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Karl Willert

Title(s)Professor In Residence, Cellular and Molecular Medicine
SchoolHealth Sciences
Address9500 Gilman Drive #0695
La Jolla CA 92093
Phone858-822-3235
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    Collapse Biography 
    Collapse Education and Training
    University of California San Francisco, San FranciscoPhD06/1996Biochemistry
    University of California San Diego, La JollaB.A.06/1989Biochemistry

    Collapse Overview 
    Collapse Overview
    Dr. Karl Willert is an expert in stem cell and developmental biology. Dr. Willert obtained a Bachelor of Science in Biochemistry from the University of California, San Diego (1989) and earned a Ph.D. in Biochemistry (1996) from the University of California, San Francisco under the guidance of Dr. Harold Varmus. Subsequently, he worked as a post-doctoral fellow with Dr. Roeland Nusse at Stanford University in Palo Alto, California. In 2008, after working in the biotechnology sector for a few years, Dr. Willert was recruited as an Assistant Professor in the Department of Cellular & Molecular Medicine at the University of California, San Diego.

    Underlying Dr. Willert’s research interests has been the study of WNT proteins, a family of secreted growth factors that regulate embryonic development and tissue homeostasis and impact a large number of human diseases, including neurodegeneration and cancer. Throughout his scientific career, Dr. Willert has incorporated biochemical, genetic and cell biological approaches to study these proteins and their signaling cascades. In a seminal study, Dr. Willert purified WNT proteins (he holds the patent on the “Composition of Active WNT protein”) and demonstrated that they harbor potent stem cell growth factor activities. His current research focuses on how WNT proteins regulate self renewal and differentiation of human pluripotent stem cells. He is actively collaborating on multiple projects, including with colleagues Dr. Dennis Carson (WNT-FZD signaling in cancer), Dr. Terry Gaasterland (genome-wide approaches to understanding WNT signaling), Dr. David Traver (development of hematopoietic stem cells in zebrafish and human pluripotent stem cells), and Dr. Dan Kaufman (targeting FZD7-expressing tumors with CAR-NK cells).

    Dr. Willert has authored over 50 peer-reviewed publications, invited book chapters and review articles in the areas of WNT biochemistry and signaling and stem cell biology. Dr. Willert has been awarded 3 grants from the California Institute of Regenerative Medicine totaling $7.1 million. His research has also been supported by The National Institutes of Health (NIGMS, NHLBI, NIDDK) and the National Foundation for Ectodermal Dysplasias.

    Research Focus Areas:

    Developmental Biology | Signal Transduction | Stem Cell Biology

    Collapse Research 
    Collapse Research Activities and Funding
    Exploring Wnt-Fzd signaling specificities
    NIH/NIGMS R35GM134961Jan 1, 2020 - Dec 31, 2024
    Role: Principal Investigator
    2019 Wnt Signaling Gordon Research Conference
    NIH/NICHD R13HD097874Jan 10, 2019 - Dec 31, 2019
    Role: Principal Investigator
    Incucyte Zoom Live Cell Analysis System
    NIH S10OD025060Apr 3, 2018 - Apr 2, 2019
    Role: Principal Investigator
    Wnt signaling in hematopoietic development
    NIH/NHLBI R01HL135205Jan 1, 2017 - Dec 31, 2020
    Role: Principal Investigator
    Analyzing the role of Wnt signaling in pluripotency
    NIH/NIGMS R01GM110304Apr 10, 2014 - Feb 28, 2023
    Role: Principal Investigator
    Wnt Signaling in Development and Disease
    NIH/NCI R13CA128448Jun 1, 2007 - May 31, 2008
    Role: Principal Investigator

    Collapse ORNG Applications 
    Collapse Websites

    Collapse Bibliographic 
    Collapse Publications
    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Researchers can login to make corrections and additions, or contact us for help. to make corrections and additions.
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    Altmetrics Details PMC Citations indicate the number of times the publication was cited by articles in PubMed Central, and the Altmetric score represents citations in news articles and social media. (Note that publications are often cited in additional ways that are not shown here.) Fields are based on how the National Library of Medicine (NLM) classifies the publication's journal and might not represent the specific topic of the publication. Translation tags are based on the publication type and the MeSH terms NLM assigns to the publication. Some publications (especially newer ones and publications not in PubMed) might not yet be assigned Field or Translation tags.) Click a Field or Translation tag to filter the publications.
    1. EGFR is required for Wnt9a-Fzd9b signalling specificity in haematopoietic stem cells. Nat Cell Biol. 2019 06; 21(6):721-730. Grainger S, Nguyen N, Richter J, Setayesh J, Lonquich B, Oon CH, Wozniak JM, Barahona R, Kamei CN, Houston J, Carrillo-Terrazas M, Drummond IA, Gonzalez D, Willert K, Traver D. PMID: 31110287.
      View in: PubMed   Mentions: 1     Fields:    Translation:HumansAnimalsCells
    2. Mechanisms of Wnt signaling and control. Wiley Interdiscip Rev Syst Biol Med. 2018 Mar 30; e1422. Grainger S, Willert K. PMID: 29600540.
      View in: PubMed   Mentions: 5     Fields:    
    3. WNT9A Is a Conserved Regulator of Hematopoietic Stem and Progenitor Cell Development. Genes (Basel). 2018 Jan 29; 9(2). Richter J, Stanley EG, Ng ES, Elefanty AG, Traver D, Willert K. PMID: 29382179.
      View in: PubMed   Mentions:    Fields:    
    4. Wnt Signaling in Hematological Malignancies. Prog Mol Biol Transl Sci. 2018 01; 153:321-341. Grainger S, Traver D, Willert K. PMID: 29389522.
      View in: PubMed   Mentions: 3     Fields:    Translation:HumansAnimalsCells
    5. The WNT target SP5 negatively regulates WNT transcriptional programs in human pluripotent stem cells. Nat Commun. 2017 10 18; 8(1):1034. Huggins IJ, Bos T, Gaylord O, Jessen C, Lonquich B, Puranen A, Richter J, Rossdam C, Brafman D, Gaasterland T, Willert K. PMID: 29044119.
      View in: PubMed   Mentions: 4     Fields:    Translation:HumansCells
    6. Wnt/ß-catenin signaling during early vertebrate neural development. Dev Neurobiol. 2017 11; 77(11):1239-1259. Brafman D, Willert K. PMID: 28799266.
      View in: PubMed   Mentions: 5     Fields:    Translation:AnimalsCells
    7. Intercellular transmission of the unfolded protein response promotes survival and drug resistance in cancer cells. Sci Signal. 2017 Jun 06; 10(482). Rodvold JJ, Chiu KT, Hiramatsu N, Nussbacher JK, Galimberti V, Mahadevan NR, Willert K, Lin JH, Zanetti M. PMID: 28588081.
      View in: PubMed   Mentions: 14     Fields:    Translation:HumansAnimalsCells
    8. The role of Wnt signaling in hematopoietic stem cell development. Crit Rev Biochem Mol Biol. 2017 08; 52(4):414-424. Richter J, Traver D, Willert K. PMID: 28508727.
      View in: PubMed   Mentions: 4     Fields:    Translation:HumansAnimalsCells
    9. A Wnt5 Activity Asymmetry and Intercellular Signaling via PCP Proteins Polarize Node Cells for Left-Right Symmetry Breaking. Dev Cell. 2017 03 13; 40(5):439-452.e4. Minegishi K, Hashimoto M, Ajima R, Takaoka K, Shinohara K, Ikawa Y, Nishimura H, McMahon AP, Willert K, Okada Y, Sasaki H, Shi D, Fujimori T, Ohtsuka T, Igarashi Y, Yamaguchi TP, Shimono A, Shiratori H, Hamada H. PMID: 28292423.
      View in: PubMed   Mentions: 11     Fields:    Translation:AnimalsCells
    10. CRISPR Guide RNA Validation In Vitro. Zebrafish. 2017 08; 14(4):383-386. Grainger S, Lonquich B, Oon CH, Nguyen N, Willert K, Traver D. PMID: 27829120.
      View in: PubMed   Mentions: 1     Fields:    Translation:Animals
    11. Wnt9a Is Required for the Aortic Amplification of Nascent Hematopoietic Stem Cells. Cell Rep. 2016 11 01; 17(6):1595-1606. Grainger S, Richter J, Palazón RE, Pouget C, Lonquich B, Wirth S, Grassme KS, Herzog W, Swift MR, Weinstein BM, Traver D, Willert K. PMID: 27806298.
      View in: PubMed   Mentions: 8     Fields:    Translation:AnimalsCells
    12. Prodigiosin inhibits Wnt/ß-catenin signaling and exerts anticancer activity in breast cancer cells. Proc Natl Acad Sci U S A. 2016 11 15; 113(46):13150-13155. Wang Z, Li B, Zhou L, Yu S, Su Z, Song J, Sun Q, Sha O, Wang X, Jiang W, Willert K, Wei L, Carson DA, Lu D. PMID: 27799526.
      View in: PubMed   Mentions: 23     Fields:    Translation:HumansAnimalsCells
    13. Prenatal ß-catenin/Brn2/Tbr2 transcriptional cascade regulates adult social and stereotypic behaviors. Mol Psychiatry. 2016 10; 21(10):1417-33. Belinson H, Nakatani J, Babineau BA, Birnbaum RY, Ellegood J, Bershteyn M, McEvilly RJ, Long JM, Willert K, Klein OD, Ahituv N, Lerch JP, Rosenfeld MG, Wynshaw-Boris A. PMID: 26830142.
      View in: PubMed   Mentions: 16     Fields:    Translation:HumansAnimalsCells
    14. Methods to Manipulate and Monitor Wnt Signaling in Human Pluripotent Stem Cells. Methods Mol Biol. 2016; 1481:161-81. Huggins IJ, Brafman D, Willert K. PMID: 27590161.
      View in: PubMed   Mentions:    Fields:    Translation:HumansCells
    15. Generation of an expandable intermediate mesoderm restricted progenitor cell line from human pluripotent stem cells. Elife. 2015 Nov 10; 4. Kumar N, Richter J, Cutts J, Bush KT, Trujillo C, Nigam SK, Gaasterland T, Brafman D, Willert K. PMID: 26554899.
      View in: PubMed   Mentions: 7     Fields:    Translation:HumansCells
    16. Daple is a novel non-receptor GEF required for trimeric G protein activation in Wnt signaling. Elife. 2015 Jun 30; 4:e07091. Aznar N, Midde KK, Dunkel Y, Lopez-Sanchez I, Pavlova Y, Marivin A, Barbazán J, Murray F, Nitsche U, Janssen KP, Willert K, Goel A, Abal M, Garcia-Marcos M, Ghosh P. PMID: 26126266.
      View in: PubMed   Mentions: 35     Fields:    Translation:HumansCells
    17. A rare human syndrome provides genetic evidence that WNT signaling is required for reprogramming of fibroblasts to induced pluripotent stem cells. Cell Rep. 2014 Dec 11; 9(5):1770-1780. Ross J, Busch J, Mintz E, Ng D, Stanley A, Brafman D, Sutton VR, Van den Veyver I, Willert K. PMID: 25464842.
      View in: PubMed   Mentions: 9     Fields:    Translation:HumansCells
    18. Endogenous WNT signaling regulates hPSC-derived neural progenitor cell heterogeneity and specifies their regional identity. Stem Cell Reports. 2014 Dec 09; 3(6):1015-28. Moya N, Cutts J, Gaasterland T, Willert K, Brafman DA. PMID: 25458891.
      View in: PubMed   Mentions: 17     Fields:    Translation:HumansCells
    19. The WNT receptor FZD7 is required for maintenance of the pluripotent state in human embryonic stem cells. Proc Natl Acad Sci U S A. 2014 Jan 28; 111(4):1409-14. Fernandez A, Huggins IJ, Perna L, Brafman D, Lu D, Yao S, Gaasterland T, Carson DA, Willert K. PMID: 24474766.
      View in: PubMed   Mentions: 30     Fields:    Translation:HumansAnimalsCells
    20. WNT5A encodes two isoforms with distinct functions in cancers. PLoS One. 2013; 8(11):e80526. Bauer M, Bénard J, Gaasterland T, Willert K, Cappellen D. PMID: 24260410.
      View in: PubMed   Mentions: 11     Fields:    Translation:HumansCells
    21. Analysis of SOX2-expressing cell populations derived from human pluripotent stem cells. Stem Cell Reports. 2013; 1(5):464-78. Brafman DA, Moya N, Allen-Soltero S, Fellner T, Robinson M, McMillen ZL, Gaasterland T, Willert K. PMID: 24286033.
      View in: PubMed   Mentions: 9     Fields:    Translation:HumansAnimalsCells
    22. Use of a molecular genetic platform technology to produce human Wnt proteins reveals distinct local and distal signaling abilities. PLoS One. 2013; 8(3):e58395. Green JL, Bauer M, Yum KW, Li YC, Cox ML, Willert K, Wahl GM. PMID: 23516471.
      View in: PubMed   Mentions: 9     Fields:    Translation:AnimalsCells
    23. Regulation of endodermal differentiation of human embryonic stem cells through integrin-ECM interactions. Cell Death Differ. 2013 Mar; 20(3):369-81. Brafman DA, Phung C, Kumar N, Willert K. PMID: 23154389.
      View in: PubMed   Mentions: 33     Fields:    Translation:HumansCells
    24. Wnt proteins. Cold Spring Harb Perspect Biol. 2012 Sep 01; 4(9):a007864. Willert K, Nusse R. PMID: 22952392.
      View in: PubMed   Mentions: 109     Fields:    Translation:HumansAnimalsCells
    25. Arrayed cellular microenvironments for identifying culture and differentiation conditions for stem, primary and rare cell populations. Nat Protoc. 2012 Mar 15; 7(4):703-17. Brafman DA, Chien S, Willert K. PMID: 22422316.
      View in: PubMed   Mentions: 16     Fields:    Translation:HumansCells
    26. A protein kinase A and Wnt-dependent network regulating an intermediate stage in epithelial tubulogenesis during kidney development. Dev Biol. 2012 Apr 01; 364(1):11-21. Gallegos TF, Kouznetsova V, Kudlicka K, Sweeney DE, Bush KT, Willert K, Farquhar MG, Nigam SK. PMID: 22290330.
      View in: PubMed   Mentions: 13     Fields:    Translation:AnimalsCells
    27. Wnt signaling: the ß-cat(enin)'s meow. Genes Dev. 2012 Jan 15; 26(2):105-9. Bauer M, Willert K. PMID: 22279043.
      View in: PubMed   Mentions: 3     Fields:    Translation:Animals
    28. Secreted Wingless-interacting molecule (Swim) promotes long-range signaling by maintaining Wingless solubility. Proc Natl Acad Sci U S A. 2012 Jan 10; 109(2):370-7. Mulligan KA, Fuerer C, Ching W, Fish M, Willert K, Nusse R. PMID: 22203956.
      View in: PubMed   Mentions: 52     Fields:    Translation:AnimalsCells
    29. Long-term human pluripotent stem cell self-renewal on synthetic polymer surfaces. Biomaterials. 2010 Dec; 31(34):9135-44. Brafman DA, Chang CW, Fernandez A, Willert K, Varghese S, Chien S. PMID: 20817292.
      View in: PubMed   Mentions: 41     Fields:    Translation:HumansCells
    30. Myristoylated Naked2 antagonizes Wnt-beta-catenin activity by degrading Dishevelled-1 at the plasma membrane. J Biol Chem. 2010 Apr 30; 285(18):13561-8. Hu T, Li C, Cao Z, Van Raay TJ, Smith JG, Willert K, Solnica-Krezel L, Coffey RJ. PMID: 20177058.
      View in: PubMed   Mentions: 27     Fields:    Translation:HumansAnimalsCells
    31. Ethacrynic acid exhibits selective toxicity to chronic lymphocytic leukemia cells by inhibition of the Wnt/beta-catenin pathway. PLoS One. 2009 Dec 14; 4(12):e8294. Lu D, Liu JX, Endo T, Zhou H, Yao S, Willert K, Schmidt-Wolf IG, Kipps TJ, Carson DA. PMID: 20011538.
      View in: PubMed   Mentions: 32     Fields:    Translation:HumansCells
    32. Defining long-term maintenance conditions of human embryonic stem cells with arrayed cellular microenvironment technology. Stem Cells Dev. 2009 Oct; 18(8):1141-54. Brafman DA, Shah KD, Fellner T, Chien S, Willert K. PMID: 19327010.
      View in: PubMed   Mentions: 35     Fields:    Translation:HumansCells
    33. Impaired Wnt-beta-catenin signaling disrupts adult renal homeostasis and leads to cystic kidney ciliopathy. Nat Med. 2009 Sep; 15(9):1046-54. Lancaster MA, Louie CM, Silhavy JL, Sintasath L, Decambre M, Nigam SK, Willert K, Gleeson JG. PMID: 19718039.
      View in: PubMed   Mentions: 84     Fields:    Translation:AnimalsCells
    34. Investigating the role of the extracellular environment in modulating hepatic stellate cell biology with arrayed combinatorial microenvironments. Integr Biol (Camb). 2009 Sep; 1(8-9):513-24. Brafman DA, de Minicis S, Seki E, Shah KD, Teng D, Brenner D, Willert K, Chien S. PMID: 20023766.
      View in: PubMed   Mentions: 18     Fields:    Translation:AnimalsCells
    35. Efficient siRNA delivery into primary cells by a peptide transduction domain-dsRNA binding domain fusion protein. Nat Biotechnol. 2009 Jun; 27(6):567-71. Eguchi A, Meade BR, Chang YC, Fredrickson CT, Willert K, Puri N, Dowdy SF. PMID: 19448630.
      View in: PubMed   Mentions: 78     Fields:    Translation:HumansAnimalsCells
    36. Inhibition of GSK-3beta promotes survival and proliferation of megakaryocytic cells through a beta-catenin-independent pathway. Cell Signal. 2008 Dec; 20(12):2317-23. Soda M, Willert K, Kaushansky K, Geddis AE. PMID: 18804163.
      View in: PubMed   Mentions: 7     Fields:    Translation:HumansCells
    37. Isolation and application of bioactive Wnt proteins. Methods Mol Biol. 2008; 468:17-29. Willert KH. PMID: 19099243.
      View in: PubMed   Mentions: 21     Fields:    Translation:AnimalsCells
    38. Wnt signaling: is the party in the nucleus? Genes Dev. 2006 Jun 01; 20(11):1394-404. Willert K, Jones KA. PMID: 16751178.
      View in: PubMed   Mentions: 201     Fields:    Translation:AnimalsCells
    39. Differential inhibition of Wnt-3a by Sfrp-1, Sfrp-2, and Sfrp-3. Dev Dyn. 2006 Mar; 235(3):681-90. Galli LM, Barnes T, Cheng T, Acosta L, Anglade A, Willert K, Nusse R, Burrus LW. PMID: 16425220.
      View in: PubMed   Mentions: 44     Fields:    Translation:AnimalsCells
    40. Integration of Notch and Wnt signaling in hematopoietic stem cell maintenance. Nat Immunol. 2005 Mar; 6(3):314-22. Duncan AW, Rattis FM, DiMascio LN, Congdon KL, Pazianos G, Zhao C, Yoon K, Cook JM, Willert K, Gaiano N, Reya T. PMID: 15665828.
      View in: PubMed   Mentions: 268     Fields:    Translation:HumansAnimalsCells
    41. A proliferative role for Wnt-3a in chick somites. Dev Biol. 2004 May 15; 269(2):489-504. Galli LM, Willert K, Nusse R, Yablonka-Reuveni Z, Nohno T, Denetclaw W, Burrus LW. PMID: 15110715.
      View in: PubMed   Mentions: 17     Fields:    Translation:Animals
    42. Wnt proteins are lipid-modified and can act as stem cell growth factors. Nature. 2003 May 22; 423(6938):448-52. Willert K, Brown JD, Danenberg E, Duncan AW, Weissman IL, Reya T, Yates JR, Nusse R. PMID: 12717451.
      View in: PubMed   Mentions: 628     Fields:    Translation:AnimalsCells
    43. A role for Wnt signalling in self-renewal of haematopoietic stem cells. Nature. 2003 May 22; 423(6938):409-14. Reya T, Duncan AW, Ailles L, Domen J, Scherer DC, Willert K, Hintz L, Nusse R, Weissman IL. PMID: 12717450.
      View in: PubMed   Mentions: 598     Fields:    Translation:AnimalsCells
    44. The status of Wnt signalling regulates neural and epidermal fates in the chick embryo. Nature. 2001 May 17; 411(6835):325-30. Wilson SI, Rydström A, Trimborn T, Willert K, Nusse R, Jessell TM, Edlund T. PMID: 11357137.
      View in: PubMed   Mentions: 64     Fields:    Translation:AnimalsCells
    45. Wnt signaling regulates B lymphocyte proliferation through a LEF-1 dependent mechanism. Immunity. 2000 Jul; 13(1):15-24. Reya T, O'Riordan M, Okamura R, Devaney E, Willert K, Nusse R, Grosschedl R. PMID: 10933391.
      View in: PubMed   Mentions: 125     Fields:    Translation:AnimalsCells
    46. A Drosophila Axin homolog, Daxin, inhibits Wnt signaling. Development. 1999 Sep; 126(18):4165-73. Willert K, Logan CY, Arora A, Fish M, Nusse R. PMID: 10457025.
      View in: PubMed   Mentions: 40     Fields:    Translation:AnimalsCells
    47. Wnt-induced dephosphorylation of axin releases beta-catenin from the axin complex. Genes Dev. 1999 Jul 15; 13(14):1768-73. Willert K, Shibamoto S, Nusse R. PMID: 10421629.
      View in: PubMed   Mentions: 101     Fields:    Translation:Cells
    48. WNT signaling in the control of hair growth and structure. Dev Biol. 1999 Mar 01; 207(1):133-49. Millar SE, Willert K, Salinas PC, Roelink H, Nusse R, Sussman DJ, Barsh GS. PMID: 10049570.
      View in: PubMed   Mentions: 70     Fields:    Translation:AnimalsCells
    49. Two members of the Tcf family implicated in Wnt/beta-catenin signaling during embryogenesis in the mouse. Mol Cell Biol. 1998 Mar; 18(3):1248-56. Korinek V, Barker N, Willert K, Molenaar M, Roose J, Wagenaar G, Markman M, Lamers W, Destree O, Clevers H. PMID: 9488439.
      View in: PubMed   Mentions: 111     Fields:    Translation:HumansAnimalsCells
    50. Beta-catenin: a key mediator of Wnt signaling. Curr Opin Genet Dev. 1998 Feb; 8(1):95-102. Willert K, Nusse R. PMID: 9529612.
      View in: PubMed   Mentions: 158     Fields:    Translation:HumansAnimalsCells
    51. Casein kinase 2 associates with and phosphorylates dishevelled. EMBO J. 1997 Jun 02; 16(11):3089-96. Willert K, Brink M, Wodarz A, Varmus H, Nusse R. PMID: 9214626.
      View in: PubMed   Mentions: 80     Fields:    Translation:AnimalsCells
    52. Cell culture and whole animal approaches to understanding signaling by Wnt proteins in Drosophila. Cold Spring Harb Symp Quant Biol. 1997; 62:185-90. Nusse R, Samos CH, Brink M, Willert K, Cadigan KM, Wodarz A, Fish M, Rulifson E. PMID: 9598351.
      View in: PubMed   Mentions: 4     Fields:    Translation:HumansAnimalsCells
    53. The Wnt-1 proto-oncogene induces changes in morphology, gene expression, and growth factor responsiveness in PC12 cells. Neuron. 1993 Nov; 11(5):865-75. Shackleford GM, Willert K, Wang J, Varmus HE. PMID: 8240810.
      View in: PubMed   Mentions: 10     Fields:    Translation:AnimalsCells
    54. Efficient incorporation of human CD4 protein into avian leukosis virus particles. Science. 1990 Dec 07; 250(4986):1421-3. Young JA, Bates P, Willert K, Varmus HE. PMID: 2175047.
      View in: PubMed   Mentions: 42     Fields:    Translation:HumansAnimalsCells
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