Universiteit Leiden

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Research project

Environmental and genetic drivers of wood and lignin formation in flowering plants

In this project, we will study the genetic and environmental drivers of woodiness and stem lignification at the level of plant‐to‐gene‐to‐molecule.

2023 - 2028
Remko Offringa

Prof Dr. Katalin Barta (University of Graz)

Dr. Richard Gosselink (Wageningen Research)

Three fundamental questions are central in our project

1) What are the genes and their interactions that control lignification and/or wood formation in stems of flowering plants and can we disentangle the woodiness phenotype from coupled longevity phenotypes?

2) Does increased woodiness or lignification lead to enhanced drought resilience both in the short term (adaptation) and the long term (evolution)?

3) What is the relationship among the lignin chemical structure, the cell wall structure of lignified tissues and drought resilience, and what is the impact on the use of lignin as valued biobased material from left‐over plant parts?


Lignin is a plant biopolymer that is highly concentrated in so-called lignified cell walls and represents a major component in lignocellulosic plant biomass. Although lignified cell walls can be found in all kinds of vegetative and reproductive plant organs, cell wall lignification is pronounced within the wood tissue, which is developed by a lateral meristem (vascular cambium) and is responsible for most of the lateral stem and root growth in woody shrubs and trees. In flowering plants, hundreds of non-woody (herbaceous) lineages independently gave rise to new woody species, suggesting that: i) woodiness provides a fitness advantage in (seasonally) dry environments, and ii) only few genetic key regulatory changes are required for the transition from herb to shrub.

Interestingly, lignin production in herbaceous stems can also be enhanced in response to drought stress, where it is assumed to strengthen cell walls, facilitate root-to-shoot water transport, and impede water loss. As such, increased lignin deposition in plant cell walls as well as increased wood formation in stems and roots have been suggested by a range of pilot studies as a key strategy for drought tolerance during plant evolution. Drought tolerance is currently a key trait in crop breeding, as traditionally productive agricultural areas suffer heavily from recurring and more intensive drought cycles across the globe. At the same time, as lignified wood tissue is a major constituent of agricultural residues, scientists should take advantage of its value in the biobased economy in terms of residue valorisation. Indeed, lignin has high potential in applications such as glues and asphalt, meaning that increased lignin content provides opportunities for replacement of fossil derived products by left-over plant parts of more drought resilient crops.

Two Arabidopsis thaliana accessions after 17days of withholding water show a contrasting phenotype: the more drought tolerant one (right) has a more lignified inflorescence stem (arrow points to lignified stem tissue) compared to the most sensitive accession (left) with a less lignified stem (arrow). Owner: Ajaree Thonglim, Frederic Lens

In IBL, three PIs (Remko Offringa, Salma Balazadeh and Frederic Lens) will investigate how plants regulate wood development or cell lignification in stems under long evolutionary time scales as well as in response to short-term drought. PIs outside IBL will focus on disentangling the woodiness phenotype from the late flowering phenotype, analysing lignin chemical structure and the cell wall structure of lignified tissue in relation to drought tolerance, and assessing utilisation of stems as biobased material in left‐over plant parts. In order to do this, we have established a novel multidisciplinary consortium uniquely bridging expertise in plant anatomy, evolution, development, physiology, genetics, molecular biology and lignin chemistry. With promising genetic key regulators for woodiness and lignification at hand and a comprehensive understanding of factors determining lignin quality, we anticipate fundamental scientific breakthroughs and impactful future applications for a more sustainable agriculture.

Two tomato genotypes showing a similar phenotype after a seven day drought treatment compared to Arabidopsis: the more drought tolerant one (right) has more wood formation in the stem (arrow points to wood tissue) compared to the most sensitive accession (left) with a less woody stem (arrow). Owner: Salma Balazadeh, Frederic Lens


Rahimi A, Karami O, Lestari AD, de Werk T, Amakorova P, Shi D, Novak O, Greb T, Offringa R. 2022. Control of cambium initiation and activity in Arabidopsis by the transcriptional regulator AHL15. Current Biology 32: 1-12.

Thonglim, A, Delzon, S, Larter, M, Karami, O, Rahimi, A, Offringa, R, Keurentjes, JJB, Balazadeh S, Smets E, Lens F. 2021. Intervessel pit membrane thickness best explains variation in embolism resistance amongst stems of Arabidopsis thaliana accessions. Annals of Botany 128: 171–182.

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