For a complete list of publications from NCBI, please click here.



Endothelin promotes colorectal tumorigenesis by activating YAP/TAZ.
Wang Z, Liu P, Zhou X, Wang T, Feng X, Sun YP, Xiong Y, Yuan HX, Guan KL.
Cancer Res. (2017). pii: canres.3229.2016

L2hgdh deficiency accumulates L-2-hydroxyglutarate with progressive leukoencephalopathy and neurodegeneration.
Ma S, Sun R, Jiang B, Gao J, Deng W, Liu P, He R, Cui J, Ji M, Yi W, Yang P, Wu X, Xiong Y, Qui Z, Ye D, Guan KL.
Mol Cell Biol. (2017). pii: MCB.00492-16

MTORC1-mediated NRBF2 phosphorylation functions as a switch for the class III PtdIns3K and autophagy.
Ma X, Zhang S, He L, Rong Y, Brier LW, Sun Q, Liu R, Fan W, Chen S, Yue Z, Kim J, Guan KL, Li D, Zhong Q.
Autophagy. (2017). 6:1-16

NLK mediates the osmotic stress signal to phosphorylate and activate YAP.
Hong AW, Meng Z, Yuan HX, Plouffe SW, Moon S, Kim W, Jho EH, Guan KL.
EMBO Rep. (2017). 18, 72-86

A YAP-IL6ST auto regulatory loop activated on APC loss controls colonic tumorigenesis.
Taniguchi K, Moroishi T, de Jong PR, Krawczyk, Grebbin BM, Luo H, Xu R, Glob-Schwarz N, Schweiger C, Wang K, Di Caro G, Feng Y, Fearon E, Raz E, Farin HF, Guan KL, Hayback J, Datz C, Zhang K, Karin M.
PNAS. (2017). 114, 1644-8



KDM2B targets c-Fos for ubiquitylation and degradation in response to mitogenic stimulation.
Han X, Zha Z, Yuan H, Feng X, Xio YK, Lei Q, Guan KL, Xiong Y.
Oncogene. (2016). 35, 4179-90

Hypertension-associated C825T polymorphism impairs the function of Gb3 to target GRK2 ubiquitination.
Zha Z, Han XR, Smith MD, Lei QY, Guan KL, Xiong Y.
Cell Discovery. (2016). 2, 16005

A new class of temporarily phenotypic enhancers identified by CRISPR/Cas9 mediated genetic screening.
Diao Y, Li B, Meng Z, Jung I, Lee AY, Dixon J. Maliskova L, Guan KL, Shen Y, Ren B.
Genome Res. (2016). 26, 397-405

SIRT5 promotes IDH2 desuccinylation and G6PD deglutarylation to enhance cellular antioxidant defense.
Zou L, Wang F, Sun R, Chen X, Zhang M, Xu Q, Wang Y, Wang S, Xiong Y, guan KL, Yang P, Yu H, Ye D.
EMBO Rep. (2016). 17, 811-22

MST1 shuts off cytosolic antiviral defense through IRF3 phosphorylation.
Meng F, Zhou R, Wu S, Zhang Q, Jin Q, Zhou Y, Plouffe SW, Liu S, Song H, Xia Z, Zhao B, Yu S, Feng XH, Guan KL, Zou J, Xu P.
Genes and Dev. (2016). 30, 1086-1100

Thromboxane A2 activates YAP/TAZ to induce vascular smooth cell proliferation and migration.
Feng X, Liu P, Zhou X, i MT, Li FL, Wang Z, Meng ZP, Sun YP, Yu Y, Xiong Y, Yuan HX, Guan KL.
J. Biol. Chem. (2016). 291, 18947-58

Flow-dependent YAP/TAZ activities regulate endothelial phenotypes and atherosclerosis.
Wang KC, Yeh YT, Nguyen P, Limqueco E, Lopez J, Thorossian S, Guan KL, Li YS, Chien S.
PNAS. (2016). 113, 11525-30

Acetylation destabilizes fatty acid synthase to inhibit de novo lipogenesis and tumor cell growth.
Lin HP, Cheng ZL, He RY, Song L, Tian MX, Zhou LS, Groh BS, Liu WR, i MB, Ding C, Shi YH, Guan KL, Yu D, Xiong Y.
Cancer Res. (2016). in press

Characterization of Hippo pathway components by gene inactivation.
Plouffe SW, Meng Z, Lin KC, Hong AW, Chun JV, Guan KL.
Mol. Cell. (2016). 64, 993-08

The Hippo pathway kinases LATS1/2 suppress cancer immunity.
Moroishi T, Hayashi T, Pan W, Fujita Y, Holt MV, Qin J, Carson DA, Guan KL.
Cell. (2016). 167, 1525-39

Mechanisms of the Hippo pathway regulation.
Meng Z, Moroishi T, Guan KL.
Genes and Dev. (2016). 30, 1-17

The Hippo pathway in intestinal regeneration and disease.
Hong AW, Meng Z, Guan KL.
Nat. Rev. Gastroenterol. and Hepatol. (2016). 13, 324-37

Structural insights of mTOR complex 1.
Yuan HX, Guan KL.
Cell Research. (2016). 26, 267-8

Guidelines for the use and interpretation of assays for monitoring autophagy.
Klionsky DJ, et al.
Autophagy. (2016). 12, 1-222

Glycoholics anonymous: Cancer sobers up with mTORC1
Fu V, Moroishi T, Guan KL.
Cancer Cell. (2016). 29, 432-434



2-Hydroxyglutate is essential for maintaining oncogenic property of mutant IDH-containing cancer cells but dispensable for cell growth.
Ma S, Jiang B, Deng W, Gu ZK, Wu FZ, Li T, Xia Y, Yang H, Ye D, Xiong Y, Guan KL.
Oncotarget. (2015). 6, 8606-20

Estrogen regulates Hippo signaling via GPER in breast cancer.
Zhou X, Wang S, Wang Z, Feng X, Liu P, Lv X, Li F, Yu FX, Sun Y, Yuan H, Zhu H, Xiong Y, Lei QY, Guan KL.
J. Clin. Invest. (2015). 125, 2123-35

A non-canonical function of Gb as a subunit of E3 ligase in targeting GRK2 ubiquitylation.
Zha Z, Han X, Smith MD, Liu Y, Giguere PM, Kopanja D, Raychaudhuri P, Siderovski DP, Guan KL, Lei QYY, Xiong Y.
Mol. Cell. (2015). 58, 794-803

Netrin-1 exerts oncogenic activities through enhancing YAP stability.
Qi Q, Li DY, Luo HR, Guan KL, Ye K.
Proc. Natrl. Acad. Sci. (2015). 112, 7255-60

PARD3 induces TAZ activation and cell growth by promoting LATS1 and PP1 interaction.
Lv X, Liu CY, Wang Z, Sun YP, Xiong Y, Lei QY, Guan KL.
EMBO Rep. (2015). 16, 975-85

Opposing roles of conventional and novel PKC in the Hippo-YAP pathway regulation.
Gong R, Yu FX, Hong AW, Plouffe SW, Zhao B, Liu G, Xu Y, Guan KL.
Cell Res. (2015). 25, 985-8

Regulation of PGK1 acetylation by insulin and mTOR pathway modulates glycolytic ATP production and cellular redox potential.
Wang S, Jiang B, Zhang T, Liu L, Wang Y, Wang YP, Chen X, Lin H, Zhou L, Chen L, Yang C, Xiong Y, Ye D, Guan KL.
PLOS Biology. (2015). 13, e1002243

MAP4K family kinases act in parallel to MST1/2 to activate LATS1/2 in the Hippo pathway.
Meng Z, Moroishi T, Mottier-Pavie V, Plouffe SW, Hansen CG, Hong AW, Park HW, Mo JS, Lu W, Lu S, Flores F, Yu FX, Halder G, Guan KL.
Nature Commun. (2015). 6, 8357

Class III PI3K regulates organismal glucose homeostasis by providing negative feedback on hepatic insulin signaling.
Nemazanyy I, Montagnac G, Russell RC, Morzyglod L, Burnol AF, Guan KL, Pende M, Panasyuk G.
Nature Commun. (2015). 6, 8283

Atg5-independent autophagy regulates mitochondrial clearance and is essential for iPSC reprogramming.
Ma T, Li J, Xu Y, Yu C, Xu T, Wang H, Liu K, Cao N, Nie BM, Zhu S, Xu S, Li K, Wei WG, Guan KL, Ding S.
Nature Cell Biol. (20105). 17, 1379-87

Oncometabolite D-2-hydroxyglutarate inhibits ALKBH DNA repair enzymes and sensitizes IDH-mutated cells to alkylating agents.
Wang P, Wu J, Ma S, Zhang L, Yao J, Hoadley KA, Wilderson MD, Perou CM, Guan KL, Ye D, Xiong Y.
Cell Rep. (2015). 13, 2353-61

NLK phosphorylates Raptor to mediate stress-induced mTORC1 inhibition.
Yuan HX, Wang Z, Yu FX, Li F, Russell RC, Jewell JL, Guan KL.
Genes and Dev. (2015). 29, 2362-76

The Hippo pathway effector YAP and TAZ promote cell growth by modulating amino acid signaling to mTORC1.
Hansen CG, Ng YLD, Lam WM, Plouffe SW, Guan KL.
Cell Res. (2015). 25, 1299-313

Hippo pathway in regulation of gastrointestinal tissues.
Yu F, Meng Z, Plouffe S, Guan KL.
Ann. Rev. Physiol. (2015). 77, 201-27

The Hippo pathway in human cancer.
Moroishi T, Hansen C, Guan KL.
Nature Rev. Cancer. (2015). 15, 73-9

The Hippo pathway in heart development, regeneration, and diseases.
Zhou Q, Li L, Zhao B, Guan KL.
Circulation Res. (2015). 116, 1431-1447

YAP and TAZ: a nexus for Hippo signaling and beyond.
Hansen C, Moroishi T, Guan KL.
Trends in Cell Bio. (2015). 25, 499-513

AMPK and autophagy in glucose/glycogen metabolism.
Ha J, Guan KL, Kim J.
Mol. Aspects Med. (2015). 46, 46-62

Hippo pathway in organ size control, tissue homeostasis, and cancer.
Yu FX, Zhao B, Guan KL.
Cell. (2015). 163, 811-828

YAP as oncotarget in uveal melanoma.
Yu FX, Zhang K, Guan KL.
Oncoscience. (2015). 1, 480-1

Glutathione-S-transferase (GST)-fusion assays for studying for protein-protein interactions.
Vikis HG, Guan KL.
Methods Mol. Biol. (2015). 1278, 353-64

The Sin1 PH domain connects mTORC2 to PI3K.
Yuan HX, Guan KL.
Cancer Discovery. (2015). 5, 1127-9

Alternative Wnt Signaling Activates YAP/TAZ.
Park HW, Kim YC, Yu B, Moroishi T, Mo JS, Plouffe SW, Meng Z, Lin KC, Yu F, Alexander CM, Wang C, Guan KL.
Cell. (2015). in press

A YAP/TAZ-induced feedback mechanism regulates Hippo pathway homeostasis
Moroishi T, Park HW, Qin B, Chen Q, Meng Z, Plouffe SW, Taniguchi K, Yu F, Karin M, Pan D, Guan KL.
Genes Dev. (2015). in press

SIRT3-dependent GOT2 acetylation enhances the malate-aspartate NADH shuttle activity to promote pancreatic tumor growth.
Yang H, Zhou L, Zhao Y, Sun Y, Zhang J, Wang Y, Guan KL, Xiong Y, Ye D.
EMBO J. (2015). 34, 1110-25

A gp130-Src-YAP Module links intestinal injury  and inflammation to epithelial regeneration.
Taniguchi K, Wu LW, Grivennikov SI, de Jong PR, Lian I, Yu FX, Wang K, Ho SB, Boland BS, Chang JT, Sandborn WJ, Hardiman G, Raz E, Maehara Y, Yoshimura A, Zucman-Rossi J, Guan KL, Karin M.
Nature. (2015). 519, 57-62

WT1 recruits TET2 to regulate its target gene expression and suppress leukemia cell proliferation.
Wang Y, Xiao M, Chen X, Xu Y, Chen L, Wang P, Yang H, Ma S, Jiang B, Ye D, Guan KL, Xiong Y.
Mol. Cell. (2015). 57, 1-12

Kaposi sarcoma-associated herpesvirus promotes tumorigenesis by modulating the Hippo pathway.
Liu G, Yu FX, Kim YC, Meng Z, Naipauer J, Looney DJ, Lu X, Gutkind JS, Mesri EA, Guan KL.
Oncogene. (2015). 34,3536-46

YAP inhibits squamous transdifferentiation of Lkb1-deficient lung adenocarcinoma through ZEB2-dependent DNp63 repression.
Gao, Zhang Y, Han X, Li F, Wang X, Wang R, Fang Z, Tong X, Yao S, Li F, Feng Y, Sun Y, Hou Y, Yang Z, Guan KL, Chen H, Zhang L, Ji H.
Nature Comm. (2015). 5, 4629

The Hippo Pathway in Heart Development, Regeneration, and Diseases.
Zhou Q, Li L, Zhao B, Guan KL.
Circ Res. (2015). 116(8):1431-1447

Sestrin2 inhibits mTORC1 through modulation of GATOR complexes.
Kim JS, Ro SH, Kim M, Park HW, Semple IA, Park H, Cho US, Wang W, Guan KL, Karin M, Lee JH.
Sci Rep. (2015). 5:9502

Glutathione-S-Transferase (GST)-Fusion Based Assays for Studying Protein-Protein Interactions.
Vikis HG, Guan KL.
Methods Mol Biol. (2015). 1278:353-64

Cellular energy stress induces AMPK-mediated regulation of YAP and the Hippo pathway.
Mo JS, Meng Z, Kim YC, Park HW, Hansen CG, Kim S, Lim DS, Guan KL.
Nat Cell Biol. (2015). 17(4):500-10

Disease implications of the Hippo/YAP pathway.
Plouffe SW, Hong AW, Guan KL.
Trends Mol Med. (2015). 21(4):212-222

Micro(RNA) managing by mTORC1.
Jewell JL, Flores F, Guan KL.
Mol Cell. (2015). 57(4):575-6

mTOR: a pharmacologic target for autophagy regulation.
Kim YC, Guan KL.
J Clin Invest. (2015). 125(1):25-32

Differential regulation of mTORC1 by leucine and glutamine.
Jewell JL, Kim YC, Russell RC, Yu F, Park HW, Plouffe SW, Tagliabracci VS, Guan KL.
Science. (2015). pii: 1259472



Glyceraldehyde-3-phosphate Dehydrogenase Is Activated by Lysine 254 Acetylation in Response to Glucose Signal
Li T, Liu M, Feng X, Wang Z, Das I, Xu Y, Zhou X, Sun Y, Guan KL, Xiong Y, Lei QY.
J. Biol. Chem. (2014). 289, 3775-85

mTORC1 Promotes Denervation-Induced Muscle Atrophy Through a Mechanism Involving the Activation of FoxO and E3 Ubiquitin Ligases.
Tang H, Inoki K, Lee M, Wright M, Khuong A, Sugiarto A, Garner M, Oaik J, Dpinho R, Goldman R, Guan KL, Shrager JB.
Science Signaling. (2014). 7, ra18, 1-10

Both Decreased and Increased SRPK1 Levels Promote Cancer by Interfering with PHLPP-Mediated Dephosphorylation of Akt. 
Wang P, Zhou Z, Hu A, Ponte de Albuquerque C, Zhou Y, Hong L, Sierecki E, Ajiro M, Kruhlak M, Harris C, Guan KL, Zheng Z, Newton AC, Sun P, Zhou H, Fu XD.
Mol. Cell. (2014). in press

Mutant Gq/11 promote uveal melanoma tumorigenesis by activating YAP. 
Yu FX, Luo J, Mo JS, Liu G, Kim YC, Meng Z, Zhao L, Peyman G, Ouyang H, Jiang W, Zhao J, Chen X, Zhang L, Bastian BC, Zhang K, Guan KL.
Cancer Cell. (2014). in press

Rag GTPases are cardioprotective by regulating lysosomal function
Kim YC, Park HW, Sciarretta S, Mo JS, Jewell JL, Russell RC, Wu X, Sadoshima J, Guan KL.
Nature Comm. (2014).

Oxidative stress activates SIRT2 to deacetylate and stimulate phosphoglycerate mutase. 
Xu Y, Li F, Li T, Zhou X, Deng C, Guan KL, Lei, QY, Xiong Y.
Cancer Research. (2014). in press

Regulation of G6PD acetylation by KAT9/SIRT2 modulates NADPH homeostatis and cell survival during oxidative stress.
Wang YP, Zhou LS, Zhao YZ, Wang SW, Chen LL, Liu LX, Ling ZQ, Hu FJ, Sun YP, Zhang JY, Yang C, Yang Y, Xiong Y, Guan KL, Ye D.
EMBO J. (2014). in press

Transcription and processing: multilayer controls of RNA biogenesis by the Hippo pathway,
Yu F, Guan KL.
EMBO J. (2014). in press



ULK1 induces autophagy by phosphorylating Beclin-1 and activating Vps34 lipid kinase
Russell RC, Tian Y, Yuan H, Park HW, Chang YY, Kim J, Kim H, Neufeld TP, Dillin A, Guan KL.
Nat Cell Biol. (2013). 15(7): 10.1038/ncb2757

Regulation of the Hippo pathway and implications for anticancer drug development.
Park HW, Guan KL. 
Trends Pharmacol Sci. (2013). S0165-6147(13) 155-7

Defects of Vps15 in skeletal muscles lead to autophagic vacuolar myopathy and lysosomal disease
Nemazanny I, Blaauw B, Paolini C, Caillaud C, Protasi F, Mueller A, Proikas-Cezanne T, Russel RC, Guan KL, Nishino I, Sandri M, Pende M, Panasyuk G.
EMBO Mol Med. (2013). 5(6): 870-890

LDH-A Acetylation: Implication in Cancer
Zhao D, Xiong Y, Lei QY, Guan KL. 
Oncotarget. (2013). 4(6): 802-803

Lysine-5 Acetylation Negatively Regulates Lactate Dehydrogenase A and Is Decreased in Pancreatic Cancer
Zhao D, Zou SW, Liu Y, Zhou X, Mo Y, Wang P, Xu YH, Dong B, Xiong Y, Lei QY, Guan KL. 
Cancer Cell. (2013). 23(4):10.1016

Nutrient Signaling to mTOR and Cell Growth
Jewell JL, Guan KL. 
Trends Biochem Sci. (2013). 38(5): 233-242

Amino acid signalling upstream of mTOR
Jewell JL, Russell RC, Guan KL. 
Nat Rev Mol Cell Biol. (2013). 14(3): 133-139

Microtubule-associated Protein/Microtubule Affinity-regulating Kinase 4 (MARK4) Is a Negative Regulator of the Mammalian Target of Rapamycin Complex 1 (mTORC1)
Li L, Guan KL.
J Biol Chem. (2013). 288(1): 703-708

Tumor development is associated with dramatic decreases of TET gene expression and global 5-hydroxymethylcytosine.  
Yang H, Liu Y, Bai F, Zhang JY, Ma SH, Liu J, Xu Z, Zhu HG, Ling ZQ, Ye D, Guan KL, Xiong Y.
Oncogene. (2013). 32, 663-669

Mutations in isocitrate dehydrogenase 1 and 2 occur frequently in intrahepatic cholangiocarcinomas and share hypermethylation targets with glioblastomas.
Wang P, Dong Q, et al… Ye D, Guan KL, Xiong Y, Qin LX, Chiang DY.
Oncogene. (2013). in press

Differential regulation of distinct Vps34 complexes by AMPK in nutrient stress and autophagy. 
Kim J, Kim YC, Fang C, Russell RC, Kim JH, Fan W, Liu R, Zhong Q, Guan KL.
Cell. (2013). 152, 290-303

Regulation of PIK3C3/VPS34 complexes by mTOR in nutrient stress-induced autophagy.
Yuan HX, Russell R, Guan KL.
Autophagy. (2013). 12, 1983-95

Protein kinase A activates the Hippo pathway to modulate cell proliferation and differentiation.
Yu FX, Zhang Y, Park HW, Jewell JL, Chen Q, Pan DJ, Taylor SS, Lai ZC, Guan KL.
Genes & Dev. (2013). 27, 1223-32

Acetylation stabilizes ATP-citrate lyase to promote lipid biosynthesis and tumor growth.
Lin R, Xue RT, Gao X, Li T, Zhou X, Guan KL, Xiong Y, Lei QY.
Mol. Cell. (2013). 51, 506-518

Mitogenic and oncogenic signals stimulate K433 acetylation to promote PKM2 protein kinase activity and nuclear localization. 
Lv L, Xu Y, Zhao D, Li F, Wang W, Sasaki N, Jiang Y, Zhou X, Li T, Guan KL, Lei Q, Xiong Y.
Mol. Cell. (2013). 52, 340-352

Phosphorylation of angiomotin by Lats1/2 kinases inhibits F-actin binding, cell migration and angiogenesis.
Dai X, She P, Chi F, Feng Y, Liu H, Jin D, Zhao Y, Guo X, Jiang D, Guan KL, Zhong TP, Zhao B.
J. Biol. Chem. (2013). 288, 34041-51

LATS2 suppressed oncogenic Wnt signaling by disrupting b-catenin/BCL9 interaction.
Li J, Chen X, Ding X, Cheng Y, Zhao B, Lai ZC, Al Hezaimi K, Hakem R, Guan KL, Wang CY.
Cell Rep. (2013). 5, 1650-63


Upstream regulators of the Hippo pathway
Yu FX, Mo JS, Guan KL. 
Cell Cycle. (2012). 11(22): 4097–4098

IDH1 and IDH2 mutations in tumorigenesis: mechanistic insights and clinical perspectives
Yang H, Ye D, Guan KL, Xiong Y.
Clin Cancer Res. (2012). 18(20): 5562–5571

Regulation of the Hippo–YAP pathway by protease-activated receptors(PARs)
Mo JS, Yu FX, Gong R, Brown JH, Guan KL.
Genes Dev. (2012). 26(19): 2138–2143

YAP mediates crosstalk between the Hippo and PI3K-TOR pathways by suppressing PTEN via miR-29. 
Tumaneng K, Schlegelmilch K, Russell RC, Yimlamai D, Basnet H, Mahadevan N, Fitamant J, Bardeesy N, Camargo F, Guan KL.
Nature Cell Biol. (2012). 14, 1322-1329

Temporal changes in PTEN and mTORC2 regulation of hematopoietic stem cell self-renewal and leukemia suppression
Magee JA, Ikenoue T, Nakada D, Lee JY, Guan KL, Morrison SJ.
Cell Stem Cell. (2012). 11(3): 415–428

The YAP and TAZ transcription coactivators: key downstream effectors of the mammalian Hippo pathway
Hong W, Guan KL. 
Semin Cell Dev Biol. (2012). 23(7): 785–793

Regulation of the Hippo-YAP pathway by G-protein coupled receptor signaling
Yu FX, Zhao B, Panupinthu N, Jewell JL, Lian I, Wang LH, Zhao J, Yuan H, Tumaneng K, Li H, Fu XD, Mills GB, Guan KL. 
Cell. (2012). 150(4): 780–791

A Critical Role for Rictor in T-lymphopoiesis
Tang F, Wu Q, Ikenoue T, Guan KL, Liu Y, Zheng P.
J Immunol. (2012). 189(4): 1850–1857

Down Syndrome Cell Adhesion Molecule (DSCAM) Associates with Uncoordinated-5C (UNC5C) in Netrin-1-mediated Growth Cone Collapse
Purohit AA, Li W, Qu C, Dwyer T, Shao Q, Guan KL, Liu G.
J Biol Chem. (2012). 287(32): 27126–27138

The N-terminal Phosphodegron Targets TAZ/WWTR1 Protein for SCF?-TrCP-dependent Degradation in Response to Phosphatidylinositol 3-Kinase Inhibition
Huang W, Lv X, Liu CY, Zha A, Zhang H, Jiang Y, Xiong Y, Lei QY, Guan KL.
J Biol Chem. (2012). 287(31): 26245–26253

Mechanistic insights into the regulation of metabolic enzymes by acetylation
Xiong Y, Guan KL.
J Cell Biol.
 (2012). 198(2): 155–164

Inhibition of α-KG-dependent histone and DNA demethylases by fumarate and succinate that are accumulated in mutations of FH and SDH tumor suppressors
Xiao M, Yang H, Xu W, Ma S, Lin H, Zhu H, Liu L, Liu Y, Yang C, Xu Y, Zhao S, Ye D, Xiong Y, Guan KL.
Genes Dev. (2012). 26(12): 1326–1338

The mechanisms of IDH mutations in tumorigenesis
Ye D, Xiong Y, Guan KL. 
Cell Res. (2012). 22(7): 1102–1104

Alterations of Metabolic Genes and Metabolites in Cancer
Oermann EK, Wu J, Guan KL, Xiong Y.
Semin Cell Dev Biol. (2012). 23(4): 370–380

Organ Size Control by Hippo and TOR Pathways
Tumaneng K, Russell RC, Guan KL.
Curr Biol. (2012). 22(9): R368–R379

The Vam6 and Gtr1–Gtr2 pathway activates TORC1 in response to amino acids in fission yeast
Valbuena N, Guan KL, Moreno S.
J Cell Sci. (2012). 125(8): 1920–1928

Guidelines for the use and interpretation of assays for monitoring autophagy
Klionsky DJ, et al.
Autophagy. (2012). 8(4): 445–544

Acetylation negatively regulates glycogen phosphorylase by recruiting protein phosphatase 1
Zhang T, Wang S, Lin Y, Xu W, Ye D, Xiong Y, Zhao S, Guan KL.
Cell Metab. (2012). 15(1): 75–87

Cell detachment activates the Hippo pathway via cytoskeleton reorganization to induce anoikis
Zhao B, Li L, Wang L, Wang CY, Yu J, Guan KL.
Genes Dev. (2012). 26(1): 54–68


The TSC1 and TSC2 tumor suppressors are required for proper ER stress response and protect cells from ER stress-induced apoptosis. 
Kang YJ, Lu MK, Guan KL.
Cell Death & Differentiation. (2011). 18, 133-44

Angiomotin is a novel Hippo pathway component that inhibits YAP oncoprotein. 
Zhao B, Li L, Lu O, Wang LH, Liu C, Lei Q, Guan KL.
Genes & Dev. (2011). 25, 51-63

Oncometabolite 2-hydroxyglutarate is a competitive inhibitor of a-ketoglutarate-dependent dioxygenases. 
Xu W, Yang H, Liu Y, et al… Guan KL, Zhao S, Xiong Y.
Cancer Cell. (2011). 19, 17-30

PP1 Cooperates with ASPP2 to dephosphorylate and activate TAZ. 
Liu C, Lv X, Li T, Xu Y, Zhou X, Zhao S, Xiong Y, Lei Q, Guan KL.
J. Biol. Chem. (2011). 286, 5558-66

AMPK and mTOR regulate autophagy via direct phosphorylation of ULK1. 
Kim J, Kundu M, Viollet B, Guan KL.
Nature Cell Biol. (2011). 13, 132-141

Sirt3 Promotes the Urea Cycle and Fatty Acid Oxidation during Dietary Restriction.  
Hallows WC, Yu W, Smith BC, Devires MK, Ellinger JJ, Someya S, Shortreed MR, Prolla T, Markley JL, Smith LM, Zhao S, Guan KL, Denu JM.
Mol. Cell. (2011). 41, 139-49

mTORC1 activation in podocytes is a critical step in the development of diabetic nephropathy.  
Inoki K, Mori H, Wang J, Suzuki T, Hong S, Yoshida S, Blattner SM, Ikenoue T, Rüegg RA, Hall MN, Kwiatkowski DJ, Rastaldi MP, Huber  TB, Kretzler M, Holzman LB, Wiggins RC, Guan KL.
J. Clin. Invest. (2011). 121, 181-196

Inactivation of Rheb by PRAK-mediated phosphorylation is essential for energy-depletion-induced suppression of mTORC1.  
Zheng M, Wang YH, Wu XN, Wu SQ, Lu BJ, Dong MQ, Zhang H, Sun P, Lin SC, Guan KL, Han J.
Nature Cell Biol. (2011). 13, 263-272

IkappaB kinase epsilon and TANK-binding kinase 1 activate AKT by direct phosphorylation.
Xie X, Zhang D, Zhao B, Lu MK, You M, Condorelli G, Wang CY, Guan KL.
Proc. Natl. Acad. Sci. (2011). 108, 6474-9

Tumor suppressor SIRT3 deacetylates and activates manganese superoxide dismutase (SOD2) to scavenge ROS.  
Chen Y, Zhang J, Lin Y, Lei Q, Guan KL, Zhao S, Xiong Y.
EMBO Report. (2011). 12, 534-41

Acetylation regulates gluconeogenesis by promoting PEPCK1 degradation via recruiting the UBR5 ubiquitin ligase.  
Jiang W, Wang S, Xiao M, Lin Y, Lei QY, Xiong Y, Guan KL, Zhao S.
Mol. Cell. (2011). 43, 33-44

Acetylation Targets the M2 Isoform of Pyruvate Kinase for Degradation through Chaperone-Mediated Autophagy and Promotes Tumor Growth.
Lv L, Li D, Zhao D, Lin R, Chu Y, Zhang H, Zha Z, Xu Y, Li Z, Wang G, Huang Y, Zhao S, Xiong Y, Guan KL, Lei QY.
Mol. Cell. (2011). 42, 719-30

The Tuberous Sclerosis Complex-Mammalian Target of Rapamycin Pathway Maintains the Quiescence and Survival of Naive T Cells.  
Wu Q, Liu Y, Chen C, Ikenoue T, Qiao Y, Li CS, Li W, Guan KL, Liu Y, Zheng P.
J. Immunol. (2011). 187, 1106-12

Crystal Structure of Gtr1p-Gtr2p complex reveals new insights into the amino acid-induced TORC1 activation.
Gong R, Li L, Liu Y, Wang P, Yang H, Wang L, Cheng J, Guan KL, Xu Y.
Genes & Dev. (2011). 25, 1668-73

Redox regulates mTORC1 activity by modulating the TSC1/TSC2-Rheb GTPase pathway. 
Yoshida S, Hong S, Suzuki T, Nada S, Mannan AM, Wang J, Okada M, Guan KL, Inoki K.
J. Biol. Chem. (2011). 286, 32651-60

Regulation of intermediate metabolism by protein acetylation. 
Guan KL, 
Xiong Y.
TIBS. (2011). 36, 108-116

Amino acid signaling in mTOR activation. 
Kim J, Guan KL.
Ann. Rev. Biochem. (2011). 80, 1001-32

The Hippo tumor suppressor pathway in organ size control, tissue regeneration and stem cell self-renewal.
Zhao B, Tumaneng K, Guan KL.
Nature Cell Biol. (2011). 13, 877-83

An emerging role for TOR signaling in mammalian tissue and stem cell physiology. 
Russell RC, Fang C, Guan KL.
Development. (2011). 138, 3343-56

AMPK and mTOR in cellular energy homeostasis and drug targets.
Inoki K, Kim J, Guan KL.
Ann. Rev. Pharmacol. Toxicol. (2011). 52, 381-400



A coordinated phosphorylation by Lats and CK1 regulates YAP stability through SCF(beta-TRCP).
Zhao B, Li L, Wang CY, Guan KL.
Genes & Dev. (2010). 24, 72-85

Wnt/GSK3 signaling regulates ATF4 and its implication in canonical Wnt signaling-induced osteogenesis.
Zhang X, Li Y, Liu K, Wang F, Eathma JL, Lee CH, Patrene K, Eoodman GD, Inoki K, MacDougald OA, Guan KL, Ouyang H.
J. Clin. Inves. (2010). in press

Structural insights into the YAP and TEAD complex. 
Li Z, Zhao B, Wang P, Chen F, Dong Z, Yang H, Guan KL, Xu Y.
Genes & Dev. (2010). 24, 235-240

Regulation of cellular metabolism by protein lysine acetylation.  
Zhao S, Xu W, Jiang W, Yu W, Lin Y, Zhang T, Yao J, Zeng Y, Li H, Li Y, Shi. J, An W, Hancock SM, He F, Qin L, Chin J, Yang P, Chen X, Lei Q, Xiong Y, Guan KL.
Science. (2010). 327, 1000-1004

Acetylation of metabolic enzymes coordinates carbon source utilization and metabolic flux.  

Wang Q, Zhang Y, Yang C, Xiong H, Lin Y, Yao J, Li H, Xie L, Zhao W, Yao Y, Ning ZB, Zeng R, Xiong Y, Guan KL, Zhao S, Zhao GP.
Science. (2010). 327, 1004-1007

ATM signals to TSC2 in the cytoplasm to regulate mTORC1 in response to ROS.
Alexander A, Cai SL, Kim J, Nanez A, Sahin M, MacLean KH, Inoki K, Guan KL, Shen J, Person MD, Kusewitt D, Mills GB, Kastan MB, Walker CL.
Proc. Nat. Acad. Sci. (2010). 107, 4153-8

The role of YAP transcription co-activator in regulating stem cells self-renewal and differentiation. 
Liang I, Kim J, Okazawa H, Zhao J, Zhao B, Yu J, Chinnaiyan A, Israel M, Goldstein L, Abujarour R, Ding S, Guan KL.
Genes & Dev. (2010). 24, 1106-1118

Regulation of mTORC1 by the Rab and ARF GTPases.
Li L, Kim E, Yuan H, Inoki K, Goraksha-Hicks, Schiesher RL,  Neufeld TP, Guan KL.
J. Biol. Chem. (2010). 285, 19705-9

Generation of acetyllysine antibodies and affinity enrichment of acetylated peptides.
Guan KL, Yu W, Lin Y, Xiong Y, Zhao S.
Nature Protocol. (2010).  5, 1583-1595

Regulation of integrin b1 recycling to lipid rafts by Rab1a to promote cell migration. 
Wang C, Yoo Y, Fan H, Kim E, Guan KL, Guan JL.
J. Biol. Chem. (2010). 285, 29389-405

MTORC1 regulates cardiac function and myocyte survival through 4E-BP1 inhibition in mice.
Zhang D, Contu R, Latronico MV, Zhang JL, Rizzi R, Catalucci D, Miyamoto S, Huang K, Ceci M, Gu Y, Dalton ND, Peterson KL, Guan KL, Brown JH, Chen J, Sonenberg N, Condorelli G.
J. Clin. Invest. (2010). 120, 2805-16

The Hippo tumor pathway promotes TAZ degradation by phosphorylating a phosphodegron and recruiting the SCFβ-TrCP E3 ligase.
Liu CY, Zha ZY, Zhang H, Zhou X, Huang W, Zhao D, Chan SW, Lim CJ, Hong S, Zhao S, Xiong Y, Lei, QY, Guan KL.
J. Biol. Chem. (2010). 285, 37159-69


Amino acid signaling to TOR activation: Vam6 functioning as a Gtr1 GEF. 
Li L, Guan KL.
Mol. Cell. (2009). 35, 543-545

Rag GTPases in TORC1 activation and nutrient signaling.
Li L, Guan KL.
Enzymes. (2009). in press

Harness the Power: New Insights into the Inhibition of YAP/ Yorkie
Zhao B, Lei Q, Guan KL.
Dev. Cell. (2009). 16, 321-322

MST Out and HCC In.
Zhao B, Lei Q, Guan KL.
Cancer Cell. (2009). 16, 363-364

Both TEAD binding and WW domains are required for the growth stimulation and oncogenic transformation activity of YAP. 
Zhao B, Kim J, Ye X, Lai ZC, Guan KL.
Cancer Research. (2009). 69, 1089-1098

DSCAM functions as a netrin receptor in commissural axon pathfinding.
Liu G, Li W, Wang L, Kar A, Guan KL, Rao Y, Wu JY.
Proc. Nat. Acad. Sci. USA. (2009). 106, 2951-2956

Critical role for hypothalamic mTOR activity in energy balance. 
Mori H, Inoki K, Münzberg H, Opland D, Faouzi M, Villanueva EC, Ikenoue T, Kwiatkowski D, MacDougald OA, Myers MG, Guan KL.
Cell Metabolism. (2009). 9, 362-74

Glioma-derived Mutations in IDH1 Dominantly Inhibit IDH1 Catalytic Activity and Induce HIF-1a.
Zhao S, Lin Y, Xu W, Jiang W, Zha Z, Wang P, Yu W, Li Z, Gong L, Peng Y, Ding J, Lei Q, Guan KL, Xiong Y.
Science. (2009). 324, 261-5

Inhibition of Osteoblast functions by IKK/NFkB in osteoporosis
Chang J, Wang Z, Tang E, Fan Z, McCauley L, Franceschi R, Guan KL, Kresbach PH, Wang CY.
Nature Med. (2009). 15, 682-9

The mTOR pathway is highly activated in diabetic nephropathy and rapamycin has a strong therapeutic potential.
Mori H, Inoki K, Masutani K, Wakabayashi Y, Komai K, Nakagawa R, Guan KL, Yoshimura A.
Biochem. Biophys. Res. Comm. (2009). 384, 471-5

Lysine 88 Acetylation Negatively Regulates Ornithine Carbamoyltransferase Activity in Response to Nutrient Signals.
Yu W, Lin Y, Yao J, Huang W, Lei Q, Xiong Y, Zhao S, Guan KL.
J. Biol. Chem. (2009). 284, 13669-75

TEAD transcription factors mediate the function of TAZ in cell growth and epithelial-mesenchymal transition.
Zhang H, Liu CY, Zha ZY, Zhao B, Yao J, Zhao SM, Xiong Y, Lie QY, Guan KL.
J. Biol. Chem. (2009). 284, 13355-62

Rheb controls misfolded protein metabolism by inhibiting aggresome and autophagy.
Zhou X, Ikenoue T, Chen X, Li L, Inoki K, Guan KL.
Proc. Nat. Acad. Sci. (2009). 106, 8923-8

Critical roles for the TSC-mTOR pathway in b-cell function. 
Mori H, Inoki K, Opland D, Münzberg H, Villanueva EC, Faouzi M, Ikenoue T, Kwiatkowski D, MacDougald OA, Myers MG, Guan KL.
American J. Physiol. Endocrinol Metab. (2009). 297, E1013-1022


Spatial regulation APPLies to Akt substrate selectivity.
Zhao S, Guan KL.
Cell. (2008). 133, 399-400

TAZ promotes cell proliferation and EMT and is inhibited by the Hippo pathway. 
Lei QY, Zhang H, Zhao B, Zha Y, Bai F, Pei XH, Zhao S, Xiong Y, Guan KL.
Mol. Cell Biol. (2008). 28, 2426-36

TEAD mediates YAP dependent gene induction and growth control.
Zhao B, Ye X, Yu J, Li L, Li W, Li S, Lin JD, Wang CY, Chinnaiyan AM, Lai ZC, Guan KL.
Genes & Development. (2008). 22, 1962-71

Essential Function of TORC2 in PKC and Akt Turn Motif Phosphorylation, Maturation, and Signaling. 
Ikenoue T, Inoki K, Yang Q, Zhou X, Guan KL.
EMBO J. (2008). 27, 1919-1931

Regulation of TORC1 by Rag GTPases in nutrient response. 
Kim E, Goraksha P, Li L, Neufeld TP, Guan KL.
Nature Cell Biol. (2008). 10, 935-945

PTEN Acetylation Modulates its Interaction with PDZ Domain.
Ikenoue T, Inoki K, Zhao B, Guan KL.
Cancer Research. (2008). 68, 6908-12

A GSK-3, TSC2, mTOR pathway regulates glucose uptake and GLUT1 glucose transporter expression. 
Buller C, Loberg RD, Fan MH, Zhu Q, Park JL, Vesely E, Inoki K, Guan KL, Brosius FC.
Am J Physiol Cell Physiol. (2008). 295, C836-43

AMP-activated protein kinase contributes to UV- and H2O2-induced apoptosis in human skin keratinocytes. 
Cao C, Lu S, Kivlin R, Wallin B, Card E, Bagdasarian A, Tamakloe T, Chu WM, Guan KL, Wan Y.
J. Biol. Chem. (2008). 283, 28897-908

TSC-mTOR maintains quiescence and function of hematopoietic stem cells by repressing mitochondrial biogenesis and reactive oxygen species.
Chen C, Liu Y, Liu R, Ikenoue T, Guan KL, Liu Y, Zheng P.
J Exp. Med. (2008). 205, 2397-2408


p130CAS Is Required for Netrin Signaling and Commissural Axon Guidance.
Liu G, Li W, Li X, Park HT, Jurgensen C, Hanks SK, He ML, Wu J, Guan KL, Rao Y.
J. Neurosci. (2007). 27, 957-68

Adiponectin sensitizes insulin signaling by reducing p70 S6 kinase-mediated serine phosphorylation of IRS-1.  
Wang C, Mao X, Wang L, Liu M, Wetzel MD, Guan KL, Dong LQ, Liu F.
J. Biol. Chem. (2007). 282, 7991-96

Inactivation of YAP oncoprotein by the Hippo pathway is involved in cell contact inhibition and tissue growth control. 
Zhao B, Wei X, et al… Guan KL.
Genes & Development. (2007). 21, 2762-74

Activation of mTOR sensitizes cells to p53 dependent apoptosis.
Lee CH, Inoki K, Karbowniczek M, Petroulakis E, Sonenberg N, Henske EP, Guan KL.
The EMBO J. (2007). 26, 4812-23

Bnip3 mediates the hypoxia-induced inhibition on mTOR by interacting with Rheb.
Li Y, Wang Y, Kim E, Beemiler P, Wang CY, Swanson J, You M, Guan KL.
J. Biol. Chem. (2007). 282, 35803-13


Semaphorin-4D activates the MAPK pathway downstream of plexin-B1. 
Aurandt J, Li W, Guan KL.
Biochem. J. (2006). 394, 459-64

FAK and Src kinases are required for netrin-induced tyrosine phosphorylation of UNC5. 
Li W, Aurandt J, Jürgensen C,  Rao Y, Guan KL.
J. Cell Science. (2006). 119, 47-55

TSC1 stabilizes TSC2 by inhibiting the interaction between TSC2 and the HERC1 ubiquitin ligase. 
Chong-Kopera H, Inoki K, Li Y, Zhu T, Garcia-Gonzalo FR, Rosa JL, Guan KL.
J. Biol. Chem. (2006). 281, 8316-6

TSC1/TSC2 and Rheb have different effects on TORC1 and TORC2 activity. 
Yang Q, Inoki K, Kim E, Guan KL.
Proc. Natl. Acad. Sci. USA. (2006). 103(18):6811-6

TSC2 integrates Wnt and cellular energy signals through a coordinated phosphorylation by AMPK and GSK3 to regulate cell growth.
Inoki K, Ouyang H, Zhu T, Lindvall C, Wang Y, Yang Q, Bennett C, Harada Y, Stankunas K, Wang C, He H, MacDougald O, You M, Williams B, Guan KL.
Cell. (2006). 126, 955-68

Identification of Sin1 as an essential TORC2 component required for complex formation and kinase activity.
Yang Q, Inoki K, Ikenoue T, Guan KL.
Genes & Dev. (2006). 20, 2833-47

Muscle atrophy in transgenic mice expressing a human TSC1 transgene.
Wan M, Wu X, Guan KL, Han M, Zhuang Y, Xu T.
FEBS Lett. (2006). 580, 5621-7


Measurements of TSC2 GAP Activity Toward Rheb.
Li Y, Inoki K, Vikis H, Guan KL.
Methods Enzymol. (2005). 407, 46-54

Expression and Phenotype Analysis of the Nephrocystin-1 and Nephrocystin-4 Homologs in Caenorhabditis elegans.
Wolf MT, Lee J, Panther F, Otto EA, Guan KL, Hildebrandt F.
J Am Soc Nephrol. (2005). 16, 676-687

The stress-inducted proteins RTP801 and RTP801L are negative regulators of the mammalian target of rapamycin pathway. 
Corradetti M, Inoki K, Guan KL.
J. Biol. Chem. (2005). 280, 9769-72

Structural basis for the unique biological function of small GTPase human Rheb.
Yu Y, Li S, Xu X, Li Y, Guan KL, Arnold E, Ding J.
J. Biol. Chem. (2005). 280, 17093-00

SRC-1 mediates UNC-5 signaling in C. elegans
Lee J, Li WQ. Guan KL.
Mol. Cell. Biol. (2005). 25, 6485-95

Identification of FIP200 interaction with TSC1-TSC2 complex and its role in regulation of cell size control. 
Gan B, Melkoumian ZK, Wu X, Guan KL, Guan JL.
J. Cell Biol. (2005). 170, 379-89


Polo-like kinase 3 (Plk3) is Golgi-localized and involved in regulating Golgi fragmentation during the cell cycle.
Ruan Q, Xie S, Fang Y, Darzynkiewicz Z, Guan KL, Uniyal MJ, Dai W.
Experimental Cell Res. (2004). 294, 51-59

Attenuation of the Ras/MAPK signal transduction pathway via zinc ion mediated inhibition of a convergent regulatory input.
Yoder JH, Chong H, Guan KL, Han M.
EMBO J. (2004). 23, 111-119

The zinc finger mutation C417R of I-kappa B kinase gamma impairs lipopolysaccharide- and TNF-mediated NF-kappa B activation through inhibiting phosphorylation of the I-kappa B kinase beta activation loop.
Yang F, Yamashita J, Tang E, Wang HL, Guan KL, Wang CY.
J Immunol. (2004). 172, 2446-2452

Pol iota is a candidate for the mouse pulmonary adenoma resistance 2 locus, a major modifier of chemically induced lung neoplasia.  
Wang M, Devereux T, Vikis H, McCulloch S, Holiday AC, Wang Y, Debenek, Kunkel T, Guan KL, You M.
Cancer Res. (2004). 64, 1924-1931

MEK1-induced Golgi dynamics during cell cycle progression is partly mediated by Polo-like kinase-3.
Xie S, Wang Q, Ruan Q, Liu T, Jhanwar-Uniyal M, Guan KL, Dai W.
Oncogene. (2004). 6, 3822-9

The Down syndrome cell adhesion molecule (DSCAM) interacts with and activates PA. 
Li W, Guan KL.
J. Biol. Chem.
 (2004). 279, 32824-31

Regulation of the TSC pathway by LKB1:  Evidence of a molecular link between Tuberous Sclerosis Complex and Peutz-Jeghers Syndrome.  
Corradetti M, Inoki K, Bardeesy N, DePinho R, Guan KL.
Genes & Dev. (2004). 18, 1533-1538

Activation of FAK and Src are receptor proximal events required for netrin signaling.
Li W, Lee J, Vikis H, et al… Guan KL.
Nature Neuroscience. (2004). 7, 1213-1221

Class IV Semaphorins Promote Angiogenesis by Stimulating Rho-Initiated Pathways through Plexin-B. 
Basile J, Barac A, Zhu T, Guan KL, Gutkind JS.
Cancer Research. (2004). 64, 5212-24

Biochemical and functional characterization of small GTPase Rheb and TSC2 GAP activity.
Li Y, Inoki K, Guan KL.
Mol. Cell. Biol. (2004). 24, 7965-75

Transformation potential of Ras isoforms correlates with activation of PI3K but not ERK.  
Li W, Zhu T, Guan KL.
J. Biol. Chem. (2004). 279, 37398-406

Mapping netrin-receptor binding reveals domains of Unc5 regulating its activity.
Kruger R, Lee J, Li W, Guan KL.
J. Neuroscience. (2004). 24, 10826-10834

Glutathione-S-transferase-fusion based assays for studying protein-protein interactions.
Vikis H, Guan KL.
Methods Mol. Biol. (2004). 261, 175-186


The p38 and MK2 kinase cascade phosphorylates tuberin, the tuberous sclerosis 2 (TSC2) gene product,  and enhances its interaction with 14-3-3. 
Li Y,  Inoki K, Vacrasis P, Guan KL.
J. Biol. Chem. (2003). 278, 13663-13671

Insulin Signaling Inhibits the 5-HT2C Receptor in Choroid Plexus via MAP kinase.
Hurley JH, Szhang S,  Bye L,  Marshall MS, DePaoli-Roach A, Guan KL, Fox AP, Yu L.
BMC Neuroscience. (2003). 4, 10


IKKbeta Plays an Essential Role in the Phosphorylation of RelA/p65 on Serine 536 Induced by Lipopolysaccharide.
Yang F, Tang E, Guan KL, Wang CY.
J. Immuno. (2003). 170, 11

Rheb GTPase is a direct target of TSC2 GAP activity and regulates mTOR signaling.
Inoki K, Li Y, Xu T, Guan KL.
Genes & Development. (2003). 17, 1829-1834

Human Homologue of Drosophila CNK interacts with Ras effector proteins, Raf and Rlf. 
Lanigan T, Liu A, Huang YZ, Mei L, Margolis B, Guan KL.
FASEB J. (2003). 17, 2048-60

A role for NF-kB essential modifier/IkB kinase-g (NEMO/IKKg) ubiquitination in the activation of the IkB kinase complex by tumor necrosis factor-a.
Tang ED, Wang C, Xiong Y, Guan KL.
J. Biol. Chem. (2003). 278, 37297-305

Roles of homotypic interaction and transautophosphorylation in IKKb activation. 
Tang ED, Inohara N, Wang C, Nunez G, Guan KL.
J. Biol. Chem. (2003). 278, 38566-70

Positional cloning of the major quantitative trait locus underlying lung tumor susceptibility in mice.
Zhang Z, Futamura M, Vikis H, Wang M, Li J, Wang Y, Guan KL, You M.
PNAS. (2003). 100, 12642-47

Regulation of Raf through phosphorylation and N-terminus-C-terminus interaction. 
Chong H, Guan KL.
J. Biol. Chem. (2003). 278, 36269-76

CD24 is a genetic modifier for risk and progression of multiple sclerosis.
Zhou Q, Rammohan K, Lin S, Robinson N, Li Q, Liu X,  Bai X, Yin L, Scarberry B,  Du P,  You M, Guan KL, Zheng P, Liu Y.
Proc. Natl. Acad. Sci. (2003). 100, 15041-15046

TSC2 mediates cellular energy response to control cell growth and survival. 
Inoki K, Zhu T, Guan KL.
Cell. (2003). 115, 577-590

Control of the Ras-MAPK signaling pathway by phosphorylation and scaffolding proteins.
Lanigan T, Vikis H, Guan KL.
Res. Signpost. (2003). 37, 661-615


SmgGDS displays differential binding and exchange activity towards different Ras isoforms. 
Vikis HG, Stewart S, Guan KL.
Oncogene. (2002). 21, 2425-32

Wnt signaling promotes oncogenic transformation by inhibiting c-Myc-induced apoptosis. 
You Z, Saims D, Chen S, Zhang Z, Guttridge DC, Guan, KL, MacDougald OA, Brown AM, Evan G, Kitajewski J, Wang CY.
J. Cell Biol. (2002). 157, 429-40

The Plexin-B1/Rac interaction inhibits PAK activation and enhances Sema4D ligand binding. 
Vikis HG, Li, W, He Z, Guan KL.
Genes & Development. (2002). 16, 836-845

Intracrine hepatopoietin potentiates AP-1 activity through JAB1 independent of MAPK pathway. 
Lu C, Li Y, Zhao Y, Xing G, Tang F, Wang Q, Sun Y, Wei H, Yang X, Wu C, Chen J, Guan, KL, Zhang C, Chen H, He F.
FASEB J. (2002). 16, 90-2

TSC2 is phosphorylated and inhibited by AKT and suppresses mTOR signaling. 
Inoki K, Li Y, Zhu T, Wu J, Guan KL.
Nature Cell Biol. (2002). 4, 648-657

The semaphorin receptor plexin-B1 signals through a direct interaction with the Rho specific nucleotide exchange factor, LARG.  
Aurandt J, Vikis HG, Gutkind S, Ahn N, Guan KL.
Proc. Natl. Acad. Sci. USA. (2002). 99, 12085-12090

Regulation of TSC2 by 14-3-3 binding.
Li Y,  Inoki K, Yeung R, Guan KL.
J. Biol. Chem.
 (2002). 277,44593-44596

Regulation of the Ras-MAPK pathway at the level of Ras and Raf. 
Vikis H, Guan KL.
Genetic Engineering (Jane Setlow eds). (2002). 24, 49-66


p53CP is p51/p63, the third member of the p53 gene family: partial purification and characterization. 
Tan M, Bian J, Guan KL, Sun Y.
Carcinogenesis. (2001). 22, 295-300

Positive and negative regulation of Raf kinase activity and function by phosphorylation.
Chong H, Lee J, Guan KL.
EMBO J. (2001). 20: 3716-3727

Wild type ras can inhibit lung carcinogenesis in mice.  
Zhang Z, Wang Y, Vikis HG, Anderson MW, Sills RC, Holliday W, Devereux TR, Guan KL, You M.
Nature Genetics. (2001). 29, 25-33

Serum and glucocorticoid-inducible kinase SGK phosphorylates and negatively regulates B-Raf. 
Zhang BH, Tang E, Zhu T, Greenberg M, Vojtek A, Guan KL.
J. Biol. Chem. (2001). 276, 31620-31626

Function of the Rho family GTPases in Ras stimulated Raf activation. 
Li W, Chong H, Guan KL.
J. Biol. Chem. (2001). 276, 34728-37

Suppression of tumor necrosis factor-mediated apoptosis by NF-kB-independent BMP/Smad signaling.
Chen S, Guttridge D, Tang E, Shi ST, Guan KL, Wang CY.
J. Biol. Chem.
 (2001). 276, 39259-39263


MAPK pathways activate and phosphorylate the osteoblast-specific transcription factor, Cbfa1. 
Xiao G, Jiang D, Thomas P, Benson MD, Guan KL, Karsenty G, Franceschi RT.
J. Biol. Chem. (2000). 275, 4453-4459

PTGF-beta, a type beta transforming growth factor (TGF-beta) superfamily member, is a p53 target gene that inhibits tumor cell growth via TGF-beta signaling pathway.
Tan M, Wang Y, Guan KL, Su Y.
Proc. Natl. Acad. Sci. USA. (2000). 97, 109-114

Essential Functions of Protein Tyrosine Phosphatases Ptp2 and Ptp3 and Rim11 Tyrosine Phosphorylation in Saccharomyces cerevisiae Meiosis and Sporulation  
Zhan X, Hong Y, Zhu T, Mitchell AP, Deschenes RJ, Guan KL.
Mol. Biol. Cell. (2000). 11, 663-676

The dominant negative Ras mutant, N17Ras, can inhibit signaling independently of Ras activation. 
Stewart S, Guan KL.
J. Biol. Chem.
 (2000). 275, 8854-8862

The leucine-rich repeat protein Sur-8 enhances MAP kinase activation and forms a complex with Ras and Raf.
Li W, Han M, Guan KL.
Genes & Dev. (2000). 14, 895-900

Negative regulation of the serine/threonine kinase B-Raf by Akt. 
Guan KL,
 Figueroa C, Brtva T, Zhu T, Taylor J, Barber T, Vojtek A.
J. Biol. Chem. (2000). 275, 27354-27359

Regulation of STAT3 by Direct Binding to the Rac1 GTPase..  
Simon AR, Vikis HG, Fanburg BL, Stewart S, Cochran BH, Guan KL.
Science. (2000). 290, 144-147

A Caenorhabditis elegans MAP kinase kinase, MEK-1, is involved in stress responses.
Koga M, Zwaal R, Guan KL, Avery L, Ohshima Y.
EMBO J. (2000). 19, 5148-5156

Activation of B-Raf kinase requires phosphorylation of the conserved residues Thr598 and Ser601. 
Zhang BH, Guan KL.
EMBO J. (2000). 19, 5429-5439

The semaphorin receptor plexin-B1 specifically interacts with active Rac in a ligand dependent manner. 
Vikis HG, Li W, He Z, Guan KL.
Proc. Natl. Acad. Sci. (2000). 97, 12457-12462

Stimulation of the Mitogen-activated Protein Kinase Cascade and Tyrosine Phosphorylation of the Epidermal Growth Factor Receptor by Hepatopoietin.
Li Y, Li M, Xing G, Hu Z, Wang Q, Dong C, Wei H, Fan G, Chen J, Yang X, Zhao S, Chen H, Guan KL, Wu C, Zhang C, He FC.
J. Biol. Chem. (2000). 275, 37443-37447


Identification of functional elements of p18INK4C essential for binding and inhibition of cyclin-dependent kinase (CDK)4 and CDK6. 
Noh SJ, Li Y, Xiong Y, Guan KL.
Cancer Research. (1999). 59, 558-564

Transcriptional activation of the human S100A2 promoter by wild type p53.
Tan M, Heizmann CW, Guan KL, Schafer BW, Sun Y.
FEBS Lett. (1999). 445, 265-268

Transcriptional activation of the human glutathione peroxidase promoter by p53. 
Tan M, Li S, Swaroop M, Guan KL, Oberley LW, Sun Y.
J. Biol. Chem. (1999). 274, 12061-12066

Negative regulation of the forkhead transcription factor FKHR by AKT.
Tang E, Nunez G, Bar D, Guan KL. 
J. Biol. Chem.
 (1999). 274, 16741-16746

Kinase Suppressor of Ras (KSR) forms a multi-protein signaling complex and modulates MEK localization.
Stewart S, Sundaram M, Zhang Y, Lee J, Han M, Guan KL.
Mol. Cell. Biol. (1999). 19, 5523-5534

Cdkn2a encodes functional variation of p16ink4a but not p19ARF, which confers selection in mouse lung tumorigenesis.
Herzog CR, Noh S, Lantry LE, Guan KL, You, M.
Mol. Carcinogenesis. (1999). 25, 92-98

Specific protein-protein interaction accounts for the in vivo substrate selectivity of Ptp3 towards the Fus3 MAP kinase.
Zhan XL, Guan KL.
Genes & Dev. (1999). 13, 2811-2817


Altering the nucleophile specificity of a protein-tyrosine phosphatase catalyzed reaction: probing the function of the invariant glutamine residues.
Zhao Y, Wu L, Noh SJ, Guan KL, Zhang ZY.
J. Biol. Chem. (1998). 273, 5484-5492

The kinase suppressor of Ras (KSR) modulates growth factor and Ras signaling by uncoupling Elk-1 phosphorylation from MAP kinase activation.
Sugimoto T, Scott, S, Guan KL.
The EMBO J. (1998). 17, 1717-1727

Cloning and characterization of GETS-1, a goldfish Ets family member that functions as a transcriptional repressor in muscle.
Goldman, D, Sapru MK, Stewart S, Plotkin J, Libermann TA, Wasylyk B, Guan KL.
Biochem. J. (1998). 335, 267-275

Growth hormone stimulates phosphorylation and activation of Elk-1 and expression of c-fos, egr-1, and junB through activation of extracellular signal-regulated kinases 1 and 2
Hodge C, Liao JF, Stofega M, Guan KL, Carter-Su C, Schwartz J.
J. Biol. Chem.
 (1998). 273, 31327-31336



Differential effect of glucose deprivation on MAPK activation in drug sensitive human breast carcinoma MCF-7 and multidrug resistant MCF-7/ADR cells.
Gupta AK, Lee YL, Galoforo SS, Berns CM, Martinez AA, Corry PM, Wu X, Guan KL.
Mol. Cell. Biochem. (1997). 170, 23-30

Structure-function studies of p38 MAP kinase: loop12 influences substrate specificity and autophosphorylation but not upstream kinase selection.
Jiang Y, Li Z, Scharz EM, Guan KL, Ulevitch RJ, Han J.
J. Biol. Chem. (1997). 272, 11096-1102

Diminished activation of the MAP kinase Pathway in CD3-stimulated T lymphocytes from old mice.  
Gorgas G, Butch E, Guan KL, Miller RA.
Mechanism of Aging and Dev. (1997). 94, 71-83

Differential regulation of FUS3 MAP kinase by tryosine specific phosphatases PTP2/PTP3 and dual specificity phosphatase MSG5 in S. cerevisiae
Zhan X, Deschenes RJ, Guan KL.
Genes & Dev. (1997). 11, 1690-1702

Inducible expression of a mutant form of MEK1 in Swiss3T3 cells
Syu LJ, Guan KL, Saltiel AR.
J. Cell. Biochem. (1997). 67, 367-377

Cholecystokinin activates a MAP kinase cascade by multiple mechanisms in rat acinar cells
Dabrowski A, Groblewski GE, Schaefer C, Guan KL, Williams JA.
Am. J. Physiol. (1997). 273, C1472-1479

Suppression of W256 carcinosarcoma cell apoptosis by arachidonic acid and other polyunsaturated fatty acids.
Tang DG, Guan KL, Li L, Honn KV, Chen YQ, Rice RL, Taylor JD, Porter AT.
Int. J. Cancer. (1997). 72, 1078-1087

Signaling molecules involved in coupling growth hormone receptor to MAP kinase activation. 
VanderKuur JA, Butch E, Waters SB, Pessin JE, Guan KL, Carter-Su, C.
Endocrinology. (1997). 138,  4301-4307

The calcium/calmodulin dependent protein phosphatase calcineurin is the major Elk-1 phosphatase.
Sugimoto T, Scott S, Guan KL.
J. Biol. Chem. (1997). 272,  29415-29418


Elevated levels of ERK2 in human breast carcinoma MCF-7 cells transfected with protein kinase C. 
Gupta AK, Galoforo SS, Berns CM, Martinez AA, Corry PM, Guan KL, Lee YJ.
Cell Prolif. (1996). 29, 655-663

Molecular cloning of the CDK6 inhibitor p20 and demonstration of two distinct mechanisms of interaction with CDK proteins for two families of CDK inhibitors.
Guan KL
, Jenkins CW, Li Y, Zariwala M, Noh S, Wu X, Xiong Y.
Mol. Biol. Cell. (1996).  7, 57-70

Atrial natriuretic peptide induces the expression of MKP-1, a mitogen activated protein kinase phosphatase, in glomerular mesangial cells.
Sugimoto T, Haneda M, Togawa M, Isono M, Shikano T,  Araki SI, Nakahiko T, Kashiwagi S, Guan KL, Kikkawa R.
J. Biol. Chem. (1996). 271, 544-547

Selective activation of MEK-1 but not MEK-2 by A-raf from epidermal growth factor stimulated Hela cells.
Wu X, Noh S, Zhou G, Dixon JE, Guan KL.
J. Biol. Chem. (1996). 271, 3265-3271

Characterization of ERK1 activation site mutants and the effect on recognition by MEK1 and EMK2.  
Butch E, Guan KL. 
J. Biol. Chem.
 (1996). 4230-4235

Contributions of the mitogen-activated protein (MAP) kinase backbone and phosphorylation loop to MEK specificity.: 
Robinson MJ,  Cheng M,  Khokhlatchev A,  Ebert D,  Ahn N,  Guan KL,  Stein B, Goldsmith E, Cobb MH.
J. Biol Chem. (1996). 271, 29734-29739

Regulation and function of the mitogen activated protein kinase (MAPK) signal transduction pathway.
Wang Y, Guan KL.
Bio/Pharma Quaterly.  (1996). 1(4), 23-26


Isolation and characterization of a novel dual specific phosphatase, HVH2, which selectively dephosphorylates the mitogen activated protein kinase. 
Guan KL, Butch E.
J. Biol. Chem.
 (1995). 270, 7197-7203

Nerve growth factor stimulates a novel protein kinase in PC-12 cells that phosphorylates and activates MEK. 
Pang R, Zheng CF, Guan KL, Saltiel AR.
Biochem. J. (1995). 307, 513-519

Characterization of MEK1 phosphorylation by v-Mos protein.
Pham CD, Arlinghaus RB, Zheng CF, Guan KL, Singh B.
Oncogene (1995). 10, 1683-1688

Mek-2, a Caenorhabditis elegans MAP kinase kinase, functions in Ras-mediated vulval induction and other development events.
Wu Y, Han M, Guan KL.
Genes & Dev. (1995). 
9, 742-755

The Caenorhabditis elegans gene mek-2 is required for vulval induction and encodes a protein similar to the protein kinase MEK. 
Kornfeld K, Guan KL, Horvitz HR.
Genes & Dev. (1995). 9, 756-768

Activation of MAP kinase kinase (MEK) and Ras by cholecystokinin in rat pancreatic acini.
Rui D, Zheng CF, Guan KL, Williams JA.
Am. J. Physiol. (1995). 268, G1060-1065

Phosphorylation and activation of the ATP-Mg-Dependent Protein Phosphatase by the Mitogen Activated Protein Kinase.
Wang QM, Guan KL, Roach PJ, Depaoli-Roach AA.
J. Biol. Chem. (1995). 
270, 18352-18358

Purification and characterization of the low molecular weight protein tyrosine phosphatase, Stp1, from fission yeast S. pombe
Zhang ZY, Zhou G, Denu JM, Wu L, Tang X, Mondesert O, Russell P, Butch E, Guan KL.
Biochemistry. (1995). 34, 10560-10568

Three genes of the MAP kinase cascade, mek-2, mpk-1/sur-1 and let-60 ras, are required for meiotic cell cycle progression in Caenorhabditis elegans.
Church DL, Guan KL, Lambie EJ.
Development. (1995). 121, 2525-2535

Desensitization of Ras activation by a feedback disassociation of the SOS-Grb2 complex.  
Waters SB,  Holt KH,  Ross SE,  Syu LJ,  Guan KL,  Saltiel AR,  Koretzky GA, Pessin JE.
J. Biol. Chem.  (1995). 270, 20883-20886


Activation of MEK family kinases requires phosphorylation of two conserved Ser/Thr residues.
Zheng CF, Guan KL.
The EMBO J. (1994). 13, 1123-1131

Growth factor induced MEK activation is primarily mediated by an activator different from c-raf.  
Zheng CF, Ohmichi M, Saltiel AR, Guan KL.
Biochemistry. (1994). 33, 5595-5599

Cytoplasmic localization of the mitogen activated protein kinase activator, MEK.
Zheng CF, Guan, KL.
J. Biol. Chem. (1994). 269, 19947-19952

Simian virus 40 small T antigen cooperates with mitogen activated kinases to stimulate AP-1 activity.

Frost JA, Alberts AS, Sontag E, Guan KL, Mumby MC, Feraminsco JR.
Mol. Cell. Biol. (1994). 14, 6244-6252

Growth suppression by p18, a p16INK4/MTS1- and p14INK4B/MTS2-related CDK6 inhibitor, correlates with wild-type pRb function. 
Guan KL
, Jenkins CW, Li Y, Nichols MA, Wu X, Matera G, Xiong Y.
Genes & Dev. (1994). 8, 2939-2952


Improved vectors for expression and purification of recombinant proteins.
Dixon JE, Hakes D, Guan KL.
Methods in Genetics. (1993). 2, 44-53


Bacterial and viral protein tyrosine phosphatases.
Guan KL, Dixon JE.
Seminars in Cell Biology. (1993). 4, 389-396

A eukaryotic homolog to viral Ser/Thr/Tyr phosphatase.
Guan KL, Hakes D, Park HD, Cooper TG, Dixon JE.
Trends in Biochem. Sci. (1993). 18, 6

Cloning and characterization of two distinct human ERK activator kinases, MEK1 and MEK2,
Zheng CF, Guan KL.
J. Biol. Chem. (1993). 268, 11435-11439

Dephosphorylation and inactivation of the mitogen activated protein kinases by a mitogen induced protein phosphatase. 
Zheng CF, Guan KL.
J. Biol. Chem. (1993). 268, 16116-16119

Normal mitochondrial structure and genome maintenance in yeast requires the dynamine like product of the MGM1 gene. 
Guan KL, Farh L, Marshall TK, Deschenes RJ.
Current Genetics. (1993). 24, 141-148

Properties of MEKs, the kinases that phosphorylate and activate the extracellular signal-regulated kinases.
Zheng CF, Guan KL.
J. Biol. Chem. (1993). 268, 23933-23939


Isolation and characterization of a second protein tyrosine phosphatase gene, PTP2, from Saccharomyces cerevisiae
Guan KL, Deschenes RJ, Dixon JE.
J. Biol. Chem. (1992). 267, 10024-10020

A yeast protein phosphatase related to the vaccinia virus VH1 phosphatase is induced by nitrogen starvation. 
Guan KL
, Hakes D, Wang Y, Park HD, Cooper TG, Dixon JE.
Proc. Natl. Acad. Sci. USA. (1992). 89, 12175-12179

Negative regulation of mitosis by functionally overlapping PTPases in fission yeast.
Millar JBA, Russell P, Dixon JE, Guan KL.
The EMBO J. (1992). 11, 4943-4952


Evidence for protein tyrosine phosphatase catalysis proceeding Via a cysteine-phosphate intermediate.
Guan KL, Dixon JE.
J. Biol. Chem. (1991). 266, 17026-17030

Expression of rat PTP1 in normal and transformed cells. 
Woodford TA, Guan KL, Dixon JE.
Adv. Prot. Phosphatases. (1991). 6, 503-524

One step purification of eukaryotic proteins expressed in E. coli:  An improved thrombin cleavage and purification procedure of fusion proteins with glutathione-S-transferase.
Guan KL, Dixon JE.
Anal. Biochem. (1991). 192, 262-267

Tyrosine phosphate hydrolysis of proteins by an essential Yersiniavirulence determinant.  
Bliska JB, Guan KL, Dixon JE, Falkow SA.
Proc. Natl. Acad. Sci. USA (1991). 88, 1187-1191

A Tyr/Ser protein phosphatase encoded by vaccinia virus.
Guan KL, Broyles SS, Dixon JE.
Nature. (1991). 350, 359-362

Cloning and expression of a yeast protein tyrosine phosphatase.
Guan KL, Deschenes, RJ, Qiu H, Dixon JE.
J. Biol. Chem. (1991). 266, 12964-12970

De novo purine nucleotide biosynthesis:  cloning and expression of a eukaryotic cDNA encoding 5-aminoimidazole-4-carboxamide-ribonucleotide transformylase-IMP cyclohydrolase.
Ni LY, Guan KL, Zalkin H, Dixon JE.
Gene. (1991). 106, 197-20


Sequence of the precursor of bovine liver mitochondrial aldehyde dehydrogenase as determined from its cDNA and its gene and its functionality.
Guan KL
, Weiner H.
Arch. Biophys. Biochem. (1990). 277, 351-360

Cloning and expression of a protein-tyrosine-phosphatase.  
Guan KL, Haun RS, Watson SJ, Geahlen RL, Dixon JE.
Proc. Natl. Acad. Sci. USA. (1990). 87, 1501-1505

Protein tyrosine phosphatase activity of an essential virulence determinant in Yersinia
Guan KL,
 Dixon JE.
Science. (1990). 249, 553-556


Primary structures of rat and bovine liver mitochondrial aldehyde dehydrogenase deduced from cDNA sequences.  
Farres J, Guan KL, Weiner H.
Eur. J. Biochem. (1989). 180, 67-74

Influence of the 5’-end region of aldehyde dehydrogenase mRNA on translational efficiency: potential secondary structure inhibition of translation in vitro.  
Guan, K-L., and Weiner, H. (1989)
J. Biol. Chem. 264, 17764-17769.


Signal peptide sequence of rat liver mitochondrial aldehyde dehydrogenase deduced from cDNA.
Farres J, Guan KL, Weiner H.
Biochem. Biophys. Res. Comm. (1988). 150, 1083-1087

Purification and characterization of beef and pig liver aldehyde dehydrogenases.
Guan K., Pak YK, Tu GC, Cao, QN, Weiner H.
Alcoholism: Clin. Exp. Res. (1988). 12, 713-719


Molecular biology studies of beef liver aldehyde dehydrogenase in Enzymology and Molecular Biology of Carbonyl Metabolism: aldehyde dehydrogenase, aldo-keto reductase and alcohol dehydrogenase
Guan KL, Weiner H.
Weiner, H. & Flynn, T.G., eds. (1987). pp. 15-24, Alan R, Liss, Inc., New York.