Table 3.

Summary of published reports using 3D organoids for in vitro T cell differentiation

Gene defectStarting cellsUse of the 3D organoidsResultsReference
Healthy donor (HD) CD34+ Original protocol for the generation of ATOs starting from human CD34+ cells Establishment of a protocol for the efficient in vitro generation of mature T cells, fully functional and with a diverse TCR repertoire Seet et al., 2017  
HD iPSCs, embryonic stem cells Original protocol for the generation of ATOs starting from human iPSCs and ES Establishment of a protocol for the efficient in vitro generation of mature T cells, fully functional and with a diverse TCR repertoire Montel-Hagen et al., 2019  
AK2, IL2RG, DGS, RAG1/2, ADA CD34+ Test ATOs to evaluate block in differentiation in patients with known gene defects (1) Definition of T cell developmental block in patients with hematopoietic-intrinsic defects; (2) discrimination between hematopoietic-intrinsic and -extrinsic defects Bosticardo et al., 2020  
IL2RG, TBX1, DGS, RAG1/2, ADA CD34+ Test 3D aggregates to evaluate block in differentiation in patients with known gene defects (1) Definition of T cell developmental block in patients with hematopoietic-intrinsic defects; (2) discrimination between hematopoietic-intrinsic and -extrinsic defects Bifsha et al., 2020  
POLD1 CD34+ Evaluation of T cell differentiation in vitro POLD1-mutant CD34+ cells showed a block in differentiation, resulting in a reduced number of CD4+CD8+ DP cells and almost complete absence of TCRαβ+CD3+ cells Nichols-Vinueza et al., 2021  
RAG2 iPSCs Test efficacy of CRISPR/Cas9-mediated gene editing (GE) approach GE-iPSCs of a RAG2-mutant patient overcome block at CD4+CD8+ DP stage and show normal differentiation to mature TCRαβ+CD3+ cells Gardner et al., 2021  
SASH3 CD34+ Evaluation of T cell differentiation in vitro SASH3-mutant CD34+ cells were able reach full maturation, but frequency and absolute count of TCRαβ+CD3+ cells were significantly decreased, due to impaired cell cycle progression and increased apoptosis Delmonte et al., 2021  
MAGT1 CD34+ Test efficacy of CRISPR/Cas9-mediated GE approach (1) GE CD34+ cells from XMEN patients showed full T maturation; (2) restoration of NKG2D expression in CD3+ cells Brault et al., 2021  
HD CD34+ Understanding how PNP inactivation affects T cell differentiation from iPSCs (1) PNP inactivation causes decreased viability of CD34+ cells and accumulation in the CD4CD8 DN stage; (2) effects of PNP inactivation were mediated through the downregulation of the dNTP tri-phosphohydrolase SAMHD1 Abt et al., 2022  
HD iPSCs Test hypothesis for T-T–mediated selection of unconventional T cells PLZF-expressing unconventional T cells originate from positive selection on other T cells, as opposed to conventional T cells, which are selected by cTECs Suo et al., 2022  
FOXI3 CD34+ Evaluation of T cell differentiation in vitro (1) CD34+ cells isolated from a patient with FOXI3 haploinsufficiency show efficient differentiation to mature TCRαβ+CD3+ cells; (2) these results support the thymic-intrinsic nature of T cell lymphopenia in FOXI3-mutant patients Ghosh et al., 2022  
HD CD34+, TSP Evaluation the T cell differentiation potential of the different TSP (TSP1, TSP2, TSP3) TSP1 are a more quiescent population; TSP3 cells are primed to develop much more rapidly into T cells compared with TSP1 and TSP2 cells Cordes et al., 2022  
IL2RG CD34+ Test efficacy of lentiviral vector– and CRISPR/Cas9-mediated GE approaches Both treatments overcome the block at the CD34+ Pro T stage and show restoration of surface expression of IL2Rgc and its mediated signaling Brault et al., 2023  
TRIM37 CD34+ Evaluation of T cell differentiation in vitro Normal mature TCRαβ+CD3+ cell generation, excluding an early hematopoietic-intrinsic defect of T cell development Gazzin et al., 2023  
CD3D CD34+ Test efficacy of CRISPR/Cas9-mediated BE approach Edited CD3D-mutant patients’ CD34+ cells differentiated to fully mature T cells, displaying diverse TCR repertoires and TCR-dependent functions McAuley et al., 2023  
HD iPSCs Understanding how CARs affect T cell differentiation from iPSCs iPSCs transduced with a CD19-targeted CAR showed a diversion of T cell differentiation from conventional T cell to ILC2 lineage Li et al., 2023  
WAS CD34+ Evaluation of T cell differentiation in vitro (1) Partial block at the positive selection checkpoint, especially on the generation of CD8+ SP cells; (2) no effects on the generation of a diverse TCR repertoire Pille et al., 2023  
PAX1 CD34+ Evaluation of T cell differentiation in vitro Normal mature TCRαβ+CD3+ cells generation, excluding an early hematopoietic-intrinsic defect of T cell development Yakici et al., 2023  
RAG1 CD34+ Test efficacy of CRISPR/Cas9-mediated GE approach (1) Integration into intron 1 of RAG1 gene achieved suboptimal correction and didn't overcome the block in differentiation at the CD4+CD8+ DP stage; (2) in-frame insertion into exon 2 led to efficient generation of mature TCRαβ+CD3+ cells Castiello et al., 2024  
HD iPSCs Evaluation of the lymphoid differentiation potential of BMO-derived HSPC BMO-derived HSPCs can differentiate to mature TCRαβ+CD3+ cells Frenz-Wiessner et al., 2024  
PTCRA CD34+ Evaluation of T cell differentiation in vitro Selective defect in TCRαβ+CD3+ cell production, directly correlating with the severity of the genetic mutation, while the ability of producing TCRγδ+CD3+ cells was preserved Materna et al., 2024  
Gene defectStarting cellsUse of the 3D organoidsResultsReference
Healthy donor (HD) CD34+ Original protocol for the generation of ATOs starting from human CD34+ cells Establishment of a protocol for the efficient in vitro generation of mature T cells, fully functional and with a diverse TCR repertoire Seet et al., 2017  
HD iPSCs, embryonic stem cells Original protocol for the generation of ATOs starting from human iPSCs and ES Establishment of a protocol for the efficient in vitro generation of mature T cells, fully functional and with a diverse TCR repertoire Montel-Hagen et al., 2019  
AK2, IL2RG, DGS, RAG1/2, ADA CD34+ Test ATOs to evaluate block in differentiation in patients with known gene defects (1) Definition of T cell developmental block in patients with hematopoietic-intrinsic defects; (2) discrimination between hematopoietic-intrinsic and -extrinsic defects Bosticardo et al., 2020  
IL2RG, TBX1, DGS, RAG1/2, ADA CD34+ Test 3D aggregates to evaluate block in differentiation in patients with known gene defects (1) Definition of T cell developmental block in patients with hematopoietic-intrinsic defects; (2) discrimination between hematopoietic-intrinsic and -extrinsic defects Bifsha et al., 2020  
POLD1 CD34+ Evaluation of T cell differentiation in vitro POLD1-mutant CD34+ cells showed a block in differentiation, resulting in a reduced number of CD4+CD8+ DP cells and almost complete absence of TCRαβ+CD3+ cells Nichols-Vinueza et al., 2021  
RAG2 iPSCs Test efficacy of CRISPR/Cas9-mediated gene editing (GE) approach GE-iPSCs of a RAG2-mutant patient overcome block at CD4+CD8+ DP stage and show normal differentiation to mature TCRαβ+CD3+ cells Gardner et al., 2021  
SASH3 CD34+ Evaluation of T cell differentiation in vitro SASH3-mutant CD34+ cells were able reach full maturation, but frequency and absolute count of TCRαβ+CD3+ cells were significantly decreased, due to impaired cell cycle progression and increased apoptosis Delmonte et al., 2021  
MAGT1 CD34+ Test efficacy of CRISPR/Cas9-mediated GE approach (1) GE CD34+ cells from XMEN patients showed full T maturation; (2) restoration of NKG2D expression in CD3+ cells Brault et al., 2021  
HD CD34+ Understanding how PNP inactivation affects T cell differentiation from iPSCs (1) PNP inactivation causes decreased viability of CD34+ cells and accumulation in the CD4CD8 DN stage; (2) effects of PNP inactivation were mediated through the downregulation of the dNTP tri-phosphohydrolase SAMHD1 Abt et al., 2022  
HD iPSCs Test hypothesis for T-T–mediated selection of unconventional T cells PLZF-expressing unconventional T cells originate from positive selection on other T cells, as opposed to conventional T cells, which are selected by cTECs Suo et al., 2022  
FOXI3 CD34+ Evaluation of T cell differentiation in vitro (1) CD34+ cells isolated from a patient with FOXI3 haploinsufficiency show efficient differentiation to mature TCRαβ+CD3+ cells; (2) these results support the thymic-intrinsic nature of T cell lymphopenia in FOXI3-mutant patients Ghosh et al., 2022  
HD CD34+, TSP Evaluation the T cell differentiation potential of the different TSP (TSP1, TSP2, TSP3) TSP1 are a more quiescent population; TSP3 cells are primed to develop much more rapidly into T cells compared with TSP1 and TSP2 cells Cordes et al., 2022  
IL2RG CD34+ Test efficacy of lentiviral vector– and CRISPR/Cas9-mediated GE approaches Both treatments overcome the block at the CD34+ Pro T stage and show restoration of surface expression of IL2Rgc and its mediated signaling Brault et al., 2023  
TRIM37 CD34+ Evaluation of T cell differentiation in vitro Normal mature TCRαβ+CD3+ cell generation, excluding an early hematopoietic-intrinsic defect of T cell development Gazzin et al., 2023  
CD3D CD34+ Test efficacy of CRISPR/Cas9-mediated BE approach Edited CD3D-mutant patients’ CD34+ cells differentiated to fully mature T cells, displaying diverse TCR repertoires and TCR-dependent functions McAuley et al., 2023  
HD iPSCs Understanding how CARs affect T cell differentiation from iPSCs iPSCs transduced with a CD19-targeted CAR showed a diversion of T cell differentiation from conventional T cell to ILC2 lineage Li et al., 2023  
WAS CD34+ Evaluation of T cell differentiation in vitro (1) Partial block at the positive selection checkpoint, especially on the generation of CD8+ SP cells; (2) no effects on the generation of a diverse TCR repertoire Pille et al., 2023  
PAX1 CD34+ Evaluation of T cell differentiation in vitro Normal mature TCRαβ+CD3+ cells generation, excluding an early hematopoietic-intrinsic defect of T cell development Yakici et al., 2023  
RAG1 CD34+ Test efficacy of CRISPR/Cas9-mediated GE approach (1) Integration into intron 1 of RAG1 gene achieved suboptimal correction and didn't overcome the block in differentiation at the CD4+CD8+ DP stage; (2) in-frame insertion into exon 2 led to efficient generation of mature TCRαβ+CD3+ cells Castiello et al., 2024  
HD iPSCs Evaluation of the lymphoid differentiation potential of BMO-derived HSPC BMO-derived HSPCs can differentiate to mature TCRαβ+CD3+ cells Frenz-Wiessner et al., 2024  
PTCRA CD34+ Evaluation of T cell differentiation in vitro Selective defect in TCRαβ+CD3+ cell production, directly correlating with the severity of the genetic mutation, while the ability of producing TCRγδ+CD3+ cells was preserved Materna et al., 2024  

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