Orderly mitosis shapes interphase genome architecture

  1. Krishnendu Guin
  2. Adib Keikhosravi
  3. Raj Chari
  4. Gianluca Pegoraro
  5. Tom Misteli  Is a corresponding author
  1. National Cancer Institute, NIH, United States
  2. High Throughput Imaging Facility (HiTIF), National Cancer Institute, NIH, United States
  3. Genome Modification Core (GMC), Frederick National Lab for Cancer Research, United States
6 figures, 1 table and 8 additional files

Figures

Figure 1 with 1 supplement
Spatial organization of centromeres is cell-type specific in human cell lines.

(a) Representative images of CENP-C (green) and DAPI (gray) stained nuclei in indicated human cell lines. Scale bar: 10 µm. (b, c) Spatial organization of centromeres quantified using Ripley K’s clustering score (b), CENP-C spot count (c). (d) Nuclear area and (e) mean radial distance in human cell lines. Statistical significance of differences between cell lines for clustering score, spot count, mean radial distance, and nuclear area was tested using analysis of variance (ANOVA) (p-value or ‘Pr(>F)’<2e-16) following Tukey’s HSD test to compare means of all pairs of cell lines. Box plots represent the interquartile range (IQR) between the first and third quartiles (box), the median (horizontal bar), and the whiskers that extend to the highest and lowest data points within 1.5 times the IQR. Values are from one representative experiment with at least 7 technical replicates. At least 1000 cells were analyzed in each category per experiment.

Figure 1—figure supplement 1
Quantification of centromere clustering using CENP-A and CENP-C as centromere markers.

(a) Co-staining of HCT116 cells with CENP-A (red), CENP-C (green), and DAPI (gray). Scale bar: 10 µm. (b) Representative image showing segmentation of DAPI-stained nuclei (gray) and CENP-C-stained centromere spots (green) in high-throughput imaging data using HiTIPS. Red lines around the DAPI-stained nuclei indicate nuclear segmentation, the green circles around CENP-C spots indicate segmentation of centromeres. Zoomed images of the same nucleus are shown, with yellow and pink borders, respectively, indicating before and after spot segmentation was applied. Scale bar: 10 µm. (c) Quantification of spot count and (d) clustering score using CENP-A (red) and CENP-C (blue) as centromere markers in HCT116 cells. Values are from two replicates, with at least 2000 cells analyzed for each experimental condition.

Figure 2 with 4 supplements
Identification of the molecular determinants of spatial centromere distribution across cell types.

(a) Schematics showing three stages of high-throughput imaging-based arrayed CRISPR knockout screen employed to identify molecular determinants of spatial centromere distribution. (b) Changes in spot count (mean Z-score of two replicates, y-axis) and clustering score (mean Z-score of two replicates, x-axis) for each of the 1064 sgRNAs. The most prominent hits were labeled and color-coded as in d. Non-hits are colored in gray. (c) Changes in clustering score in HCT116 (mean Z-score of two replicates, x-axis) and in RPE1 (mean Z-score of two replicates, y-axis) cells for each of the 1064 sgRNAs. Hits and non-hits are color-coded and labeled as in d. A linear trend line (gray) was fitted to the data, and Pearson’s correlation coefficient calculated is indicated at the top-left corner of the plot. (d) Network diagram with lines between 52 common hits drawn based on known physical and/or genetic interactions generated by the STRING database. The thickness of the lines indicates higher strength of data supporting the interaction. Broad categories are color-coded as indicated. (e) Plot shows changes in clustering (clustered or unclustered), count (higher or lower), and direction between two cell lines (same or opposite) for each of the common genes that are color-coded based on their category as in d. Counts of genes in each subcategory are indicated. Values represent two biological replicates. Typically, 200–500 cells were analyzed for each target gene per experiment.

© 2025, BioRender Inc. Figure 2a was created with BioRender and is published under a Creative Commons Attribution License [https://creativecommons.org/licenses/by/4.0/]. Further reproductions must adhere to the terms of this license

Figure 2—figure supplement 1
CRISPR knockout screens for centromere distribution phenotypes in HCT116 cells are reproducible.

(a) Mean and standard deviation for phenotypic separation between control sgRNAs with individual data points representing mean values per well for number of spots per nucleus and (b) clustering score in two biological replicates. (c) Scatter plot showing changes in clustering score or (d) spot count for replicate 1 (x-axis) and replicate 2 (y-axis) in HCT116 cells for each of the 1068 sgRNAs. A linear regression line (gray) was fitted to the data, and Pearson’s correlation coefficient calculated is indicated at the top-left corner of the plot. (e) Scatter plot and linear regression line correlating changes in clustering score (y-axis) and (f) spot count (y-axis) with nuclear area (x-axis). Pearson’s correlation coefficients and corresponding p-values are indicated at the top-left corner. Values are from 2 biological replicates. Typically, 200–500 cells were imaged for each target gene per experiment. Box plots represent the interquartile range (IQR) between the first and third quartiles (box), the median (horizontal bar), and the whiskers that extend to the highest and lowest value within 1.5 times the IQR.

Figure 2—figure supplement 2
Identification of the molecular determinants of spatial centromere distribution in RPE1 cells.

Changes in spot count (mean Z-score of two replicates, y-axis) and clustering score (mean Z-score of two replicates, x-axis) for each of the 1064 sgRNAs. The most prominent hits were labeled and color-coded as in Figure 2d. Non-hits are colored in gray. Values are from two biological replicates. Typically, 200–500 cells were analyzed for each target gene per experiment.

Figure 2—figure supplement 3
Validation of screen hits, cell-cycle analysis of clustering factor knockdown, and their effect on clustering score.

(a) Clustering scores of targets (x-axis) after siRNA knockdown in HCT116 cells. Two control siRNAs for siNCAPH2 are in blue, and siScrambled are in yellow. Upon siRNA knockdown, the clustering score or spot count for the targets (x-axis) labeled in green changes in the same direction as in the CRISPR-KO screens and is compared to the mean value for siScrambled as depicted by a horizontal yellow dotted line by performing pairwise t-tests with Bonferroni correction. Significantly different pairs are labeled with stars, where * indicates p≤0.05, ** indicates p≤0.01, *** indicates p≤0.001, and **** indicates p<0.0001. Targets in gray disagree with either clustering score or spot count or both parameters compared to data in CRISPR-KO screens. Three separate siRNAs were used per target. (b) Fraction of cells in each cell-cycle stage (x-axis) after knockdown of select targets as indicated (y-axis). Individual subpopulations of the cell cycle are color-coded as identified using DAPI and EdU fluorescence intensity measurement, and their percentages are indicated. (c) Bar plots showing the fraction of G1, S, and G2/M cells (y-axis) after siRNA knockdown of select targets (red) and scrambled siRNA control (blue). Statistical significance of difference (p<0.05) was tested using t-test with FDR correction as compared to the scrambled control, and significantly different targets are labeled as stars, where * indicates p≤0.05, ** indicates p≤0.01, *** indicates p≤0.001. A higher number of stars indicate a lower p-value. (d) Clustering score (y-axis) for select targets (x-axis) at G1 (brown), S (gray), and G2/M (green) stages. Values are from one representative experiment. Typically, 200–500 cells were analyzed per gene per experiment. Box plots represent the interquartile range (IQR) between the first and third quartiles (box), the median (horizontal bar), and the whiskers that extend to the highest and lowest value within 1.5 times the IQR.

Figure 2—figure supplement 4
Comparative analysis of cell lines confirms common molecular determinants of spatial centromere distribution.

(a) 52 genes (black mesh) that are hits in both HCT116 and RPE1 cells. A total of 113 hits were selected in RPE1 cells for either clustering score (pink, 45) or spot count (blue, 87), and 111 hits in HCT116 cells for either clustering score (gold, 89) or spot count (black, 45). White non-shaded areas indicate unique hits in HCT116 (51) and RPE1 (53) cells. Values are from one representative experiment. Typically, 200–500 cells were analyzed for each target gene per experiment. (b) Z-scores (x-axis) of spot count and clustering score for the 52 common hits (y-axis) in HCT116 and RPE1 cells. Genes are color-coded based on their category as indicated in Figure 2e.

Figure 3 with 1 supplement
Changes in spatial organization of centromeres require progression through the cell cycle.

(a) HCT116 and RPE1 cells in G1, S, or G2/M phases stained with CENP-C (green), DAPI (gray), and EdU (red). Scale bar: 10 µm. (b) EdU intensity (y-axis) and DAPI intensity (x-axis) showing separation between G1 (brown), S (gray), and G2/M (green) subpopulations in cycling HCT116 cells. Comparison of cells in G1, S, or G2/M (x-axis) for their clustering score (c), spot count (d), mean radial distance (e), or nuclear area (f). Statistical significance of differences was tested by pairwise t-test with Bonferroni correction. Asterisks indicate level of significance between a given pair reflecting the corresponding p-value of that comparison. (g) A linear regression line (red) fitted through the single-cell data for nuclear area (y-axis) and clustering score (x-axis) in cells in different cell-cycle phases in HCT116 and RPE1 cells. Pearson’s correlation coefficient and respective adjusted p-values are indicated at the top of each panel. (h) Experimental outline to test cell-cycle stage-specific effect of knocking down select hits. (i) Effect of siRNA knockdown for a panel of genes (x-axis) using three individual siRNAs per gene in HCT116 cells that are either arrested at G/S and G2 or cycling. Two control siRNAs for siNCAPH2 are in blue and siScrambled in yellow. The mean value for siScrambled is depicted by a horizontal yellow dotted line. Statistical significance of differences was tested by performing pairwise t-tests with Bonferroni correction using siScrambled as control group. Box plots represent the interquartile range (IQR) between the first and third quartiles (box), the median (horizontal bar), and the whiskers that extend to the highest and lowest value within 1.5 times the IQR. Values are from one representative experiment. Typically, 200–500 cells were analyzed in each category. Statistical significance of difference was denoted by stars where * indicates p≤0.05, ** indicates p≤0.01, *** indicates p≤0.001, and **** indicates p<0.0001.

© 2025, BioRender Inc. Figure 3h was created with BioRender and is published under a Creative Commons Attribution License [https://creativecommons.org/licenses/by/4.0/]. Further reproductions must adhere to the terms of this license

Figure 3—figure supplement 1
Validation of protein depletion in siRNA knockdown.

(a) Number of cells per well at 72 hrs after transfection of siDEATH and siScrambled control siRNAs in cycling (red), G1/S (green), or G2/M (blue) synchronized HCT116 cells as indicated. Values are from one representative experiment with 7 technical replicates. Box plots represent the interquartile range (IQR) between the first and third quartiles (box), the median (horizontal bar), and the whiskers that extend to the highest and lowest value within 1.5 times the IQR. (b) Western blots showing NCAPH2 protein levels after 72 hrs of siRNA knockdown using siNCAPH2 and siScrambled in cycling, G1/S, or G2/M synchronized HCT116 cells as indicated. NCAPH2 levels across samples were normalized using β-actin, and quantitated band intensities of NCAPH2 in siNCAPH2 samples were expressed as a fraction of the corresponding siScrambled samples in cycling, G1/S, and G2/M cells.

Figure 3—figure supplement 1—source data 1

Original western blot images used in Figure 3—figure supplement 1.

https://cdn.elifesciences.org/articles/108410/elife-108410-fig3-figsupp1-data1-v1.zip
Figure 3—figure supplement 1—source data 2

PDF file containing annotated western blot images used in Figure 3—figure supplement 1, with figure legend explaining relevant details.

https://cdn.elifesciences.org/articles/108410/elife-108410-fig3-figsupp1-data2-v1.zip
Figure 4 with 4 supplements
Progression through S-phase in the absence of select clustering factors does not alter interphase genome organization.

(a) Experimental outline to compare centromere distribution during progression through S-phase in the presence or absence of clustering factors. (b, c) Clustering score in all cells (b) or G2/M cells (c) in the presence (blue) or absence (red) of indicated clustering factors before and after S-phase release from G1/S arrest. Pairwise comparisons were performed using t-tests with Bonferroni correction, and the level of significance is indicated by asterisks if any. Pairs without significant difference are not labeled. Box plots represent the interquartile range (IQR) between the first and third quartiles (box), the median (horizontal bar), and the whiskers that extend to the highest and lowest value within 1.5 times the IQR. Values are from one representative experiment with three technical replicates. Typically, 200–500 cells were analyzed in each category.

Figure 4—figure supplement 1
Construction and genotyping of FLAG-dTAG-SPC24 and NUF2-dTAG-FLAG cell lines.

(a, b) CRISPR knock-in strategy for homozygous tagging of SPC24 (a) and NUF2 (b) with the dTAG-FLAG epitope. Horizontal black arrows indicate positions of primers used for PCR confirmation of the tagged allele. (c, d) PCR genotyping of single-cell clone for FLAG-dTAG-SPC24 (c) and NUF2-dTAG-FLAG based on the strategy explained in a and b.

Figure 4—figure supplement 1—source data 1

Original western blot images used in Figure 4—figure supplement 1.

https://cdn.elifesciences.org/articles/108410/elife-108410-fig4-figsupp1-data1-v1.zip
Figure 4—figure supplement 1—source data 2

PDF file containing annotated western blot images used in Figure 4—figure supplement 1, with figure legend explaining relevant details.

https://cdn.elifesciences.org/articles/108410/elife-108410-fig4-figsupp1-data2-v1.zip
Figure 4—figure supplement 2
Characterization of FLAG-dTAG-SPC24 and NUF2-dTAG-FLAG cell lines.

(a, b) Representative images of knock-in cell lines expressing FLAG-dTAG-SPC24 (a), and NUF2-dTAG-FLAG (b) stained with DAPI (gray), CENP-C (green), and FLAG (red). Scale bar: 10 µm. (c, d) Western blot images showing levels of FLAG-dTAG-SPC24 (c) and NUF2-dTAG-FLAG (d) at indicated time points after incubation with dTAG ligands and the relative ratios of dTAG-SPC24 or NUF2-dTAG to tubulin control compared to the level at the beginning of depletion (0 hr) are indicated below. (e, f) Western blots showing comparative levels of SPC24 and FLAG-dTAG-SPC24 (e) and NUF2 and NUF2-dTAG-FLAG (f) proteins in the indicated cell lines. Relative intensity ratio of FLAG-dTAG-SPC24 or NUF2-dTAG-FLAG proteins in the respective cell lines to the untagged SPC24 or NUF2 protein level in HCT116 cells is indicated.

Figure 4—figure supplement 2—source data 1

Original western blot images used in Figure 4—figure supplement 2.

https://cdn.elifesciences.org/articles/108410/elife-108410-fig4-figsupp2-data1-v1.zip
Figure 4—figure supplement 2—source data 2

PDF file containing annotated western blot images used in Figure 4—figure supplement 2, with figure legend explaining relevant details.

https://cdn.elifesciences.org/articles/108410/elife-108410-fig4-figsupp2-data2-v1.zip
Figure 4—figure supplement 3
Quantification of the cell-cycle stage-specific depletion of FLAG-dTAG-SPC24 and NUF2-dTAG-FLAG.

(a, b) Western blots showing degron-based depletion of FLAG-dTAG-SPC24 in G1/S synchronized and cycling cells (a) and G2/M synchronized and cycling HCT116 cells (b). (c, d) Western blots showing degron-based depletion of SPC24-dTAG-FLAG in G1/S synchronized and cycling cells (c) and G2/M synchronized and cycling HCT116 cells (d). Middle lanes contain protein size markers largely invisible in the chemiluminescence images.

Figure 4—figure supplement 3—source data 1

Original western blot images used in Figure 4—figure supplement 3.

https://cdn.elifesciences.org/articles/108410/elife-108410-fig4-figsupp3-data1-v1.zip
Figure 4—figure supplement 3—source data 2

PDF file containing annotated western blot images used in Figure 4—figure supplement 3, with figure legend explaining relevant details.

https://cdn.elifesciences.org/articles/108410/elife-108410-fig4-figsupp3-data2-v1.zip
Figure 4—figure supplement 4
Quantification of cell-cycle stages during G1 and mitotic release.

(a) Fraction of G1 (blue), S (green), and G2/M (orange) cells were quantified before (0 hr) and after (6 hr) release from double thymidine block in the presence or absence of indicated clustering factors. (b) Fraction of G1 (blue), S (green), and G2/M (orange) cells were quantified before (0 hr) and after (6 hr) release from G2/M block in the presence or absence of indicated clustering factors. Values are from one representative experiment containing three technical replicates. Typically, 200–500 cells were analyzed per sample. (c and d) show percent of G1, S, and G2/M cells (x-axis) in presence (blue) and absence (red) of indicated mitotic factors before (0 hr) and after (6 hr) release from G1/S block (c) and G2/M block (d). Statistical significance of difference was tested using t-test, and significantly (p<0.05) different pairs are indicated by stars, where * indicates p≤0.05, ** indicates p≤0.01. A higher number of stars indicate a lower p-value. Box plots represent the interquartile range (IQR) between the first and third quartiles (box), the median (horizontal bar), and the whiskers that extend to the highest and lowest value within 1.5 times the IQR.

Figure 5 with 1 supplement
Orderly progression through mitosis is required for normal centromere distribution.

(a) Experimental outline to compare centromere distribution during mitotic progression in the presence or absence of clustering factors. (b, c) Clustering score in all cells (b) or G1 cells (c) in the presence (blue) and absence (red) of indicated clustering factors before (0 hr) and after (6 hr) mitotic release from G2 arrest. Pairwise comparisons were performed using t-tests with Bonferroni correction, and the level of significance is indicated by asterisks. Pairs without significant differences are not labeled. (d) Representative images showing G1 nuclei stained with DAPI (gray) and CENP-C (green) in the presence or absence of indicated factors. Scale bar: 10 µm. (e) Schematics for co-depletion of indicated factors. (f) Clustering score (y-axis) in G1 cells after siRNA knockdown of indicated factors (x-axis) in presence (blue) or absence (red) of SPC24, KI67, or NCAPH2 as indicated. Statistical significance of difference between indicated pairs was tested by performing t-test with Bonferroni corrections for multiple comparisons and denoted by stars, where * indicates p≤0.05, ** indicates p≤0.01, *** indicates p≤0.001, and **** indicates p<0.0001. Comparisons between individual siRNA groups (x-axis) and inter-group comparisons in the presence and absence of the given degron tagged protein are shown in black, blue (presence), and red (absence), respectively. (g) Clustering score (y-axis) in cells that were depleted (red) or depleted of indicated factors and then re-expressed (green) or remained unperturbed (blue). Statistical significance of difference between indicated pairs was tested by performing t-test with Bonferroni corrections for multiple comparisons and denoted by stars, where a higher number of stars indicate higher confidence levels. Box plots represent the interquartile range (IQR) between the first and third quartiles (box), the median (horizontal bar), and the whiskers that extend to the highest and lowest value within 1.5 times the IQR. Values are from one representative experiment containing three technical replicates. Typically, 200–500 cells were analyzed for each category. Statistical significance of difference was denoted by stars where * indicates p≤0.05, ** indicates p≤0.01, *** indicates p≤0.001, and **** indicates p<0.0001.

Figure 5—figure supplement 1
Mitotic defects in the absence of SPC24.

(a–d) Representative images showing examples of mitotic defects, including karyokinesis defect (a), mis-segregation (b), micronuclei formation (c), and aberrant metaphase alignment (d) observed at 6 hrs after G2/M arrested HCT116 cells, were released in the absence of SPC24. Cells were immunofluorescently labeled with CENP-C (green) and DAPI (gray). Scale bar: 10 µm.

Mitotic events shape interphase genome organization.

A model showing defective loading of outer kinetochore (inset; green and red box) in late G2 leads to uncoordinated metaphase alignment and aberrant migration toward spindle pole during anaphase that lowers chances of interactions between centromeres during telophase, and the lack of homotypic adhesion results in dispersion of centromeres in the daughter nuclei.

© 2025, BioRender Inc. Figure 6 was created with BioRender and is published under a Creative Commons Attribution License [https://creativecommons.org/licenses/by/4.0/]. Further reproductions must adhere to the terms of this license

Tables

Key resources table
Reagent type (species) or resourceDesignationSource or referenceIdentifiersAdditional information
Cell line (Homo sapiens)Cell line FLAG-dTAG-SPC24 S8This studyParental cell line
HCT116 Cas9
Cell line (Homo sapiens)Cell line NUF2-dTAG-FLAG N10This studyParental cell line
HCT116 Cas9
Cell line (Homo sapiens)Cell line NCAPH2-mAChTakagi et al., 2018Parental cell line
HCT116
Cell line (Homo sapiens)Cell line
KI-67-mACl
Takagi et al., 2018Parental cell line
HCT116
Cell line (Homo sapiens)Cell line HCT116 Cas9Hart et al., 2015Human colon cancer
Cell line (Homo sapiens)Cell line
A375 Cas9
Hart et al., 2015Near triploid, Human malignant melanoma
Cell line (Homo sapiens)Cell line hTERT-RPE1 Cas9Hart et al., 2015Immortalized human Retinal pigment epithelia
Cell line (Homo sapiens)Cell line
WTC-11
Coriell InstituteGM25256Human induced pluripotent stem cell (iPSC)
Cell line (Homo sapiens)Cell line
MDA-MB-231 Cas9
Horizon DiscoveryHD Cas9-014Human triple-negative breast cancer
Cell line (Homo sapiens)Cell line
HAP1 Cas9
Horizon DiscoveryHD Cas9-011Near haploid, chronic myelogenous leukemia
Cell line (Homo sapiens)Cell line
A549 Cas9
Horizon DiscoveryHD Cas9-001Human lung adenocarcinoma
Cell line (Homo sapiens)Cell line
HFF-hTERT
Clone 6,
Dekker lab; 4DNucleome project cell line
Immortalized human foreskin fibroblast
Recombinant DNA reagentPlasmid pJT142This studyAvailable from AddgeneCas9 and sgRNA targeting SPC24
Recombinant DNA reagentPlasmid pMG1040This studyAvailable from AddgeneCas9 and sgRNA targeting NUF2
Recombinant DNA reagentPlasmid pJT152This studyAvailable from AddgeneDonor for FLAG-dTAG-SPC24
Recombinant DNA reagentPlasmid pMG1064This studyAvailable from AddgeneDonor for NUF2-dTAG-FLAG
AntibodyGuinea pig polyclonal anti-CENP-CMBL BiosciencesPD0301:1000 dilution for IF staining
AntibodyMouse monoclonal anti-CENP-AAbcamAB139391:1000 dilution for IF staining
AntibodyMouse monoclonal anti-FLAGSigmaF31651:250 dilution for IF staining
AntibodyMouse monoclonal anti-β-ActinMillipore SigmaA22281:25,000 dilution for western blotting
AntibodyRabbit polyclonal anti-SPC24McCleland et al., 2004Gift from Todd Stukenberg lab1:8000 dilution for western blotting
AntibodyRabbit monoclonal anti-NUF2Abcamab1765561:2000 dilution for western blotting
Chemical compoundRO-3306Millipore SigmaCat. No. 217721Working concentration 9 μM
Chemical compoundAuxinMillipore SigmaI3750-25G-AWorking concentration 500 nM
Chemical compounddTAG13Tocris BioscienceCat. No. 6605Working concentration 1 μM
Chemical compounddTAGV-1Tocris BioscienceCat. No. 6914Working concentration 1 μM
Chemical compoundThymidineSigmaCat. No. T9250-5GWorking concentration 2 mM
Chemical compoundLipofectamineThermo Fisher ScientificCat. No. 13778075Transfection reagent

Additional files

Supplementary file 1

Source, culture condition, and growth media used for cell lines used in this study.

https://cdn.elifesciences.org/articles/108410/elife-108410-supp1-v1.xlsx
Supplementary file 2

Results of statistical analysis for CENP-C spot count, clustering score, radial position, and nuclear area as shown in Figure 1.

https://cdn.elifesciences.org/articles/108410/elife-108410-supp2-v1.xlsx
Supplementary file 3

Sequences of sgRNAs used to target 1064 genes in the CRISPR-KO screens.

https://cdn.elifesciences.org/articles/108410/elife-108410-supp3-v1.csv
Supplementary file 4

Measurement data for CENP-C spots and nuclei obtained from CRISPR-KO screens in HCT116 Cas9 cells.

https://cdn.elifesciences.org/articles/108410/elife-108410-supp4-v1.csv
Supplementary file 5

Measurement data for CENP-C spots and nuclei obtained from CRISPR-KO screens in hTERT-RPE1 Cas9 cells.

https://cdn.elifesciences.org/articles/108410/elife-108410-supp5-v1.csv
Supplementary file 6

Sequences of sgRNAs and DNA oligonucleotides used to generate SPC24 and NUF2 dTAG cell lines.

https://cdn.elifesciences.org/articles/108410/elife-108410-supp6-v1.xlsx
Supplementary file 7

Details of siRNAs used for validation of CRISPR-KO screen hits.

https://cdn.elifesciences.org/articles/108410/elife-108410-supp7-v1.xlsx
MDAR checklist
https://cdn.elifesciences.org/articles/108410/elife-108410-mdarchecklist1-v1.pdf

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  1. Krishnendu Guin
  2. Adib Keikhosravi
  3. Raj Chari
  4. Gianluca Pegoraro
  5. Tom Misteli
(2026)
Orderly mitosis shapes interphase genome architecture
eLife 14:RP108410.
https://doi.org/10.7554/eLife.108410.3