Cited by 269PI3K/AKT/mTOR signaling in cancerMetabolism, Diabetes, and CancerTuberous Sclerosis Complex Research

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Diverse signaling mechanisms of mTOR complexes: mTORC1 and mTORC2 in forming a formidable relationship

Jhanwar-Uniyal, Meena, Wainwright, John V., Mohan, Avinash L., Tobias, Michael E., Murali, Raj, Gandhi, Chirag D., Schmidt, Meic H.·2019·Advances in Biological Regulation

DOI 10.1016/j.jbior.2019.03.003

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Abstract

Activation of Mechanistic target of rapamycin (mTOR) signaling plays a crucial role in tumorigenesis of numerous malignancies including glioblastoma (GB). The Canonical PI3K/Akt/mTOR signaling cascade is commonly upregulated due to loss of the tumor suppressorm PTEN, a phosphatase that acts antagonistically to the kinase (PI3K) in conversion of PIP2 to PIP3. mTOR forms two multiprotein complexes, mTORC1 and mTORC2 which are composed of discrete protein binding partners to regulate cell growth, motility, and metabolism. These complexes are sensitive to distinct stimuli, as mTORC1 is sensitive to nutrients while mTORC2 is regulated via PI3K and growth factor signaling. The main function of mTORC1 is to regulate protein synthesis and cell growth through downstream molecules: 4E-BP1 (also called EIF4E-BP1) and S6K. On the other hand, mTORC2 is responsive to growth factor signaling by phosphorylating the C-terminal hydrophobic motif of some AGC kinases like Akt and SGK and it also plays a crucial role in maintenance of normal and cancer cells through its association with ribosomes, and is involved in cellular metabolic regulation. mTORC1 and mTORC2 regulate each other, as shown by the fact that Akt regulates PRAS40 phosphorylation, which disinhibits mTORC1 activity, while S6K regulates Sin1 to modulate mTORC2 activity. Allosteric inhibitors of mTOR, rapamycin and rapalogs, remained ineffective in clinical trials of Glioblastoma (GB) patients, in part due to their incomplete inhibition of mTORC1 as well as unexpected activation of mTOR via the loss of negative feedback loops. In recent years, novel ATP binding inhibitors of mTORC1 and mTORC2 suppress mTORC1 activity completely by total dephosphorylation of its downstream substrate pS6K Ser235/236, while effectively suppressing mTORC2 activity, as demonstrated by complete dephosphorylation of pAKT Ser473. Furthermore by these novel combined mTORC1/mTORC2 inhibitors reduced the proliferation and self-renewal of GB cancer stem cells. However, a search of more effective way to target mTOR has generated a third generation inhibitor of mTOR, "Rapalink", that bivalently combines rapamycin with an ATP-binding inhibitor, which effectively abolishes the mTORC1 activity. All in all, the effectiveness of inhibitors of mTOR complexes can be judged by their ability to suppress both mTORC1/mTORC2 and their ability to impede both cell proliferation and migration along with aberrant metabolic pathways.

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Background
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Background
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Background
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Background
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“It is intriguing that upregulation of the active form of abovementioned signals (FGFR1, mTOR, p70S6K, p-S473-AKT, and p-S262-Cx43) is also linked to cell proliferation (Doble et al. 2004; Jhanwar-Uniyal et al. 2019; Sheikh et al. 1999).”
Background
A preclinical report of a cobimetinib-inspired novel anticancer small-molecule scaffold of isoflavones, NSC777213, for targeting PI3K/AKT/mTOR/MEK in multiple cancers.
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Background
The female syndecan-4−/− heart has smaller cardiomyocytes, augmented insulin/pSer473-Akt/pSer9-GSK-3β signaling, and lowered SCOP, pThr308-Akt/Akt and GLUT4 levels
Støle, T. P., Lunde, M., Shen, X. M., Martinsen, M., Lunde, P., Li, J., Lockwood, F., Sjaastad, I., Louch, W. E., Aronsen, J. M., et al. (2022) · Frontiers in Cell and Developmental Biology · doi:10.3389/fcell.2022.908126 · Cited by 9
“mTORc1 is a rapamycin-sensitive protein complex consisting of mTOR, mLST8 and raptor, while mTORc2, upstream of Akt activation, includes mTOR, mLST8, rictor, mSin1 and protor (Jhanwar-Uniyal et al., 2019).”
Background
Combination Treatment Targeting mTOR and MAPK Pathways Has Synergistic Activity in Multiple Myeloma
Sun, K., Jin, L., Karolová, J., Vorwerk, J., Hailfinger, S., Opalka, B., Zapukhlyak, M., Lenz, G., & Khandanpour, C. (2023) · Cancers · doi:10.3390/cancers15082373 · Cited by 9
“As a crucial downstream effector of the PI3K pathway, mTOR exists in two different forms: mTOR complex 1 (mTORC1) and mTORC2, which have distinct functions [14].”
Background
Circular RNAs: characteristics, functions, mechanisms, and potential applications in thyroid cancer
Wu, M., Yuan, H., Zou, W., Xu, S., Liu, S., Gao, Q., Guo, Q., Han, Y. P., & An, X. (2023) · Clinical & Translational Oncology · doi:10.1007/s12094-023-03324-0 · Cited by 12
“Subsequently, Akt activation translates to the phosphorylation of downstream effectors, such as mTOR [92].”
Design, Synthesis and Gene Modulation Insights into Pigments Derived from Tryptophan-Betaxanthin, Which Act against Tumor Development in Caenorhabditis elegans
Henarejos‐Escudero, P., Méndez-García, F. F., Hernández‐García, S., Martínez‐Rodríguez, P., & Gandía‐Herrero, F. (2023) · International Journal of Molecular Sciences · doi:10.3390/ijms25010063 · Cited by 8
“In fact, the down-regulation of RICTOR and/or RAPTOR was shown to be a promising approach for cancer therapy [29].”
Background
Exploiting new strategies in combating head and neck carcinoma: A comprehensive review on phytochemical approaches passing through PI3K /Akt/ mTOR signaling pathway
Iranpanah, A., Majnooni, M. B., Biganeh, H., Amirian, R., Rastegari‐Pouyani, M., Filosa, R., Cheang, W. S., Fakhri, S., & Khan, H. Z. (2024) · Phytotherapy Research · doi:10.1002/ptr.8228 · Cited by 8
“…(Hua et al., 2019). mTORC1 and mTORC2 act downstream of PI3K/Akt signaling orchestrating essential cellular processes (Figlin et al., 2013; Jhanwar-Uniyal et al., 2019). mTORC2 provokes the activation of Akt, protein kinase C alpha (PKC α ), eukaryotic translation initiation factor 4E…”
Deciphering the role of cell signaling pathways in gout pathogenesis and the therapeutic potential of phytoconstituents in their modulation
Dey, H., Ali, S. S., & Sethi, V. A. (2025) · Inflammopharmacology · doi:10.1007/s10787-025-01741-x · Cited by 9
“…actuates the activation of two separate complexes, mTORC1 and mTORC2, which regulate protein synthesis, cell proliferation, and immune responses (Jhanwar-Uniyal et al. 2019). mTORC1 plays a role in the production of pro-inflammatory cytokines (IL-1β, TNF-α, or IL-6) necessary to fuel the…”
Synergistic pathways in Parkinson's disease: The promise of FGF21 and ACE2
Liu, T., Li, J., Yang, Z., & Wei, J. (2025) · Ageing Research Reviews · doi:10.1016/j.arr.2025.102804 · Cited by 8
Targeted Deletion of PTEN in Kisspeptin Cells Results in Brain Region- and Sex-Specific Effects on Kisspeptin Expression and Gonadotropin Release
Negrón, A. L., Yu, G., Boehm, U., & Acosta‐Martínez, M. (2020) · International Journal of Molecular Sciences · doi:10.3390/ijms21062107 · Cited by 8
“mTORC2 downstream targets include Akt and the serumand glucocorticoid-related kinase (SGK), which are part of growth factor signaling pathways [35].”
Background
Identification of glioblastoma‑specific prognostic biomarkers via an integrative analysis of DNA methylation and gene expression
Mao, Y. X., Liu, Z., & Cai, L. (2020) · Oncology Letters · doi:10.3892/ol.2020.11729 · Cited by 9
“Despite great efforts, little progress has been made in the treatment of GBM over the past decade (6,7).”
Background
Gene and protein expression of mTOR and LC3 in hepatocellular carcinoma, colorectal liver metastasis and “normal” liver tissues
Bortolami, M., Comparato, A., Benna, C., Errico, A., Maretto, I., Pucciarelli, S., Cillo, U., & Farinati, F. (2020) · PLoS ONE · doi:10.1371/journal.pone.0244356 · Cited by 8
Fluorescence Imaging of Epidermal Growth Factor Receptor Tyrosine Kinase Inhibitor Resistance in Non-Small Cell Lung Cancer
Martin-Fernandez, M. L. (2022) · Cancers · doi:10.3390/cancers14030686 · Cited by 8
Rictor, an mTORC2 Protein, Regulates Murine Lymphatic Valve Formation Through the AKT-FOXO1 Signaling
Banerjee, R., Knauer, L. A., Iyer, D., Barlow, S. E., Shalaby, H., Dehghan, R., Scallan, J. P., & Yang, Y. (2024) · Arteriosclerosis Thrombosis and Vascular Biology · doi:10.1161/atvbaha.124.321164 · Cited by 7
“23 RIC-TOR is a critical component in the multiprotein mTORC2 complex, and its interaction with the mTOR protein is crucial for Ser473 (Serine 473) phosphorylation of AKT.”
Background
mTOR in the Development of Hypoxic Pulmonary Hypertension Associated with Cardiometabolic Risk Factors
Flores, K., Almeida, C., Arriaza, K., Pena, E., & Alam, S. E. (2024) · International Journal of Molecular Sciences · doi:10.3390/ijms252011023 · Cited by 9
The TXNIP–mTOR–Autophagy axis in diabetic kidney disease: mechanistic insights and therapeutic implications
Zhu, Q., Wu, X., Di, Y., Liu, L., & Liu, Y. (2025) · Molecular Biology Reports · doi:10.1007/s11033-025-10946-w · Cited by 8
“…mTORC2 is mainly activated by growth factors such as insulin and plays critical roles in regulating cytoskeletal dynamics, glucose metabolism, and cell survival. mTORC2 activates members of the AGC kinase family, particularly Akt (via Ser473 phosphorylation), SGK1, and certain PKC isoforms [ 46].”
Quercetin prevents sarcopenia by reversing oxidative stress and mitochondrial damage
Sun, J., Liu, H., Yan, Y., & Fang, F. (2025) · Journal of Molecular Histology · doi:10.1007/s10735-025-10411-9 · Cited by 7
“It’s important to note that mTOR exists in two main complexes, mTORC1 and mTORC2, which may have distinct functions and expression patterns within the PI3K/AKT signaling pathway(Jhanwar-Uniyal et al. 2019 ).”
Transcriptomic analysis reveals that mTOR pathway can be modulated in macrophage cells by the presence of cryptococcal cells
Piffer, A. C., dos Santos, F. M., Thomé, M. P. M., Diehl, C., Garcia, A. W. A., Kinskovski, U. P., Schneider, R., Gerber, A. L., Feltes, B. C., Schrank, A., et al. (2021) · Genetics and Molecular Biology · doi:10.1590/1678-4685-gmb-2020-0390 · Cited by 7
Mammalian Target of Rapamycin as the Therapeutic Target of Vascular Proliferative Diseases: Past, Present, and Future
Huang, W.-Q., Zou, Y., Tian, Y., Ma, X.-F., Zhou, Q.-Y., Li, Z.-Y., Gong, S.-X., & Wang, A.-P. (2022) · Journal of Cardiovascular Pharmacology · doi:10.1097/fjc.0000000000001208 · Cited by 7
The self-renewal dental pulp stem cell microtissues challenged by a toxic dental monomer
Kaufman, G., Kiburi, N. M., & Škrtić, D. (2020) · Bioscience Reports · doi:10.1042/bsr20200210 · Cited by 7
“The suppression leads to a decrease in cell growth rate and a reduction of cell volume [49].”
Insulin degrading enzyme (IDE) expressed by Chinese hamster ovary (CHO) cells is responsible for degradation of insulin in culture media
Louie, S., Lakkyreddy, J., Castellano, B. M., Haley, B., Dang, A. N., Lam, C., Tang, D., Lang, S. E., Snedecor, B., & Misaghi, S. (2020) · Journal of Biotechnology · doi:10.1016/j.jbiotec.2020.04.016 · Cited by 7
The GLUT1–mTORC1 axis affects odontogenic differentiation of human dental pulp stem cells
Liu, L., Xie, H., Zhao, S., & Huang, X. (2022) · Tissue and Cell · doi:10.1016/j.tice.2022.101766 · Cited by 7
The genomic mosaic of mitochondrial dysfunction: Decoding nuclear and mitochondrial epigenetic contributions to maternally inherited diabetes and deafness pathogenesis
Donato, L., Scimone, C., Alibrandi, S., Vadalà, M., Castellucci, M., Bonfiglio, V. M. E., Scalinci, S. Z., Abate, G., D’Angelo, R., & Sidoti, A. (2024) · Heliyon · doi:10.1016/j.heliyon.2024.e34756 · Cited by 6
mTOR Regulates Mineralocorticoid Receptor Transcriptional Activity by ULK1-Dependent and -Independent Mechanisms
Ali, Y. M., Gómez-Sánchez, C. E., Plonczynski, M. W., Náray‐Fejes‐Tóth, A., Fejes‐Tóth, G., & Gómez-Sánchez, E. P. (2024) · Endocrinology · doi:10.1210/endocr/bqae015 · Cited by 6
Acute Conditioning of Antigen-Expanded CD8+ T Cells via the GSK3β-mTORC Axis Differentially Dictates Their Immediate and Distal Responses after Antigen Rechallenge
Táborská, P., Stakheev, D., Svobodová, H., Strizova, Z. O., Bartůňková, J., & Smrž, D. (2020) · Cancers · doi:10.3390/cancers12123766 · Cited by 6
“GSK3β is known to be a regulator of mTOR signaling complexes, mTOR complex 1 (mTORC1) and mTOR complex 2 (mTORC2) [40].”
Background
A scoping review regarding reproductive capacity modulation based on alpha-ketoglutarate supplementation
Doroftei, B., Ilie, O., Timofeiov, S., Dabuleanu, A., Scripcariu, I. S., Micu, R., & Tataranu, E. (2024) · Reproduction · doi:10.1530/rep-24-0137 · Cited by 4
“Both play a role in growth and metabolism, primarily found in the nucleus and cytoplasm (Saxton & Sabatini 2017, Jhanwar-Uniyal et al. 2019), while AMPK serves as a key energy sensor in aging and lifespan (Hardie et al. 2012, Huang et al. 2013), collectively promoting lysosomal translocation…”
Interaction between the PI3K/AKT pathway and mitochondrial autophagy in macrophages and the leukocyte count in rats with LPS-induced pulmonary infection
Wu, C., Guo, L., Muhataer, X., Li, Q., Lian, Z., Li, Y., Wang, W., Ding, W., Zhou, Y. M., Yang, X., et al. (2023) · Open Life Sciences · doi:10.1515/biol-2022-0588 · Cited by 6
AMPK–mTORC1 pathway mediates hepatic IGFBP-1 phosphorylation in glucose deprivation: a potential molecular mechanism of hypoglycemia-induced impaired fetal growth
Kakadia, J. H., Khalid, M. U., Heinemann, I. U., & Han, V. K. M. (2024) · Journal of Molecular Endocrinology · doi:10.1530/jme-23-0137 · Cited by 5
“…and mTORC1 was inhibited by glucose deprivation. mTORC1 is a regulator of protein synthesis, mitochondrial function, nutrient transport and autophagy in response to signaling by nutrients, oxygen and growth factors, thereby modulating cell proliferation and growth (Jhanwar-Uniyal et al. 2019).”
ITGB2 fosters the cancerous characteristics of ovarian cancer cells through its role in mitochondrial glycolysis transformation
Li, G., Qiu, J., Lu, X., & Jin, Y. (2024) · Aging · doi:10.18632/aging.205529 · Cited by 5
Cerebellar phenotypes in germline PTEN mutation carriers
Gambini, D., Ferrero, S., Bulfamante, G., Pisani, L., Corbo, M., & Kuhn, E. (2024) · Neuropathology and Applied Neurobiology · doi:10.1111/nan.12970 · Cited by 5
“A synthetic view of the components involved in the pathway and their relative functions is summarised in Figure 2 [106 – 108].”
Background
mTOR inhibition by TAK‐228 is effective against growth, survival and angiogenesis in preclinical retinoblastoma models
Tang, L., Fu, Y., Song, J., Hu, T., Li, K., & Li, Z. (2022) · Pharmacology Research & Perspectives · doi:10.1002/prp2.930 · Cited by 5
Rapamycin attenuated podocyte apoptosis via upregulation of nestin in Ang II-induced podocyte injury
Shi, H., Zhao, Y., He, T., Wen, X., Qu, G., Li, S., Gan, W., & Zhang, A. (2022) · Molecular Biology Reports · doi:10.1007/s11033-021-07029-x · Cited by 6
“mTOR is the catalytic subunit of two distinct protein complexes, mTORC1 and mTORC2, which can be distinguished by their unique composition and different substrates [28].”
Background
Mammalian Target of Rapamycin Inhibition Decreases Angiotensin II-Induced Steroidogenesis in HAC15 Human Adrenocortical Carcinoma Cells
Ali, Y. M., Gómez-Sánchez, E. P., & Gómez-Sánchez, C. E. (2022) · Endocrinology · doi:10.1210/endocr/bqac185 · Cited by 5
Does β-Hydroxy-β-Methylbutyrate Have Any Potential to Support the Treatment of Duchenne Muscular Dystrophy in Humans and Animals?
Gorji, A. E., Ostaszewski, P., Urbańska, K., & Sadkowski, T. (2023) · Biomedicines · doi:10.3390/biomedicines11082329 · Cited by 6
Programmed Cell Death and Its Therapeutic Implications in Parkinson's Disease: A Review of Apoptosis, Autophagy, Ferroptosis, and Necroptosis
Erekat, N. S. (2025) · Clinical Anatomy · doi:10.1002/ca.70057 · Cited by 4
Crosstalk between dysregulated amino acid sensing and glucose and lipid metabolism in colorectal cancer
Wang, D., Zou, S., Ding, J., Gao, C., Wang, J., & Tang, Z. (2025) · Frontiers in Oncology · doi:10.3389/fonc.2025.1665056 · Cited by 5
Insulin Receptor Genetic Variants Causal Association with Type 2 Diabetes: A Mendelian Randomization Study
Soliman, G. A., & Schooling, C. M. (2022) · Current Developments in Nutrition · doi:10.1093/cdn/nzac044 · Cited by 5
The role of leucine in the activation of cellular metabolism: a large integrative review
Gebrin, A. S., & Filho, I. J. Z. (2022) · International Journal of Nutrology · doi:10.54448/ijn22s201 · Cited by 4
Liver impact of growth hormone (GH) intermittent treatment during the growth period in mice
Piazza, V. G., Cicconi, N. S., Martinez, C. S., Dominici, F. P., Miquet, J. G., & Sotelo, A. I. (2023) · Molecular and Cellular Endocrinology · doi:10.1016/j.mce.2023.111911 · Cited by 4
Recent advances to Neuroprotection: repurposing drugs against neuroinflammatory disorders
Bilal, W., Khawar, M. B., Afzal, A., Naseer, A., Hamid, S. E., Shahzaman, S., & Qamar, F. (2023) · Molecular Biology Reports · doi:10.1007/s11033-023-08490-6 · Cited by 4
Redistribution of SOD3 expression due to R213G polymorphism affects pulmonary interstitial macrophage reprogramming in response to hypoxia
Lewis, C., Garcia, A. M., Burciaga, S. D., Posey, J. N., Jordan, M., Nguyen, T.-T. N., Stenmark, K. R., Mickael, C. S., Sul, C., Delaney, C. A., et al. (2024) · Physiological Genomics · doi:10.1152/physiolgenomics.00078.2024 · Cited by 4
Effects of air exposure time on tissue morphology, energy metabolism, and non-specific immunity of the juvenile Chinese mitten crab, Eriocheir sinensis
Zhao, A., Zhang, Q., Sun, Y., & Cheng, Y. (2025) · Comparative Biochemistry and Physiology Part B Biochemistry and Molecular Biology · doi:10.1016/j.cbpb.2025.111090 · Cited by 4
Reconsidering phosphorylation in the control of inducible CARD11 scaffold activity during antigen receptor signaling
Pomerantz, J. L. (2020) · Advances in Biological Regulation · doi:10.1016/j.jbior.2020.100775 · Cited by 4
“Many critical signaling pathways rely on regulated protein phosphorylation for the precise temporal control of the signaling output that determines the cellular response to external stimuli (please see for example (Hirsch et al., 2020; Jhanwar-Uniyal et al., 2019; Rahimova et al., 2020)).”
Background
Proteomics and Phospho-Proteomics Profiling of the Co-Formulation of Type I and II Interferons, HeberFERON, in the Glioblastoma-Derived Cell Line U-87 MG
Vázquez‐Blomquist, D., Hardy-Sosa, A., Baez, S. D. L. C., Besada, V., Palomares, S., Guirola, O., Ramos, Y., Wiśniewski, J. R., González, L. J., & Rivero, I. B. (2022) · Cells · doi:10.3390/cells11244068 · Cited by 4
“TSC/Rheb signaling promotes mTOR S1261 phosphorylation, then promotes mTORC1-mediated autophosphorylation and substrate phosphorylation to stimulate cell growth [47, 48].”
Background
Comparative genomic landscape of lower-grade glioma and glioblastoma
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“AKT/mTOR pathway promotes proliferation, survival and migration [58].”
Background
The contribution of mTOR to the immunopathology of bipolar disorder.
Bertollo, A. G., Jordan, V. A., Capitanio, M., & Ignácio, Z. M. (2025) · Journal of Neuroimmunology · doi:10.1016/j.jneuroim.2025.578744 · Cited by 2
Rictor maintains endothelial integrity under shear stress
Li, H., Zhou, W., Liu, Y., Xia, Y., Xia, C., Pan, D.-R., Li, Z., Shi, Y., Chen, S., & Zhang, J. (2022) · Frontiers in Cell and Developmental Biology · doi:10.3389/fcell.2022.963866 · Cited by 3
“Relative to mTORC1, which has been extensively studied in the control of cell growth, proliferation, autophagy, and glycolipid metabolism (Jhanwar-Uniyal et al., 2019), relatively little research has been performed on mTORC2.”
Background

Authors: Jhanwar-Uniyal, M., Wainwright, J. V., Mohan, A. L., Tobias, M. E., Murali, R., Gandhi, C. D., & Schmidt, M. H.
Affiliations: Westchester Medical Center (US)
ORCID: Wainwright, John V.
Gandhi, Chirag D.
Schmidt, Meic H.
Venue: Advances in Biological Regulation
Publisher: Elsevier BV
Published: 2019-05
Volume: 72
Pages: 51-62
ISSN: 2212-4926 (ISSN-L)
DOI: 10.1016/j.jbior.2019.03.003
OpenAlex: W2939056166
Cited by: 268
References: 94
Citations by year: 2026: 20 · 2025: 32 · 2024: 37 · 2023: 38 · 2022: 52 · 2021: 44 · 2020: 40 · 2019: 5
FWCI: 3.10
Type: journal article
Language: en
Update notices: erratum (2021-12-01)
Open access: closed
Licence: www.elsevier.com/tdm/userlicense/1.0/ (tdm)
www.elsevier.com/legal/tdmrep-license (tdm)
Locations: Advances in Biological Regulation (publishedVersion)
PubMed (publishedVersion)
Touro Scholar (Touro College) (submittedVersion)
Funders: Advanced Research Foundation
Topics: Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › PI3K/AKT/mTOR signaling in cancer
Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Metabolism, Diabetes, and Cancer
Health Sciences › Medicine › Genetics (clinical) › Tuberous Sclerosis Complex Research
Keywords: mTORC2, PI3K/Akt signaling, glioblastoma, PTEN loss, 4E-BP1, S6K, PRAS40, SIN-1, rapamycin, cancer stem cells, drug resistance, cell proliferation, cell motility, cell migration, cell invasion, cell adhesion
MeSH terms: Mechanistic Target of Rapamycin Complex 1, Mechanistic Target of Rapamycin Complex 2, Animals, Humans, Signal Transduction, Cell Proliferation, Proto-Oncogene Proteins c-akt

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Diverse signaling mechanisms of mTOR complexes: mTORC1 and mTORC2 in forming a formidable relationship

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