Abstract:SUMMARY a-Kleisins are core components of meiotic and mitotic cohesin complexes. Arabidopsis contains four genes that encode a-kleisin proteins: SYN1, SYN2, SYN3 and SYN4. SYN1, a REC8 ortholog, is essential for meiosis, while SYN2 and SYN4 appear to be SCC1 orthologs and function in mitosis. SYN3 is essential for megagametogenesis and is enriched in the nucleolus of meiotic and mitotic cells. In this study the role of SYN3 during meiosis was investigated by characterization of plants that express SYN3-RNAi co… Show more
“…They facilitate long-distance DNA interactions (Dorsett and Merkenschlager, 2013) and have been shown to affect long-range interactions between binding sites of the transcriptional repressor CTCF in a number of genes in animal cells (Bonora et al, 2014). Consistent with this, alterations in cohesin protein levels can result in tissue-specific alterations in transcript levels for a number of genes in Arabidopsis (this study; Yuan et al, 2012;Bolaños-Villegas et al, 2013;Liu and Makaroff, 2015). For example, transgenic plants expressing a pro35S:AtCTF7ΔB construct and ctf7 plants contain elevated levels of transcripts for epigenetically regulated transposable elements, including MU1, COPIA 28, and solo LTR, and alterations in transcript levels of several small interfering RNA-associated genes (Bolaños-Villegas et al, 2013;Liu and Makaroff, 2015).…”
Section: Ctf7 and Wapl Are Essential For Embryo Development But Not supporting
Sister chromatid cohesion, which is mediated by the cohesin complex, is essential for the proper segregation of chromosomes during mitosis and meiosis. Stable binding of cohesin with chromosomes is regulated in part by the opposing actions of CTF7 (CHROMOSOME TRANSMISSION FIDELITY7) and WAPL (WINGS APART-LIKE). In this study, we characterized the interaction between Arabidopsis thaliana CTF7 and WAPL by conducting a detailed analysis of wapl1-1 wapl2 ctf7 plants. ctf7 plants exhibit major defects in vegetative growth and development and are completely sterile. Inactivation of WAPL restores normal growth, mitosis, and some fertility to ctf7 plants. This shows that the CTF7/WAPL cohesin system is not essential for mitosis in vegetative cells and suggests that plants may contain a second mechanism to regulate mitotic cohesin. WAPL inactivation restores cohesin binding and suppresses ctf7-associated meiotic cohesion defects, demonstrating that WAPL and CTF7 function as antagonists to regulate meiotic sister chromatid cohesion. The ctf7 mutation only had a minor effect on wapl-associated defects in chromosome condensation and centromere association. These results demonstrate that WAPL has additional roles that are independent of its role in regulating chromatin-bound cohesin.
“…They facilitate long-distance DNA interactions (Dorsett and Merkenschlager, 2013) and have been shown to affect long-range interactions between binding sites of the transcriptional repressor CTCF in a number of genes in animal cells (Bonora et al, 2014). Consistent with this, alterations in cohesin protein levels can result in tissue-specific alterations in transcript levels for a number of genes in Arabidopsis (this study; Yuan et al, 2012;Bolaños-Villegas et al, 2013;Liu and Makaroff, 2015). For example, transgenic plants expressing a pro35S:AtCTF7ΔB construct and ctf7 plants contain elevated levels of transcripts for epigenetically regulated transposable elements, including MU1, COPIA 28, and solo LTR, and alterations in transcript levels of several small interfering RNA-associated genes (Bolaños-Villegas et al, 2013;Liu and Makaroff, 2015).…”
Section: Ctf7 and Wapl Are Essential For Embryo Development But Not supporting
Sister chromatid cohesion, which is mediated by the cohesin complex, is essential for the proper segregation of chromosomes during mitosis and meiosis. Stable binding of cohesin with chromosomes is regulated in part by the opposing actions of CTF7 (CHROMOSOME TRANSMISSION FIDELITY7) and WAPL (WINGS APART-LIKE). In this study, we characterized the interaction between Arabidopsis thaliana CTF7 and WAPL by conducting a detailed analysis of wapl1-1 wapl2 ctf7 plants. ctf7 plants exhibit major defects in vegetative growth and development and are completely sterile. Inactivation of WAPL restores normal growth, mitosis, and some fertility to ctf7 plants. This shows that the CTF7/WAPL cohesin system is not essential for mitosis in vegetative cells and suggests that plants may contain a second mechanism to regulate mitotic cohesin. WAPL inactivation restores cohesin binding and suppresses ctf7-associated meiotic cohesion defects, demonstrating that WAPL and CTF7 function as antagonists to regulate meiotic sister chromatid cohesion. The ctf7 mutation only had a minor effect on wapl-associated defects in chromosome condensation and centromere association. These results demonstrate that WAPL has additional roles that are independent of its role in regulating chromatin-bound cohesin.
“…8). Overexpression of cohesin proteins affects meiotic cell division in Arabidopsis (Yuan et al, 2012). Therefore, we established lines expressing kleisin subunits under the control of the ABI3 promoter, which has a relatively low and localized activity (Ng et al, 2004).…”
Section: Atscc4 Is Essential For Plant Cohesin Immobilization In the mentioning
confidence: 99%
“…The Arabidopsis genome encodes four kleisin subunits (Fig. 1A): the meiosis-specific SYN1 (Cai et al, 2003) and SYN3, involved in gene expression of meiotic genes, but also expressed in somatic cells (Yuan et al, 2012;Jiang et al, 2007), and SYN2 and SYN4, which have been suggested to participate in mitotic cell division (da Costa-Nunes et al, 2006).…”
Factors regulating dynamics of chromatin structure have direct impact on expression of genetic information. Cohesin is a multi-subunit protein complex that is crucial for pairing sister chromatids during cell division, DNA repair and regulation of gene transcription and silencing. In non-plant species, cohesin is loaded on chromatin by the Scc2-Scc4 complex (also known as the NIBPL-MAU2 complex). Here, we identify the Arabidopsis homolog of Scc4, which we denote Arabidopsis thaliana (At)SCC4, and show that it forms a functional complex with AtSCC2, the homolog of Scc2. We demonstrate that AtSCC2 and AtSCC4 act in the same pathway, and that both proteins are indispensable for cell fate determination during early stages of embryo development. Mutant embryos lacking either of these proteins develop only up to the globular stage, and show the suspensor overproliferation phenotype preceded by ectopic auxin maxima distribution. We further establish a new assay to reveal the AtSCC4-dependent dynamics of cohesin loading on chromatin in vivo. Our findings define the Scc2-Scc4 complex as an evolutionary conserved machinery controlling cohesin loading and chromatin structure maintenance, and provide new insight into the plant-specific role of this complex in controlling cell fate during embryogenesis.
“…However, further studies point to SYN3 acting as a significant player during female meiosis [Yuan et al, 2012]. In SYN3 RNAi plants, male gametogenesis is only slightly affected, with a marginal reduction in pollen viability [Yuan et al, 2012]. Yet, these plants have significantly reduced fertility, resulting from the abortion of megaspore mother cells [Yuan et al, 2012].…”
Section: Sex-specific Cohesins In Arabidopsismentioning
confidence: 99%
“…However, further studies point to SYN3 acting as a significant player during female meiosis [Yuan et al, 2012]. In SYN3 RNAi plants, male gametogenesis is only slightly affected, with a marginal reduction in pollen viability [Yuan et al, 2012].…”
Section: Sex-specific Cohesins In Arabidopsismentioning
Chromosome structure is important for many meiotic processes. Here, we outline 3 main determinants of chromosome structure and their effects on meiotic processes in plants. Cohesins are necessary to hold sister chromatids together until the first meiotic division, ensuring that homologous chromosomes and not sister chromatids separate during anaphase I. During meiosis in maize, Arabidopsis, and rice, cohesins are needed for establishing early prophase chromosome structure and recombination and for aligning bivalents at the metaphase plate. Condensin complexes play pivotal roles in controlling the packaging of chromatin into chromosomes through chromatin compaction and chromosome individualization. In animals and fungi, these complexes establish a meiotic chromosome structure that allows for proper recombination, pairing, and synapsis of homologous chromosomes. In plants, information on the role of condensins in meiosis is limited, but they are known to be required for successful completion of reproductive development. Therefore, we speculate that they play roles similar to animal and fungal condensins during meiosis. Plants generally have large and complex genomes due to frequent polyploidy events, and likely, condensins and cohesins organize chromosomes in such a way as to ensure genome stability. Hexaploid wheat has evolved a unique mechanism using a Ph1 locus-controlled chromosome organization to ensure proper chromosome pairing in meiosis. Altogether, studies on meiotic chromosome structure indicate that chromosome organization is not only important for chromatin packaging but also fulfills specific functions in facilitating chromosome interactions during meiosis, including pairing and recombination.
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