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Papers & Reviews


Takeo, M., Toyoshima, K., Fujimoto, R., Iga, T., Takase, M., Ogawa, M., Tsuji, T. Cyclical dermal micro-niche switching governs the morphological infradian rhythm of mouse zigzag hair. Nat. Commun. 4;14(1):4478, 2023. [PubMed] [Full Text]

Raja E, Changarathil G, Oinam L, Tsunezumi J, Ngo YX, Ishii R, Sasaki T, Imanaka-Yoshida K, Yanagisawa H*, Sada A*: The extracellular matrix fibulin 7 maintains epidermal stem cell heterogeneity during skin aging. EMBO Rep. e55478, 2022. [PubMed] [Full Text]

Ngo YX, Haga K, Suzuki A, Kato H, Yanagisawa H, Izumi K, Sada A*: Isolation and culture of primary oral keratinocytes from the adult mouse palate. J Vis Exp, (175): e62820, 2021. [PubMed] [Full Text]


Kimura K, Ramirez K, Nguyen TAV, Yamashiro Y, Sada A and Yanagisawa H: Contribution of PDGFRα-positive cells in maintenance and injury responses in mouse large vessels. Sci Rep, 11(1):8683, 2021. [PubMed] [Full Text]

Ichijo R, Kabata M, Kidoya H, Muramatsu F, Ishibashi R, Abe K, Tsutsui K, Kubo H, Iizuka Y, Kitano S, Miyachi H, Kubota Y, Fujiwara H, Sada A, Yamamoto T, Toyoshima F: Vasculature-driven stem cell population coordinates tissue scaling in dynamic organs. Science Advances, 7(7): eabd2575, 2021.[PubMed] [Full Text]

Takeo, M., Asakawa, K., Toyoshima, K., Ogawa, M., Tong, JJ., Irié, T., Yanagisawa, M., Sato, A. and Tsuji, T: Expansion and characterization of epithelial stem cells with organ-inductive potential for long-term cyclical hair regeneration. Sci Rep, 11(1):1173, 2021. [PubMed] [Full Text]

Ishii R, Yanagisawa H*, Sada A*: Defining compartmentalized stem cell populations with distinct cell division dynamics in the ocular surface epithelium. 

*Corresponding authors. ​

Development, 147(24):dev197590, 2020. [PubMed] [Full Text]

Oinam L, Changarathil G, Raja E, Yen Ngo YX, Tateno H, Sada A*, Yanagisawa H: Glycome profiling by lectin microarray reveals dynamic glycan alterations during epidermal stem cell aging. 

* Corresponding author

Aging Cell, e13190, 2020. [PubMed] [Full Text] [EurekAlert! AAAS]

Kang S, Long K, Wang S, Sada A, and Tumbar T: Histone H3 K4/9/27 trimethylation levels affect wound healing and stem cell dynamics in adult skin. 

Stem Cell Reports 14(1): 34-48, 2020 [PubMed] [Full Text]


Changarathil G, Ramirez K, Isoda H, Sada A*, Yanagisawa H: Wild-type and SAMP8 mice show age-dependent changes in distinct stem cell compartments of the interfollicular epidermis. 

* Corresponding author

PLoS One 14(5): e0215908​, 2019 [PubMed] [Full Text] [Summary]

Oinam L, Changarathil G, Ngo YX, Yanagisawa H, Sada A*: Epidermal stem cell lineages. 

* Corresponding author

Advances in Stem Cells and their Niches, 2019 (A book chapter) [Full Text]

Tsunezumi J, Sugiura H, Oinam L, Ali A, Thang BQ, Sada A, Yamashiro Y, Kuro-O M, and Yanagisawa H: Fibulin-7, a heparin binding matricellular protein, promotes renal tubular calcification in mice.

Matrix Biol 74: 5-20, 2018 [PubMed] [Full Text]

Sada A, Jain P, Wang S, Leung E, Tumbar T: Slc1a3-CreER as a targeting tool for the K6+ epithelial stem cell niche and its precursors during mouse hair follicle cycle.

J Invest Dermatol 137(7): 1569-1571, 2017 [PubMed] [Full Text] [Commentary]

Sada A, Jacob F, Leung E, Wang S, White BS, Shalloway D and Tumbar T: Defining the cellular lineage hierarchy in the interfollicular epidermis of adult skin.

Nat Cell Biol 18: 619-631, 2016. Selected for F1000 prime. [PubMed] [Full Text] [News@Weill Cornell]

Takeo, M., Lee, W., Rabbani, R., Sun, Q., Hu, H., Lim, CH, Manga, P. and Ito, M. EdnrB governs regenerative response of melanocyte stem cells by crosstalk with Wnt signaling. Cell Reports 15(6):1291-302. 2016.

Takeo, M., Hale, C.S, and Ito, M. Epithelium-derived Wnt ligands are essential for maintenance of underlying digit bone. J Invest Dermatol. 136(7):1355-63. 2016.

Takeo, M., Chou, W.C., Sun, Q., Lee, W., Rabbani, P., Loomis, C., M. Taketo, M.M, and Ito. M. Wnt activation in nail epithelium couples nail growth to digit regeneration. Nature. 499:228-32. 2013

Rabbani, P.*, Takeo, M.*, Chou., W.C., Myung, P., Taketo, M.M., Chin, L., Bosenberg, M., and Ito, M. Ito.  Coordinated activation of Wnt signaling in epithelial and melanocyte stem cells initiates pigmented hair follicle regeneration. Cell. 145:941-55. 2011. (*co-first author)

Zhou Z, Shirakawa T, Ohbo K, Sada A, Wu Q, Hasegawa K, Saba R and Saga Y: RNA binding protein Nanos2 organizes post-transcriptional buffering system to retain primitive state of mouse spermatogonial stem cells. 

Dev Cell 34(1): 96-107, 2015 [PubMed] [Full Text]

Lee SE, Sada A, Zhang M, McDermitt DJ, Lu SY, Kemphues KJ and Tumbar T: High Runx1 levels promote a reversible, more-differentiated cell state in hair-follicle stem cells during quiescence. 

Cell Rep 6(3): 499-513, 2014 [PubMed] [Full Text]

Sada A and Tumbar T: New insights into mechanisms of stem cell daughter fate determination in regenerative tissues.

Int Rev Cell Mol Biol 300: 1-50, 2013 (A book chapter) [PubMed] [Full Text]

Sada A, Hasegawa K, Pin PH and Saga Y: NANOS2 acts downstream of glial cell line-derived neurotrophic factor signaling to suppress differentiation of spermatogonial stem cells. 

Stem Cells 30(2): 280-291, 2012 [PubMed] [Full Text]

Suzuki H, Saba R, Sada A and Saga Y: The Nanos3-3'UTR is required for germ cell specific NANOS3 expression in mouse embryos. 

PLoS One 5(2): e9300, 2010 [PubMed] [Full Text]

Suzuki H, Sada A, Yoshida S and Saga Y: The heterogeneity of spermatogonia is revealed by their topology and expression of marker proteins including the germ cell-specific proteins Nanos2 and Nanos3. 

Dev Biol 336(2): 222-231, 2009 [PubMed] [Full Text]

Sada A, Suzuki A, Suzuki H and Saga Y: The RNA-binding protein NANOS2 is required to maintain murine spermatogonial stem cells. 

Science 325(5946): 1394-1398, 2009 [PubMed] [Full Text]

Yamasaki H, Sada A, Iwata T, Niwa T, Tomizawa M, Xanthopoulos KG, Koike T and Shiojiri N: Suppression of C/EBPalpha expression in periportal hepatoblasts may stimulate biliary cell differentiation through increased Hnf6 and Hnf1b expression. 

Development 133(21): 4233-4243, 2006 [PubMed] [Full Text]

INVITED TALKS (international meeting) 

Aiko Sada: Stem cell dynamics underlying skin regeneration and resilience, 2023 ISRB Inaugural meeting, Vienna, September 3-6, 2023


Aiko Sada: Stem cell division dynamics underlying skin regeneration and aging, International Symposium on Skin Stem Cell Dynamics, Tokyo, May 14, 2023


Aiko Sada: Unraveling the mechanisms of skin regeneration and aging: from the perspective of stem cell proliferative heterogeneity, 3rd SFBD-JSDB conference, Strasbourg, November 7-10, 2022

Aiko Sada: Elucidating the cellular and molecular mechanisms of epidermal stem cell aging. ISSCR/JSRM 2021 Tokyo International Symposium, Online, October 27-29, 2021


Sada A: Skin stem cells shuffle sugars as they age: Glycome profiling reveals dynamic glycan alterations during epidermal stem cell aging, Singapore International Skin Conference 2021, Online, February 24-26, 2021

Sada A: Maintenance of heterogeneous epidermal stem cell populations by the distinct niche, Japan-Singapore International Skin Conference 2019, Singapore, April 10-12, 2019

Sada A: Defining the stem cell lineages in adult skin epidermis: CDB Symposium, Kobe, Japan, March 26-28, 2018

Sada A: Stem Cell Lineages of the Interfollicular Epidermis. Gordon Research Conference, Tissue Niches & Resident Stem Cells in Adult Epithelia, Hong Kong, China, August 7-12, 2016



Japan Agency for Medical Research and Development (AMED)/AMED-PRIME (22gm6710003), October 2022 – March 2026

Title: Elucidating the stem cell aging process caused by glycosylation abnormalities

Japan Agency for Medical Research and Development (AMED) /The Program for Technological Innovation of Regenerative Medicine, June 2021 – March 2024

Title: Development and evaluation of bioengineered 3D skin equivalent

Grant-in-Aid for Scientific Research/Grant-in-Aid for Scientific Research (B) (20H03266), April 2020 – March 2024

Title: Compartmentalization of epithelial stem cells in skin development and homeostasis


Grant-in-Aid for Scientific Research/ Grant-in-Aid for Challenging Research (Exploratory) (20K21431), July 2020 – March 2022

Title: Elucidating the roles of cell surface glycans on epithelial stem cell aging


Grant-in-Aid for Scientific Research/Early-Career Scientists (22K15126), PI: Nguyen Thi Kim Nguyen, April 2022 – March 2025

Title: Elucidating the role of IL-1R2, a decoy receptor of IL-1, in inflammation-induced stem cell aging

Japan Agency for Medical Research and Development (AMED)/Interstellar Initiative Beyond,  August 2022 – March 2023

Title: Defining mechanobiological effects on epidermal stem cell aging using skin on a chip model

The Nakatomi Foundation, April 2022 – March 2023

Title: Mechanism of epidermal stem cell regulation by the anti-aging matrix fibulin-7


Koyanagi Foundation, April 2022 – March 2023

Title: Stem cell heterogeneity as a key mechanism of skin aging

The NOVARTIS Foundation (Japan) for the Promotion of Science, April 2022 – March 2023

Title: Mechanism of epidermal stem cell regulation by the anti-aging matrix fibulin-7

Japan Agency for Medical Research and Development (AMED) /AMED-PRIME (21bm0704067), October 2018 – March 2022

Title: Elucidating the cellular and molecular mechanisms of epithelial stem cell aging

Grant-in-Aid for Scientific Research/for Early-Career Scientists (18K14709), April 2018 – March 2020

Title: Oral epithelial stem cells: cellular dynamics and future use in regenerative therapy


Grant-in-Aid for Scientific Research/Research Activity Start-up (20K22659), PI: Erna Raja, September 2020 – March 2022

Title: Defining the mechanism of epidermal stem cell heterogeneity in skin aging

Grant-in-Aid for Scientific Research/on Innovative Areas “Stem Cell Aging and Disease” (17H05631), April 2017 – March 2019

Title: Stem cell aging in the skin epidermis: If stem cells divide more, do they age more?


Grant-in-Aid for Scientific Research/for Research Activity Start-up (16H06660), October 2016 – March 2017

Title: Epidermal stem cells: from homeostasis to aging


Nanotech CUPAL (Career-up Allicance) /Nanotech Research Professional (N.R.P.) course, April 2018 – March 2019

Title: Comprehensive glycome profiling of skin stem cells during aging

Japan Agency for Medical Research and Development (AMED) /Interstellar Initiative, September 2020 – March 2021

Title: Elucidating the Cellular and Molecular Mechanisms of Stem Cell Aging


Naito Foundation/Research grant, April 2021 – March 2024

Title: Contribution of distinct stem cell populations in age-related skin dysfunction

Astellas Foundation for Research on Metabolic Disorders/Research grant, October 2020 – September 2022

Title: Elucidating the roles of cell surface glycans on skin stem cell aging


The Senri Life Science Foundation/Research grant, April 2021 – March 2022

Title: Mechanisms of glycomeshift in epidermal stem cell aging


Uehara Memorial Foundation/Research grant, January 2021 – March 2022

Title: Elucidating the role of stem cell heterogeneity and bioengineering of 3D skin


Tokyo Biochemical Research Foundation/Research grant, April 2021 – March 2022

Title: Elucidating the roles of cell surface glycans on skin stem cell aging


The Mitsubishi Foundation/Research grant, October 2019 – March 2021

Title: Dynamic changes in protein glycosylation in skin regeneration and aging


The Sumitomo foundation/Research grant, October 2019 – November 2021

Title: Identification of lectin biomarkers to define dynamic glycan alterations during epidermal stem cell aging


The Ichiro Kanehara foundation/Research grant, October 2019 – September 2020, Aiko Sada, 700,000 yen

Title: Stem cell dynamics on the ocular surface epithelium

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