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In plants, DNA methylation occurs in distinct sequence contexts, including CG, CHG, and CHH. Thus, plants have developed a surprisingly diverse set of DNA methylation readers to cope with an extended repertoire of methylated sites. The Arabidopsis genome contains twelve Methyl-Binding Domain proteins (MBD), and nine SET and RING finger-associated (SRA) domain containing proteins belonging to the SUVH clade, in addition to three homologs of UHRF1, namely VIM1-3, all containing SRA domains. In this review, we will highlight several research questions that remain unresolved with respect to the function of plant DNA methylation readers, which can have both de novo demethylase and maintenance activity. We argue that maintenance of CG methylation in plants likely involved actors not found in their mammalian counterparts, and that new evidence suggests significant reprogramming of DNA methylation during plant reproduction as an important new development in the field. Image 1 • Plants acquired DNA methylation readers to cope with a large repertoire of methylated sites. • This diversity is reflected throughout land plant evolution. • There are evidences for some level of conservation between animals and plants. • Plants have evolved unique features, including dedicated polymerases and plant-specific protein complexes. • Plant SUVH proteins can have both demethylase and methyltransferase activity. • CG maintenance in plants differs from the mammalian model, with VIM proteins bearing atypical motifs. [ABSTRACT FROM AUTHOR]

Summary: Methylation of DNA is an epigenetic mechanism for the control of gene expression. Alterations in the regulatory pathways involved in the establishment, perpetuation and removal of DNA methylation can lead to severe developmental alterations. Our understanding of the mechanistic aspects and relevance of DNA methylation comes from remarkable studies in well‐established angiosperm plant models including maize and Arabidopsis. The study of plant models positioned at basal lineages opens exciting opportunities to expand our knowledge on the function and evolution of the components of DNA methylation. In this Tansley Insight, we summarize current progress in our understanding of the molecular basis and relevance of DNA methylation in the liverwort Marchantia polymorpha. [ABSTRACT FROM AUTHOR]

Apomixis in plants covers a variety of cloning systems through seeds of great potential for plant breeding. Among long-standing approaches for crop improvement is the attempt to exploit wild relatives as natural, vast reservoirs for novel genetic variation. With regard to apomixis, maize possesses an apomictic wild relative, Tripsacum, which we used to produce advanced maize- Tripsacum hybrid generations. However, introgression of apomixis in maize has failed so far. In order to understand the how's and why's, we undertook characterization of seed development and inheritance studies in these materials. We show that apomictic seeds suffer from epigenetic loads. Both seed tissues, the endosperm and the embryo, displayed developmental defects resulting from imbalanced parental genomic contributions and aberrant methylation patterns, respectively. Progeny characterization of several maize- Tripsacum hybrid generations allowed significant progress toward the unraveling of the genetics of apomixis. First, chromosome deletion mapping showed that expression of apomixis requires one single Tripsacum chromosome. However, inheritance studies revealed that female gametes inheriting this segment were unequivalent carriers depending on their origin: unreduced gametes transmit a functional segment, whereas progeny derived from reduced ones reproduced sexually. Finally, chromosomal or genomic dosage variation barely affected the apomictic phenotype suggesting no dependency for ploidy in these materials. We conclude that epigenetic information imposes constraints for apomictic seed development and seems pivotal for transgenerational propagation of apomixis. The nature of the triggering mechanisms remains unknown as-yet, but it certainly explains the modest success relative to the development of apomictic maize thus far. [ABSTRACT FROM AUTHOR]

Plants' ability to respond to environmental stimuli and developmental cues depends upon changes in gene expression. In eukaryotes, genetic information encoded by DNA is packed in a highly regulated and dynamic nucleoprotein complex known as chromatin that is subject to epigenetic modifications. Historically, several biological phenomena relying on epigenetic mechanisms were first characterized in plants. Seminal discoveries such as paramutation and silencing of transposable elements were made in maize (Zea mays). Later rice (Oryza sativa) was selected as a good model for monocotyledons owing to its relatively small genome size and well-annotated sequenced genome. In the past few years an increasing number of epigenetic and epigenomic studies were performed in both maize and rice. In this chapter we will first compare the basic knowledge acquired on epigenetic regulation in rice and maize versus Arabidopsis, then we will describe cereals-specific aspects of epigenetic regulations. [ABSTRACT FROM AUTHOR]

Cytosine methylation is a key epigenetic mark in many organisms, important for both transcriptional control and genome integrity. While relatively stable during somatic growth, DNA methylation is reprogrammed genome-wide during mammalian reproduction. Reprogramming is essential for zygotic totipotency and to prevent transgenerational inheritance of epimutations. However, the extent of DNA methylation reprogramming in plants remains unclear. Here, we developed sensors reporting with single-cell resolution CG and non-CG methylation in Arabidopsis. Live imaging during reproduction revealed distinct and sex-specific dynamics for both contexts. We found that CHH methylation in the egg cell depends on DOMAINS REARRANGED METHYLASE 2 (DRM2) and RNA polymerase V (Pol V), two main actors of RNA-directed DNA methylation, but does not depend on Pol IV. Our sensors provide insight into globalDNAmethylation dynamics at the single-cell level with high temporal resolution and offer a powerful tool to track CG and non-CG methylation both during development and in response to environmental cues in all organisms with methylated DNA, as we illustrate in mouse embryonic stem cells. [ABSTRACT FROM AUTHOR]