From:  Targeting tumor transition windows

 Biological mechanisms supporting drug-tolerant persister survival during therapy.

Adaptive survival mechanismCellular processes involvedKey signaling pathwaysExperimental evidenceTherapeutic targeting strategies
Oxidative stress adaptationUpregulation of antioxidant defenses allowing tumor cells to tolerate therapy-induced reactive oxygen species and oxidative damageNRF2 signaling, glutathione metabolism, redox homeostasis regulators [90, 92]Observed in drug-tolerant persister populations in melanoma, lung cancer, and colorectal cancer models [93, 94]Targeting redox balance using reactive oxygen species (ROS)-inducing agents or inhibitors of antioxidant pathways
UPRActivation of proteostasis mechanisms that restore endoplasmic reticulum function and prevent accumulation of misfolded proteins under treatment stressProtein kinase R-like endoplasmic reticulum kinase (PERK)-eukaryotic translation initiation factor 2 alpha (eIF2α) pathway, activating transcription factor 4 (ATF4) signaling, inositol-requiring enzyme 1 (IRE1)-X-box binding protein 1 (XBP1) pathway [91, 95]Therapy-induced proteotoxic stress and UPR activation reported in multiple targeted therapy models [91, 95]Inhibition of ER stress response pathways or proteostasis regulators
Autophagy activationRecycling of intracellular components to sustain energy production and remove damaged organelles during therapeutic stressAMPK signaling, mTOR suppression, autophagy initiation complexes [84, 96]Increased autophagy activity documented in persister cells following chemotherapy and targeted therapy [84, 96]Autophagy inhibitors combined with targeted therapy or chemotherapy
Metabolic rewiringAdaptive metabolic reprogramming enabling tumor cells to maintain energy production under therapy-induced metabolic stressAMPK signaling, mitochondrial stress pathways, altered glucose and glutamine metabolism [68, 97]Therapy-induced metabolic dependencies observed in targeted therapy-resistant tumor models [90, 91]Targeting metabolic dependencies such as mitochondrial metabolism or nutrient utilization pathways
Adaptive signaling rewiringActivation of compensatory signaling pathways that restore survival signaling after inhibition of oncogenic driversReceptor tyrosine kinase activation, PI3K-AKT signaling, MAPK pathway reactivation [87, 98]Observed in EGFR-mutant lung cancer, BRAF-mutant melanoma, and other targeted therapy models [87, 88]Sequential or combination therapies targeting compensatory pathways
Chromatin remodeling and epigenetic adaptationEpigenetic reprogramming enabling reversible drug-tolerant states through transcriptional reorganizationHistone modification enzymes, chromatin remodeling complexes, enhancer reprogramming [68, 99]Chromatin remodeling associated with reversible drug-tolerant persister states across multiple tumor models [68, 99]Targeting epigenetic regulators such as histone modifiers or chromatin remodeling complexes

AMPK: AMP-activated protein kinase; BRAF: B-Raf proto-oncogene; EGFR: epidermal growth factor receptor; NRF2: nuclear factor erythroid 2-related factor 2; MAPK: mitogen-activated protein kinase; PI3K-AKT: phosphoinositide 3-kinase-protein kinase B; UPR: unfolded protein response.