bims-imseme Biomed News
on Immunosenescence and T cell metabolism
Issue of 2026–07–05
27 papers selected by
Pierpaolo Ginefra, Ludwig Institute for Cancer Research



  1. Proc Natl Acad Sci U S A. 2026 Jul 07. 123(27): e2602385123
      Although A2AR is a key immunoregulatory receptor that suppresses CD8+ T cell activation in response to elevated extracellular adenosine in inflamed or hypoxic microenvironments, its role in CD8+ T cell differentiation and cell-fate decisions during chronic viral infection and cancer remains poorly understood. Using A2AR-eGFP reporter mice, we show that A2AR expression is rapidly induced by TCR stimulation and persists under chronic antigen exposure and hypoxia, with sustained expression strongly associated with terminal exhaustion via the canonical Gαs-cAMP-PKA pathway. Paradoxically, A2AR loss does not alleviate exhaustion but instead accelerates differentiation toward the terminally exhausted state. Single-cell multiomics profiling revealed that A2AR deficiency activates CD122 (IL-2Rβ)-dependent signaling, driving T cell exhaustion. Genetic deletion of CD122 in A2AR-deficient CD8+ T cells reduced terminal exhaustion, identifying CD122 signaling as a key mediator of A2AR loss-driven exhaustion. Intriguingly, both sustained A2AR expression and A2AR loss converge to promote T cell exhaustion differentiation through distinct mechanisms. These findings uncover a paradoxical role of A2AR in shaping CD8+ T cell fate choices during chronic infection and cancer.
    Keywords:  A2AR; CD8(+) T cell differentiation; exhaustion; immunotherapy
    DOI:  https://doi.org/10.1073/pnas.2602385123
  2. bioRxiv. 2026 Jun 17. pii: 2026.06.14.732148. [Epub ahead of print]
      AML is an aggressive blood cancer associated with poor clinical outcomes. Chemotherapy remains the standard of treatment, but unfortunately relapse is very common, highlighting the need for alternative therapies. T cell dysfunction and exhaustion are prominent in AML and may represent a barrier to effective immunotherapy yet remains poorly studied in AML. DNA methylation is a major driver of T cell exhaustion and inhibition of de novo methylation can block exhaustion and restore T cell function in chronic viral infections and other cancers but is understudied in AML. Here, we investigated the impact of azacytidine (Aza), an FDA-approved hypomethylating agent, on T cell exhaustion in AML. Using a spontaneous AML mouse model and samples from patients with AML, we found that Aza treatment modulates T cell function. In vivo Aza-treatment of AML-bearing mice decreased tumor burden and reshaped CD8+ T cell states, with increases in frequencies of memory subsets and decreases in regulatory T cells (Tregs). Functionally, Aza treatment overcame the impaired proliferation displayed by both CD4 and CD8+ T cells in our model. DNA methylation sequencing of T cells after Aza treatment revealed hypomethylation and increased expression of stem-like precursor gene TCF7 and E2F2 , a regulator of cell cycle progression and proliferation. Similar changes in phenotypes were observed in cultures of AML patient samples treated with Aza. Collectively, we show that Aza remodels epigenetic and functional states in AML and has the potential to reverse T cell exhaustion, with enhanced memory and proliferation capacity. Our work generates a mechanistic framework that provides rationale of combining hypomethylating agents with T cell-based immunotherapies in this lethal disease.
    Data Sharing Statement: RRBS data is available in GEO under the accession number GSE328721. For original data please contact Dr. Evan F. Lind.
    Key Points: Azacytidine mediated epigenetic modulation can alleviate T cell exhaustion in AML.
    Translational Relevance: Immune therapy has shown limited efficacy in AML, despite increasing evidence of T cell dysfunction in this malignancy. Azacytidine (Aza) is an FDA approved drug for AML, but patients develop therapy resistance and relapse. Studies have mainly focused on Aza's tumor intrinsic effects. In this study, we investigated the impact of Aza on immune function, especially T cell exhaustion in AML, since exhaustion is a major mechanism of disease resistance. We demonstrated that Aza can modulate T cell phenotype and restore T cell proliferation. Mechanistically, Aza induces epigenetic reprogramming in T cells and increases the expression of a stem-like precursor marker, TCF7. By shifting the focus on T cell biology, our study provides a rationale for combining Aza with other immunotherapies that can enhance durable immune responses in this malignancy.
    DOI:  https://doi.org/10.64898/2026.06.14.732148
  3. Front Immunol. 2026 ;17 1791098
       Background: Vitamin D (VitD) is an important immunometabolic regulator of T cell function. Its active form, 1,25-dihydroxyvitamin D3, signals through the VitD receptor (VDR), which is highly expressed in activated CD4+ T cells. Although VDR signaling suppresses glycolysis by reducing glucose uptake and glycolytic enzyme expression, early T cell expansion is preserved, suggesting the involvement of alternative metabolic pathways. Since glutaminolysis is essential for T cell activation and proliferation, we investigated whether VitD modulates glutamine metabolism during early CD4+ T cell activation.
    Methods: Human CD4+ T cells were stimulated with αCD3/CD28 for four days in the presence or absence of VitD, and analyzed using complementary metabolic, proteomic, and functional approaches.
    Results: VitD-treated cultures exhibited increased cell numbers despite reduced glucose uptake and lactate production, indicating proliferation partially independent of classical glycolytic metabolism. Proteomic analysis revealed increased expression of glutaminase, glutamate dehydrogenase, and CD38, together with enrichment of Selenium Metabolism and Selenoproteins and Nicotinate and Nicotinamide Metabolism, suggesting enhanced glutaminolysis and NAD+ remodeling. Consistently, tritiated and ¹³C-glutamine tracing demonstrated increased glutamine uptake and incorporation into glutamate, α-ketoglutarate, glucose, and inositol-related metabolites, supporting a glutaminolysis-dependent anabolic program rather than oxidative phosphorylation. Pharmacological inhibition of VDR (MeTC7, 1 nM), glutamine uptake (GPNA, 250 µM), or glutaminase activity (BPTES and compound 968, 5 µM) significantly reduced T cell expansion, highlighting glutamine metabolism as essential for the VitD-mediated cell expansion. Interestingly, prolonged cultures showed that VitD ultimately restricted proliferation at day 7; however, supplementation with glutamine and VitD restored cell expansion, suggesting that VitD promotes a metabolically restrained but adaptive proliferative state.
    Discussion: Overall, our findings identify glutaminolysis as a central metabolic pathway supporting VitD-induced CD4+ T cell expansion independently of canonical glycolytic and OXPHOS-associated programs, while promoting metabolic resilience and inflammatory restraint.
    Keywords:  T cell proliferation; T cell regulation; glutaminolysis; immunometabolism; vitamin D
    DOI:  https://doi.org/10.3389/fimmu.2026.1791098
  4. bioRxiv. 2026 Jun 19. pii: 2026.06.17.732005. [Epub ahead of print]
      Two prominent mechanisms by which tumors fend off immune control are by constraining the ability of T cells and CAR T cells to survive and expand in the tumor, and by restraining their ability to sustain full cytotoxic capacity. We identified IκBδ, encoded by Nfkbid , a poorly characterized IκB family member, as a molecular lever that overcomes both of these constraints on anti-tumor CD8 + tumor-infiltrating lymphocytes (TILs). Nfkbid is an NFAT target gene that is expressed in CD8 + effector T cells and, at modest levels, in CD8 + TILs. We found that Nfkbid depletion impaired TIL accumulation, exacerbating the growth of solid tumors. On the other hand, ectopic IκBδ overexpression enhanced TIL expansion, reduced the expression of exhaustion-associated transcription factors and inhibitory receptors, and elevated cytotoxic molecule production, leading to enhanced tumor control. IκBδ has a shorter protein isoform that is identical in a core region spanning the ankyrin-repeat domain known to interact with NFκB proteins, but that lacks the ∼150-residue N-terminal region. We showed that the shared core region is sufficient to drive T cell accumulation, whereas the N-terminal peptide region is required for robust effector function and to counter exhaustion, underscoring that tumor-infiltrating CD8 + T cell accumulation and effector differentiation are separable programs. Our current study provides evidence that IκBδ, an atypical member of the NFκB family, is a lever to overcome two cardinal deficits that limit CD8 + TIL anti-tumor efficacy: impaired accumulation in the tumor and diminished effector function.
    DOI:  https://doi.org/10.64898/2026.06.17.732005
  5. Front Immunol. 2026 ;17 1874242
       Introduction: Osteoporosis weakens the skeleton by lowering bone mass and damaging bone microarchitecture. A growing body of evidence points to disruptions in osteoimmune homeostasis as a key driver of this disease. In particular, apoptosis of bone cells and immune cells appears to be a critical contributing mechanism. Within the bone marrow, CD8+ T cells display a surprising functional duality: whether they protect bone or promote its loss depends heavily on local signals. Under healthy conditions or after mechanical loading, activated CD8+ T cells help maintain bone mass. They do so mainly by secreting interferon‑γ (IFN‑γ), which blocks osteoclast formation through interference with NF‑κB and MAPK signaling. However, this protective role is not fixed. Pathological cues such as estrogen deficiency can flip CD8+ T cells into a bone‑destroying phenotype. For example, when estrogen drops, muscle‑derived IL‑33 decreases. That drop prompts bone marrow CD8+ T cells to release large amounts of CCL5. CCL5 then binds to CCR3 on osteoclast precursors and activates the ERK pathway, thereby accelerating bone loss in postmenopausal osteoporosis. Aging and chronic inflammation add another layer of complexity. They drive the accumulation of senescent CD28- CD8+ T cells. These aged cells no longer produce enough IFN‑γ to protect bone; instead, they adopt a senescence‑associated secretory phenotype (SASP) that suppresses osteoblast differentiation and reduces mesenchymal stromal cell viability. Moreover, apoptosis of osteoblasts and osteocytes-triggered by TNF‑α, IFN‑γ, or glucocorticoids-worsens bone loss. New findings suggest that CD8+ T cells may fuel this apoptotic process through Fas/FasL interactions and the granzyme/perforin pathway. This review brings together these context‑dependent mechanisms. We place special emphasis on how hormonal shifts, metabolic changes, and inflammatory mediators converge to decide whether CD8+ T cells support skeletal integrity or drive bone resorption.
    Methods: We performed a narrative review of recent studies on CD8+ T cells in osteoporosis. We searched PubMed and Web of Science using keywords such as "osteoporosis", "osteoimmunology", "CD8+ T cells", "IFN‑γ", "CCL5", "cellular senescence", "SASP", "Fas/FasL", and "granzyme". From the retrieved articles, we selected original research (including in vitro co‑culture systems, animal models, and clinical samples) and relevant reviews. We then extracted and integrated key mechanistic evidence regarding the dual functionality of CD8+ T cells and its regulatory pathways.
    Results: CD8+ T cells have two opposing functions. Under healthy conditions or mechanical loading, activated CD8+ T cells secrete IFN‑γ, which inhibits osteoclast formation by interfering with NF‑κB and MAPK signaling, thus maintaining bone mass. In contrast, when estrogen is deficient, reduced muscle‑derived IL‑33 causes bone marrow CD8+ T cells to release large amounts of CCL5. CCL5 binds to CCR3 on osteoclast precursors and activates the ERK pathway, accelerating bone loss. Aging and chronic inflammation shift the balance. Aging and chronic inflammation lead to the accumulation of senescent CD28- CD8+ T cells. These cells no longer produce sufficient protective IFN‑γ. Instead, they exhibit a senescence‑associated secretory phenotype (SASP) that suppresses osteoblast differentiation and reduces mesenchymal stromal cell viability. CD8+ T cells may promote bone cell apoptosis through multiple pathways. TNF‑α, IFN‑γ, or glucocorticoids can trigger apoptosis of osteoblasts and osteocytes. Newer evidence suggests that CD8+ T cells amplify this process via Fas/FasL interactions and the granzyme/perforin pathway. Two newly identified CD8+ T cell‑derived molecules deserve attention. Granzyme K protects against bone loss, whereas CCL5 promotes bone loss. These two molecules offer fresh opportunities for biomarkers and therapies.
    Discussion: Our synthesis shows that CD8+ T cells act as a double‑edged sword in osteoporosis: their functional switch depends on the integration of hormonal, metabolic, and inflammatory signals. Under normal conditions or after mechanical loading, the anti‑osteoclastogenic effect of IFN‑γ dominates. But when estrogen levels fall, the IL‑33/CCL5/CCR3/ERK axis becomes activated, turning CD8+ T cells toward a bone‑resorbing phenotype. During aging, the buildup of CD28⁻ CD8+ T cells and their SASP further weakens bone formation and worsens bone cell death through apoptotic pathways. These observations point to a more precise therapeutic strategy. Instead of broadly modulating T cells, one could specifically target pathogenic pathways-for example, by interfering with the IL 33/ST2/CCL5 cascade or by clearing immune senescent CD28⁻ subsets. Such approaches might help restore osteoimmune balance. Two recently discovered CD8+ T cell derived molecules, granzyme K and CCL5, are particularly interesting because they respectively protect against and promote bone loss. They could serve as novel biomarkers or even therapeutic targets. Future studies should examine the cell type specific regulation of these pathways in vivo and test the safety and efficacy of targeted interventions.
    Keywords:  CD8+ T cells; cellular senescence; immunotherapy; interferon-γ; osteoimmunology; osteoporosis
    DOI:  https://doi.org/10.3389/fimmu.2026.1874242
  6. Front Immunol. 2026 ;17 1838764
      Cytoskeletal remodelling is central to naive T cell fitness, organizing receptor-proximal signaling and mechanotransduction during TCR engagement. However, how cytoskeletal dynamics are coordinated with TCR signaling to preserve naive T cell fitness remains incompletely defined. Here, we identify the Sterile 20-family member Thousand and One Kinase 3 (TAOK3) as a kinase-dependent regulator of naive CD8+ T cell maintenance that couples TCR signal integration to cytoskeletal control. Genetic deletion or kinase inactivation of TAOK3 resulted in a profound, cell-intrinsic loss of naive CD8+ T cells. Despite enhanced sensitivity to TCR ligation and enhanced downstream signaling, proliferating CD8+ T cells did not survive in vitro and anti-viral CD8+ T cell immunity was compromised in vivo in the absence of TAOK3. Unbiased phospho-proteomic analysis of Taok3-deficient mice revealed altered phosphorylation of the Rac regulators Dedicator of Cytokinesis DOCK8 and DOCK10, alongside actin-membrane scaffolding proteins. Consistent with this, Taok3-deficient naive CD8+ T cells exhibited elevated basal actin polymerisation, excessive reactive oxygen species accumulation, mitochondrial hyperpolarisation, and reduced spare respiratory capacity. Pharmacologic Rac inhibition normalised cytoskeletal dynamics, corrected the heightened TCR sensitivity, and preferentially restored mitochondrial membrane potential. Collectively, these findings identify TAOK3 as a coordinator of membrane-proximal organisation and cytoskeletal regulation that calibrates Rac-dependent signaling, thereby linking TCR signal integration to mitochondrial fitness and long-term maintenance of the naive CD8+ T cell pool.
    Keywords:  Rac (Rac GTPase); TCR - T cell receptor; cytoskeleton; naive CD8+ T cells; naive T cell homeostasis
    DOI:  https://doi.org/10.3389/fimmu.2026.1838764
  7. Acta Pharm Sin B. 2026 Jun;16(6): 3892-3906
      The immunosuppressive tumor microenvironment (TME) profoundly limits the therapeutic efficacy of CD8+ T cells in solid tumors. While cytokine therapies have shown promise in reactivating CD8+ T cells, they fail to address the common suppressive environmental attributes (e.g., acidosis and Mg2+ deficiency) of solid tumors. Here, we report an innovative CD8+ T cell dual-functional modulator, interleukin-12-tethered nano-aluminum adjuvant (IL12@NAM), which counteracts the acidic TME to relieve acidosis and concurrently releases Mg2+ and IL12. Locally released Mg2+ and IL12 synergize in T cell infiltration and activation by promoting the phosphorylation of focal adhesion kinase and extracellular signal-regulated kinase 1/2, and enhance CD8+ T cell activation and functions via the Ca2+-nuclear factor of activated T cells 2 pathway. Furthermore, dual-functional IL12@NAM mitigates CD8+ T cell exhaustion by reducing PD1 and LAG3 expression and effectively increases the differentiation towards T helper 1 cells while decreasing regulatory T cells, creating a more favorable immune network for enhanced CD8+ T cell-mediated anti-tumor immunity. As a result, IL12@NAM has demonstrated potent therapeutic efficacy against advanced melanoma and breast cancer, and remarkably empowered adoptive T therapy of solid tumors. This study provides a paradigm for empowering cytotoxic T cells by reactivating and creating a sustainable immunoresponsive environment, offering a potential adjuvant strategy to enhance solid tumor therapy.
    Keywords:  Acidosis alleviation; Cytotoxic T cell activation; Cytotoxic T cell immune exhaustion; Cytotoxic T cell infiltration; Interleukin-12 tumor therapy; Mg2+ supplementation; Nano-aluminum adjuvants; Tumor microenvironment
    DOI:  https://doi.org/10.1016/j.apsb.2025.10.021
  8. Gut. 2026 Jul 02. pii: gutjnl-2026-338277. [Epub ahead of print]
       BACKGROUND: The application of the novel KRASG12D inhibitor in pancreatic ductal adenocarcinoma (PDAC) is currently hindered by adaptive resistance. Metabolic reprogramming is a hallmark of KRASG12D signalling, yet the mechanisms linking these alterations to immunosuppression and low therapeutic response are poorly defined.
    OBJECTIVE: To identify the key regulatory nodes connecting KRASG12D-driven metabolic adaptations to tumour microenvironment and develop a mechanistic-based combinatorial strategy.
    DESIGN: We integrated whole-exome sequencing, untargeted metabolomics and single-cell RNA sequencing of human PDAC specimens to analyse the metabolic-immune landscape. We evaluated therapeutic efficacy using the autochthonous mouse and patient-derived xenograft models.
    RESULTS: We found that KRASG12D enhanced cholesterol metabolism and promoted CD8+ T cell exhaustion, whereas KRASG12D inhibition or cholesterol synthesis blockade induced compensatory ULK1-associated autophagy. Cotargeting cholesterol metabolism and autophagy potentiated the antitumour efficacy of the KRASG12D inhibitor MRTX1133 and alleviated CD8+ T cell exhaustion. Mechanistically, KRASG12D transcriptionally upregulated USP20 via EGR1, which simultaneously deubiquitinated and stabilised 3-hydroxy-3-methylglutaryl-CoA reductase and ULK1, thereby orchestrating cholesterol metabolism and autophagy-associated survival. Genetic depletion or pharmacological inhibition of USP20 with GSK2643943A suppressed these pathways and restored CD8+ T cell function, improving responses to MRTX1133 and anti-programmed cell death protein-1 (anti-PD-1). In preclinical PDAC models, triple therapy with GSK2643943A, MRTX1133 and anti-PD-1 elicited a robust therapeutic response and induced significant tumour regression.
    CONCLUSION: USP20 acts as a critical metabolic checkpoint that orchestrates CD8+ T cell exhaustion and therapeutic response. Targeting the USP20-cholesterol-autophagy axis represents a promising strategy to reverse immune suppression and unlock the full potential of KRASG12D inhibitors in PDAC.
    Keywords:  IMMUNE RESPONSE; MOLECULAR TARGETED THERAPY; PANCREATIC CANCER
    DOI:  https://doi.org/10.1136/gutjnl-2026-338277
  9. bioRxiv. 2026 Jun 17. pii: 2026.06.14.732162. [Epub ahead of print]
      Stem-like TCF7+ CD8 T cells sustain anti-tumor responses and support immune checkpoint blockade. We systematically identified regulators of this cell state using genome-wide CRISPR screens in primary T cells in vitro . Using random barcodes to link clonal relationships with guide identity and transcriptional states in single cells, we inferred differentiation trajectories and differentiation rates of CD8 T cells in tumors, while mitigating confounding clonal bias. We found that Trim28-deficient T cells in tumors were enriched in the TCF7+ cell state, depleted in cycling and terminal effector states, and uniquely generated a tissue-resident memory (TRM)-like state with increased chromatin accessibility at known TRM loci as well as repeat elements. Despite the increase in TCF7+ CD8 T cells, loss of Trim28 did not improve tumor control, likely because of reduced effector differentiation, highlighting the need for tuning the balance and dynamics of stem-like versus effector states for effective tumor clearance.
    DOI:  https://doi.org/10.64898/2026.06.14.732162
  10. Front Aging. 2026 ;7 1736105
       Introduction: Respiratory illnesses like influenza and SARS-CoV-2 disproportionately affect older adults, leading to severe complications and high mortality rates. Age-related immune dysregulation impairs infection responses and hinders recovery. The geroscience hypothesis suggests that targeting biological aging can enhance overall healthspan. Mitochondrial dysfunction and dysregulated nutrient sensing, hallmarks of aging, profoundly affect metabolism and cellular function. Metformin, an FDA-approved diabetes drug, is a candidate anti-aging drug and has been shown to positively impact immune cell function in many contexts. However, the totality of these effects on immune cells remains under investigation. Here, we aim to determine if metformin treatment could improve immune memory responses by utilizing a heterologous flu challenge model.
    Methods: Young and aged mice were given control or metformin treated chow for 6 weeks prior to being infected with a sublethal dose of H3N2 influenza virus A/HKx31 (X31). Control and treated chow continued until 10 days post infection to examine the effects of metformin on immune memory formation. Mice were then allowed to recover and at 30 days post initial infection and were challenged with a heterologous H1N1 influenza virus A/Puerto Rico/8/34 (PR8). Mice were sacrificed on day 0 (prior to secondary flu challenge), and at 5, 7, 10, and 14 days post-secondary infection to unveil changes in the kinetics of immune responses.
    Results: Metformin altered only some aspects of immune responses during secondary flu challenge, and more so in young mice compared to aged mice. More specifically, we did not observe improved T cell memory populations in the lungs following primary flu infection in aged metformin treated mice compared to aged control treated mice. Moreover, while aged metformin treated mice had modestly improved weight loss during heterologous challenge, they had transiently increased lung viral load compared to aged control treated mice.
    Discussion: This suggests that metformin could not overcome the totality of aging to improve T cell memory responses. Thus, while metformin has been shown to have many benefits in a variety of aging conditions, its specific utility in improving age-related declines in immune memory formation during infection is unclear in our studies. More research is necessary to determine how metformin can target aging physiology and T cell function to enhance immune responses, and importantly, understand the limitations of its utility in aging populations.
    Keywords:  T cell memory; aging; geroscience; influenza; metformin
    DOI:  https://doi.org/10.3389/fragi.2026.1736105
  11. Front Immunol. 2026 ;17 1819470
      Adoptive T cell therapies have markedly improved outcomes in hematologic malignancies but their efficacy in solid tumors can be diminished by a hostile tumor microenvironment that impedes sustained therapeutic responses. Beyond challenges such as limited trafficking and antigen heterogeneity, engineered T cells face suppressive myeloid and stromal populations, inhibitory checkpoint ligand interactions, and metabolically hostile niches that collectively diminish effector function and persistence. To overcome these barriers, a new generation of fusion protein-based costimulatory strategies has emerged that couple ligand-guided sensing of the tumor microenvironment with modular control of T cell activation and fate. This review examines how conventional and non-canonical costimulatory modules, when incorporated into chimeric antigen receptor (CAR) and T cell receptor (TCR) architectures, modulate T cell differentiation and function within the tumor site. It further analyzes how membrane-anchored and secreted fusion proteins enable engineered T cells to activate dendritic cells, reprogram myeloid cells, and convert poorly inflamed tumors into treatment-responsive environments. Together, these advances establish a design framework in which fusion protein-based receptors and ligands enhance T cell function and remodel the tumor microenvironment, thereby expanding the therapeutic potential of adoptive T cell therapy for solid tumors.
    Keywords:  T cell engineering; costimulatory signaling; fusion protein; myeloid cell reprogramming; switch receptor; tumor microenvironment
    DOI:  https://doi.org/10.3389/fimmu.2026.1819470
  12. Cell. 2026 Jul 01. pii: S0092-8674(26)00709-9. [Epub ahead of print]
      In settings of persistent (self or foreign) antigen, such as autoimmunity and chronic infection, immune responses are sustained by stem-like T cells. Although TCF1 has emerged as a key transcription factor (TF) associated with stemness, the TCF1hi population is heterogeneous, raising the question of whether TCF1 exclusively defines the stem T cell (TSC) pool. Using preclinical models of autoimmune type 1 diabetes and chronic infection, we discover that a small subset of TCF1hi T cells express the TF LEF1. LEF1+ TCF1hi T cells define a true self-renewing TSC pool. TSC give rise to LEF1- TCF1hi progenitor T cells (TPRO), which lack stem functions and generate terminally differentiated TCF1lo T cells (TDIFF) (TSC→TPRO→TDIFF). We show that LEF1 is essential for T cell stemness. Autoimmune and exhausted LEF1+ TSC share a unique epigenetically encoded core program enriched for genes and pathways characteristic of embryonic and adult stem cells, including WNT/β-catenin and Notch signaling. Spatial positioning, niche signals, and migration regulate stem-cell fate; accordingly, targeting integrins or Notch signaling impairs T cell stemness and prevents disease. Our studies identify LEF1 and niche-derived factors as fundamental regulators of T cell stemness across chronic diseases.
    Keywords:  CD8 T cells; LEF1; TCF1; autoimmunity; chronic infection; differentiation; niche factors; stemness; transcription factors
    DOI:  https://doi.org/10.1016/j.cell.2026.06.022
  13. Front Tuberc. 2026 ;pii: 1783887. [Epub ahead of print]4
       Introduction: Memory CD4+ T cells are central to long-term immunity in tuberculosis (TB), yet their functional roles that define their protective capacity remain unclear. Understanding the immune mechanisms that prevent clinical progression from latent TB infection (LTBI) to active TB disease is critical for the development of next-generation vaccines and biomarkers.
    Methods: We characterized the transcriptomic, metabolic, and functional programs of Mycobacterium tuberculosis (Mtb) antigen-stimulated peripheral CD4+ T stem cell (T-SCM), central (T-CM), transitional (T-TM), and effector (T-EM) memory subsets from individuals with remote LTBI. We utilized a multi-platform validation strategy that integrated RNA-sequencing data with protein-level metabolic profiling using "Met-Flow" cytometry and functional growth restriction assays to link memory CD4+ T cell differentiation states to immunometabolism and antimycobacterial function. Finally, we evaluated the immunometabolic profiles of memory CD4+ T cell subsets in an independent, longitudinal cohort of Mtb-exposed progressors and non-progressors from Brazil (GSE112104).
    Results: We identified a differentiation gradient associated with distinct immunometabolic states. T-SCM and T-CM subsets exhibited elevated mitochondrial activity and oxidative metabolism (fatty acid oxidation), supporting their proliferative capacity. In contrast, T-TM and T-EM subsets underwent glycolytic reprogramming and engaged the pentose phosphate pathway, which fueled enhanced cytokine production and Mtb growth restriction. Importantly, we observed that non-progressors exhibit fatty acid oxidation-driven, stem/central memory-like signatures, while progressors and active TB cases display elevated exhaustion markers, glycolytic reprogramming and pro-inflammatory profiles aligned with disease progression.
    Conclusion: Collectively, findings from our proof-of-concept study suggest metabolic state as a key axis connecting Mtb antigen-induced memory T cell differentiation, restimulation-induced transcriptional programming, and durability of immune control. The findings provide the basis for future longitudinal studies to examine the dynamic metabolic and functional modulation in Mtb antigen-specific memory T cell subsets from contained infection to disease progression.
    Keywords:  central memory (T) CM; effector memory (T) EM; immunometabolism; latent TB (LTBI); memory CD4+ T cell subsets; stem cell memory (T) SCM; transitional memory (T) TM; tuberculosis
    DOI:  https://doi.org/10.3389/ftubr.2026.1783887
  14. Front Immunol. 2026 ;17 1804678
       Introduction: Adoptive cell therapy (ACT) has shown promising results in cancer treatment, however, achieving effective ex vivo expansion of potent, functionally active, and cytotoxic T cells remains challenging. To address this challenge, we recently developed artificial antigen-presenting scaffolds (Ag-scaffolds) capable of expanding antigen-specific T cells with phenotypes favorable for ACT. Here, we compared the established technology using IL2/IL21-loaded Ag-scaffolds (Ag-IL2/21) with scaffolds incorporating Neoleukin-2/15 (Ag-Neo2/15), an engineered cytokine that selectively signals via IL-2Rβ/γ complexes to enhance CD8+ T cell proliferation while limiting regulatory T cell expansion.
    Methods: Antigen-specific T cells were expanded ex vivo using Ag-IL2/21 or Ag-Neo2/15 scaffolds. Expansion efficiency, cytokine production, and cytotoxic activity were assessed. Functional and transcriptional states were profiled using single-cell sequencing, including cytotoxic and dysfunction gene signature scoring. T cell receptor (TCR) sequencing was performed to evaluate clonal expansion.
    Results: Ag-Neo2/15 scaffolds supported robust expansion of antigen-specific CD8+ T cells at levels comparable to Ag-IL2/21 scaffolds. Ag-Neo2/15-expanded CD8+ T cells exhibited increased effector cytokine production and durable cytotoxicity against tumor cells. Single-cell transcriptomic analysis revealed higher cytotoxic gene signature scores, reduced dysfunctional scores, and increased expression of genes linked to increased proliferation in Ag-Neo2/15-expanded T cells. TCR sequencing demonstrated preferential expansion of CD8+ T cell clones enriched for transcriptional features associated with antigen-experienced effector states and sustained effector function.
    Discussion: Ag-Neo2/15 scaffolds enhance the quality of ex vivo-expanded antigen-specific T cells by promoting a cytotoxic, less dysfunctional, and highly proliferative phenotype. These findings identify Ag-Neo2/15 as a promising improvement for ACT manufacturing, optimizing the balance between expansion capacity and durable effector function, with potential implications for improving cancer immunotherapy outcomes.
    Keywords:  T-cell expansion; T-cells; adoptive cell therapy; artificial antigen-presenting scaffolds; neoleukin-2/15
    DOI:  https://doi.org/10.3389/fimmu.2026.1804678
  15. Immunity. 2026 Jul 02. pii: S1074-7613(26)00254-2. [Epub ahead of print]
      B cells are highly abundant lymphocytes and central players in humoral immunity. Although T cells are well known to support humoral responses, how B cells influence T cell responses is less understood. Here, we show that B cells are critical for CD8+ T cell responses to chronic, but not acute, viral infections. In the absence of B cells, T cells responding to chronic infection exhibited severely impaired effector differentiation. This dependency on B cell help was dictated by high antigen loads and strong T cell receptor (TCR) stimulation. Loss of either B cells or interferon-I (IFN-I) signaling led to severe functional deficits in exhausted T cells, implicating B cells as key producers of IFN-I. The IFN-I-dependent T cell response to strong TCR stimulation is mediated, in part, by the transcription factor IRF1. Therefore, during chronic infection, we uncover an important role for B cell-derived IFN-I in modulating T cell responses to strong TCR stimulation.
    Keywords:  B cells; IFN-I; LCMV; T cell exhaustion; chronic infection; dysfunctional T cells; tolerance
    DOI:  https://doi.org/10.1016/j.immuni.2026.06.003
  16. J Immunother Cancer. 2026 Jun 29. pii: e003100corr1. [Epub ahead of print]14(6):
      
    DOI:  https://doi.org/10.1136/jitc-2021-003100corr1
  17. Sci Immunol. 2026 Jul 03. 11(121): eaeh4437
      Immunological memory underpins long-term protection and vaccination. Although early work established the durability and recall capacity of memory T cells, understanding of CD8 T cell memory generation, maintenance, and plasticity has advanced substantially over recent decades. Initial models viewed memory as an outcome of peak effector responses and were defined mainly by persistence. Successive waves of technological innovation, from MHC tetramers and genetic lineage-tracing approaches to single-cell and epigenomic profiling and modern high-throughput gene-perturbation screens, have reshaped this view. These approaches reveal memory as a dynamic continuum of cellular states that is actively maintained, tissue-adapted, and epigenetically programmed. CD8 T cell memory is now understood as a flexible and regulated fate rather than a static end point. In this Review, we outline the historical development of the field, highlight how emerging technologies have refined core concepts, and present a modern framework in which memory is an actively enforced yet adaptable property of the CD8 T cell lineage.
    DOI:  https://doi.org/10.1126/sciimmunol.aeh4437
  18. bioRxiv. 2026 Jun 22. pii: 2026.06.17.732787. [Epub ahead of print]
      Fibrosing skin diseases are highly morbid conditions with diverse clinical and histopathologic features. Prior work, primarily in systemic sclerosis (SSc), has yielded mixed data regarding the immune drivers of fibrosis, as well as the identity and spatial localization of pro-fibrotic fibroblast cell subsets. Here, we focus on morphea and eosinophilic fasciitis (EF), which cause more acutely inflammatory skin fibrosis. Using multimodal single-nucleus and spatial transcriptomics, we find that effector CD8+ T cells are highly enriched in fibrotic skin. These cells are particularly abundant in inflammatory tissue domains bordering fibrotic stroma, which are marked by expression of interferon-γ stimulated genes. Inflammatory domains feature a loss of local homeostatic fibroblast populations and replacement with ADAM12-expressing inflammatory fibroblasts and myofibroblasts, which co-localize closely with CD8+ T cells. All subtypes of morphea featured similar patterns of CD8+ T-cell-associated fibro-inflammatory zonation and fibroblast transformation, suggesting that shared mechanisms can drive fibrosis across stromal compartments of skin. We apply these findings to a large publicly available scleroderma dataset and find that similar processes occur in SSc. Mechanistically, ablation of CD8+ T cells in mice ameliorates bleomycin-driven inflammation and fibrosis, as does fibroblast-intrinsic abrogation of IFN-γ signaling. These data establish CD8+ T cell-driven fibrogenesis as a key feature of fibrosing skin diseases and raise the prospect of targeting CD8+ T cells in autoimmune fibrosis more broadly.
    One sentence summary: Spatial profiling of morphea-spectrum diseases reveals CD8 T cells as key drivers of fibrosis through fibroblast IFN-γ signaling.
    DOI:  https://doi.org/10.64898/2026.06.17.732787
  19. NPJ Drug Discov. 2024 Dec 04. pii: 2. [Epub ahead of print]1(1):
      ITK is a kinase involved in T cell activation, proliferation and differentiation. In mice, selective knock-out of the ITK gene produces Th1 skewing of T helper cell differentiation. Soquelitinib, a covalent ITK inhibitor, blocks ITK activity with greater than 100-fold selectivity compared to inhibition of a related kinase, RLK. We describe the chemistry and biologic effects of soquelitinib. In vitro studies with normal or malignant T cells demonstrated that soquelitinib suppresses Th2 cytokine production preferentially with relative sparing of Th1 cytokines. Soquelitinib inhibits the in vivo growth of several syngeneic murine tumors including those that do not express ITK. Treatment with soquelitinib leads to increased tumor infiltration of normal CD8+ cells that possess enhanced T effector function. Soquelitinib reduced expression of T cell exhaustion markers and was able to restore T effector function to exhausted cells. Pharmacologic selective ITK inhibition may represent a novel approach to cancer immunotherapy.
    DOI:  https://doi.org/10.1038/s44386-024-00002-1
  20. Exp Hematol Oncol. 2026 Jul 02.
      Metabolic crosstalk between cancer cells and immune cells is now recognized as a major determinant of immune escape and resistance to anticancer treatments. Cancer cells profoundly reshape the metabolic landscape of the tumor microenvironment, driving nutrient competition, hypoxia, and the accumulation of immunosuppressive oncometabolites that collectively blunt antitumor immunity. Effector T cells, NK cells, and dendritic cells are exposed to nutrient deprivation and suppressive metabolites, including lactate, adenosine, and kynurenine, resulting in impaired T cell proliferation and cytotoxic function and expansion of metabolically adapted regulatory T cells and myeloid-derived suppressor cells. Cancer-associated fibroblasts further reinforce this metabolic reprogramming through extracellular matrix remodeling, secretion of immunosuppressive metabolites, and nutrient recycling that supports tumor growth. Abnormal tumor vasculature sustains metabolic stress by causing uneven perfusion, hypoxia, and acidosis, thereby limiting immune cell infiltration, and promoting immune exhaustion. In addition, diet- and microbiome-driven metabolic cues dynamically shape cancer-immunity interactions and therapeutic responses. Targeting key metabolic checkpoints, including glycolysis, adenosine signaling, tryptophan metabolism, fatty acid oxidation, and lactate production, has emerged as a promising strategy to restore antitumor immunity. Nevertheless, metabolic heterogeneity, context-dependent immune responses, and safety concerns pose persistent challenges to its successful implementation. Recent advances in biomarker development, patient stratification, and rational combination strategies underpin the clinical translation of metabolic-immune vulnerabilities in cancer therapy. Integrating metabolic interventions with immune checkpoint blockade or adoptive cell therapies has demonstrated synergistic effects in preclinical and early clinical studies, enhancing T cell persistence and cytotoxic function within metabolically hostile tumor microenvironments. This review addresses these issues and delineates the mechanistic basis of the dynamic interplay between cancer metabolism and immune regulation. It discusses how anti-cancer therapies affect metabolic and immune pathways and highlights next-generation, metabolically targeted therapies that leverage newly uncovered, tumor-specific rewiring of glycolysis, mitochondrial function, and nutrient uptake. Special emphasis is given to the development of first-in-class inhibitors targeting glutaminase, lipid biosynthesis, one-carbon pathways, and redox homeostasis, which, when paired with immunotherapy or conventional treatments, offer unprecedented opportunities to overcome metabolic barriers, abrogate resistance, and achieve durable immune control of cancer.
    Keywords:  Cancer-associated fibroblasts; Diet and cancer; Immune cell metabolism; Immunotherapy; Microbiome and cancer; Oncometabolites; Therapy resistance; Tumor metabolism; Tumor microenvironment
    DOI:  https://doi.org/10.1186/s40164-026-00776-2
  21. Med Sci Sports Exerc. 2026 Jun 30.
       PURPOSE: Cardiorespiratory fitness (CRF) is associated with enhanced leukocyte metabolism. Whether CRF impacts the metabolism of specific leukocytes such as cytotoxic T cells (Tc) is unknown. As Tc metabolism and function, including interferon (IFN)-γ production, are linked, metabolic changes associated with higher CRF could impact Tc function. Therefore, this study aimed to determine whether CRF predicted Tc metabolism and function.
    METHODS: Forty-three adults (16 women) underwent a maximal oxygen consumption (VO2peak) test, had body fat percentage (BF%) measured, and donated resting blood. Tc mitochondrial mass, mitochondrial membrane potential, fatty acid and amino acid oxidation (FAO and AAO) capacity, glycolytic capacity, and IFN-γ expression were assessed via flow cytometry. Linear regression assessed CRF and BF% as predictors of metabolism and function. Analyses were conducted within total Tc and in Tc subsets.
    RESULTS: CRF predicted FAO and AAO capacity of Tc (ꞵ= 0.362; p= 0.017) and Tc subsets, and glycolytic capacity of Tc (ꞵ= 0.495; p< 0.001), central memory Tc (ꞵ= 0.304; p= 0.047), and effector memory re-expressing CD45RA (EMRA) Tc subsets (ꞵ= 0.451; p= 0.003). CRF also predicted IFN-γ expression by Tc (ꞵ= 0.384; p= 0.011), naïve Tc (ꞵ= 0.358; p= 0.020), and EMRA Tc subsets (ꞵ= 0.349; p= 0.022). Inclusion of BF% in models partially abrogated these relationships. CRF remained a significant predictor of Tc and EMRA Tc glycolytic capacity, and Tc function.
    CONCLUSIONS: CRF predicts Tc metabolism and function in healthy adults, where greater CRF associates with greater Tc FAO and AAO capacity, glycolytic capacity, and function. However, these relationships are in part attenuated by BF%, and Tc subsets exhibit differences in the degree to which they relate to CRF and BF%.
    Keywords:  AEROBIC CAPACITY; CD8+ T LYMPHOCYTES; IMMUNOMETABOLISM; MITOCHONDRIAL FUNCTION; SCENITH
    DOI:  https://doi.org/10.1249/MSS.0000000000004063
  22. Med Oncol. 2026 Jul 01. pii: 208. [Epub ahead of print]43(8):
      The tumor microenvironment (TME) limits durable antitumor immunity by impairing CD8+ T cell responses. Memory like CD8+ T cells are important for long-term immune control but are often restricted in the TME. Dendritic cells (DCs) are key regulators of T cell fate. Previous studies have shown that SHP1 in DCs fosters an immunosuppressive microenvironment and facilitates tumor immune escape. T cell factor-1 (TCF-1), encoded by Tcf7 gene, is required for central memory CD8+ T cell (TCM) formation and is closely linked to canonical Wnt/β-catenin signaling. However, whether SHP1 in DCs regulates TCF-1 expression and TCM formation remains unclear. To investigate the role of DC intrinsic SHP1 in T cell immunity, SHP1 deficient DC2.4 cells and primary bone marrow derived dendritic cells (BMDCs) were co-cultured with OT-1 T cells to assess proliferation, TCM formation, cytotoxic activity, and TCF-1 expression. A DC-specific SHP1 knockout mice model was used to evaluate antitumor immunity in vivo, and Tcf7 or Ctnnb1 silencing was used to probe the TCF-1/Wnt/β-catenin axis. SHP1 downregulation in DCs markedly enhanced CD8+ T cell proliferation, promoted the generation of CD62L+ CD44+ central memory T cells, and potentiated B16-F10-OVA tumor cell killing, accompanied by increased TCF-1 expression in OT-1 T cells. In DC-specific SHP1 knockout mice, EO771 tumor growth was suppressed with concurrent increases in intratumoral IFN-γ+ and TCF-1+ CD8+ T cell frequencies. Mechanistically, we found that DC SHP1 regulates TCM formation via TCF-1, as silencing Tcf7 in OT-1 T cells abrogated this effect. SHP1-deficient DCs activated Wnt/β-catenin signaling in CD8+ T cells, as shown by increased active β-catenin, total β-catenin, c-Myc and Cyclin D1, and a reduced phospho β-catenin/total β-catenin ratio. Critically, Ctnnb1 silencing in T cells abrogated the enhanced proliferation, TCM formation, and cytotoxic activity induced by SHP1-deficient DCs. DC-intrinsic SHP1 restrains central memory CD8+ T cell formation via the TCF-1/Wnt/β-catenin axis.
    Keywords:  Dendritic cells; SHP-1; TCF-1; TCM; Wnt/β-catenin
    DOI:  https://doi.org/10.1007/s12032-026-03329-z
  23. iScience. 2026 Jul 17. 29(7): 116411
      miR-139 acts as a tumor suppressor in breast cancer cells, yet its role within the tumor immune microenvironment (TIME) remains unclear. This study explores the dual functions of miR-139 in both tumor and immune cells, particularly its effect on CD8+ T cell function and chemokine-mediated immune recruitment. Bioinformatics analysis using TCGA and GEO data identified miR-139 as differentially expressed in breast cancer. Using samples from 32 patients, we detected elevated miR-139 levels in tumor-infiltrating CD8+ T cells compared with adjacent normal tissues. Further experiments confirmed that miR-139 directly targets CD28, leading to its downregulation. Concurrently, increased expression of exhaustion markers PD-1 and TIGIT was observed. Moreover, tumors with high miR-139 expression showed upregulation of T-cell-recruiting chemokines CX3CL1, CXCL12, and CXCL14. These results demonstrate that miR-139 promotes CD8+ T cell exhaustion via suppression of CD28 and facilitates chemokine-mediated T cell recruitment, highlighting its immunosuppressive role in breast cancer.
    Keywords:  Cancer; Cell biology; Immunology
    DOI:  https://doi.org/10.1016/j.isci.2026.116411
  24. bioRxiv. 2026 Jun 23. pii: 2026.06.18.733147. [Epub ahead of print]
      Transforming growth factor-β (TGF-β) regulates CD4 T cell quiescence, activation, and regulatory T cell differentiation, but its role in T cell iron metabolism is poorly defined. Here, we investigated whether TGF-β regulates iron homeostasis and how iron overload alters TGF-β responsiveness. During T cell activation, TGF-β enhanced survival but markedly reduced proliferation. These effects were accompanied by decreased CD71 expression and cytosolic iron availability, as well as increased mitochondrial iron accumulation. Genetic deletion of TGFβR1 reversed these changes, demonstrating that TGF-β regulates CD4 T cell iron homeostasis through TGFβR1-dependent signaling. Iron-overloaded CD4 T cells lacking the heme exporter FLVCR1 exhibit hypersensitivity to TGF-β, increased TGF-β secretion, and sustained TGFβR1 expression upon activation. Pharmacologic inhibition of TGFβR1restored proliferation, CD71 expression, and iron levels in FLVCR1-deficient cells. Although TGF-β selectively induced total and mitochondrial ROS levels in FLVCR1-deficient cells, antioxidant treatment or Nox2 inhibition did not rescue this phenotype, suggesting that ROS is associated with, but not sufficient to explain, TGF-β hypersensitivity. Acute FeSO 4 -induced iron overload partially recapitulated the phenotype of FLVCR1-deficient cells, although TGFβR1 expression and TGF-β production differed. Finally, regulatory T cells generated in vitro in the presence of TGF-β displayed reduced iron acquisition, and excess iron impaired FoxP3 induction. Together, this work identifies TGF-β as a context-dependent regulator of CD4 T cell iron homeostasis.
    DOI:  https://doi.org/10.64898/2026.06.18.733147
  25. bioRxiv. 2026 Jun 17. pii: 2026.06.15.732401. [Epub ahead of print]
      Here we identify NKG2C + CD27 - as a novel surface marker unifying multiple previously reported CD8 + regulatory T cell (Treg) populations. This population displays high clonality and divides into CD226 - (Treg1) and CD226 + (Treg2) subsets, with Treg2 exhibiting stronger suppressive activities. Up to 35% of CD8 + T EMRA cells are Tregs, whereas approximately 85% of CD8 + Tregs are T EMRA cells, which increase with aging. These findings establish a unified and novel cell surface marker for CD8 + Tregs and their subsets, which resolves prior heterogeneity in the field, and show that CD8 + T EMRA cells are a heterogeneous population that includes T cells with regulatory function. Our findings provide a critical framework for the isolation and in-depth functional characterization of CD8 + Tregs in health, aging, and disease.
    DOI:  https://doi.org/10.64898/2026.06.15.732401
  26. Acta Pharmacol Sin. 2026 Jun 29.
      Emerging evidence underscores the pivotal role of CD4+ effector T cells in antitumor immunity, yet their full therapeutic potential remains underexplored. This study investigates the impact of targeting the Wnt coactivators B-cell lymphoma 9 and B-cell lymphoma 9-like (BCL9/BCL9L) on CD4+ T cell-mediated antitumor immunity. We demonstrate that either genetic deletion or pharmacological inhibition of BCL9/BCL9L suppresses tumor progression and increases CD4+ T cell activation in the tumor microenvironment (TME). Critically, loss of Bcl9/Bcl9l skews CD4+ T cell differentiation toward a Th1 phenotype at the expense of Th2, Th17, and Treg lineages, thereby establishing a Th1-dominant TME and unleashing a direct cytotoxic program in these Th1 cells. Mechanistically, BCL9/BCL9L inhibition promotes Th1 polarization by suppressing TCF4-mediated regulation of PIAS family genes and enhancing STAT1/STAT4 activation, thereby driving a potent Th1-mediated antitumor response. Collectively, our findings identify BCL9/BCL9L as critical negative regulators of CD4+ T cell antitumor immunity and establish them as promising therapeutic targets for reprogramming the TME toward a Th1-supportive state.
    Keywords:  BCL9/BCL9L; CD4+ T cells; STAT1/STAT4; Th1 cells; tumor microenvironment
    DOI:  https://doi.org/10.1038/s41401-026-01845-2