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Fig 1.

CD16+ Vδ2 T cells are more likely to express cytotoxic markers and KIRs.

(A) The percentage of CD16+ and CD16-Vδ2 T cells expressing relevant cytotoxic markers are shown (n = 65; scatter plot with median and IQR). (B) Vδ2 T cells positive for 4 or any combination of 3, 2 or 1 of the cytotoxicity markers listed in A are grouped along the x-axis and their relative percentage of CD16 is shown (n = 65; scatter plot with median and IQR). (C) The percentage of CD16+ and CD16-Vδ2s expressing the listed combinations of the transcription factors Tbet and Eomes are shown. (n = 65). (D) The percentage of CD16+ and CD16-Vδ2 T cells expressing the listed KIR or KIR combinations is shown. Individuals who were genotype negative for the listed KIRs were omitted from this analysis (n = 22 for KIR3DL1 and KIR2DS3/L1/S1/S5; n = 20 for KIR2DL3). (E) The percentage of CD16+ and CD16-Vδ2 T cells expressing IL18Rα and the MFI of IL18Rα on CD16+ and CD16-Vδ2 T cells is shown (n = 10). All Vδ2+ T cells shown were gated on singlets and CD3+ events to exclude NK cells. P values for A, B, D and E determined by Wilcoxon matched pairs signed rank test.

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Fig 2.

CD16+ Vδ2 T cells from malaria-exposed individuals downregulate TCR and can be independently stimulated through CD16.

(A)The MFI of the Vδ2 and Vγ9 chains is shown for CD16+ and CD16- Vδ2+Vγ9+ T cells. Data points and accompanying p-value are paired. (n = 39; scatter plots with median and IQR). (B) The percentage of Vδ2 T cells producing IFNγ in response to anti-CD16 crosslinking antibody or isotype is compared (n = 35; box plots with median and Tukey whiskers). (C) The percentage of Vδ2 T cells producting IFNγ in response to plate-bound anti-CD16 crosslinking antibody is graphed against the percentage of Vδ2 T cells expressing CD16 prior to stimulation (n = 35). (D) The percentage of Vδ2 T cells positive for IFNγ or CD107a after stimulation with anti-CD16 crosslinking antibody or iRBC is compared (n = 10) (E) Vδ2 T cells positive for IFNγ or CD107a after iRBC stimulation from the experiment in E are grouped by CD16 expression. All Vδ2+ T cells shown were gated on singlets and CD3+ events to exclude NK cells. P values determined by Wilcoxon matched pairs signed rank test in A, B, D and D, and with Spearman correlation in C.

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Fig 3.

Stimulation through CD16 augments Vδ2 cell activation by P.falciparum.

(A) Schematic detailing stimulation conditions for data presented in panel B. (B) Percent of Vδ2 T cells producing IFNγ or positive for CD107a after stimulation of whole PBMC with uRBC, iRBC or IPP, with or without plate bound anti-CD16 crosslinking antibody (n = 9). (C). Percent of Vδ2 T cells producing IFNγ or positive for CD107a after stimulation of negatively selected γδ T cells with uRBC or iRBC, with or without plate bound anti-CD16 crosslinking antibody (n = 4). (D) Live cells positive for γδ TCR after negative selection. All Vδ2+ T cells shown in B-C were gated on singlets and CD3+ events to exclude NK cells. Samples for these experiments were selected from individuals with greater than 30% of their Vδ2 T cells expressing CD16. P values were determined by Wilcoxon matched pairs signed rank test.

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Fig 4.

Stimulation by opsonized antigen augments Vδ2 cell activation by P.falciparum.

(A) Schematic detailing the stimulation conditions for data presented in panel B. (B) Percent of Vδ2 T cells positive for IFNγ and CD107a after stimulation of whole PBMC with uRBC, iRBC, iRBC + naïve IgG, or iRBC + hyper-immune IgG. (n = 6). (C) Percent of Vδ2 T cells positive for IFNγ and CD107a after stimulation of negatively selected γδ T cells with uRBC, iRBC, iRBC + naïve IgG, or iRBC + hyper-immune IgG. (n = 9). All Vδ2+ T cells shown were gated on singlets and CD3+ events to exclude NK cells. Samples for this experiment were selected from individuals with greater than 30% of their Vδ2 T cells expressing CD16. P values were determined by Wilcoxon matched pairs signed rank test.

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Fig 5.

Blocking BTN3a1 does not inhibit stimulation of Vδ2 T cells by opsonized antigen.

Percent of Vδ2 T cells positive for IFNγ and CD107a after stimulation with iRBC + naïve IgG, or with iRBC + hyper-immune IgG, are compared in the presence or absence of a Butyrophilin 3a1 blocking antibody. (n = 10; p values determined by Wilcoxon matched pairs signed rank test). All Vδ2+ T cells shown were gated on singlets and CD3+ events to exclude NK cells. Samples for this experiment were selected from adults with greater than 30% of their Vδ2s expressing CD16.

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Fig 6.

Model of Vδ2 T cells over the course of repeated Pf infection.

As an individual experiences more episodes of Pf, the frequency of Vδ2 T cells in their peripheral blood declines. Remaining Vδ2 T cells become less responsive through their TCRs while the frequency of CD16+ expression increases. These changes occur while an individual’s anti-Pf antibody response is simultaneously growing and diversifying, presumably providing more opsonized parasite targets for CD16+ Vδ2 T cells. TCR-responsive Vδ2s are activated by engagement with BTN3A1 on another cell, probably a lymphocyte, when BTN3A1’s conformation is altered by the intracellular binding of parasite-derived phosphoantigens. CD16-responsive Vδ2 T cells are activated by opsonized parasite antigen and may have their activation threshold modulated by KIR expression and engagement. Created with Biorender.

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