- S. Tsukaharaet al., Centromere-targeted de novo integrations
of an LTR retrotransposon ofArabidopsis lyrata.Genes Dev. 26 ,
705 – 713 (2012). doi:10.1101/gad.183871.111; pmid: 22431508 - A. Kawabe, S. Nasuda, Structure and genomic organization
of centromeric repeats inArabidopsisspecies.
Mol. Genet. Genomics 272 , 593–602 (2005). doi:10.1007/
s00438-004-1081-x; pmid: 15586291 - S. J. Klein, R. J. O’Neill, Transposable elements: Genome
innovation, chromosome diversity, and centromere
conflict.Chromosome Res. 26 ,5–23 (2018). doi:10.1007/
s10577-017-9569-5; pmid: 29332159 - H. S. Malik, The centromere-drive hypothesis: A simple basis
for centromere complexity.Prog. Mol. Subcell. Biol. 48 , 33– 52
(2009). doi:10.1007/978-3-642-00182-6_2; pmid: 19521811 - D. Haig, A. Grafen, Genetic scrambling as a defence against
meiotic drive.J. Theor. Biol. 153 , 531–558 (1991). doi:10.1016/
S0022-5193(05)80155-9; pmid: 1806752
ACKNOWLEDGMENTS
This paper is dedicated to Simon Chan. We thank I. Thompson for
ATHILAanalysis, S. Henikoff for the generous gift of CENH3
antibodies, A. Shumate for help with gene Liftoff interpretation,
B. Fischer for advice on high–molecular weight DNA isolation, and
M. Pouch for assistance designing FISH probes.Funding:This
work was supported by BBSRC grants BB/S006842/1, BB/
S020012/1, and BB/V003984/1 to I.R.H.; European Research
Council Consolidator Award ERC-2015-CoG-681987
“SynthHotSpot”to I.R.H.; Marie Curie International Training
Network“MEICOM”to I.R.H.; Human Frontier Science Program
award RGP0025/2021 to T.K., M.C.S., and I.R.H.; US National
Institutes of Health grant S10OD028632-01; US National
Science Foundation grants DBI-1350041 and IOS-1732253 to
M.C.S.; Royal Society awards UF160222 and RGF/R1/180006 to
A.B.; the Czech Science Foundation grant no. 21-03909S to
M.A.L.; the Gregor Mendel Institute to F.B.; grants Fonds zur
Förderung der wissenschaftlichen Forschung (FWF) P26887,
P28320, P30802, P32054, and TAI304 to F.B. and chromatin
dynamics W1238 to A.S. and B.J.; Leverhulme Trust Research
Leadership grant RL-2012-042 to J.T.; and grants from the
Howard Hughes Medical Institute and US National Institutes of
Health (RO1GM067014) to R.A.M.Author contributions:M.N.
sequenced DNA; performed genome assembly and analysis,
ChIP-seq, and DNA methylation analysis; and wrote the
manuscript. M.A. performed genome assembly, polishing,
validation, annotation, and analysis and wrote the manuscript.
P.W. performed satellite repeat annotation and genome analysis
and wrote the manuscript. A.J.T. performed short-read
alignment and genome analysis and wrote the manuscript.
B.W.A. sequenced DNA, performed optical mapping, and
contributed to the assembly. A.S. performed chromatin
immunofluorescence analysis. B.J. provided ChIP-seq data.
C.L. and P.K. performed immunocytology. N.Y. generated the
DMC1 epitope-tagged line. N.H. and K.C. sequenced DNA and
contributed to the assembly. L.M.S., J.T., and K.S. performed
PacBio sequencing. T.K. and R.A.M. provided intellectual input.
T.M. and M.A.L. performed FISH. F.B. supervised ChIP-seq and
immunofluorescence analysis and wrote the manuscript. A.B.
performedATHILAannotation and genome analysis and wrote
the manuscript. T.P.M. supervised DNA sequencing and genome
assembly and analysis and wrote the manuscript. M.C.S.
supervised genome assembly, validation, annotation, and analysis and
wrote the manuscript. I.R.H. supervised DNA sequencing, genome
assembly, validation, annotation, and analysis and wrote the
manuscript.Competing interests:The authors have no competing
interests.Data and materials availability:The ONT sequencing
reads used for assembly are available for download at ArrayExpress
accession E-MTAB-10272 (www.ebi.ac.uk/arrayexpress/). The PacBio
HiFi reads are available for download at European Nucleotide Archive
accession number PRJEB46164 (www.ebi.ac.uk/ena/browser/view/
PRJEB46164). All data, code and materials are available in the
manuscript or the supplementary materials and athttps://github.
com/schatzlab/Col-CEN.
SUPPLEMENTARY MATERIALS
science.org/doi/10.1126/science.abi7489
Materials and Methods
Figs. S1 to S22
Tables S1 to S8
References ( 42 – 94 )
MDAR Reproducibility Checklist
31 March 2021; resubmitted 5 August 2021
Accepted 27 September 2021
10.1126/science.abi7489
Naishet al.,Science 374 , eabi7489 (2021) 12 November 2021 9of9
RESEARCH | RESEARCH ARTICLE