Primary Hallmarks (Direct Causes of Cellular Damage)
* Genomic Instability: Accumulation of DNA damage from endogenous (e.g., reactive oxygen species/ROS) and exogenous sources, including mutations, deletions, chromosomal rearrangements, and impaired DNA repair mechanisms. This leads to errors in genetic information over time.
* Telomere Attrition: Progressive shortening of telomeres (protective caps on chromosome ends) with each cell division, eventually triggering DNA damage responses, replicative senescence, or cell death. Telomerase activity often declines.
* Epigenetic Alterations: Changes in DNA methylation patterns (often global hypomethylation with site-specific hypermethylation), histone modifications (e.g., altered acetylation/methylation), and chromatin remodeling. This causes "epigenetic drift," dysregulating gene expression without altering the DNA sequence.
* Loss of Proteostasis: Decline in the systems maintaining protein quality control, including chaperone-mediated folding, ubiquitin-proteasome degradation, and autophagy. Results in accumulation of misfolded, aggregated, or damaged proteins (e.g., amyloid-like aggregates).
* Disabled Macroautophagy: Reduced efficiency of autophagy (cellular "recycling" of damaged organelles and proteins via lysosomes). This exacerbates buildup of cellular debris and dysfunctional components.