Robert shared

Since the associated SNP lies in the first intron of MYB305a, do you suspect this region contains regulatory elements that affect spatial or temporal expression rather than overall transcript abundance?

Yes, this is certainly possible, but we don’t have any clear evidence on this yet.

Do you have any insight into the downstream targets of MYB305a in the leaf epidermis?

Unfortunately, we do not have any clear insights yet!

Do you envision pavement cell shape changes as an adaptive response to drought, or as a developmental side-effect of altered growth dynamics under stress?

We suspect that pavement cell shape is indeed an adaptive response to drought, since altered growth conditions have been shown to affect pavement cell shape acquisition in several plant species. At the present time, we do not understand enough about the functions of the interdigitated cell shape to fully comprehend the advantages that variation in this shape may give a plant experiencing drought stress. However, since the puzzle shapes of pavement cells may function in relieving mechanical stress in the leaf epidermis, alterations in these shapes may help the leaf to cope with changing mechanical stresses caused by altered turgor pressure in the tissues under drought conditions.

Was there a result in this study that surprised you most or changed how you think about pavement cell morphogenesis?

We were pleasantly surprised that, despite much higher pavement cell shape complexity in leaves of juvenile greenhouse-grown aspen trees compared to leaves of adult field-grown trees, the relative level of shape complexity in the genotypes was the same. This suggests that both short-term developmental plasticity of pavement cell shape in response to local growth conditions and long-term evolution of pavement shape complexity at a genotype-level are important for the final cell shape acquired in the leaf.

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List by Barbora Knotkova et al.

End-of-year preprints – the genetics & genomics edition

In this community-driven preList, a group of preLighters, with expertise in different areas of genetics and genomics have worked together to create this preprint reading list. Categories include: 1) genomics 2) bioinformatics 3) gene regulation 4) epigenetics

 



List by Chee Kiang Ewe et al.

BSDB/GenSoc Spring Meeting 2024

A list of preprints highlighted at the British Society for Developmental Biology and Genetics Society joint Spring meeting 2024 at Warwick, UK.

 



List by Joyce Yu, Katherine Brown

BSCB-Biochemical Society 2024 Cell Migration meeting

This preList features preprints that were discussed and presented during the BSCB-Biochemical Society 2024 Cell Migration meeting in Birmingham, UK in April 2024. Kindly put together by Sara Morais da Silva, Reviews Editor at Journal of Cell Science.

 



List by Reinier Prosee

9th International Symposium on the Biology of Vertebrate Sex Determination

This preList contains preprints discussed during the 9th International Symposium on the Biology of Vertebrate Sex Determination. This conference was held in Kona, Hawaii from April 17th to 21st 2023.

 



List by Martin Estermann

Alumni picks – preLights 5th Birthday

This preList contains preprints that were picked and highlighted by preLights Alumni - an initiative that was set up to mark preLights 5th birthday. More entries will follow throughout February and March 2023.

 



List by Sergio Menchero et al.

Semmelweis Symposium 2022: 40th anniversary of international medical education at Semmelweis University

This preList contains preprints discussed during the 'Semmelweis Symposium 2022' (7-9 November), organised around the 40th anniversary of international medical education at Semmelweis University covering a wide range of topics.

 



List by Nándor Lipták

20th “Genetics Workshops in Hungary”, Szeged (25th, September)

In this annual conference, Hungarian geneticists, biochemists and biotechnologists presented their works. Link: http://group.szbk.u-szeged.hu/minikonf/archive/prg2021.pdf

 



List by Nándor Lipták

2nd Conference of the Visegrád Group Society for Developmental Biology

Preprints from the 2nd Conference of the Visegrád Group Society for Developmental Biology (2-5 September, 2021, Szeged, Hungary)

 



List by Nándor Lipták

EMBL Conference: From functional genomics to systems biology

Preprints presented at the virtual EMBL conference "from functional genomics and systems biology", 16-19 November 2020

 



List by Jesus Victorino

TAGC 2020

Preprints recently presented at the virtual Allied Genetics Conference, April 22-26, 2020. #TAGC20

 



List by Maiko Kitaoka et al.

ECFG15 – Fungal biology

Preprints presented at 15th European Conference on Fungal Genetics 17-20 February 2020 Rome

 



List by Hiral Shah

Autophagy

Preprints on autophagy and lysosomal degradation and its role in neurodegeneration and disease. Includes molecular mechanisms, upstream signalling and regulation as well as studies on pharmaceutical interventions to upregulate the process.

 



List by Sandra Malmgren Hill

Zebrafish immunology

A compilation of cutting-edge research that uses the zebrafish as a model system to elucidate novel immunological mechanisms in health and disease.

 



List by Shikha Nayar

Also in the molecular biology category:

Keystone Symposium on Stem Cell Models in Embryology 2026

The Keystone Symposium on Stem Cell Models in Embryology, 2026, was organised by Jun Wu (UT Southwestern), Jianping Fu (University of Michigan) and Miki Ebisuya (TU Dresden) and held at Asilomar Conference Grounds in California (US). The meeting discussed recent advances made in establishing stem-cell-based embryo models, what fundamental insights into developmental processes have been gleaned from them, as well as how they are beginning to be applied more widely. This prelist contains preprints by presenters at the talk and poster sessions at the conference, which our Reviews Editor in attendance spotted. Please do reach out to preLights@biologists.com if you notice any that we’ve missed.

 



List by Ingrid Tsang

SciELO preprints – From 2025 onwards

SciELO has become a cornerstone of open, multilingual scholarly communication across Latin America. Its preprint server, SciELO preprints, is expanding the global reach of preprinted research from the region (for more information, see our interview with Carolina Tanigushi). This preList brings together biological, English language SciELO preprints to help readers discover emerging work from the Global South. By highlighting these preprints in one place, we aim to support visibility, encourage early feedback, and showcase the vibrant research communities contributing to SciELO’s open science ecosystem.

 



List by Carolina Tanigushi

October in preprints – DevBio & Stem cell biology

Each month, preLighters with expertise across developmental and stem cell biology nominate a few recent developmental and stem cell biology (and related) preprints they’re excited about and explain in a single paragraph why. Short, snappy picks from working scientists — a quick way to spot fresh ideas, bold methods and papers worth reading in full. These preprints can all be found in the October preprint list published on the Node.

 



List by Deevitha Balasubramanian et al.

October in preprints – Cell biology edition

Different preLighters, with expertise across cell biology, have worked together to create this preprint reading list for researchers with an interest in cell biology. This month, most picks fall under (1) Cell organelles and organisation, followed by (2) Mechanosignaling and mechanotransduction, (3) Cell cycle and division and (4) Cell migration

 



List by Matthew Davies et al.

September in preprints – Cell biology edition

A group of preLighters, with expertise in different areas of cell biology, have worked together to create this preprint reading list. This month, categories include: (1) Cell organelles and organisation, (2) Cell signalling and mechanosensing, (3) Cell metabolism, (4) Cell cycle and division, (5) Cell migration

 



List by Sristilekha Nath et al.

June in preprints – the CellBio edition

A group of preLighters, with expertise in different areas of cell biology, have worked together to create this preprint reading lists for researchers with an interest in cell biology. This month, categories include: (1) Cell organelles and organisation (2) Cell signaling and mechanosensation (3) Genetics/gene expression (4) Biochemistry (5) Cytoskeleton

 



List by Barbora Knotkova et al.

May in preprints – the CellBio edition

A group of preLighters, with expertise in different areas of cell biology, have worked together to create this preprint reading lists for researchers with an interest in cell biology. This month, categories include: 1) Biochemistry/metabolism 2) Cancer cell Biology 3) Cell adhesion, migration and cytoskeleton 4) Cell organelles and organisation 5) Cell signalling and 6) Genetics

 



List by Barbora Knotkova et al.

Keystone Symposium – Metabolic and Nutritional Control of Development and Cell Fate

This preList contains preprints discussed during the Metabolic and Nutritional Control of Development and Cell Fate Keystone Symposia. This conference was organized by Lydia Finley and Ralph J. DeBerardinis and held in the Wylie Center and Tupper Manor at Endicott College, Beverly, MA, United States from May 7th to 9th 2025. This meeting marked the first in-person gathering of leading researchers exploring how metabolism influences development, including processes like cell fate, tissue patterning, and organ function, through nutrient availability and metabolic regulation. By integrating modern metabolic tools with genetic and epidemiological insights across model organisms, this event highlighted key mechanisms and identified open questions to advance the emerging field of developmental metabolism.

 



List by Virginia Savy, Martin Estermann

April in preprints – the CellBio edition

A group of preLighters, with expertise in different areas of cell biology, have worked together to create this preprint reading lists for researchers with an interest in cell biology. This month, categories include: 1) biochemistry/metabolism 2) cell cycle and division 3) cell organelles and organisation 4) cell signalling and mechanosensing 5) (epi)genetics

 



List by Vibha SINGH et al.

Biologists @ 100 conference preList

This preList aims to capture all preprints being discussed at the Biologists @100 conference in Liverpool, UK, either as part of the poster sessions or the (flash/short/full-length) talks.

 



List by Reinier Prosee, Jonathan Townson

February in preprints – the CellBio edition

A group of preLighters, with expertise in different areas of cell biology, have worked together to create this preprint reading lists for researchers with an interest in cell biology. This month, categories include: 1) biochemistry and cell metabolism 2) cell organelles and organisation 3) cell signalling, migration and mechanosensing

 



List by Barbora Knotkova et al.

Community-driven preList – Immunology

In this community-driven preList, a group of preLighters, with expertise in different areas of immunology have worked together to create this preprint reading list.

 



List by Felipe Del Valle Batalla et al.

January in preprints – the CellBio edition

A group of preLighters, with expertise in different areas of cell biology, have worked together to create this preprint reading lists for researchers with an interest in cell biology. This month, categories include: 1) biochemistry/metabolism 2) cell migration 3) cell organelles and organisation 4) cell signalling and mechanosensing 5) genetics/gene expression

 



List by Barbora Knotkova et al.

2024 Hypothalamus GRC

This 2024 Hypothalamus GRC (Gordon Research Conference) preList offers an overview of cutting-edge research focused on the hypothalamus, a critical brain region involved in regulating homeostasis, behavior, and neuroendocrine functions. The studies included cover a range of topics, including neural circuits, molecular mechanisms, and the role of the hypothalamus in health and disease. This collection highlights some of the latest advances in understanding hypothalamic function, with potential implications for treating disorders such as obesity, stress, and metabolic diseases.

 



List by Nathalie Krauth

BSCB-Biochemical Society 2024 Cell Migration meeting

This preList features preprints that were discussed and presented during the BSCB-Biochemical Society 2024 Cell Migration meeting in Birmingham, UK in April 2024. Kindly put together by Sara Morais da Silva, Reviews Editor at Journal of Cell Science.

 



List by Reinier Prosee

‘In preprints’ from Development 2022-2023

A list of the preprints featured in Development's 'In preprints' articles between 2022-2023

 



List by Alex Eve, Katherine Brown

CSHL 87th Symposium: Stem Cells

Preprints mentioned by speakers at the #CSHLsymp23

 



List by Alex Eve

9th International Symposium on the Biology of Vertebrate Sex Determination

This preList contains preprints discussed during the 9th International Symposium on the Biology of Vertebrate Sex Determination. This conference was held in Kona, Hawaii from April 17th to 21st 2023.

 



List by Martin Estermann

Alumni picks – preLights 5th Birthday

This preList contains preprints that were picked and highlighted by preLights Alumni - an initiative that was set up to mark preLights 5th birthday. More entries will follow throughout February and March 2023.

 



List by Sergio Menchero et al.

CellBio 2022 – An ASCB/EMBO Meeting

This preLists features preprints that were discussed and presented during the CellBio 2022 meeting in Washington, DC in December 2022.

 



List by Nadja Hümpfer et al.

EMBL Synthetic Morphogenesis: From Gene Circuits to Tissue Architecture (2021)

A list of preprints mentioned at the #EESmorphoG virtual meeting in 2021.

 



List by Alex Eve

FENS 2020

A collection of preprints presented during the virtual meeting of the Federation of European Neuroscience Societies (FENS) in 2020

 



List by Ana Dorrego-Rivas

ECFG15 – Fungal biology

Preprints presented at 15th European Conference on Fungal Genetics 17-20 February 2020 Rome

 



List by Hiral Shah

ASCB EMBO Annual Meeting 2019

A collection of preprints presented at the 2019 ASCB EMBO Meeting in Washington, DC (December 7-11)

 



List by Madhuja Samaddar et al.

Lung Disease and Regeneration

This preprint list compiles highlights from the field of lung biology.

 



List by Rob Hynds

MitoList

This list of preprints is focused on work expanding our knowledge on mitochondria in any organism, tissue or cell type, from the normal biology to the pathology.

 



List by Sandra Franco Iborra