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![HLA-A*02:01 allele-restricted peptide
presentation as a driver of intratumoral heterogeneity. Tumor cells
expressing the same HLA allele (in this case the HLA-A*02:01
allele) can present distinct repertoires of tumor-derived peptides,
generating multiple layers of intratumoral heterogeneity. (A,B)
Neoantigen-encoding somatic mutations are frequently subclonal or
unevenly expressed across tumor regions, resulting in spatially
variable peptide availability for HLA class I presentation. (C)
Even when a mutation is present, defects in antigen-processing and
presentation pathways, including proteasomal cleavage, peptide
trimming, transporter associated with antigen processing-mediated
transport, endoplasmic reticulum loading and endoplasmic reticulum
aminopeptidase editing, negatively affect the stability of
peptide-HLA class I complex and its display at the cell surface.
(D) Sine peptide presentation is HLA allele-specific,
allele-selective loss or downregulation of HLA class I molecules
(such as via loss of heterozygosity or other MHC-I alterations)
produces tumor subclones that no longer present the peptide,
enabling immune-mediated selection and spatial or temporal
segregation of presenting and non-presenting cells. Under sustained
T-cell-mediated immune pressure, tumors may acquire defects in
antigen-presentation pathways, such as (E) HLA loss or (F)
β2-microglobulin mutations, further increasing phenotypic
heterogeneity and promoting resistance to T-cell-based therapies
[Created in BioRender. Fortis, S. (2026) https://BioRender.com/mmsl3zm]. Agreement license
WF29LKNSK4. HLA, human leukocyte antigen; MHC, major
histocompatibility complex.](/article_images/ijo/69/3/ijo-69-03-05917-g00.jpg)
![HLA-A*24:02-mediated presentation of
immunogenic tumor peptides drives potent CD8+ T cell
antitumor immunity. The schematic depicts the HLA-A*24:02 allele
effectively binding and presenting tumor-derived immunogenic
peptides on the cell surface, facilitating recognition by
CD8+ T cells. Peptides produced via intracellular
antigen processing are loaded onto HLA-A24 molecules, forming
stable peptide-HLA complexes that drive strong activation, clonal
expansion and effector differentiation of tumor-specific
CD8+ T cells. This coordinated process of antigen
presentation and CD8+ T-cell activation enhances
cytotoxic activity, resulting in efficient tumor cell elimination
[Created in BioRender. Fortis, S. (2026) https://BioRender.com/3v0nh8u]. Agreement license
GD29LKO1LQ. HLA, human leukocyte antigen.](/article_images/ijo/69/3/ijo-69-03-05917-g01.jpg)
![HLA-A*24:02 may preferentially
present conserved or essential 'driver' antigens as targets with
low heterogeneity. HLA-A*24:02 may preferentially facilitate the
presentation of conserved, clonally expressed tumor peptides with
high MHC class I binding affinity, leading to the activation and
expansion of CD8+ cytotoxic T cells bearing T-cell
receptors specific for these epitopes. Consequently, tumors
expressing HLA-A*24:02 remain readily detectable by activated
cytotoxic CD8+ T cells, which can efficiently recognize
and eliminate them [Created in BioRender. Fortis, S. (2026)
https://BioRender.com/lwwc6qi].
Agreement license KY29LKV4HJ. HLA, human leukocyte antigen; MHC,
major histocompatibility complex.](/article_images/ijo/69/3/ijo-69-03-05917-g02.jpg)
![HLA-A*24:02-mediated immunogenicity
and durable immune control in PCa. Schematic representation of the
proposed mechanism by which HLA-A*24:02 expression confers
favorable clinical outcomes in PCa. HLA-A*24:02-positive tumor
cells maintain sustained presentation of tumor-derived peptides
throughout disease evolution, resulting in enhanced immunogenicity
and persistent activation of tumor-specific CD8+ T-cell
memory. This continuous immune recognition supports long-term
immunosurveillance and establishes an extended state of immune
'equilibrium,' characterized by reduced tumor growth rates and
improved OS [Created in BioRender. Fortis, S. (2026) https://BioRender.com/3dl9c85]. Agreement
license VD29LKOJ3P. HLA, human leukocyte antigen; PCa, prostate
cancer; OS, overall survival.](/article_images/ijo/69/3/ijo-69-03-05917-g03.jpg)
![Proposed mechanism of AE37
vaccine-induced antitumor immunity and epitope spreading. Schematic
illustration of the AE37 vaccine-driven immune response. (A)
Vaccination induces robust activation of preexisting AE37-specific
memory CD8+ T cells, leading to the generation of a
proinflammatory tumor microenvironment and (B) direct lysis of
autologous tumor cells expressing epitopes encompassing the AE37
amino acid sequence. (C) Tumor cell lysis results in the release of
additional tumor-derived peptides, including HER-2/neu, PSA and
TERT, which are subsequently (D) captured and cross-presented by
DCs to other preexisting tumor-specific memory CD8+ T
cells bearing cognate T-cell receptors. (E) This process promotes
secondary memory CD8+ T-cell activation, clonal
expansion and amplified tumor cell killing across both (F) clonal
and subclonal populations, thereby overcoming tumor heterogeneity
[Created in BioRender. Fortis, S. (2026) https://BioRender.com/mydl63k]. Agreement license
DI29LKOS9N. EM, effector memory; CCL, chemokine (C-C motif) ligand;
CXCL, chemokine (C-X-C motif) ligand; PSA, prostate-specific
antigen; TERT, telomerase reverse transcriptase; IL, interleukin;
DCs, dendritic cells.](/article_images/ijo/69/3/ijo-69-03-05917-g04.jpg)
![Peripheral blood prognostic
biosignatures integrating soluble factors and tumor-specific
CD8+ T cells in localized PCa. Schematic representation
of prognostic immune biosignatures in patients with localized PCa
based on circulating levels of TGFβ, IL-8 and HER-ECD in
combination with PSA (153-161)-specific CD8+ T-cell
frequencies. Low peripheral levels of TGFβ, IL-8 and HER-ECD define
a favorable biosignature associated with increased frequencies of
PSA (153-161)-specific CD8+ T cells,
enhanced antitumor immunity and improved patient survival. By
contrast, elevated circulating levels of TGFβ, IL-8 and HER-ECD
characterize an unfavorable biosignature marked by reduced
tumor-specific CD8+ T-cell frequencies, immune
suppression, tumor progression and poorer clinical outcomes. These
findings suggest that the degree of peripheral immune suppression
regulates the abundance of preexisting tumor-specific
CD8+ T cells and thereby significantly influences
disease course and prognosis [Created in BioRender. Fortis, S.
(2026) https://BioRender.com/unh610m].
Agreement license ZF29LKOZWI. PCa, prostate cancer; TGFβ,
transforming growth factor-β; IL-8, interleukin-8; HER-ECD, HER
extracellular domain; PSA, prostate-specific antigen.](/article_images/ijo/69/3/ijo-69-03-05917-g05.jpg)
