Citation: Moretti, I.F.; Lerario, A.M.;
Sola, P.R.; Macedo-da-Silva, J.;
Baptista, M.d.S.; Palmisano, G.;
Oba-Shinjo, S.M.; Marie, S.K.N. GBM
Cells Exhibit Susceptibility to
Metformin Treatment According to
TLR4 Pathway Activation and
Metabolic and Antioxidant Status.
Cancers 2023, 15, 587.
https://doi.org/10.3390/
cancers15030587
Academic Editor: Edward Pan
Received: 20 December 2022
Revised: 10 January 2023
Accepted: 14 January 2023
Published: 18 January 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
cancers
Article
GBM Cells Exhibit Susceptibility to Metformin Treatment
According to TLR4 Pathway Activation and Metabolic and
Antioxidant Status
Isabele Fattori Moretti
1,
* , Antonio Marcondes Lerario
2
, Paula Rodrigues Sola
1
, Janaína Macedo-da-Silva
3
,
Mauricio da Silva Baptista
4
, Giuseppe Palmisano
3
, Sueli Mieko Oba-Shinjo
1
and Suely Kazue Nagahashi Marie
1,
*
1
Laboratory of Molecular and Cellular Biology (LIM 15), Department of Neurology, Faculdade de Medicina
FMUSP, Universidade de Sao Paulo, Sao Paulo 01246-903, SP, Brazil
2
Department of Internal Medicine, Division of Metabolism, Endocrinology and Diabetes, University of
Michigan, Ann Arbor, MI 48108, USA
3
GlycoProteomics Laboratory, Department of Parasitology, ICB, University of Sao Paulo,
São Paulo 05508-000, SP, Brazil
4
Biochemistry Department, Institute of Chemistry, Universidade de Sao Paulo,
São Paulo 05508-900, SP, Brazil
* Correspondence: imortetti@usp.br (I.F.M.); sknmarie@usp.br (S.K.N.M.); Tel.: +55-11-3061-7458 (S.K.N.M.)
Simple Summary: An analysis of metformin (MET) treatment in combination with temozolomide
(TMZ) in two glioblastoma cell lines, U87MG and A172, stimulated with lipopolysaccharide (LPS),
a TLR4 agonist was conducted. Both cells presented blunted mitochondrial respiration leading
to oxidative stress after MET treatment, and decreased cell viability after MET + TMZ treatment.
U87MG cells presented increased apoptosis after MET + LPS + TMZ treatment by increment of ER
stress, and downregulation of BLC2. A172, with an upregulated antioxidant background, including
SOD1, exhibited cell cycle arrest after MET + TMZ treatment. The observed differential response
was associated with a distinct metabolic status: glycolytic/plurimetabolic (GPM) subtype in U87MG
and mitochondrial (MTC) in A172. TCGA-GBM-RNASeq in silico analysis showed that GPM-GBM
cases with an activated TLR4 pathway might respond to MET, but the concomitant CXCL8/IL8
upregulation may demand a combination treatment with an IL8 inhibitor. MET combined with an
antioxidant inhibitor, such as anti-SOD1, may be indicated for MTC-GBM cases.
Abstract: Glioblastoma (GBM) is an aggressive brain cancer associated with poor overall survival.
The metabolic status and tumor microenvironment of GBM cells have been targeted to improve ther-
apeutic strategies. TLR4 is an important innate immune receptor capable of recognizing pathogens
and danger-associated molecules. We have previously demonstrated the presence of TLR4 in GBM
tumors and the decreased viability of the GBM tumor cell line after lipopolysaccharide (LPS) (TLR4
agonist) stimulation. In the present study, metformin (MET) treatment, used in combination with
temozolomide (TMZ) in two GBM cell lines (U87MG and A172) and stimulated with LPS was ana-
lyzed. MET is a drug widely used for the treatment of diabetes and has been repurposed for cancer
treatment owing to its anti-proliferative and anti-inflammatory actions. The aim of the study was to
investigate MET and LPS treatment in two GBM cell lines with different metabolic statuses. MET
treatment led to mitochondrial respiration blunting and oxidative stress with superoxide production
in both cell lines, more markedly in U87MG cells. Decreased cell viability after MET + TMZ and
MET + LPS + TMZ treatment was observed in both cell lines. U87MG cells exhibited apoptosis after
MET + LPS + TMZ treatment, promoting increased ER stress, unfolded protein response, and BLC2
downregulation. LPS stimulation of U87MG cells led to upregulation of SOD2 and genes related to
the TLR4 signaling pathway, including IL1B and CXCL8. A172 cells attained upregulated antioxidant
gene expression, particularly SOD1, TXN and PRDX1-5, while MET treatment led to cell-cycle arrest.
In silico analysis of the TCGA-GBM-RNASeq dataset indicated that the glycolytic plurimetabolic
(GPM)-GBM subtype had a transcriptomic profile which overlapped with U87MG cells, suggesting
Cancers 2023, 15, 587. https://doi.org/10.3390/cancers15030587 https://www.mdpi.com/journal/cancers