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Haseloff, J.

Publications and source records attributed to Haseloff, J..

At least 19 recordsLinked to original sources

Functional diversification of ERF transcription factors in land plants

The ERF family is one of the largest families of plant-specific transcription factors, controlling diverse developmental processes and stress responses. Extensive gene duplication and redundancy in flowering plants have generated large ERF repertoires, complicating comparisons of ERF function across land-plant lineages. How ERF transcription factor repertoires and their functions have evolved across land plants remains poorly understood. Here, we reconstruct the phylogenetic relationships and expansion of ERF transcription factors across land plants and exploit the compact ERF repertoire of the model liverwort Marchantia polymorpha to investigate their functional diversification. Systematic gain-of-function analysis reveals that different clades of ERFs can drive distinct cell fate and reprogramming outcomes in Marchantia, in addition to stress-related functions. Interestingly, we found that MpERF2, the sole Group I ERF in Marchantia, phylogenetically related to the WIND clade, promotes rhizoid cell differentiation. This contrast with the role of the WIND clade in wound-induced regeneration in Arabidopsis. Heterologous expression of AtWIND1 in Marchantia was also capable of inducing rhizoid formation, supporting the hypothesis that this reprogramming capacity is conserved at the protein level. On the other hand, loss of MpERF2 reduced rhizoid coverage but did not significantly impair regeneration. These findings provide an evolutionary framework for understanding the functional evolution of ERF transcription factors and how conserved transcriptional regulators are co-opted into novel developmental programs during the radiation of land plants.

plant biology↗

Neryl diphosphate-derived monoterpene biosynthesis via a biosynthetic gene cluster in the liverwort Marchantia polymorpha

Monoterpenes (C10) are a large group of specialized metabolites important for plant interactions with the environment. Their biosynthesis is well understood in seed plants, where geranyl diphosphate serves as the canonical substrate, but knowledge of monoterpene biosynthesis outside seed plants remains very limited. Here, we report neryl diphosphate (NPP)-derived monoterpene biosynthesis via a biosynthetic gene cluster in the liverwort Marchantia polymorpha. MpMTPSL2, a microbial-type terpene synthase, converts NPP into -phellandrene and D-limonene in vitro. CRISPR knockout lines showed reduced production of both monoterpenes, providing direct genetic evidence for its in planta function. MpCPT5, a cis-prenyltransferase (CPT) family member identified through co-expression with MpMTPSL2, was confirmed to encode NPP synthase, as its knockout plants abolished -phellandrene and D-limonene production. Subcellular localization analyses in protoplasts and stable transgenic plants demonstrated that both MpCPT5 and MpMTPSL2 localize to plastids, co-localizing across all cell types with markedly stronger signals in non-green plastids of oil-body cells. Consistent with this, expression of both genes under their respective promoters was nearly abolished in oil-body-deficient mutants and strongly upregulated in a gain-of-function line for oil-body formation. MpMTPSL2 and MpCPT5 are physically linked through a shared bidirectional promoter that drives their coexpression specific to oil body cells, forming a unique biosynthetic gene cluster whose coordinated expression is maintained by PRC2-mediated H3K27me3 repression. Phylogenetic analysis implies that NPP synthases in M. polymorpha and in flowering plants evolved independently from their respective long-chain CPT ancestors. These findings provide new insights into the mechanisms and evolution of monoterpene biosynthesis in non-seed plants. Significance statementMonoterpenes are a diverse group of specialized metabolites produced widely among land plants, yet our understanding of their biosynthesis outside seed plants remains limited. Here we report that in the liverwort Marchantia polymorpha, the non-canonical substrate neryl diphosphate is used for monoterpene biosynthesis. The functions of the monoterpene synthase gene MpMTPSL2 and the neryl diphosphate synthase gene MpCPT5 were demonstrated through CRISPR knockouts. These two genes are physically linked and share a bidirectional promoter. Promoter assays show both genes function in plastids within oil-body cells, revealing cell-type specificity. These findings shew new light on the mechanisms and evolution of monoterpene biosynthesis in non-seed plants.

plant biology↗

Marchantia stem cell maintenance and re-establishment are controlled by MpPIN1-mediated auxin transport and ARF signalling

Meristem organization in land plants depends on coordinated auxin distribution, transport and response, yet how these processes are integrated during vegetative growth of the gametophyte of non-seed plants remains poorly understood. Here, we describe dissection of the auxin network in the liverwort Marchantia polymorpha using a suite of endogenous and synthetic reporters. Quantitative imaging with the R2DII auxin response sensor revealed that the apical meristem is positioned at a stable auxin minimum, with auxin levels increasing toward differentiated tissues. This spatial configuration correlates with polar localization of the auxin efflux carrier MpPIN1 along the direction of auxin flux. Disruption of MpPIN1 alters auxin distribution and compromises tissue regeneration, but does not prevent initial meristem formation, indicating that PIN-mediated transport reinforces rather than defines stem-cell niche identity. Imaging of auxin response reporters and MpARF1 and MpARF2 knock-in reporters further revealed a constrained auxin response within the meristem, characterized by low auxin signalling, dependent on the inhibitory activity of MpARF2 in the stem cell zone. During regeneration, transient, oscillatory ARF dynamics were observed before stabilization of a new auxin minimum and re-establishment of normal meristem architecture. Together, our results are consistent with auxin responses in the Marchantia meristem being shaped by an incoherent feed-forward network topology that couples auxin flux, transcriptional constraint and niche permissiveness. This architecture provides a robust framework for stem-cell maintenance and re-establishment, revealing ancestral principles of meristem regulation in land plants. SUMMARY STATEMENTBy mapping hormone distribution at cellular resolution in the liverwort Marchantia polymorpha, we reveal fundamental principles governing how plants maintain and rebuild their stem cell niches.

Plant Biology↗

Marchantia polymorpha as a simple platform for plant-based production of functional nanobodies

Recombinant proteins have transformative potential in biomedicine, but their production is often costly and carries contamination risks. Plants offer an attractive alternative, with low growth costs and reduced pathogen risk, yet their slow growth cycles limit their use for rapid protein engineering. Here, we establish Marchantia polymorpha, a genetically tractable liverwort with a short life cycle, as a new platform for recombinant protein production. Using stable Agrobacterium-mediated transformation, we expressed an anti-mCherry nanobody fused to the fluorescent protein mTurquoise2 with different purification tags. Expression levels reached up to [~]120 {micro}g/g fresh weight, and nanobody functionality was validated through a microscopy-based bead-binding assay. This yield rivals that of established systems such as Nicotiana benthamiana. Our results position M. polymorpha as a scalable, safe, and efficient chassis for protein engineering, with broad potential for applications in synthetic biology. This work opens the door to exploiting liverwort biotechnology for fast, cost-effective, and biosafe production of valuable recombinant proteins. Graphical Table of Contents O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=133 SRC="FIGDIR/small/733889v1_ufig1.gif" ALT="Figure 1"> View larger version (21K): org.highwire.dtl.DTLVardef@aabc4aorg.highwire.dtl.DTLVardef@12f9fa4org.highwire.dtl.DTLVardef@199d3e0org.highwire.dtl.DTLVardef@2533a4_HPS_FORMAT_FIGEXP M_FIG C_FIG

Plant Biology↗

Engineering auronidin production in Marchantia polymorpha enables a new class of plant-derived textile pigments

Plant-derived pigments offer sustainable alternatives to synthetic colorants, yet their practical deployment in textiles is limited by restricted chemical diversity and low abundance. Liverworts represent a source of diverse chemical compounds, and the model liverworts Marchantia polymorpha is an emerging as chassis for bioengineering and synthetic biology. Here, we report the biotechnological application of auronidins, a rare class of flavonoid pigments, as textile dyes. Using the Marchantia, we engineered enhanced auronidin production through controlled expression of the R2R3-MYB transcription factor MpMYB14. We systematically benchmarked constitutive and inducible gene expression systems, including heat-shock, glucocorticoid receptor, and {beta}-estradiol (XVE) circuits, identifying inducible strategies that decouple biomass accumulation from secondary metabolite production while achieving high pigment yields. Extracted auronidins were used to dye cotton yarn directly, demonstrating the feasibility of auronidins for textile dyeing. Our results establish Marchantia as a versatile plant chassis for programmable secondary metabolite production and introduce auronidins as a promising natural pigment platform for sustainable textile biotechnology.

synthetic biology↗

Initiation of asexual reproduction by the AP2/ERF gene GEMMIFER in Marchantia polymorpha

Plants can propagate their own clones through asexual reproduction. Genetic and hormonal factors regulating asexual reproduction have begun to be elucidated in the liverwort Marchantia polymorpha, which produce asexual propagules called gemmae within the gemma cups. Here, we report an AP2/ERF family gene, GEMMIFER (MpGMFR), as a key regulator of asexual reproduction in M. polymorpha. Suppression of MpGMFR function using genome editing and amiRNA results in the loss of gemma and gemma cup formation. In contrast, activation of MpGMFR function using a dexamethasone inducible system promotes gemma and/or gemma cup formation depending on the induction conditions. Notably, transient strong activation of MpGMFR induced gemma initial cells at the meristem, which develop into mature gemmae capable of reproducing as new individuals after detachment. These results show that MpGMFR is necessary and sufficient for the initiation of asexual reproduction at the meristem. MpGMFR expression was detected from the meristem through to the early stages of gemma development including the gemma cup floor cells. MpGMFR expression precedes the expression of GEMMA CUP-ASSOCIATED MYB1 (MpGCAM1) in early gemma development, supporting the notion that MpGMFR initiates asexual reproduction. We anticipate this finding to be a foundation to study the evolution of extra meristem formation in plant bodies that may have made a significant contribution to the prosperity of plants on land.

plant biology↗

Conservation of chromatin states and their association with transcription factors in land plants

The complexity of varied modifications of chromatin composition is integrated in archetypal combinations called chromatin states that predict the local potential for transcription. The degree of conservation of chromatin states has not been established amongst plants, and how they interact with transcription factors is unknown. Here we identify and characterize chromatin states in the flowering plant Arabidopsis thaliana and the bryophyte Marchantia polymorpha, showing a large degree of functional conservation over more than 450 million years of land plant evolution. We used this new resource of conserved plant chromatin states to understand the influence of chromatin states on gene regulation. We established the preferential association of chromatin states with binding sites and activity of transcription factors. These associations define three main groups of transcription factors that bind upstream of the transcription start site, at the +1 nucleosome or further downstream of the transcription start site and broadly associate with distinct biological functions. The association with the +1 nucleosome defines a list of candidate pioneer factors we know little about in plants, compared to their important roles in animal stem cells and early development.

genomics↗

Replicated repurposing of an ancestral transcriptional complex in land plants

Transcriptional complexes with a common composition regulate the production of flavonoid pigments, trichomes, root hairs and other epidermal traits in seed plants. These complexes are composed of transcription factors from the MYB and basic helix-loop-helix (bHLH) families along with a tryptophan-aspartate repeat (WDR) scaffold protein (MBW complexes). The MYB member has been found to be the most pathway-specific component of the complex and modifications to these MYB genes are overrepresented in studies investigating the genetic basis of changes in pigmentation phenotypes across flowering plants. Here we investigated the orthologues of the MBW complex in a divergent lineage to understand its origin and evolution. We found evidence that these transcriptional complexes also form in the liverwort Marchantia polymorpha, indicating, together with an analysis of published gene family phylogenies, that they are ancestral to land plants. The functions of each of the two orthologous MYB genes, MpMYB14 and MpMYB02, both depend on the single orthologous bHLH gene, MpbHLH12. We could not assess the functional role of the WDR genes in M. polymorpha, due to low mutant recovery suspected to be caused by pleiotropic effects on viability. We propose that the two transcriptional complexes with alternative MYB paralogues in M. polymorpha represent an ancestral function, regulation of the flavonoid pathway, and a derived function, maturation of liverwort-specific oil bodies. Our findings imply a replicated pattern by which new complexes have evolved in independent land plant lineages, through duplication of the evolutionarily labile MYB member and co-option of its interaction partners.

plant biology↗

A tRNA-gRNA multiplexing system for CRISPR genome editing in Marchantia polymorpha.

The liverwort Marchantia polymorpha is a widely used model organism for studying land plant biology, which has also proven to be a promising testbed for bioengineering. CRISPR/Cas9 technology has emerged as a transformative tool for precise genome modifications in M. polymorpha. However, a robust method for the simultaneous expression of multiple gRNAs, which is crucial for enhancing the efficiency and versatility of CRISPR/Cas9-based genome editing, has yet to be fully developed. In this study, we introduce an adaptation from the OpenPlant kit CRISPR/Cas9 tools, that facilitates expression of multiple gRNAs from a single transcript through incorporation of tRNA sequences. This approach significantly improves the efficiency and scalability of genome editing in M. polymorpha. Additionally, by combining this vector system with a simplified and optimized protocol for thallus transformation, we further streamline the generation of CRISPR/Cas9 mutants in M. polymorpha. The resulting gene- editing system offers a versatile, time-saving and straightforward tool for advancing functional genomics in M. polymorpha, enabling more comprehensive genetic modifications and genome engineering.

plant biology↗

Arsenic-sensing domain controls ACR3 transporter trafficking and function in Marchantia polymorpha

Arsenic, a toxic and carcinogenic metalloid, is a pervasive environmental contaminant that threatens human health through contaminated water and food. The efflux of As(III) via ACR3 transporters is an ancient detoxification mechanism conserved across prokaryotes, fungi, and plants, with the notable exception of angiosperms. Despite their evolutionary significance, plant ACR3s remain largely uncharacterized. Here, we demonstrate that MpACR3, the ACR3 orthologue from the liverwort Marchantia polymorpha, functions as a metalloid/proton antiporter, conferring resistance to arsenicals and moderate tolerance to antimony. Additionally, we uncover an arsenic-sensing domain within MpACR3 that regulates its intracellular trafficking. Under normal conditions, MpACR3 sorting to the plasma membrane is delayed, resulting in its retention within Golgi bodies. However, As(III) binding to three cysteine residues in the N-terminal cytosolic domain induces a conformational change that facilitates MpACR3 trafficking to the plasma membrane. Furthermore, mutational analysis of a conserved arginine-based motif reveals that the N-terminal domain not only controls MpACR3 accumulation at the plasma membrane but also modulates its transport activity. Importantly, this arsenic-sensing domain is conserved among plant ACR3 transporters, suggesting a plant-specific adaptation to arsenic toxicity.

plant biology↗

The minimal cell-cycle control system in Marchantia as a framework for understanding plant cell proliferation

The regulation of cell division is broadly conserved across eukaryotes, governed by cyclins and cyclin-dependent kinases (CDKs) to coordinate progression through the cell cycle. Plants have evolved a complex set of cell-cycle genes with unique features. The high number of cyclin-CDK pairs in flowering plants complicates functional studies due to redundancy and diversification. It is critical to study simple systems in other plant lineages to better understand the functional integration of the cell-cycle control machinery and its evolution across land plants. Through a comprehensive phylogenetic analysis, we show that non-seed plants possess a simple repertoire of cyclin and CDK proteins, suggesting that the observed complexity in seed plants is a derived trait. The liverwort Marchantia polymorpha possesses a streamlined set of core cell cycle genes with minimal redundancy during vegetative development. Using single-cell RNA-seq and fluorescent reporters, we found a precise, phase-specific expression pattern for cell cycle genes. We demonstrated in vivo that only three cyclins are active, one at a given phase, without redundancy. Functional analyses revealed that MpCYCD;1 promotes cell cycle re-entry and disrupts differentiation, while overexpression of MpCYCA or MpCYCB;1 arrests the cell cycle, consistent with their respective roles at G1, S, and G2/M progression. Our findings highlight the functional conservation of mechanisms for cell-cycle control across eukaryotes and provide insights into its ancestral state, revealing a minimal set of functional components required for multicellular development. This study advances our understanding of fundamental aspects of cell-cycle regulation and opens new possibilities for engineering plant growth.

plant biology↗

The ABCG1 transporter is required for terpene biosynthesis in the lumen of Marchantia polymorpha oil bodies.

Marchantia polymorpha oil bodies (OBs) are specialised cell structures housing a diverse array of C15-terpenes, called sesquiterpenes. These compounds are known for their roles as herbivore repellents, yet the enzymes responsible for the biosynthesis of their precursors (C5 isoprenoid units) remain poorly characterized. Discrepancies remain between enzyme localizations suggested by computational predictions and those observed in earlier experimental studies, complicating our understanding of terpene biosynthesis. We investigated the localization of isoprenoid biosynthetic enzymes using translational and transcriptional reporters, coupled with confocal microscopy. Most enzymes localized as predicted (e.g., cytosol, chloroplast and the endoplasmic reticulum), and OB cells were identified as the primary sites of terpene biosynthesis. To explore OBs as potential storage sites for terpenes, we attempted to produce exogenous but easily identifiable compounds in Marchantia, such as the diterpene taxadiene and the triterpene {beta}-amyrin. Targeting to OB cells resulted in measurable amounts of these compounds, but their yields remained unaffected by the over-expression of key precursor genes, underscoring challenges in redirecting metabolic flux. To further investigate terpene accumulation in OBs, we focused on ABCG1, an ABC transporter previously reported to localize at the OB membrane. Overexpression of ABCG1 in OB cells, alongside an exogenous sesquiterpene synthase, only increased the levels of endogenous sesquiterpenes, while CRISPR-mediated disruption of ABCG1 resulted in a dramatic reduction in sesquiterpene accumulation. These findings establish ABCG1 as a critical factor for sesquiterpene retention within OBs and provide new insights into the mechanisms governing terpene metabolism and storage in Marchantia polymorpha.

plant biology↗

Semi-automated workflow for high-throughput Agrobacterium-mediated plant transformation

High-throughput experiments in plants are hindered by long generation times and high costs. To address these challenges, we present an optimized pipeline for Agrobacterium tumefaciens transformation and simplified a protocol to obtain stable transgenic lines of the model liverwort Marchantia polymorpha, paving the way for efficient high-throughput experiments for plant synthetic biology and other applications. Our protocol involves freeze-thaw Agrobacterium transformation method in 6-well plates that can be adapted to robotic automation. Using the Opentrons open-source platform, we implemented a semi-automated protocol showing similar efficiency compared to manual manipulation. Additionally, we have streamlined and simplified the process of stable transformation and selection of M. polymorpha, reducing cost, time, and manual labour without compromising transformation efficiency. The addition of sucrose in the selection media significantly enhances the production of gemmae, accelerating the generation of isogenic plants. We believe these protocols have the potential to facilitate high-throughput screenings in diverse plant species and represent a significant step towards the full automation of plant transformation pipelines. This approach allows testing [~]100 constructs per month, using conventional plant tissue culture facilities. We recently demonstrated the successful implementation of this protocol for screening hundreds of fluorescent reporters in Marchantia gemmae.

synthetic biology↗

Control of cell fate specification and patterning by an ancestral microRNA

The formation of an organized body requires the establishment and maintenance of cells with structural and functional distinctive characteristics. A central question in developmental biology is how changes in the regulation of genes drive cell specification and patterning1. microRNAs (miRNAs) are small non-coding RNAs that regulate development through mRNA cleavage and/or translational repression2. In plants, miRNAs regulate key aspects including growth, development, stem cell maintenance, vegetative phase change, leaf morphogenesis, floral organ formation and flowering time3. Biogenesis of plant miRNAs depends on the activity of DICER-LIKE 1 (DCL1), an RNase type III endonuclease that processes double stranded RNA to give rise to mature miRNAs 4. The genomes of todays flora contain at least one bona fide copy of DCL1 5,6. Using Marchantia polymorpha -a model bryophyte that allows comparative approaches to infer characteristics of the ancestral land plant-, we demonstrate that MpDCL1a is required for the biogenesis of miRNAs and uncovered a central role for miR166/Homeodomain Zipper Class III-regulated auxin synthesis in the specification of cell identity, patterning, meristem function, laminar expansion and the development of the body in the last common ancestor of extant land plants.

plant biology↗

Synergistic control of chloroplast biogenesis by MYB-related and Golden2-like transcription factors

Chloroplast biogenesis is dependent on master regulators from the GOLDEN2-LIKE (GLK) family of transcription factors, but glk mutants contain residual chlorophyll and therefore other proteins must also be involved. Here we identify MYB-related transcription factors as regulators of chloroplast biogenesis in the liverwort Marchantia polymorpha and angiosperm Arabidopsis thaliana. In both species, double mutant alleles in MYB-Related genes show very limited chloroplast development, and photosynthesis gene expression is perturbed to a greater extent than in mutants of GLK. In M. polymorpha MYB-related genes act upstream of GLK, while in A. thaliana this relationship has been rewired. In both species, genes encoding enzymes of chlorophyll biosynthesis are controlled by MYB-related and GLK proteins whilst those allowing CO2 fixation, photorespiration and photosystem assembly and repair require the MYB-related proteins. Thus, MYB-related and GLK genes have overlapping as well as distinct targets. We conclude that together MYB-related and GLK transcription factors orchestrate chloroplast development in land plants.

plant biology↗

An Enhancer Trap system to track developmental dynamics in Marchantia polymorpha

A combination of streamlined genetics, experimental tractability and relative morphological simplicity compared to vascular plants makes the liverwort Marchantia polymorpha an ideal model system for studying many aspects of plant biology. Here we describe a transformation vector combining a constitutive fluorescent membrane marker with a nuclear marker that is regulated by nearby enhancer elements, and use this to produce a library of enhancer trap lines for Marchantia. Screening gemmae from these lines allowed for identification and characterization of novel marker lines, including markers for rhizoids and oil cells. The library allowed the identification of a margin tissue running around the thallus edge, highlighted during thallus development. Expression of this marker is correlated with auxin levels. We generated multiple markers for the meristematic apical notch region, which have different spatial expression patterns, reappear at different times during meristem regeneration following apical notch excision and have varying responses to auxin supplementation or inhibition. This reveals that there are proximodistal substructures within the apical notch that could not be observed otherwise. We employed our markers to study Marchantia sporeling development, observing meristem emergence as defining the protonema-to-prothallus stage transition, and subsequent production of margin tissue during the prothallus stage. Exogenous auxin treatment stalls meristem emergence at the protonema stage but does not inhibit cell division, resulting in callus-like sporelings with many rhizoids, whereas pharmacologically inhibiting auxin synthesis and transport does not prevent meristem emergence. This enhancer trap system presents a useful resource for the community and will contribute to future Marchantia research. Significance StatementThe liverwort Marchantia polymorpha is an emerging model species with huge potential for synthetic biology. We generated an enhancer trap library as a resource for Marchantia research and characterized marker lines for several important cell types (rhizoids, thallus margin, apical notch and oil cells). These marker lines can be used to track cellular dynamics in growing plants, exemplified by studying auxin responses and sporeling growth, revealing new insights into thallus development and cell fate specification.

plant biology↗

Mapping the landscape of transcription factor promoter activity during vegetative development in Marchantia

Transcription factors (TFs) are essential for the regulation of gene expression and cell fate determination. Characterising the transcriptional activity of TF genes in space and time is a critical step towards understanding complex biological systems. The vegetative gametophyte meristems of bryophytes share some characteristics with the shoot-apical meristems of flowering plants. However, the identity and expression profiles of TFs associated with gametophyte organization are largely unknown. With only [~]450 TF genes, Marchantia polymorpha is an outstanding model system for plant systems biology. We have generated a near-complete collection of promoter elements derived from Marchantia TF genes. We experimentally tested in planta reporter fusions for all the TF promoters in the collection and systematically analysed expression patterns in Marchantia gemmae. This allowed us to build a map of precise expression domains and identify a unique set of TFs expressed in the stem-cell zone, providing new insight into the dynamic regulation of the gametophytic meristem and its evolution. In addition, we provide an online database of expression patterns for all promoters in the collection. We expect that the promoter elements characterised here will be useful for cell-type specific expression, synthetic biology applications, and functional genomics.

plant biology↗

Streamlined regulation of chloroplast development in the liverwort Marchantia polymorpha

Photosynthesis in eukaryotic cells takes place in chloroplasts that develop from undifferentiated plastids in response to light. In angiosperms, after perception of light de-repression allows Elongated Hypocotyl 5 (HY5) transcription factor to initiate photomorphogenesis, and two families of transcription factors known as Golden2-like (GLK) and GATA are considered master regulators of chloroplast development. The MIR171-targeted SCARECROW-LIKE (SCL) GRAS transcription factors also impact on chlorophyll biosynthesis. The extent to which these proteins carry out conserved roles in non-seed plants is not known. Here we report in the model liverwort Marchantia polymorpha that GLK controls chloroplast biogenesis and HY5 shows a small conditional effect on chlorophyll content. In contrast, GATA and SCL have no detectable roles in this fundamental process. ChIP-SEQ and RNA-SEQ revealed that MpGLK regulates many photosynthetic and chloroplast development-related genes, but also has a broader set of targets than previously reported in angiosperms. This implies GLK carries out a conserved role relating to chloroplast biogenesis in land plants but also supports extensive divergence between its targets in M. polymorpha and flowering plants. The data support the hypothesis that regulation of chloroplast biogenesis in M. polymorpha is streamlined compared with angiosperms and allows us to present a core regulatory network for chloroplast biogenesis in land plants.

plant biology↗