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NKG2D Receptor and Its Ligands in Host Defense
Open accessAbstract
NKG2D is an activating receptor expressed on the surface of natural killer (NK) cells, CD8+ T cells, and subsets of CD4+ T cells, invariant NKT cells (iNKT), and γδ T cells. In humans, NKG2D transmits signals by its association with the DAP10 adapter subunit, and in mice alternatively spliced isoforms transmit signals either using DAP10 or DAP12 adapter subunits. Although NKG2D is encoded by a highly conserved gene (KLRK1) with limited polymorphism, the receptor recognizes an extensive repertoire of ligands, encoded by at least eight genes in humans (MICA, MICB, RAET1E, RAET1G, RAET1H, RAET1I, RAET1L, and RAET1N), some with extensive allelic polymorphism. Expression of the NKG2D ligands is tightly regulated at the level of transcription, translation, and posttranslation. In general, healthy adult tissues do not express NKG2D glycoproteins on the cell surface, but these ligands can be induced by hyperproliferation and transformation, as well as when cells are infected by pathogens. Thus, the NKG2D pathway serves as a mechanism for the immune system to detect and eliminate cells that have undergone “stress.” Viruses and tumor cells have devised numerous strategies to evade detection by the NKG2D surveillance system, and diversification of the NKG2D ligand genes likely has been driven by selective pressures imposed by pathogens. NKG2D provides an attractive target for therapeutics in the treatment of infectious diseases, cancer, and autoimmune diseases. Cancer Immunol Res; 3(6); 575–82. ©2015 AACR.
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“Remarkably, the candidate signature gene RAET1G is one of the few genes that could encode a ligand recognized by NKG2D proteins in response to stress and infections (Eagle and Trowsdale, 2007; Tomasec et al., 2007; Lanier, 2015).”
From Identification of Candidate Signature Genes and Key Regulators Associated With Trypanotolerance in the Sheko Breed (2019) · cites this paper as Lanier, 2015
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“In addition, we engineered the NKL cells to express the activating mouse receptor Ly49H to study another NKR signaling kinetics that uses both the ITAM-bearing adaptor DAP12 and the DAP10 adaptor, in contrast to human NKG2D receptor signaling, which solely uses the DAP10 adaptor [18,19].”
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“33,34 Expression of these ligands is regulated by different mechanisms that can be induced by virus infection, DNA damage, and oxidative stress, among others.”
The MICA deletion across different populations.
Ciriaco, V. A. D. O., Rodrigues, A., Tibúrcio, B. C. D. S., Silva, J. M., Naslavsky, M., Mendes-Junior, C., Bannwart-Castro, C., & Castelli, E. C. (2024) · Human Immunology · doi:10.1016/j.humimm.2024.111183 · Cited by 1
Absence of ATM leads to altered NK cell function in mice.
Covino, D., Desimio, M., Giovinazzo, A., de Oliveira, B. S. P., Merolle, M., Marazziti, D., Pellegrini, M., & Doria, M. (2024) · Clinical Immunology · doi:10.1016/j.clim.2024.110233 · Cited by 1
The role of immune mechanisms in the pathogenesis of hemorrhagic fever with renal syndrome
Ivanov, M. F. (2023) · Aspirantskiy Vestnik Povolzhiya · doi:10.55531/2072-2354.2023.23.3.4-12 · Cited by 1
Development of small molecule inhibitors of natural killer group 2D receptor (NKG2D).
Wang, J., Nakafuku, K., Ziff, J., Gelin, C., Gholami, H., Thompson, A. A., Karpowich, N., Limon, L., Coate, H. R., Damm-Ganamet, K., et al. (2023) · Bioorganic & Medicinal Chemistry Letters · doi:10.1016/j.bmcl.2023.129492 · Cited by 1
Wei, A., Zhang, L.-P., Ma, H.-H., Cui, L., Zhang, Q., Wang, D., Chen, S., Du, J.-Y., Zhao, X.-X., Wang, T.-Y., et al. (2023) · Journal of pediatric hematology/oncology · doi:10.1097/mph.0000000000002652 · Cited by 1
NKG2D signaling regulates IL-17A-producing γδT cells to promote cancer progressionPreprint
Curio, S., Edwards, S., Suzuki, T., McGovern, J. L., Triulzi, C., Yoshida, N., Jonsson, G., Glauner, T., Rami, D., Purcell, R., et al. (2021) · bioRxiv · doi:10.1101/2021.08.26.457761 · Cited by 1
“NKG2D is a potent immunoreceptor expressed on various innate and adaptive immune cells including γδT cells (Lanier, 2015; Spits et al., 2016).”
BackgroundThe NKG2D ligand ULBP4 is not expressed by human monocytes
Lazarova, M., Kim, Y., & Steinle, A. (2020) · bioRxiv · doi:10.1371/journal.pone.0246726 · Cited by 1
Delaune, V., Toso, C., Kahler‐Quesada, A., Slits, F., Gex, Q., Kaya, G., Lavallard, V., Orci, L. A., Peloso, A., & Lacotte, S. (2020) · Transplant International · doi:10.1111/tri.13589 · Cited by 1
“The expression of NKG2D ligands [MHC-class I chain-related proteins (MICA, MICB), and UL-16 binding proteins (ULBP1-6) in humans; ribonucleic acid export 1-like (RAE1L), and similar to retinoic acid early transcript 1L (RRLT) in rats] increases in cells exposed to stress such as hypoxia, DNA damage, exposure to cytoki…”
BackgroundZhao, J.-G., Schlösser, H., Wang, Z.-F., Qin, J., Li, J.-H., Popp, F., Popp, M., Alakus, H., Chon, S.-H., Jauch, K., et al. (2019) · Pancreatology (Print) · doi:10.1016/j.pan.2019.05.006 · Cited by 1
“and ULBP-5 are transmembrane-anchored proteins, whereas ULBP-1, ULBP-2, ULBP-3 and ULBP-6 are glycophosphatidylinositol–anchored proteins[55].”
Molfetta, R., Zitti, B., Santoni, A., & Paolini, R. (2017) · AIMS Allergy and Immunology · doi:10.3934/allergy.2017.4.164
“All are orthologous to the human ULPB family, and can be expressed as transmembrane (MULT1) or GPI-linked (Rae-1 and some H60 ligands) proteins [4,9,17].”
Parikh, B., Piersma, S. J., & Yokoyama, W. (2016) · doi:10.1016/b978-0-12-374279-7.14006-8 · Cited by 1
Promotion of Early Pregnancy Events in Mice and Humans by Uterine Natural Killer Cells
Felker, A. M., & Croy, B. (2016) · NeoReviews · doi:10.1542/neo.17-2-e93 · Cited by 1
Stiene, D. S., Osterburg, A., Corsarie, L. B., Balzarini, N. R., Medvedovic, M., & Borchers, M. (2025) · Journal of cellular immunology · doi:10.33696/immunology.7.227 · Cited by 1
“This activates other proximal immune cells, such as dendritic cells (DCs), macrophages, and other lymphocytes [22,25–29].”
InfluentialAging of Natural Killer Cells in Acute Myeloid Leukemia
Sánchez-Correa, B., Solana, R., & Tarazona, R. (2017) · Springer eBooks · doi:10.1007/978-3-319-44870-1_75-1
NK Cell Responses in Immunotherapy: Novel Targets and Applications
Pachynski, R. K., & Kohrt, H. E. K. (2016) · Springer eBooks · doi:10.1007/978-3-319-29827-6_4
Jonsson, G., Hofmann, M., Mereiter, S., Hartley‐Tassell, L., Onji, M., Sakic, I., Oliveira, T., Hoffmann, D., Novatchkova, M., Schleiffer, A., et al. (2026) · The FEBS Journal · doi:10.1111/febs.70236 · Cited by 1
“For example, the C-type lectin NKG2D recognizes polymorphic MHC class I-related stress-induced ligands and serves as a key activating receptor for NK cells [10].”
CLEC18A interacts with sulfated GAGs and controls clear cell renal cell carcinoma progression
Jonsson, G., Hofmann, M., Mereiter, S., Hartley-Tassell, L., Sakic, I., Oliveira, T., Hoffmann, D., Novatchkova, M., Schleiffer, A., & Penninger, J. M. (2024) · doi:10.1101/2024.07.08.602586 · Cited by 1
“5I ) Discussion C-type lectins are known to play essential roles in cancer ranging from recognition of damaged cells and antigen uptake [26, 27], to elimination of cancer cells through MHC I associated stress signals [10].”
BackgroundVEGFR2-targeted fusion antibody improved NK cell-mediated immunosurveillance against K562 cells
Ren, X.-Y., Xie, W., Wang, Y.-F., Xu, M., Liu, F., Tang, M., Li, C.-C., Wang, M., & Zhang, J. (2016) · Immunologic research · doi:10.1007/s12026-016-8800-3
Shivarov, V. S., & Ivanova, M. (2026) · Frontiers in Hematology · doi:10.3389/frhem.2026.1969104
Basher, F., Smith, T. A., Rubinstein, M. P., & Wu, J. D. (2026) · Biomedicines · doi:10.3390/biomedicines14102390
“A variety of NKG2D ligands have been identified, presumably due to evolutionary pressure [30,31], among which the family members of the MHC class I chain-related molecules A and B (MICA/B, collectively termed MIC) are most prevalently expressed by human solid tumors in response to genotoxic stress…”
Lauretta, C., Sorrentino, C., Sabbatino, E., Tutino, V., Souza, D. S., MIGLIACCIO, A., Castoria, G., Giovannelli, P., & Donato, M. D. (2026) · Frontiers in Cell and Developmental Biology · doi:10.3389/fcell.2026.1905416
“NKG2D is an activating receptor expressed on NK cells and cytotoxic CD8 + T lymphocytes that recognizes stress-induced ligands, including MICA/B and UL16-binding proteins (ULBPs), expressed on tumor cells (Lanier, 2015).”
Immunomodulatory Potential of KSTM in Activating Natural Killer Cells for Cancer Cell Elimination
Pianmak, T., Panya, A., Boonsatit, N., Tabtimmai, L., & Choowongkomon, K. (2026) · Pharmaceuticals · doi:10.3390/ph19101525
“Nevertheless, tumor cells expressing MHC class I can also be eliminated by NK cells through the expression of stress-induced ligands that activate NK cell receptors, including NKG2D, NKp46, and NKp44 [16–18].”
Tran, T.-D., Lamorlette, C., Gerard, L., Brouard, J., Dotti, G., Moulin, D., Reppel, L., Pochon, C., & Rubio, M. (2026) · bioRxiv · doi:10.64898/2026.08.23.746558
2B4 co-engagement promotes serial degranulation and killing by human NK cells
Kröll, L., Sandusky, M., Saretzki, M., Claus, M., Wingert, S., Niemann, J., & Watzl, C. (2026) · bioRxiv · doi:10.64898/2026.07.31.741999
“NKG2D (CD314) is an NK cell activating receptor that can bind to MICA, MICB, and members of the ULBP protein family (Lanier, 2015).”
Ginting, T. E., Larasati, Y., Cornelia, V., Hidayat, N. R., Christian, S., Zarkasie, K., & Yusuf, I. (2026) · ImmunoTargets and Therapy · doi:10.2147/itt.s614889
“14–18 In a physiological condition, there is a regulation to balance the immune response sufficiently to minimize damage caused by the response, known as an immune checkpoint.”
The Role of Natural Killer (NK) Cells in Dengue Virus Infection: A Narrative Review
Islam, F. (2026) · Health Science Reports · doi:10.1002/hsr2.72783
Host-parasite interactions of an avian blood parasite elucidated by single-cell transcriptomicsPreprint
Rapp, T. M., Lautwein, T., Peuß, R., Kurtz, J., Krüger, O., & Chakarov, N. (2026) · bioRxiv · doi:10.1101/2025.03.14.643032
Alijanizadeh, P., Shahmoradi, M., Kamroo, A., Shahmoradi, T., Tajafrooz, F., Khodaee, P., Yazdanpanah, N., Saleki, K., & Rezaei, N. (2026) · Cancer Cell International · doi:10.1186/s12935-026-04345-9
Establishment and characterization of NKMS-1, a novel mouse NK cell line
Miyakura, Y., Nagashima, T., Ito, S., Fujimori, H., Imai, T., Asada, Y., Mochizuki, M., Tamai, K., Katori, Y., Yasuda, J., et al. (2026) · Scientific Reports · doi:10.1038/s41598-026-50317-4
“NKG2D is a receptor that recognizes target cells expressing its ligands and triggers subsequent signaling cascades leading to target cell killing 40,41 .”
Fan, H., Du, C., Xu, C., Wang, H., Wang, Y., Wu, Z., Zhao, B., Chen, C., Shang, S., Sun, M.-A., et al. (2026) · BMC Genomics · doi:10.1186/s12864-026-12854-0 · Cited by 3
“Cluster T15 also highly expresses KLRK1 (also known as NKG2D ) which is an activation receptor for T and NK cells [68].”
Immunologic Combinations in Multiple Myeloma: Synergistic Stimulation with T cells and NK cells
Liu, Z.-Y., Lai, J.-X., Yang, C., Liu, H., Ding, K., Song, J., & Fu, R. (2026) · International Journal on Biological Sciences · doi:10.7150/ijbs.127871
Álvarez-Carrasco, P., Basurto-Olvera, P., Jiménez‐Hernández, E., Medina‐Sansón, A., & Maldonado‐Bernal, C. (2026) · Frontiers in Immunology · doi:10.3389/fimmu.2026.1757205
“To enhance the stringency of the identi fi cation and capture the cytotoxic effector lineage, NKG2D expression was included as a complementary marker (28, 29).”
Zhao, C.-P., Ren, H., Ke, Q.-M., Chen, Q., He, J., Tian, R., Pan, H.-D., & Liu, L. (2026) · Frontiers in Immunology · doi:10.3389/fimmu.2026.1770733
“These divergent outcomes likely arise from differences in adaptor protein usage (DAP10 vs. DAP12), local immune microenvironments, and ligand diversity (27, 28).”
Takahashi, K., Ikegami, T., Kato, A., Yamada, N., Iwanaga, T., Sakuma, T., Senoo, J., & Kamezaki, H. (2026) · Onco · doi:10.3390/onco6010015
“The NKG2D (Natural Killer Group 2D) receptor is a pivotal activating receptor expressed on NK cells, CD8+ T cells, and γδ T cells [13,14].”
Hagelstein, I., Wang, K., Holz, A., Blessing, J., & Lutz, M. S. (2026) · Journal of Ovarian Research · doi:10.1186/s13048-025-01962-2
“The natural killer group 2 member D (NKG2D; CD314), a C-type lectin-like receptor, is - among many other mechanisms - an essential component of this immune surveillance system [1, 2].”
Lu, J. Y., Ma, J., Li, K., Bao, S., Hu, Y., Yu, L., Jin, Y., Han, X., Wu, J., & Jin, Y. (2025) · International Immunopharmacology · doi:10.1016/j.intimp.2025.116117
Huang, Y., Gong, C., Yang, E., Yuan, H.-C., Lv, D., Yang, C., Chen, W.-J., Wan, Q.-Y., Tan, Z.-Y., Wang, H.-F., et al. (2025) · Cytokine · doi:10.1016/j.cyto.2025.157084
Dap10 co-stimulation enhances the anti-HCC efficacy of NKp30 chimeric antigen receptor T cells
Li, J., Chen, L., Chen, M., Lin, M. Y., Xie, Z., Wu, H., Zhou, Z., & Lin, W. (2025) · Translational Oncology · doi:10.1016/j.tranon.2025.102425
“The presence and signaling of NKG2D in T cells are contingent upon another membrane protein, namely DNAX-activating protein 10 (DAP10) [16,17].”
Chou, S.-C., Kuo, C.-Y., Ko, H.-W., Huang, P., Liu, C.-H., Wang, L.-S., & Liang, Y.-J. (2025) · Experimental and Therapeutic Medicine · doi:10.3892/etm.2025.12822
“Outcomes of NK cell cytolytic response to target cells depend on the balance between activation and inhibition signaling (7).”
Metadata
- Authors
- Lanier, L. L.
- Affiliations
- University of California, San Francisco (US)
- Corresponding author
- Lanier, Lewis L.
- ORCID
- Lanier, Lewis L.
- Venue
- Cancer Immunology Research
- Published
- 2015-06-01
- Volume
- 3
- Issue
- 6
- Pages
- 575-582
- ISSN
- 2326-6066 (ISSN-L)
- 2326-6074
- PMC
- PMC4457299
- OpenAlex
- W2142548447
- Cited by
- 599
- References
- 78
- Citations by year
- 2026: 27 · 2025: 53 · 2024: 56 · 2023: 73 · 2022: 56 · 2021: 66 · 2020: 63 · 2019: 51 · 2018: 65 · 2017: 47 · 2016: 37 · 2015: 4
- FWCI
- 4.80
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Cancer Immunology Research (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- eScholarship (California Digital Library) (submittedVersion)
- Funders
- National Cancer Institute (R01CA089189)
- National Institute of Allergy and Infectious Diseases (AI-068129, R01AI068129, R37AI066897, AI066897)
- Topics
- Life Sciences › Immunology and Microbiology › Immunology › Immune Cell Function and Interaction
- Life Sciences › Immunology and Microbiology › Immunology › interferon and immune responses
- Life Sciences › Immunology and Microbiology › Immunology › Reproductive System and Pregnancy
- Keywords
- NKG2D, NKG2D ligands, mica, MICB, natural killer cells, DAP10, DAP12, immune evasion, viral evasion, tumor immune evasion, host defense, NK cell activation
- MeSH terms
- Animals, Communicable Diseases, Humans, Killer Cells, Natural, Ligands, Lymphocyte Activation, Neoplasms, Lymphocyte Subsets, NK Cell Lectin-Like Receptor Subfamily K, Disease Resistance
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