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. 2021 Mar 31;12:659558. doi: 10.3389/fpls.2021.659558

Table 1.

Selection of cohesin/SMC genes and cofactors with potential for inclusion in breeding initiatives in major crops.

SMC gene and associated factors Functional features in the literature and public databases1,2 Locus ID as determined by TAIR, NCBI, and the Human Genome Organization (HUGO)5 Presumed homologs of interest (rice and maize) retrieved from databases3,4,6 by the respective coding sequences (CDS) Likely agricultural use
Arabidopsis HORMA-domain protein ASYNAPTIC 1 (ASY1) Distribution of COs along meiotic chromosomes in a REC8/SYN1 dependent manner At1g67370 Os09g32930, GRMZM2G035996 Knockdown alleles may reduce linkage drag in major crops
Human cohesin interacting factors BRG1/SMARCA4 and ARID2 (SWI/SNF subunits) Cohesin-mediated repression of transcriptional activity, genome maintenance HGNC:6597 and HGNC:196528 Os05g05230, GRMZM2G102625, Os07g33860, GRMZM2G009412 Promotion of seed longevity, intentional induction of inversions and translocations
Human ATP-dependent DNA helicase DEAD/H-box helicase 11 (DDX11) Synthetically lethal with human ECO1 homolog ESCO2 (HGNC:27230) and epistatic to human WAPL (HGNC:23293). Promotes cohesion in arms and centromeres during replication, probably by promotion of SMC3 acetylation, and enhanced stability of replication forks HGNC:1663 Os05g13300, GRMZM2G100067 Promotion of replication fork stability. May constitute an important layer of genome maintenance under stress across all eukaryotes
Arabidopsis N-acetylase ECO1/CTF7 Acetylation of the ATP-ase domain of SMC proteins, establishment of cohesion, homology-dependent DNA repair. Its function is antagonized by WAPL At4g31400 Os05g31230, GRMZM2G100067; and Os04g42120, GRMZM2G075145 Prevention of large-scale rearrangements, targeted manipulation of chromosome folding during MSUC and MSCI in meiocytes of papaya, gene expression.
Yeast chromatin-remodeling ATPase ISW1/YBR245C Cohesin-dependent processes such as DNA repair and transcription in promoters SGD:S000000449 Os03g22900, GRMZM2G469162 May stabilize transcription under stress across eukaryotic organisms
Cohesin subunit PRECOCIOUS DISSOCIATION OF SISTERS 5 (PDS5A/E) Cohesion-promoting cofactor; has a role in fork protection and stable DNA replication At5g47690, At1g77600, At4g31880, At1g80810, and At1g15940 For PDS5A: Os06g17840, GRMZM2G010637 May stabilize DNA replication under stress, may be a target for the manipulation of DNA looping and chromosome folding during MSUC and MSCI in meiocytes of papaya
Arabidopsis SCC2 Initial CO formation during meiosis, 3D-chromosome folding, loading of cohesin complexes, chromosome looping and DNA repair At5g15540 Os07g01940, GRMZM2G132504, Carica papaya evm.model.supercontig_2744.1 Promotion of crossover resolution, targeted manipulation of chromosome folding during MSUC and MSCI in meiocytes of papaya
Arabidopsis SCC3 Regulates release of α-kleisins. Regulates meiotic orientation of kinetochores in Arabidopsis, and may have a role in the prevention of large-scale rearrangements during replication, dimerization of cohesin, and chromosome looping At2g47980 Os05g09620, GRMZM2G131443 May prevent chromosomal rearrangements during replication, in plants, it may provide a valuable layer of genome maintenance under stress
Maize SCC4/DEK15 Mitotic chromosome segregation, endosperm and embryo formation, transcriptional regulation of kernel development in maize GRMZM2G079796 Os04g28010 Promotion of embryo and endosperm development
Arabidopsis condensin subunit STRUCTURAL MAINTENANCE OF CHROMOSOMES 4 (SMC4) Silencing of pericentromeric TEs, DNA methylation, transcription of genes related to translesion synthesis, male gamete development, tolerance to Boron toxicity At5g48600 Os05g41750, GRMZM2G416501 May lead to enhanced tolerance to exogenous DNA damage, and improved genome maintenance in crops
Arabidopsis EXTRA SPINDLE POLES (AESP1), Separase Cleavage of the REC8/SYN1 α-kleisin during anaphase I and anaphase II, allowing disjunction of homologs and sister centromeres, respectively. Proper assembly of radial microtubule system after telophase II, embryo development, cellularization of the endosperm, and vesicle trafficking At4g22970 Os02g53120, GRMZM2G300624 Promotion of crossovers resolution and proper tetrad formation. May facilitate breeding of fertile cultivars in major crops
Arabidopsis SWITCH1/DYAD (SWI1) Sister chromatid cohesion and meiotic chromosome organization, assembly of the chromosome axis At5g51330 Known as AMEIOTIC (OsAM1) in rice: Os03g44760, and AMEIOTIC (AM1) in maize (possibly GRMZM5G883855) May facilitate the identification of new alternative meiotic α-kleisins in crops
Arabidopsis WINGS APART-LIKE PROTEIN 1 (WAPL1/2) Release of meiotic α-kleisin REC8/SYN1 during prophase I. Cohesin removal by WAPL is required to complete DNA synthesis under conditions of persistent DNA replication stress. Regulates chromosome folding At1g11060 and At1g61030 Os10g35380, GRMZM2G034276; Os10g35380, GRMZM2G034276 New alleles may facilitate DNA replication under persistent damage. May confer tolerance to chronic DNA damage in crops

MSUC, meiotic silencing of unsynapsed chromatin; MSCI, meiotic sex chromosome inactivation; TE, transposable elements.

1

Database consulted was The Arabidopsis Information Resource (TAIR): https://www.arabidopsis.org/.

2

Database consulted was National Center for Biotechnology Information (NCBI): https://www.ncbi.nlm.nih.gov/.

3

Database consulted was Phytozome v12.1: https://phytozome.jgi.doe.gov/pz/portal.html.

4

Database consulted was Maize GDB: https://www.maizegdb.org/.

5

Database consulted was the Human Genome Organization (HUGO): http://www.hugo-international.org/.

6

Database consulted was the Rice Genome Annotation Project: http://rice.plantbiology.msu.edu/cgi-bin/gbrowse/rice/.