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. 2019 Aug 20;13:381. doi: 10.3389/fncel.2019.00381

TABLE 1.

List of genes involved in bRG generation.

Gene/Protein Function Model Manipulation Effect(s) on progenitors Cell mechanism described Other phenotype(s) described
ARHGAP11B ? Mouse (IUE+mRNA microinjection) Expression IPs↗, basal mitoses↗ Cell fate change in aRG to produce more BPs, Proliferation↗ Thicker SVZ, Cortical folds, cortical expansion
Ferret (IUE) Expression IPs↘, bRGs↗ Oblique/horizontal division in aRGs, Proliferation↗ Cortical expansion, upper-layer neurons↗
Aspm MT associated protein Ferret (transgenic line) KO bRGs↗ Early delamination of aRGs Thinner VZ, microcephaly with conserved cortical layers
Mouse (transgenic line) KO ? Mislocalized centrosomes and aberrant ventricular lining
Cdh1 Cell adhesion molecule Ferret (IUE) Expression of dominant-negative form aRGs↘, bRGs↗ Cell adhesion↘, Oblique/horizontal division in aRGs
FGFR1, MEK, ERK, ETV4 RTK, MAPK, TF Mouse (IUE) Pathway activation (constitutively active form of FGFR1, Mek, Etv4) bRGs↗ Proliferation↗ Upper-layer neurons↗
FGFRs RTK Ferret (IUE) Pathway inhibition bRGs↘ Proliferation↘ in bRGs Loss of cortical folds
GPSM2 (LGN) Regulator of mitotic apparatus Mouse (IUE/transgenic line) Expression of dominant-negative form bRGs↗ Oblique/horizontal division in aRGs
Hif1α TF Mouse Regulation of the pathway (Hyperoxia during pregnancy) IPs↗, bRGs↗ ? OSVZ-like region
INSM1 TF Mouse (IUE) Overexpression IPs↗ in the VZ, bRGs↗ Cell detachment from VZ, disruption of adherens junctions
Pax6 TF Mouse (IUE/transgenic line) Forced expression in BPs IPs↘, bRGs↗ Oblique/horizontal division in aRGs, Proliferation↗ OSVZ-like region, thicker cortex, upper-layer neurons↗
Mouse (transgenic line) KO bRGs↗ Cell adhesion↘, mostly non-cell autonomous
PDGFD/PDGFR signaling PDGF receptor/ligand Human brain slices Pathway inhibition (treatment with PDGFRβ inhibitor) IPs↘, bRGs↘ ?
Mouse (IUE+PDGF-DD injection) Expression (constitutively active form of PDGFRβ)/Pathway activation bRGs↗
PTEN Phosphatase Human organoids Pathway activation (CRISPR/Cas9 targeting of PTEN) bRGs↗ ? Increased volume and cortical folds
Mouse organoids ? Increased volume but no cortical folds
SMARCC1 (Baf155) Chromatin remodeling factor Mouse (IUE/transgenic line) KO IPs↘ (ectopic IPs in IZ), bRGs↗ Oblique/horizontal division in aRGs, regulates Pax6, cell adhesion↘ -
SmoM2 Shh pathway activator Mouse (transgenic line) Pathway activation (constitutively active form of SmoM2) IPs↗, bRGs↗ Proliferation↗ Cortical folds, brain expansion
TBC1D3 RABGAP Mouse (IUE/transgenic line) Expression aRGs↗, IPs↗, bRGs↗ Cell adhesion↘, proliferation↗ Cortical folds
Human brain slices (electroporation) KD bRGs↘ ?
TMEM14B Kinase? Mouse (IUE) Expression IPs↗, bRGs↗ Oblique/horizontal division in aRGs, Proliferation↗ OSVZ-like region, cortical folds
Trnp1 TF Mouse (IUE) KD aRGs↘, IPs↗, bRGs↗ Cell adhesion? Cortical folds

In this table are indicated the genes, their function, the model in which they were tested, the type of experiment performed, the effect on progenitor cells, the cellular mechanisms indicated in each study and other phenotypes associated with bRG regulation. IUE, in utero electroporation; aRGs↘, IPs↘, bRGs↘, diminution of the proportion of aRGs, IPs, and/or bRGs; aRGs↗, IPs↗, and bRGs↗, increase of the proportion of aRGs, IPs, and/or bRGs; Cell adhesion↘, diminution of cell adhesion (generally associated with loss or diminution of expression of adherens junction proteins or other cell adhesion molecules). Proliferation↗, increase in the proportion of mitotic and/or cycling cells; upper-layer neurons↗, increase in the proportion of upper-layer neurons in the cortical plate.