Larner College of Medicine

Stein Lab Research

Nuclear Structure and Function

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Regulatory machinery for vital cellular processes, including transcription, replication, repair, and apoptosis, as well as biological control of proliferation, lineage commitment, phenotype, tumor promotion, and tumor suppression are organized in distinct nuclear microenvironments. Our group has identified molecular and epigenetic mechanisms that govern localization of lineage-determining regulatory proteins to subnuclear sites of transcription. Applying a combination of genetic, biochemical, and in situ cell biological techniques supported by advanced bioinformatic analysis, we have identified unique trafficking signals in Runx proteins, master regulators of osteogenesis, hematopoiesis, and neurogenesis/gastro-intestinal development. Disruption of physiological localization of proteins to distinct subnuclear sites (e.g., ‘Runx foci’) results in altered transcriptional programs ultimately leading to compromised cell growth and differentiation. Clinical relevance of these findings is emphasized by aberrant subnuclear targeting of these proteins under a variety of pathological conditions that include acute myelogenous leukemia, as well as breast and prostate cancers. Currently, we are using multiomic genomic (high-throughput DeepSeq and single cell analysis) and proteomic Bio ID and high-resolution mass spec approaches to define mechanisms that relate subnuclear organization of regulatory factors and gene loci with biological control of cell function and pathological disruption in cancer. Nuclear microenvironments are being explored that support the function of Runx proteins. We are investigating subnuclear organelles including Histone Locus Bodies (see Genetic and Epigenetic Regulation of Gene Expression). Utilizing a recently developed degron approach to rapidly and selectively degrade the RUNX1 transcription factor, which is an obligatory tumor suppressor in normal mammary epithelial cells, we are, for the first time, defining initial parameters of control that epigenetically mediate an epithelial to mesenchymal transition and initiation of breast cancer tumor genesis. 

Our laboratory examines the chromatin structure and higher-order chromatin organization of genes that mediate cell cycle control (e.g., histone genes) and osteoblast differentiation (e.g., osteocalcin and Runx2 genes) (see Musculoskeletal Biology and Pathology).

Nuclear Structure and Function Papers

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  • Glencer A; Ramalingam K; Schindler N; Hidetoshi M; Ghule P; Lee K; Nachmanson D; Officer A; Harismendy O; Stein JL; Stein GS;  Evans M; Weaver D; Yau C; Hirst GL; Campbell MJ; Esserman LJ; Borowsky AD. Tumor microenvironmental determinants of high-risk DCIS progression. Submitted.
  • Gordon JAR, Tye CE, Banerjee B, Ghule PN, van Wijnen AJ, Kabala FS, Page NA, Falcone MM, Stein JL, Stein GS, Lian JB. LINC01638 sustains human mesenchymal stem cell self-renewal and competency for osteogenic cell fate. Sci Rep. 2023 Nov 20;13(1):20314. doi: 10.1038/s41598-023-46202-z. PMID: 37985890; PMCID: PMC10662126.
  • Ghule PN, Boyd JR, Kabala F, Fritz AJ, Bouffard NA, Gao C, Bright K, Macfarlane J, Seward DJ, Pegoraro G, Misteli T, Lian JB, Frietze S, Stein JL, van Wijnen AJ, Stein GS. Spatiotemporal higher-order chromatin landscape of human histone gene clusters at histone locus bodies during the cell cycle in breast cancer progression. Gene. 2023 Jul 1;872:147441. doi: 10.1016/j.gene.2023.147441. Epub 2023 Apr 23. Erratum in: Gene. 2023 May 11;873:147469. PMID: 37094694; PMCID: PMC10370284.
  • Gordon JAR, Evans MF, Ghule PN, Lee K, Vacek P, Sprague BL, Weaver DL, Stein GS, Stein JL. Identification of molecularly unique tumor-associated mesenchymal stromal cells in breast cancer patients. PLoS One. 2023 Mar 20;18(3):e0282473. doi: 10.1371/journal.pone.0282473. PMID: 36940196; PMCID: PMC10027225.
  • Tye CE, Ghule PN, Gordon JAR, Kabala FS, Page NA, Falcone MM, Tracy KM, van Wijnen AJ, Stein JL, Lian JB, Stein GS. LncMIR181A1HG is a novel chromatin-bound epigenetic suppressor of early stage osteogenic lineage commitment. Sci Rep. 2022 May 11;12(1):7770. doi: 10.1038/s41598-022-11814-4. PMID: 35546168; PMCID: PMC9095685.
  • Nachmanson D, Officer A, Mori H, Gordon J, Evans MF, Steward J, Yao H, O'Keefe T, Hasteh F, Stein GS, Jepsen K, Weaver DL, Hirst GL, Sprague BL, Esserman LJ, Borowsky AD, Stein JL, Harismendy O. The breast pre-cancer atlas illustrates the molecular and micro-environmental diversity of ductal carcinoma in situ. NPJ Breast Cancer. 2022 Jan 13;8(1):6. doi: 10.1038/s41523-021-00365-y. PMID: 35027560; PMCID: PMC8758681.
  • Andrew J. Fritz, Mohammed El Dika, Rabail H. Toor, Princess D. Rodriguez, Stephen J. Foley, Rahim Ullah, Daijing Nie, Bodhisattwa Banerjee, Dorcas Lohese, Karen C. Glass, Seth Frietze, Prachi N. Ghule, Jessica L. Heath, Anthony N. Imbalzano, Andre van Wijnen, Jonathan Gordon, Jane B. Lian, Janet L. Stein, Gary S. Stein. Epigenetic-mediated regulation of gene expression for biological control and cancer: cell and tissue structure, function, and phenotype, Nuclear, Chromosomal, and Genomic Architecture in Biology and Medicine. Switzerland: Springer Nature. 2022. In print.
  • Nirala NK, Li Q, Ghule PN, Chen HJ, Li R, Zhu LJ, Wang R, Rice NP, Mao J, Stein JL, Stein GS, van Wijnen AJ, Ip YT. Hinfp is a guardian of the somatic genome by repressing transposable elements. Proc Natl Acad Sci U S A. 2021 Oct 12;118(41):e2100839118. doi: 10.1073/pnas.2100839118. PMID: 34620709; PMCID: PMC8521681.
  • Tracy KM, Tye CE, Ghule PN, Malaby HLH, Stumpff J, Stein JL, Stein GS, Lian JB. Mitotically-Associated lncRNA (MANCR) Affects Genomic Stability and Cell Division in Aggressive Breast Cancer. Mol Cancer Res. 2018 Apr;16(4):587-598. doi: 10.1158/1541-7786.MCR-17-0548. Epub 2018 Jan 29. PMID: 29378907; PMCID: PMC5882506.
  • Zaidi SK, Young DW, Javed A, Pratap J, Montecino M, van Wijnen A, Lian JB, Stein JL, Stein GS. Nuclear microenvironments in biological control and cancer. Nat Rev Cancer. 2007 Jun;7(6):454-63.
  • Vradii D, Zaidi SK, Lian JB, van Wijnen AJ, Stein JL, Stein GS. Point mutation in AML1 disrupts subnuclear targeting, prevents myeloid differentiation, and effects a transformation-like phenotype. Proc Natl Acad Sci U S A. 2005 May 17;102(20):7174-9.
  • Javed A, Barnes GL, Pratap J, Antkowiak T, Gerstenfeld LC, van Wijnen AJ, Stein JL, Lian JB, Stein GS. Impaired intranuclear trafficking of Runx2 (AML3/CBFA1) transcription factors in breast cancer cells inhibits osteolysis in vivo. Proc Natl Acad Sci U S A. 2005 Feb 1;102(5):1454-9.
  • Young DW, Zaidi SK, Furcinitti PS, Javed A, van Wijnen AJ, Stein JL, Lian JB, Stein GS. Quantitative signature for architectural organization of regulatory factors using intranuclear informatics. J Cell Sci. 2004 Oct 1;117(Pt 21):4889-96.
  • McNeil S, Zeng C, Harrington KS, Hiebert S, Lian JB, Stein JL, van Wijnen AJ, Stein GS. The t(8;21) chromosomal translocation in acute myelogenous leukemia modifies intranuclear targeting of the AML1/CBFalpha2 transcription factor. Proc Natl Acad Sci U S A. 1999 Dec 21;96(26):14882-7.
     

 

Cell Cycle, Cancer and Aging

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Cell division requires faithful duplication of the genome as chromatin, and subsequent partitioning of chromosomes into progeny cells at mitosis. Cell cycle progression is stringently regulated and mechanisms that ensure the fidelity of this process are often deregulated in cancer. Normal cells are growth factor dependent and commit to initiate DNA replication at the G1/S phase transition. This growth-factor dependent control of the cell cycle is abrogated in cancer cells. Our studies have established that pluripotent stem cells have an abbreviated cell cycle due to a reduced G1 phase.

All dividing cells must package newly replicated DNA into chromatin during S phase, which necessitates the coordinated expression of multiple histone genes that encode the core nucleosomal proteins. Because transcription of histone genes is tightly coupled with the onset of DNA replication at the G1/S phase transition, we have pioneered the use of histone H4 genes as a paradigm for cell cycle control of transcription. These studies have resulted in the definition of a new cell cycle regulatory signaling pathway (Cyclin E-CDK2-NPAT-HiNFP axis). This pathway converges on subnuclear structures referred to as Histone Locus Bodies that contain histone gene clusters and the regulatory machinery for transcribing and processing of histone mRNAs (see Nuclear Structure and Function). We are examining cell cycle control in vivo during the earliest stages of embryonic development from the zygote to the blastocyst using conditional null mouse models lacking critical gene regulators. Recent findings from our laboratory have identified modifications in the higher-order organization of histone genes during breast cancer initiation and progression. In utilizing chromosome conformation capture and multispectral imaging strategies we are mechanistically characterizing modifications in genome structure and function that are functionally associate with cancer-compromised cell cycle control and regulation of proliferation. 

Beyond defining basic mechanisms of cell cycle control in cancer cells and pluripotent embryonic stem cells, we are also addressing broader physiological processes linked to cancer and regenerative medicine. For example, tumor progression and cancer metastasis are examined in immune-deficient mouse models and organoid cultures (see Musculoskeletal Biology and Pathology). Moreover, we are examining mechanisms of stem cell self-renewal and expansion of lineage-committed cells as it relates to regenerative medicine and aging (see Stem Cells and Regenerative Medicine). Recent initiatives are advancing understanding of the distinction between ductal carcinoma in situ breast tumors that will remain indolent and those that will progress and metastasize.  

Cell Cycle, Cancer and Aging Papers

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Genetic and Epigenetic Regulation of Gene Expression

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While genetic information encoded by DNA provides a framework for gene expression, epigenetic regulation is equally critical for physiological responsiveness to extracellular cues. Our laboratory pioneered examining the chromatin structure of several key gene loci (e.g., histone H4, osteocalcin, Runx2) to understand how architectural and topological constraints on gene promoters and enhancers at distal regulatory sites in chromatin contribute to epigenetic control of gene expression. These studies have evolved into detailed molecular analyses of histone modifications (e.g., H4 acetylation, H3 methylation), SWI/SNF related chromatin remodeling and genome-wide transcription factor binding analyses using ChIP-on-chip and ChIP-Seq approaches (see Nuclear Structure and Function).

Beyond the conventional epigenetic mechanisms that include post-transcriptional modifications of histones, and methylation of CpG moieties in gene promoters, we have identified a unique epigenetic mechanism, where phenotypic transcription factors remain associated with target genes during mitosis. This association, termed architectural epigenetics (‘Bookmarking’) ensures that the progeny cells maintain the cell growth and proliferative potential, as well as remain committed to lineage of the parent cells. We are actively exploring mechanisms that ensure sustained architectural epigenetics during symmetrical division of committed cells and during asymmetrical mitosis of stem cells.

Another epigenetic mechanism that has become a focus of our research group is the involvement of non-coding RNAs, microRNAs, and long non-coding RNAs, in regulating osteoblast- and hematopoiesis-related gene expression. We have carried out genome-wide screens for the expression of miRs and long non-coding RNAs under various physiological conditions and have identified miR signatures that have the potential to predict a biological response or a pathological outcome. Our group is further dissecting roles of these non-coding RNA signatures in osteoblastogenesis (see Musculoskeletal Biology and Pathology) and hematopoeisis, as well as in human leukemia. We have identified a mitotically associated long non-coding RNA in triple-negative breast cancer cells designated MANCR that is required to stabilize the breast cancer-compromised genome. Based on our discovery that knock-out of MANCR in triple-negative breast cancer cells results in apoptosis/cell death we are investigating MANCR knock-out as a targeted strategy for triple-negative breast cancer in vivo utilizing a xenograft mouse mode. 

Our research group was among the first to investigate cancer-compromised epigenetic control of gene expression during the onset of progression of breast cancer. In a tumor progression breast cancer cell culture model we have leverage chromosome conformation capture approaches to identify cancer-compromised modifications in higher-order chromatin organization. We have directly confirmed breast cancer-compromised inter and intrachromosomal interaction by multispectral imaging. We are mechanistically defining epigenetic control of aberrant chromatin organization in breast cancer cells by genomic strategies that identify histone modifications and DNA methylation. Initial epigenetic responses to degron ablation of the RUNX1 tumor suppressor in mammary epithelial cells is providing the first direct indication of regulatory consequences that are functionally linked with breast cancer initiation. We are pursuing clinically relevant validation of our findings by multi-omic and spatial transcriptomic analysis of patient-derived breast tumors.

Genetic and Epigenetic Regulation of Gene Expression Papers

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  • Tye CE, Ghule PN, Gordon JAR, Kabala FS, Page NA, Falcone MM, Tracy KM, van Wijnen AJ, Stein JL, Lian JB, Stein GS. LncMIR181A1HG is a novel chromatin-bound epigenetic suppressor of early stage osteogenic lineage commitment. Sci Rep. 2022 May 11;12(1):7770. doi: 10.1038/s41598-022-11814-4. PMID: 35546168; PMCID: PMC9095685.
  • Gordon JAR, Tye CE, Banerjee B, Ghule PN, van Wijnen AJ, Kabala FS, Page NA, Falcone MM, Stein JL, Stein GS, Lian JB. LINC01638 sustains human mesenchymal stem cell self-renewal and competency for osteogenic cell fate. Sci Rep. 2023 Nov 20;13(1):20314. doi: 10.1038/s41598-023-46202-z. PMID: 37985890; PMCID: PMC10662126.
  • Ghule PN, Boyd JR, Kabala F, Fritz AJ, Bouffard NA, Gao C, Bright K, Macfarlane J, Seward DJ, Pegoraro G, Misteli T, Lian JB, Frietze S, Stein JL, van Wijnen AJ, Stein GS. Spatiotemporal higher-order chromatin landscape of human histone gene clusters at histone locus bodies during the cell cycle in breast cancer progression. Gene. 2023 Jul 1;872:147441. doi: 10.1016/j.gene.2023.147441. Epub 2023 Apr 23. Erratum in: Gene. 2023 May 11;873:147469. PMID: 37094694; PMCID:
  • Fritz AJ, Ghule PN, Toor R, Dillac L, Perelman J, Boyd J, Lian JB, Gordon JAR, Frietze S, Van Wijnen A, Stein JL, Stein GS. Spatiotemporal Epigenetic Control of the Histone Gene Chromatin Landscape during the Cell Cycle. Crit Rev Eukaryot Gene Expr. 2023;33(3):85-97. doi: 10.1615/CritRevEukaryotGeneExpr.2022046190. PMID: 37017672; PMCID: PMC10826887.
  • Messier TL, Boyd JR, Gordon JAR, Tye CE, Page NA, Toor RH, Zaidi SK, Komm BS, Frietze S, Stein JL, Lian JB, Stein GS. Epigenetic and transcriptome responsiveness to ER modulation by tissue selective estrogen complexes in breast epithelial and breast cancer cells. PLoS One. 2022 Jul 21;17(7):e0271725. doi: 10.1371/journal.pone.0271725. PMID: 35862394; PMCID: PMC9302754. PMC10370284.
  • Hong D, Fritz AJ, Zaidi SK, van Wijnen AJ, Nickerson JA, Imbalzano AN, Lian JB, Stein JL, Stein GS. Epithelial-to-mesenchymal transition and cancer stem cells contribute to breast cancer heterogeneity. J Cell Physiol. 2018 Dec;233(12):9136-9144. doi: 10.1002/jcp.26847. Epub 2018 Jul 3. PMID: 29968906; PMCID: PMC6185773.
  • Zaidi SK, Young DW, Montecino MA, Lian JB, van Wijnen AJ, Stein JL, Stein GS. Mitotic bookmarking of genes: a novel dimension to epigenetic control.Nat Rev Genet. 2010 Aug;11(8):583-9.
  • Gordon JA, Hassan MQ, Saini S, Montecino M, van Wijnen AJ, Stein GS, Stein JL, Lian JB. Pbx1 represses osteoblastogenesis by blocking Hoxa10-mediated recruitment of chromatin remodeling factors. Mol Cell Biol. 2010 Jul;30(14):3531-41.
  • Ali SA, Zaidi SK, Dacwag CS, Salma N, Young DW, Shakoori AR, Montecino MA, Lian JB, van Wijnen AJ, Imbalzano AN, Stein GS, Stein JL. Phenotypic transcription factors epigenetically mediate cell growth control. Proc Natl Acad Sci U S A. 2008 May 6;105(18):6632-7.
  • Gutierrez J, Paredes R, Cruzat F, Hill DA, van Wijnen AJ, Lian JB, Stein GS, Stein JL, Imbalzano AN, Montecino M. Chromatin remodeling by SWI/SNF results in nucleosome mobilization to preferential positions in the rat osteocalcin gene promoter. J Biol Chem. 2007 Mar 30;282(13):9445-57.
  • Young DW, Hassan MQ, Yang XQ, Galindo M, Javed A, Zaidi SK, Furcinitti P, Lapointe D, Montecino M, Lian JB, Stein JL, van Wijnen AJ, Stein GS. Mitotic retention of gene expression patterns by the cell fate-determining transcription factor Runx2. Proc Natl Acad Sci U S A. 2007 Feb 27;104(9):3189-94. Epub 2007 Feb 20.
  • Young DW, Hassan MQ, Pratap J, Galindo M, Zaidi SK, Lee SH, Yang X, Xie R, Javed A, Underwood JM, Furcinitti P, Imbalzano AN, Penman S, Nickerson JA, Montecino MA, Lian JB, Stein JL, van Wijnen AJ, Stein GS. Mitotic occupancy and lineage-specific transcriptional control of rRNA genes by Runx2. Nature. 2007 Jan 25;445(7126):442-6.
  • Zaidi SK, Young DW, Pockwinse SM, Javed A, Lian JB, Stein JL, van Wijnen AJ, Stein GS. Mitotic partitioning and selective reorganization of tissue-specific transcription factors in progeny cells. Proc Natl Acad Sci U S A. 2003 Dec 9;100(25):14852-7.
  • Javed A, Gutierrez S, Montecino M, van Wijnen AJ, Stein JL, Stein GS, Lian JB. Multiple Cbfa/AML sites in the rat osteocalcin promoter are required for basal and vitamin D-responsive transcription and contribute to chromatin organization. Mol Cell Biol. 1999 Nov;19(11):7491-500.
  • Montecino M, Frenkel B, van Wijnen AJ, Lian JB, Stein GS, Stein JL. Chromatin hyperacetylation abrogates vitamin D-mediated transcriptional upregulation of the tissue-specific osteocalcin gene in vivo. Biochemistry. 1999 Jan 26;38(4):1338-45.

Musculoskeletal Biology and Pathology

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A main focus of our research for several decades has been exploration of molecular mechanisms that regulate skeletal development and remodeling. These studies center on commitment of mesenchymal stem cells to the osteoblast lineage, and the subsequent growth and differentiation of pre-committed osteoblasts. Our lab has identified several unique parameters of transcriptional control that regulate phenotype-restricted gene expression in the context of nuclear architecture (see Nuclear Structure and Function). We identified nuclear matrix protein 2 (NMP2) as an osteoblast-restricted, nuclear matrix-associated, transcription factor. NMP2, identified by other labs as AML3 or Cbfa1 (and later designated Runx2), is essential for bone formation in vivo and is mutated in Cleidocranial Dysplasia, an autosomal bone disorder.

We have since continued our quest to identify biological roles of Runx2 in osteoblast proliferation and differentiation and have demonstrated its central place in balancing growth with differentiation in osteoblasts. Using genomics, we have discovered many target genes for Runx2, and through proteomic approaches, we have identified several co-regulatory proteins that contribute to Runx2-mediated gene expression. Of direct clinical relevance is our discovery that Runx2 is highly expressed in breast and prostate cancer cells metastasizing to bone. Exploration of these new dimensions to Runx-mediated control of gene expression represents a major focus of our research group.

We have also established the essential role of non-coding RNAs in the bone-lineage using a conditional Dicer null mouse model which has a high bone mass phenotype. We have examined the biological function of microRNAs at multiple stages during differentiation in various mesenchymal lineages (e.g., osteoblasts, chondrocytes and myoblasts), as well as identified a network of microRNAs that controls the osteoblast-related master regulator Runx2 (see Genetic and Epigenetic Regulation of Gene Expression). We are investigating the contribution of long non-coding RNAs to epigenetic control of osteoblast phenotype commitment. Genomic targets are being identified by ChIP-Seq. Functional consequences of conditional activation and suppression of long non-coding RNAs are being explored by CRISPR and antisense approaches. 

Musculoskeletal Biology and Pathology Papers

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Stem Cells and Regenerative Medicine

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Human embryonic stem cells (hESCs) have a tremendous potential to be used for both therapeutic applications as well as for understanding basic biological processes. The potential to differentiate into any cell lineage and potential for unlimited cell division are unique characteristics of hESCs. We have discovered that this unlimited proliferative potential is linked to an abbreviated cell cycle; the embryonic stem cells have a shortened G1 phase of the cell cycle. This finding provides a paradigm to further explore cell growth regulatory mechanisms in the physiological context of pluripotency (see Cell Cycle, Cancer and Aging). We are examining mechanisms that mediate lengthening of the cell cycle and early stage of cell fate determination as pluripotent stem cells undergo differentiation.

In addition, we are examining osteogenic differentiation from mesenchymal stem cells to define the molecular basis of skeletal regenerative medicine (see Musculoskeletal Biology and Pathology).

Stem Cells and Regenerative Medicine Papers

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Cell Signaling and Regulatory Networks

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Osteoblasts and hematopoietic cells respond to a variety of extracellular signals. Fidelity of signaling responses is essential for biological outcomes in both lineages. Our laboratory has identified many cell signaling pathways that culminate at Runx target gene promoters, enhancers, and upstream regulatory elements. These include TGFβ/BMP2, Wnt, and Src tyrosine kinase signaling pathways in osteoblasts and the MAPK pathway in hematopoiesis that support protein/protein interactions of Runx proteins with critical co-factors. We discovered that these pathways are integrated by Runx proteins at target gene loci associated with distinct nuclear microenvironments (see Nuclear Structure and Function). Integration of signaling pathways at subnuclear sites of transcription provides a mechanistic model for regulatory communication from the extracellular milieu via the cell surface to the nucleus where gene regulation occurs. Using CRISPR-modified human cells in a xenograft model, we are now delineating the in vivo roles of these pathways linked to higher-order chromatin organization and the subnuclear functions of Runx proteins during the epithelial to mesenchymal transition associated with breast cancer tumorigenesis as well as for bone cell development and fracture repair.

Cell Signaling and Regulatory Networks Papers

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  • Fritz AJ, Ghule PN, Toor R, Dillac L, Perelman J, Boyd J, Lian JB, Gordon JAR, Frietze S, Van Wijnen A, Stein JL, Stein GS. Spatiotemporal Epigenetic Control of the Histone Gene Chromatin Landscape during the Cell Cycle. Crit Rev Eukaryot Gene Expr. 2023;33(3):85-97. doi: 10.1615/CritRevEukaryotGeneExpr.2022046190. PMID: 37017672; PMCID: PMC10826887.
  • Fritz AJ, El Dika M, Toor RH, Rodriguez PD, Foley SJ, Ullah R, Nie D, Banerjee B, Lohese D, Tracy KM, Glass KC, Frietze S, Ghule PN, Heath JL, Imbalzano AN, van Wijnen A, Gordon J, Lian JB, Stein JL, Stein GS. Epigenetic-Mediated Regulation of Gene Expression for Biological Control and Cancer: Cell and Tissue Structure, Function, and Phenotype. Results Probl Cell Differ. 2022;70:339-373. doi: 10.1007/978-3-031-06573-6_12. PMID: 36348114; PMCID: PMC9753575.
  • Fritz AJ, Hong D, Boyd J, Kost J, Finstaad KH, Fitzgerald MP, Hanna S, Abuarqoub AH, Malik M, Bushweller J, Tye C, Ghule P, Gordon J, Frietze S, Zaidi SK, Lian JB, Stein JL, Stein GS. RUNX1 and RUNX2 transcription factors function in opposing roles to regulate breast cancer stem cells. J Cell Physiol. 2020 Oct;235(10):7261-7272. doi: 10.1002/jcp.29625. Epub 2020 Mar 17. PMID: 32180230; PMCID: PMC7415511.
  • Fritz AJ, Gillis NE, Gerrard DL, Rodriguez PD, Hong D, Rose JT, Ghule PN, Bolf EL, Gordon JA, Tye CE, Boyd JR, Tracy KM, Nickerson JA, van Wijnen AJ, Imbalzano AN, Heath JL, Frietze SE, Zaidi SK, Carr FE, Lian JB, Stein JL, Stein GS. Higher order genomic organization and epigenetic control maintain cellular identity and prevent breast cancer. Genes Chromosomes Cancer. 2019 Jul;58(7):484-499. doi: 10.1002/gcc.22731. Epub 2019 Mar 15. PMID: 30873710; PMCID: PMC6549233.
  • Hong D, Fritz AJ, Zaidi SK, van Wijnen AJ, Nickerson JA, Imbalzano AN, Lian JB, Stein JL, Stein GS. Epithelial-to-mesenchymal transition and cancer stem cells contribute to breast cancer heterogeneity. J Cell Physiol. 2018 Dec;233(12):9136-9144. doi: 10.1002/jcp.26847. Epub 2018 Jul 3. PMID: 29968906; PMCID: PMC6185773.
  • Hong D, Fritz AJ, Finstad KH, Fitzgerald MP, Weinheimer A, Viens AL, Ramsey J, Stein JL, Lian JB, Stein GS. Suppression of Breast Cancer Stem Cells and Tumor Growth by the RUNX1 Transcription Factor. Mol Cancer Res. 2018 Dec;16(12):1952-1964. doi: 10.1158/1541-7786.MCR-18-0135. Epub 2018 Aug 6. PMID: 30082484; PMCID: PMC6289193.
  • Hong D, Fritz AJ, Gordon JA, Tye CE, Boyd JR, Tracy KM, Frietze SE, Carr FE, Nickerson JA, Van Wijnen AJ, Imbalzano AN, Zaidi SK, Lian JB, Stein JL, Stein GS. RUNX1-dependent mechanisms in biological control and dysregulation in cancer. J Cell Physiol. 2019 Jun;234(6):8597-8609. doi: 10.1002/jcp.27841. Epub 2018 Dec 4. PMID: 30515788; PMCID: PMC6395522.
  • Teplyuk NM, Galindo M, Teplyuk VI, Pratap J, Young DW, Lapointe D, Javed A, Stein JL, Lian JB, Stein GS, van Wijnen AJ. Runx2 regulates G protein-coupled signaling pathways to control growth of osteoblast progenitors. J Biol Chem. 2008 Oct 10;283(41):27585-97.
  • Pratap J, Wixted JJ, Gaur T, Zaidi SK, Dobson J, Gokul KD, Hussain S, van Wijnen AJ, Stein JL, Stein GS, Lian JB. Runx2 transcriptional activation of Indian Hedgehog and a downstream bone metastatic pathway in breast cancer cells. Cancer Res. 2008 Oct 1;68(19):7795-802.
  • Lengner CJ, Steinman HA, Gagnon J, Smith TW, Henderson JE, Kream BE, Stein GS, Lian JB, Jones SN. Osteoblast differentiation and skeletal development are regulated by Mdm2-p53 signaling. J Cell Biol. 2006 Mar 13;172(6):909-21.
  • Gaur T, Lengner CJ, Hovhannisyan H, Bhat RA, Bodine PV, Komm BS, Javed A, van Wijnen AJ, Stein JL, Stein GS, Lian JB. Canonical WNT signaling promotes osteogenesis by directly stimulating Runx2 gene expression. J Biol Chem. 2005 Sep 30;280(39):33132-40.
  • Zaidi SK, Sullivan AJ, Medina R, Ito Y, van Wijnen AJ, Stein JL, Lian JB, Stein GS. Tyrosine phosphorylation controls Runx2-mediated subnuclear targeting of YAP to repress transcription. EMBO J. 2004 Feb 25;23(4):790-9.
  • Zaidi SK, Sullivan AJ, van Wijnen AJ, Stein JL, Stein GS, Lian JB. Integration of Runx and Smad regulatory signals at transcriptionally active subnuclear sites. Proc Natl Acad Sci U S A. 2002 Jun 11;99(12):8048-53.
  • Aslam F, McCabe L, Frenkel B, van Wijnen AJ, Stein GS, Lian JB, Stein JL. AP-1 and vitamin D receptor (VDR) signaling pathways converge at the rat osteocalcin VDR element: requirement for the internal activating protein-1 site for vitamin D-mediated trans-activation. Endocrinology. 1999 Jan;140(1):63-70.