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Characterization of chromatin replication associated with stem cell fate decision

Planned Research

Project AreaMechanisms underlying replication of non-genomic codes that mediate plasticity and robustness for cellular inheritance
Project/Area Number 19H05746
Research Category

Grant-in-Aid for Scientific Research on Innovative Areas (Research in a proposed research area)

Allocation TypeSingle-year Grants
Review Section Biological Sciences
Research InstitutionThe University of Tokyo

Principal Investigator

Iwama Atsushi  東京大学, 医科学研究所, 教授 (70244126)

Co-Investigator(Kenkyū-buntansha) 北村 俊雄  東京大学, 大学院薬学系研究科(薬学部), 名誉教授 (20282527)
Project Period (FY) 2019-06-28 – 2024-03-31
Project Status Completed (Fiscal Year 2023)
Budget Amount *help
¥168,220,000 (Direct Cost: ¥129,400,000、Indirect Cost: ¥38,820,000)
Fiscal Year 2023: ¥33,670,000 (Direct Cost: ¥25,900,000、Indirect Cost: ¥7,770,000)
Fiscal Year 2022: ¥31,850,000 (Direct Cost: ¥24,500,000、Indirect Cost: ¥7,350,000)
Fiscal Year 2021: ¥30,030,000 (Direct Cost: ¥23,100,000、Indirect Cost: ¥6,930,000)
Fiscal Year 2020: ¥33,670,000 (Direct Cost: ¥25,900,000、Indirect Cost: ¥7,770,000)
Fiscal Year 2019: ¥39,000,000 (Direct Cost: ¥30,000,000、Indirect Cost: ¥9,000,000)
Keywords造血幹細胞 / 運命制御 / 自己複製 / エピジェネティクス / クロマチン複製 / 幹細胞 / 運命決定 / クロマチン高次構造 / 非ゲノム情報
Outline of Research at the Start

造血幹細胞とその分裂によって生じる2 つの娘細胞のシングルセル解析を行い、それらのクロマチン複製様式を明らかにする。シングルセルATAC-Seq によるオープンクロマチン領域解析、シングルセルHi-C 解析によるトポロジカル関連ドメイン解析、ならびにシングルセルRNA-seq を行い、造血幹細胞の分裂様式をクロマチン複製と遺伝子発現の観点から理解し、娘細胞のシングルセル骨髄移植によって規定される造血幹細胞の生物学的な自己複製・分化との対応を明らかにする。さらに、本領域研究において同定されるクロマチン複製制御因子群の生物学的意義を、造血幹細胞の自己複製における機能の検証を通して理解する。

Outline of Final Research Achievements

We succeeded to perform Hi-C and RNA-seq analysis using the same single cells. We applied this method to two daughter cells from hematopoietic cells. The obtained data suggested the combination of transcriptome and higher order chromatin structure data is powerful to define symmetric versus asymmetric stem cell division. Further analysis would clarify the epigenetic mechanism that determines the fate decision of hematopoietic stem cells: self-renewal versus differentiation.

Academic Significance and Societal Importance of the Research Achievements

造血幹細胞の分裂により生じる2つの娘細胞の解析において、Hi-CとRNA-seqデータを同一1細胞から同時取得が可能となった。RNA-seqと高次クロマチン構造データの統合を進めることにより、造血幹細胞のクロマチン複製様式とその自己複製・分化の運命制御機構が明らかになるものと期待される。また、骨髄系血液腫瘍において頻用されるDNAメチル化阻害剤の耐性に関わる分子として、ユビキチンE3リガーゼTOPORSを同定した。DNAメチル化阻害剤とTOPORSの機能抑制の併用が有望な新規治療戦略となりうることが示され、新規創薬につながることが期待される。

Report

(6 results)
  • 2023 Annual Research Report   Final Research Report ( PDF )
  • 2022 Annual Research Report
  • 2021 Annual Research Report
  • 2020 Annual Research Report
  • 2019 Annual Research Report
  • Research Products

    (50 results)

All 2024 2023 2022 2021 2020 2019 Other

All Int'l Joint Research (10 results) Journal Article (20 results) (of which Int'l Joint Research: 13 results,  Peer Reviewed: 20 results,  Open Access: 19 results) Presentation (15 results) (of which Int'l Joint Research: 7 results,  Invited: 15 results) Remarks (5 results)

  • [Int'l Joint Research] オックスフォード大学/ロンドンがん研究所(英国)

    • Related Report
      2023 Annual Research Report
  • [Int'l Joint Research] マルセイユがんセンター(フランス)

    • Related Report
      2023 Annual Research Report
  • [Int'l Joint Research] コロンビア大学(米国)

    • Related Report
      2023 Annual Research Report
  • [Int'l Joint Research] McGill University(カナダ)

    • Related Report
      2022 Annual Research Report
  • [Int'l Joint Research] Marseille University(フランス)

    • Related Report
      2022 Annual Research Report
  • [Int'l Joint Research] Oxford University(英国)

    • Related Report
      2022 Annual Research Report
  • [Int'l Joint Research] マルセイユがんセンター(フランス)

    • Related Report
      2021 Annual Research Report
  • [Int'l Joint Research] PTC Therapeutics/Memorial Sloan-Kettering Cancer Center(米国)

    • Related Report
      2020 Annual Research Report
  • [Int'l Joint Research] マルセイユがんセンタ-/Cleveland Clinic(フランス)

    • Related Report
      2020 Annual Research Report
  • [Int'l Joint Research] ルンド大学(スウェーデン)

    • Related Report
      2020 Annual Research Report
  • [Journal Article] Polycomb repressive complex 1.1 coordinates homeostatic and emergency myelopoiesis2023

    • Author(s)
      Nakajima-Takagi Yaeko、Oshima Motohiko、Takano Junichiro、Koide Shuhei、Itokawa Naoki、Uemura Shun、Yamashita Masayuki、Andoh Shohei、Aoyama Kazumasa、Isshiki Yusuke、Shinoda Daisuke、Saraya Atsunori、Arai Fumio、Yamaguchi Kiyoshi、Furukawa Yoichi、Koseki Haruhiko、Ikawa Tomokatsu、Iwama Atsushi
    • Journal Title

      eLife

      Volume: 12

    • DOI

      10.7554/elife.83004

    • Related Report
      2023 Annual Research Report
    • Peer Reviewed / Open Access
  • [Journal Article] UTX inactivation in germinal center B cells promotes the development of multiple myeloma with extramedullary disease2023

    • Author(s)
      Rizq Ola、Mimura Naoya、Oshima Motohiko、Momose Shuji、Takayama Naoya、Itokawa Naoki、Koide Shuhei、Shibamiya Asuka、Miyamoto-Nagai Yurie、Rizk Mohamed、Nakajima-Takagi Yaeko、Aoyama Kazumasa、Wang Changshan、Saraya Atsunori、Ito Ryoji、Seimiya Masanori、et al
    • Journal Title

      Leukemia

      Volume: 37 Issue: 9 Pages: 1895-1907

    • DOI

      10.1038/s41375-023-01928-7

    • Related Report
      2023 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] Exposure to microbial products followed by loss of Tet2 promotes myelodysplastic syndrome via remodeling HSCs2023

    • Author(s)
      Yokomizo-Nakano T, Hamashima A, Kubota S, Bai J, Sorin S, Sun Y, Kikuchi K, Iimori M, Morii M, Kanai A, Iwama A, Huang G, Kurotaki D, Takizawa H, Matsui H, Sashida G.
    • Journal Title

      Journal of Experimental Medicine (JEM)

      Volume: 220 (7) Issue: 7 Pages: 9999-9999

    • DOI

      10.1084/jem.20220962

    • Related Report
      2023 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] Mesenchymal loss of p53 alters stem cell capacity and models human soft tissue sarcoma traits.2023

    • Author(s)
      Sorimachi Y, Kobayashi H, Shiozawa Y, Koide S, Nakato R, Shimizu Y, Okamura T, Shirahige K, Iwama A, Goda N, Takubo K, Takubo K.
    • Journal Title

      Stem Cell Reports

      Volume: S2213-6711(23) Issue: 5 Pages: 00097-8

    • DOI

      10.1016/j.stemcr.2023.03.009

    • Related Report
      2023 Annual Research Report
    • Peer Reviewed / Open Access
  • [Journal Article] The pathogenetic significance of exhausted T cells in a mouse model of mature B cell neoplasms2023

    • Author(s)
      1.Shibamiya A, Miyamoto-Nagai Y, Koide S, Oshima M, Rizq O, Aoyama K, Nakajima-Takagi Y, Kato R, Kayamori K, Isshiki Y, Oshima-Hasegawa N, Muto T, Tsukamoto S, Takeda Y, Koyama-Nasu R, Chiba T, Honda H, Yokote K, Iwama A, Sakaida E, Mimura N.
    • Journal Title

      Cancer Immunol Immunother

      Volume: - Issue: 8 Pages: 2635-2648

    • DOI

      10.1007/s00262-023-03447-x

    • Related Report
      2023 Annual Research Report
    • Peer Reviewed / Open Access
  • [Journal Article] Epigenetic traits inscribed in chromatin accessibility in aged hematopoietic stem cells2022

    • Author(s)
      Itokawa Naoki、Oshima Motohiko、Koide Shuhei、Takayama Naoya、Kuribayashi Wakako、Nakajima-Takagi Yaeko、Aoyama Kazumasa、Yamazaki Satoshi、Yamaguchi Kiyoshi、Furukawa Yoichi、Eto Koji、Iwama Atsushi
    • Journal Title

      Nature Communications

      Volume: 13 Issue: 1 Pages: 2691-2691

    • DOI

      10.1038/s41467-022-30440-2

    • Related Report
      2022 Annual Research Report 2021 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] YAP1/TAZ activity maintains vascular integrity and organismal survival2022

    • Author(s)
      Uemura Shun、Yamashita Masayuki、Aoyama Kazumasa、Yokomizo-Nakano Takako、Oshima Motohiko、Nishio Miki、Masuko Masayoshi、Takizawa Jun、Sone Hirohito、Yamada Yasuhiro、Suzuki Akira、Iwama Atsushi
    • Journal Title

      Biochemical and Biophysical Research Communications

      Volume: 619 Pages: 117-123

    • DOI

      10.1016/j.bbrc.2022.06.050

    • Related Report
      2022 Annual Research Report
    • Peer Reviewed / Open Access
  • [Journal Article] A high prevalence of myeloid malignancies in progeria with Werner syndrome is associated with p53 insufficiency2022

    • Author(s)
      Kato Hisaya、Ogawa Seishi, et al.
    • Journal Title

      Experimental Hematology

      Volume: 109 Pages: 11-17

    • DOI

      10.1016/j.exphem.2022.02.005

    • Related Report
      2022 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] Unraveling unique features of plasma cell clones in POEMS syndrome with single-cell analysis.2022

    • Author(s)
      Isshiki Y, Oshima M, Mimura N, Kayamori K, Miyamoto-Nagai Y, Seki M, Nakajima-Takagi Y, Kanamori T, Iwamoto E, Muto T, Tsukamoto S, Takeda Y, Ohwada C, Misawa S, Ikeda JI, Sanada M, Kuwabara S, Suzuki Y, Sakaida E, Nakaseko C, Iwama A.
    • Journal Title

      JCI Insight

      Volume: 7 Issue: 20 Pages: 1-13

    • DOI

      10.1172/jci.insight.151482

    • Related Report
      2022 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] PCGF1-PRC1 links chromatin repression with DNA replication during hematopoietic cell lineage commitment2022

    • Author(s)
      Takano J, Ito S, Dong Y, Sharif J, Nakajima-Takagi Y, Umeyama T, Han YW, Isono K, Kondo T, Iizuka Y, Miyai T, Koseki Y, Ikegaya M, Sakihara M, Nakayama M, Ohara O, Hasegawa Y, Hashimoto K, Arner E, Klose RJ, Iwama A, Koseki H, Ikawa T
    • Journal Title

      Nat Commun.

      Volume: 13 (1) Issue: 1 Pages: 7159-7159

    • DOI

      10.1038/s41467-022-34856-8

    • Related Report
      2022 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] Insufficiency of non-canonical PRC1 synergizes with JAK2V617F in the development of myelofibrosis2021

    • Author(s)
      Shinoda Daisuke、Nakajima-Takagi Yaeko、Oshima Motohiko、Koide Shuhei、Aoyama Kazumasa、Saraya Atsunori、Harada Hironori、Rahmutulla Bahityar、Kaneda Atsushi、Yamaguchi Kiyoshi、Furukawa Yoichi、Koseki Haruhiko、Shimoda Kazuya、Tanaka Tomoaki、Sashida Goro、Iwama Atsushi
    • Journal Title

      Leukemia

      Volume: 36 Issue: 2 Pages: 452-463

    • DOI

      10.1038/s41375-021-01402-2

    • Related Report
      2021 Annual Research Report
    • Peer Reviewed / Open Access
  • [Journal Article] Limited rejuvenation of aged hematopoietic stem cells in young bone marrow niche2021

    • Author(s)
      Kuribayashi W, Oshima M, Itokawa N, Koide S, Nakajima-Takagi Y, Yamashita M, Yamazaki S, Rahmutulla B, Miura F, Ito T, Kaneda A, Iwama A.
    • Journal Title

      J Exp Med

      Volume: 218 Issue: 3

    • DOI

      10.1084/jem.20192283

    • Related Report
      2020 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] DHODH inhibition synergizes with DNA-demethylating agents in the treatment of myelodysplastic syndromes2021

    • Author(s)
      Kayamori Kensuke、Nagai Yurie、Zhong Cheng、Kuribayashi Wakako、Oshima Motohiko、Nakajima-Takagi Yaeko、Yamashita Masayuki、Mimura Naoya、Becker Hans Jiro、Izawa Kiyoko、Yamazaki Satoshi、Iwano Satoshi、Ito Ryoji、Lennox William、Sheedy Josephine、Yokote Koutaro、Iwama Atsushi
    • Journal Title

      Blood Advances

      Volume: 5 Issue: 2 Pages: 438-450

    • DOI

      10.1182/bloodadvances.2020001461

    • Related Report
      2020 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] Mutant ASXL1 induces age-related expansion of phenotypic hematopoietic stem cells through activation of Akt/mTOR pathway2021

    • Author(s)
      Fujino Takeshi、Goyama Susumu、Sugiura Yuki、Inoue Daichi、Asada Shuhei、Yamasaki Satoshi、Matsumoto Akiko、Yamaguchi Kiyoshi、Isobe Yumiko、Tsuchiya Akiho、Shikata Shiori、Sato Naru、Morinaga Hironobu、Fukuyama Tomofusa、Tanaka Yosuke、Fukushima Tsuyoshi、Takeda Reina、Yamamoto Keita、Honda Hiroaki et al
    • Journal Title

      Nature Communications

      Volume: 12 Issue: 1 Pages: 1826-1826

    • DOI

      10.1038/s41467-021-22053-y

    • Related Report
      2020 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] HHEX promotes myeloid transformation in cooperation with mutant ASXL1.2020

    • Author(s)
      Takeda R, Asada S, Park SJ, Yokoyama A, Becker H, Kanai A, Visconte V, Hershberger C, Hayashi Y, Yonezawa T, Tamura M, Fukushima T, Tanaka Y, Fukuyama T, Matsumoto A, Yamasaki S, Nakai K, Yamazaki S, Inaba T, Shibata T, Inoue D, Honda H, Goyama S, Maciejewski J, Kitamura T.
    • Journal Title

      Blood

      Volume: 136 Pages: 1670-1684

    • DOI

      10.1182/blood.2019004613

    • Related Report
      2020 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] Plant homeodomain finger protein 6 in the regulation of normal and malignant hematopoiesis.2020

    • Author(s)
      Miyagi S, Iwama A.
    • Journal Title

      Curr Opin Hematol.

      Volume: - Issue: 4 Pages: 248-253

    • DOI

      10.1097/moh.0000000000000588

    • Related Report
      2019 Annual Research Report
    • Peer Reviewed
  • [Journal Article] Bmi1 counteracts hematopoietic stem cell aging by repressing target genes and enforcing the stem cell gene signature.2020

    • Author(s)
      Nitta E, Itokawa N, Yabata S, Koide S, Hou LB, Oshima M, Aoyama K, Saraya A, Iwama A.
    • Journal Title

      Biochem Biophys Res Commun

      Volume: 521 Issue: 3 Pages: 612-619

    • DOI

      10.1016/j.bbrc.2019.10.153

    • Related Report
      2019 Annual Research Report
    • Peer Reviewed / Open Access
  • [Journal Article] Akt inhibition synergizes with polycomb repressive complex 2 inhibition in the treatment of multiple myeloma.2019

    • Author(s)
      Rizk M, Rizq O, Oshima M, Nakajima-Takagi Y, Koide S, Saraya A, Isshiki Y, Chiba T, Yamazaki S, Ma A, Jin J, Iwama A, Mimura N.
    • Journal Title

      Cancer Sci

      Volume: 110 Issue: 12 Pages: 3695-3707

    • DOI

      10.1111/cas.14207

    • Related Report
      2019 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] KDM2B in polycomb repressive complex 1.1 functions as a tumor suppressor in the initiation of T-cell leukemogenesis.2019

    • Author(s)
      Isshiki Y, Nakajima-Takagi Y, Oshima M, Aoyama K, Rizk M, Kurosawa S, Saraya A, Kondo T, Sakaida E, Nakaseko C, Yokote K, Koseki H, Iwama A.
    • Journal Title

      Blood Adv

      Volume: 3 Issue: 17 Pages: 2537-2549

    • DOI

      10.1182/bloodadvances.2018028522

    • Related Report
      2019 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Journal Article] The chromatin binding protein Phf6 restricts the self-renewal of hematopoietic stem cells.2019

    • Author(s)
      Miyagi S, Sroczynska P, Kato Y, Nakajima-Takagi Y, Oshima M, Rizq O, Takayama N, Saraya A, Mizuno S, Sugiyama F, Takahashi S, Matsuzaki Y, Christensen J, Helin K, and Iwama A.
    • Journal Title

      Blood

      Volume: - Issue: 23 Pages: 2495-2506

    • DOI

      10.1182/blood.2019000468

    • Related Report
      2019 Annual Research Report
    • Peer Reviewed / Open Access / Int'l Joint Research
  • [Presentation] Aging of hematopoietic stem cells2024

    • Author(s)
      Atsushi Iwama
    • Organizer
      International Symposium on “Signals for Human, Animal and Planetary Health: From Metabolites To Biological Interactions”
    • Related Report
      2023 Annual Research Report
    • Int'l Joint Research / Invited
  • [Presentation] Targeting Hippo pathway in bone marrow niche promotes hematopoietic regeneration.2024

    • Author(s)
      Atsushi Iwama
    • Organizer
      The 21st IRCMS Symposium on The Dynamics and Development of Normal and Malignant Hematopoietic Stem Cells.
    • Related Report
      2023 Annual Research Report
    • Int'l Joint Research / Invited
  • [Presentation] TOPORS ubiquitin ligase promotes degradation of DNMT1 in the presence of DNA hypomethylating agents2023

    • Author(s)
      Atsushi Iwama
    • Organizer
      Tsuruoka Conference “Role of epigenetic factors in cancer”
    • Related Report
      2023 Annual Research Report
    • Int'l Joint Research / Invited
  • [Presentation] Chromatin accessibility in stem cells unveils progressive transcriptional reprogramming in myelodysplastic syndrome2023

    • Author(s)
      Atsushi Iwama
    • Organizer
      Symposia ”Revisiting epigenetics in cancer” The 82th Annual Meeting of the Japanese Cancer Association
    • Related Report
      2023 Annual Research Report
    • Int'l Joint Research / Invited
  • [Presentation] TOPORS ubiquitin ligase promotes DNMT1 degradation upon DNA hypomethylating agents2023

    • Author(s)
      Atsushi Iwama
    • Organizer
      The 2nd International Symposium on REPLICATION of NON GENOME 2023
    • Related Report
      2023 Annual Research Report
    • Int'l Joint Research / Invited
  • [Presentation] Chromatin accessibility in MDS stem and progenitor cells. Symposium “Recent advances in leukemia stem cell research”2022

    • Author(s)
      Iwama A
    • Organizer
      The 84th Annual Meeting of the Japanese Society of Hematology
    • Related Report
      2022 Annual Research Report
    • Invited
  • [Presentation] 「造血幹細胞特性の加齢変化とエピジェネティックメモリー」シンポジウム「100歳を目指した健康寿命延伸医療の実現」2022

    • Author(s)
      岩間厚志
    • Organizer
      日本心血管作動部質学会
    • Related Report
      2022 Annual Research Report
    • Invited
  • [Presentation] Chromatin accessibility in MDS stem and progenitor cells. Hematopoiesis2021

    • Author(s)
      Atsushi Iwama
    • Organizer
      Keystone Symposia
    • Related Report
      2021 Annual Research Report
    • Int'l Joint Research / Invited
  • [Presentation] Non-canonical PRC1 regulates homeostatic and emergency hematopoiesis by restricting myeloid differentiation program2021

    • Author(s)
      Atsushi Iwama
    • Organizer
      The Hematologic Malignancies Conference, FASEB Meeting
    • Related Report
      2021 Annual Research Report
    • Int'l Joint Research / Invited
  • [Presentation] KDM6A inactivation in germinal center B cells promotes the development of plasma cell neoplasms2020

    • Author(s)
      Atsushi Iwama
    • Organizer
      日本がん学会総会
    • Related Report
      2020 Annual Research Report
    • Invited
  • [Presentation] 造血幹細胞のエピジェネティックプロファイリング2020

    • Author(s)
      岩間厚志
    • Organizer
      共同利用・共同研究拠点「マルチオミックスによる遺伝子発現制御の先端的医学共同研究拠点」シンポジウム
    • Related Report
      2019 Annual Research Report
    • Invited
  • [Presentation] Epigenetic alterations in aged hematopoietic stem cells and age-associated malignancies2019

    • Author(s)
      Iwama A
    • Organizer
      第13回日本エピジェネティクス研究会年会
    • Related Report
      2019 Annual Research Report
    • Invited
  • [Presentation] 造血幹細胞の加齢変化を規定するエピゲノム特性2019

    • Author(s)
      岩間厚志
    • Organizer
      第42回日本基礎老化学会大会
    • Related Report
      2019 Annual Research Report
    • Invited
  • [Presentation] Epigenetic alterations in aged hematopoietic stem cells and age-associated malignancies2019

    • Author(s)
      Iwama A
    • Organizer
      第62回日本放射線影響学会大会
    • Related Report
      2019 Annual Research Report
    • Invited
  • [Presentation] Epigenetic alterations in aged hematopoietic stem cells and age-associated malignancies2019

    • Author(s)
      岩間厚志
    • Organizer
      第42回日本分子生物学会年会
    • Related Report
      2019 Annual Research Report
    • Invited
  • [Remarks] https://www.ims.u-tokyo.ac.jp/molmed/

    • Related Report
      2023 Annual Research Report
  • [Remarks] 幹細胞分子医学分野ホームページ

    • URL

      https://www.ims.u-tokyo.ac.jp/molmed/

    • Related Report
      2022 Annual Research Report 2020 Annual Research Report
  • [Remarks] 幹細胞分子医学分野

    • URL

      https://www.ims.u-tokyo.ac.jp/molmed/

    • Related Report
      2021 Annual Research Report
  • [Remarks] 細胞療法分野ホームページ

    • URL

      https://www.ims.u-tokyo.ac.jp/clinical_oncol/index.html

    • Related Report
      2020 Annual Research Report
  • [Remarks] 幹細胞分子医学分野HP

    • URL

      https://www.ims.u-tokyo.ac.jp/molmed/

    • Related Report
      2019 Annual Research Report

URL: 

Published: 2019-07-04   Modified: 2025-01-30  

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