Naive T cell explained

In immunology, a naive T cell (Th0 cell) is a T cell that has differentiated in the thymus, and successfully undergone the positive and negative processes of central selection in the thymus. Among these are the naive forms of helper T cells (CD4+) and cytotoxic T cells (CD8+). Any naive T cell is considered immature and, unlike activated or memory T cells, has not encountered its cognate antigen within the periphery. After this encounter, the naive T cell is considered a mature T cell.

Phenotype

Naive T cells are commonly characterized by the surface expression of L-selectin (CD62L) and C-C Chemokine receptor type 7 (CCR7); the absence of the activation markers CD25, CD44 or CD69; and the absence of memory CD45RO isoform.[1] [2] They also express functional IL-7 receptors, consisting of subunits IL-7 receptor-α, CD127, and common-γ chain, CD132. In the naive state, T cells are thought to require the common-gamma chain cytokines IL-7 and IL-15 for homeostatic survival mechanisms. [3] While naive T cells are regularly regarded as a developmentally synchronized and fairly homogeneous and quiescent cell population, only differing in T cell receptor specificity, there is increasing evidence that naive T cells are actually heterogeneous in phenotype, function, dynamics and differentiation status, resulting in a whole spectrum of naive cells with different properties. For instance, some non-naive T cells express surface markers similar to naive T cells (Tscm, stem cell memory T cells;[4] Tmp, memory T cells with a naive phenotype[5]), some antigen-naive T cells have lost their naive phenotype,[6] and some T cells are incorporated within the naive T cell phenotype but are a different T cell subset (Treg, regulatory T cells; RTE, Recent Thymic emigrant). It is important to appreciate these differences when assessing naive T cells. Majority of human naive T cells are produced very early in life when infant's thymus is large and functional. Decrease in naive T cell production due to involution of the thymus with age is compensated by so called "peripheral proliferation" or "homeostatic proliferation" of naive T cells which have emigrated from the thymus earlier in life. The homeostatic proliferation causes change to naive T cell gene expression and i.e. is manifested by acquisition of CD25 surface protein expression.

Function

Naive T cells can respond to novel pathogens that the immune system has not yet encountered. Recognition by a naive T cell clone of its cognate antigen results in the initiation of an immune response. In turn, this results in the T cell acquiring an activated phenotype seen by the up-regulation of surface markers CD25+, CD44+, CD62Llow, CD69+ and may further differentiate into a memory T cell.

Having adequate numbers of naive T cells is essential for the immune system to continuously respond to unfamiliar pathogens.

Mechanism of activation

When a recognized antigen binds to the T cell antigen receptor (TCR) located in the cell membrane of Th0 cells, these cells are activated through the following "classical" signal transduction cascade:[7]

An alternative "non-classical" pathway involves activated Zap70 directly phosphorylating the p38 MAPK that in turn induces the expression of the vitamin D receptor (VDR). Furthermore, the expression of PLC-γ1 is dependent on VDR activated by calcitriol.[7] Naive T cells have very low expression of VDR and PLC-γ1. However, activated TCR signaling through p38 upregulates VDR expression and calcitriol activated VDR, in turn, upregulates PLC-γ1 expression. Hence the activation of naive T cells is crucially dependent on adequate calcitriol levels.[7]

In summary, activation of T cells first requires activation through the non-classical pathway to increase expression of VDR and PLC-γ1 before activation through the classical pathway can proceed. This provides a delayed response mechanism where the innate immune system is allowed time (~48 hrs) to clear an infection before the inflammatory T cell mediated adaptive immune response kicks in.[7]

See also

Notes and References

  1. De Rosa SC, Herzenberg LA, Herzenberg LA, Roederer M . 11-color, 13-parameter flow cytometry: identification of human naive T cells by phenotype, function, and T-cell receptor diversity . Nat. Med. . 7 . 2 . 245–8 . February 2001 . 11175858 . 10.1038/84701 . 25144260 .
  2. van den Broek. Theo. Borghans. José A. M.. van Wijk. Femke. 2018-03-08. The full spectrum of human naive T cells. Nature Reviews. Immunology. 18 . 6 . 363–373 . 10.1038/s41577-018-0001-y. 1474-1741. 29520044. 256745422 .
  3. Jeffrey Rathmell . Rathmell . Jeffrey C. . Farkash . Evan A. . Gao . Wei . Thompson . Craig B. . IL-7 Enhances the Survival and Maintains the Size of Naive T Cells . The Journal of Immunology . 15 December 2001 . 167 . 12 . 6869–6876 . 10.4049/jimmunol.167.12.6869 . 11739504 . free .
  4. Gattinoni. Luca. Lugli. Enrico. Ji. Yun. Pos. Zoltan. Paulos. Chrystal M.. Quigley. Máire F.. Almeida. Jorge R.. Gostick. Emma. Yu. Zhiya. 2011-09-18. A human memory T cell subset with stem cell-like properties. Nature Medicine. 17. 10. 1290–1297. 10.1038/nm.2446. 1546-170X. 3192229. 21926977.
  5. Pulko. Vesna. Davies. John S.. Martinez. Carmine. Lanteri. Marion C.. Busch. Michael P.. Diamond. Michael S.. Knox. Kenneth. Bush. Erin C.. Sims. Peter A.. August 2016. Human memory T cells with a naive phenotype accumulate with aging and respond to persistent viruses. Nature Immunology. 17. 8. 966–975. 10.1038/ni.3483. 1529-2916. 4955715. 27270402.
  6. White. Jason T.. Cross. Eric W.. Kedl. Ross M.. June 2017. Antigen-inexperienced memory CD8+T cells: where they come from and why we need them. Nature Reviews. Immunology. 17. 6. 391–400. 10.1038/nri.2017.34. 1474-1741. 5569888. 28480897.
  7. von Essen MR, Kongsbak M, Schjerling P, Olgaard K, Odum N, Geisler C . Vitamin D controls T cell antigen receptor signaling and activation of human T cells . Nat. Immunol. . 11 . 4 . 344–9 . April 2010 . 20208539 . 10.1038/ni.1851 . 6119729 . 2010-12-26 . https://web.archive.org/web/20140912193535/http://www.microbio.uab.edu/CMIJournalClub/March17.pdf . 2014-09-12 . dead .
  8. Rudd CE, Trevillyan JM, Dasgupta JD, Wong LL, Schlossman S . Pillars article: the CD4 receptor is complexed in detergent lysates to a protein-tyrosine kinase (pp58) from human T lymphocytes . J. Immunol. . 185 . 5 . 2645–9 . September 2010 . 20724730 . 3791413.