(Hartenstein, 1993; Pereanu et al., 2007). A ganglionic branch (one per segment) is a primary epithelial tube made by seven cells (Samakovlis et al., 1996). During budding and elongation, a single cell at the tip of the emerging structure leads the migratory process towards the CNS. This process starts during embryogenesis around 10 h after egg laying and is guided by the presence of clusters of epidermal cells expressing *branchless*, a member of the Fibroblast Growth Factor family (Sutherland et al., 1996). Within the next two hours, the leading cell at the tip of the ganglionic branch reaches, migrates along and finally enters the ventral nerve cord laterally and ventrally (Samakovlis et al., 1996; Englund et al., 1999). By the beginning of the larval stage, a ring-like rudimentary tracheal plexus (much like the mammalian perineural vascular plexus) is set up around the neuropile by the invasion, extension/migration and fusion of segmental ganglionic branches. From there, this plexus will grow and branch throughout larval stages to form a complex network mostly localized close to the neuropile (Pereanu et al., 2007; Baccino-Calace et al., 2020). Their growth appears constrained by glial cells (Pereanu et al., 2007) and involves some proliferative mechanisms at late stages (Rao et al., 2015). Little is known about the coordination between trachea growth, function and interplay with other cell types of the niche. One common sensitivity is to nutrition-induced PI3K/Akt signalling at early stage, which fuels tracheal growth. In turn, tracheal growth seems to favour NSC reactivation (Yuan et al., 2020). Whether this role is linked to the supply of oxygen or of other signalling factors is not known. Interestingly, the distance between the tracheal network and NSCs increases during larval stage along with NSC lineage growth (Bailey et al., 2015).

### 5.3.5 The fat body: an example of a distant niche

The fat body is a crucial player in NSC reactivation, secreting nutrition-responsive factors, yet unknown, which reach the CNS and trigger insulin signalling from the glial cells (Britton and Edgar, 1998; Chell and Brand, 2010; Sousa-Nunes et al., 2011; Spéder and Brand, 2014). Fat body development (reviewed in (Zheng et al., 2016; Li et al., 2019; Parra-Peralbo et al., 2021)) starts with the progressive apparition of mesodermal cell clusters between embryonic stages 10-14 (5-10 h after egg laying). These fat body primordia undergo mitosis to reach around 2,200 cells and shortly afterwards at stage 16 (13-16 h after egg laying) fuse to generate one continuous monolayer of fat cells. These cells keep growing through endoreplication, until entering quiescence in late embryogenesis. Fat cells will resume endoreplication around 8 h after larval hatching, paralleling the timing of NSC reactivation, and grow throughout larval stages without a change in number, while storing various nutrients.

## 6 Conclusion

### 6.1 Take home messages across all model systems

Niche morphogenesis is a coordinated process between different cell types and acellular components to ensure that the needs of the tissue to develop and acquire specialized functions are met in the right time and place. The timing of growth, adhesion and intercellular communication will generate a specific, yet dynamic, niche

architecture, which in turn is a powerful means to regulate the reach of signalling.

The scales of both niche morphogenesis and niche architecture are multifold. Niche morphogenesis spans time (from early embryonic development to early post-natal growth, and even adulthood) and tissues (neural, hematopoietic, and immune compartments). Niche architecture is built on cell-to-cell adhesions (ECM, adherens junctions, tight junctions, surface immunoglobulins, gap junctions), which will act to generate local structures (e.g., astrocyte/blood vessel/RGC basal process unit; cortex glia chamber) and signalling (e.g., primary cilia of RGCs; ECM/integrin signalling), but which also can reach to farther regions of the tissue (e.g., gap junction coupling in the subperineurial glia; attachment of RGCs' basal process to blood vessels; ECM as a mechanical constraint). Some of these molecular and cellular processes can also be regional (e.g., ependymal pinwheel structure grouping few RGCs together; mosaic of cortex glia), producing "mini-niches" within the niche.

### 6.2 Where the neurogenic niche field should go next

Stem cell niches have emerged as powerful and plastic regulators of stem cell behaviour. The neurogenic niche is no exception yet distinguishes itself by its especially heterogeneous and complex composition, the inclusion of components with unique role at the organism level (i.e., the blood-brain barrier) and its multiscale architecture.

While the field has made critical progress in the signalling aspect of the neurogenic niche, and despite the growing understanding that mechanical cues and physical constraints have powerful implications, little is still known on niche architecture.

The first question which has been explored in recent years is how the niche forms and acquired its “final” (mature) architecture (at least in homeostatic conditions), since such understanding is a prerequisite to its manipulation for probing its function. Deciphering niche formation includes the identification of the morphogenetic processes at play, their cellular and molecular players, and the adhesive properties eventually set in place. The mouse and the fruit fly have been leading this aspect, with different focuses, and with still much left to comprehend. Niche morphogenesis is pretty much unknown in the CNS of other organisms, and one of the further challenges of the field is to bring this comparative understanding to the table.

Nearly all is left to achieve on the other aspects of niche architecture, covering scales, rules, and functions. Our own perception of the neurogenic niche, and its relationship with other fields of biological research, would suggest concentrating on the following questions.

1. Interplay between architecture and signalling, including physical/ mechanical cues both as a signal *per se* and as a modulator of signalling range

2. Relationship between local, regional and tissue scales, especially in relationship with the cellular coordination or independence within the NSPC population

3. Plasticity in response to time (from development to ageing), signals (metabolic status), and conditions (pathologies)
(Hartenstein, 1993; Pereanu et al., 2007). A ganglionic branch (one per segment) is a primary epithelial tube made by seven cells (Samakovlis et al., 1996). During budding and elongation, a single cell at the tip of the emerging structure leads the migratory process towards the CNS. This process starts during embryogenesis around 10 h after egg laying and is guided by the presence of clusters of epidermal cells expressing *branchless*, a member of the Fibroblast Growth Factor family (Sutherland et al., 1996). Within the next two hours, the leading cell at the tip of the ganglionic branch reaches, migrates along and finally enters the ventral nerve cord laterally and ventrally (Samakovlis et al., 1996; Englund et al., 1999). By the beginning of the larval stage, a ring-like rudimentary tracheal plexus (much like the mammalian perineural vascular plexus) is set up around the neuropile by the invasion, extension/migration and fusion of segmental ganglionic branches. From there, this plexus will grow and branch throughout larval stages to form a complex network mostly localized close to the neuropile (Pereanu et al., 2007; Baccino-Calace et al., 2020). Their growth appears constrained by glial cells (Pereanu et al., 2007) and involves some proliferative mechanisms at late stages (Rao et al., 2015). Little is known about the coordination between trachea growth, function and interplay with other cell types of the niche. One common sensitivity is to nutrition-induced PI3K/Akt signalling at early stage, which fuels tracheal growth. In turn, tracheal growth seems to favour NSC reactivation (Yuan et al., 2020). Whether this role is linked to the supply of oxygen or of other signalling factors is not known. Interestingly, the distance between the tracheal network and NSCs increases during larval stage along with NSC lineage growth (Bailey et al., 2015).

### 5.3.5 The fat body: an example of a distant niche

The fat body is a crucial player in NSC reactivation, secreting nutrition-responsive factors, yet unknown, which reach the CNS and trigger insulin signalling from the glial cells (Britton and Edgar, 1998; Chell and Brand, 2010; Sousa-Nunes et al., 2011; Spéder and Brand, 2014). Fat body development (reviewed in (Zheng et al., 2016; Li et al., 2019; Parra-Peralbo et al., 2021)) starts with the progressive apparition of mesodermal cell clusters between embryonic stages 10-14 (5-10 h after egg laying). These fat body primordia undergo mitosis to reach around 2,200 cells and shortly afterwards at stage 16 (13-16 h after egg laying) fuse to generate one continuous monolayer of fat cells. These cells keep growing through endoreplication, until entering quiescence in late embryogenesis. Fat cells will resume endoreplication around 8 h after larval hatching, paralleling the timing of NSC reactivation, and grow throughout larval stages without a change in number, while storing various nutrients.

## 6 Conclusion

### 6.1 Take home messages across all model systems

Niche morphogenesis is a coordinated process between different cell types and acellular components to ensure that the needs of the tissue to develop and acquire specialized functions are met in the right time and place. The timing of growth, adhesion and intercellular communication will generate a specific, yet dynamic, niche

architecture, which in turn is a powerful means to regulate the reach of signalling.

The scales of both niche morphogenesis and niche architecture are multifold. Niche morphogenesis spans time (from early embryonic development to early post-natal growth, and even adulthood) and tissues (neural, hematopoietic, and immune compartments). Niche architecture is built on cell-to-cell adhesions (ECM, adherens junctions, tight junctions, surface immunoglobulins, gap junctions), which will act to generate local structures (e.g., astrocyte/blood vessel/RGC basal process unit; cortex glia chamber) and signalling (e.g., primary cilia of RGCs; ECM/integrin signalling), but which also can reach to farther regions of the tissue (e.g., gap junction coupling in the subperineurial glia; attachment of RGCs' basal process to blood vessels; ECM as a mechanical constraint). Some of these molecular and cellular processes can also be regional (e.g., ependymal pinwheel structure grouping few RGCs together; mosaic of cortex glia), producing "mini-niches" within the niche.

### 6.2 Where the neurogenic niche field should go next

Stem cell niches have emerged as powerful and plastic regulators of stem cell behaviour. The neurogenic niche is no exception yet distinguishes itself by its especially heterogeneous and complex composition, the inclusion of components with unique role at the organism level (i.e., the blood-brain barrier) and its multiscale architecture.

While the field has made critical progress in the signalling aspect of the neurogenic niche, and despite the growing understanding that mechanical cues and physical constraints have powerful implications, little is still known on niche architecture.

The first question which has been explored in recent years is how the niche forms and acquired its “final” (mature) architecture (at least in homeostatic conditions), since such understanding is a prerequisite to its manipulation for probing its function. Deciphering niche formation includes the identification of the morphogenetic processes at play, their cellular and molecular players, and the adhesive properties eventually set in place. The mouse and the fruit fly have been leading this aspect, with different focuses, and with still much left to comprehend. Niche morphogenesis is pretty much unknown in the CNS of other organisms, and one of the further challenges of the field is to bring this comparative understanding to the table.

Nearly all is left to achieve on the other aspects of niche architecture, covering scales, rules, and functions. Our own perception of the neurogenic niche, and its relationship with other fields of biological research, would suggest concentrating on the following questions.

1. Interplay between architecture and signalling, including physical/mechanical cues both as a signal *per se* and as a modulator of signalling range

2. Relationship between local, regional and tissue scales, especially in relationship with the cellular coordination or independence within the NSPC population

3. Plasticity in response to time (from development to ageing), signals (metabolic status), and conditions (pathologies)
The answers to all these questions bid exciting times for stem cell biology, converging with other outstanding questions in the field touching their regulation under stress and their double-edged potential for curative (e.g., regeneration) and pathological (e.g., tumourigenesis) outcomes.

## Author contributions

GV: Writing-original draft, Writing-review and editing. PS: Funding acquisition, Writing-original draft, Writing-review and editing,

## Funding

The author(s) declare financial support was received for the research, authorship, and/or publication of this article. This work has been funded by a core funding from Institut Pasteur to PS. GV was supported by a post-doctoral fellowship from the LabEx Revive (ANR-10-LABX-0073).

## References

* Ackerman, S. D., and Monk, K. R. (2016). The scales and tales of myelination: using zebrafish and mouse to study myelinating glia. *Brain Res.* 1641, 79-91. doi:10.1016/J.BRAINRES.2015.10.011

* Adolf, B., Chapouton, P., Lam, C. S., Topp, S., Tannhäuser, B., Strähle, U., et al. (2006). Conserved and acquired features of adult neurogenesis in the zebrafish telencephalon. *Dev. Biol.* 295, 278-293. doi:10.1016/J.YDBIO.2006.03.023

* Agirman, G., Broix, L., and Nguyen, L. (2017). Cerebral cortex development: an outside-in perspective. *FEBS Lett.* 591, 3978-3992. doi:10.1002/1873-3468.12924

* Alliot, F., Godin, I., and Pessac, B. (1999). Microglia derive from progenitors, originating from the yolk sac, and which proliferate in the brain. *Dev. Brain Res.* 117, 145-152. doi:10.1016/S0165-3806(99)00113-3

* Altenhein, B., Cattenzop, P. B., and Giangrande, A. (2015). The early life of a fly glial cell. *Wiley Interdiscip. Rev. Dev. Biol.* 5, 67-84. doi:10.1002/wdev.200

* Altman, J., and Das, G. D. (1965). Autoradiographic and histological evidence of postnatal hippocampal neurogenesis in rats. *J. Comp. Neurol.* 124, 319-335. doi:10.1002/CNE.901240303

* Alunni, A., and Bally-Cuif, L. (2016). A comparative view of regenerative neurogenesis in vertebrates. *Development* 143, 741-753. doi:10.1242/dev.122796

* Araya, C., Ward, L. C., Girdler, G. C., and Miranda, M. (2016). Coordinating cell and tissue behavior during zebrafish neural tube morphogenesis. *Dev. Dyn.* 245, 197-208. doi:10.1002/DVDY.24304

* Armulik, A., Genové, G., and Betsholtz, C. (2011). Pericytes: developmental, physiological, and pathological perspectives, problems, and promises. *Dev. Cell* 21, 193-215. doi:10.1016/J.DEVCEL.2011.07.001

* Arnò, B., Grassivaro, F., Rossi, C., Bergamaschi, A., Castiglioni, V., Furlan, R., et al. (2014). Neural progenitor cells orchestrate microglia migration and positioning into the developing cortex. *Nat. Commun.* 5, 5611. doi:10.1038/NCOMMS6611

* Avet-Rochex, A., Kaul, A. K., Gatt, A. P., McNeill, H., and Bateman, J. M. (2012). Concerted control of gliogenesis by InR/TOR and FGF signalling in the Drosophila post-embryonic brain. *Development* 139, 2763-2772. doi:10.1242/dev.074179

* Kawasaki, T., Lai, S.-L., Ito, K., and Lee, T. (2008). Organization and postembryonic development of glial cells in the adult central brain of Drosophila. *J. Neurosci.* 28, 13742-13753. doi:10.1523/JNEUROSCI.4844-08.2008

* Atwatramanil, R., Soriano, P., Rodriguez, C., Mai, J. J., and Dymecki, S. M. (2003). Cryptic boundaries in roof plate and choroid plexus identified by intersectional gene activation. *Nat. Genet.* 35, 70-75. doi:10.1038/NG1228

* Babatz, F., Faffin, E., and Klämbt, C. (2018). The Drosophila blood-brain barrier adapts to cell growth by unfolding of pre-existing septate junctions. *Dev. Cell* 47, 697-710. doi:10.1016/j.devcel.2018.10.002

## Acknowledgments

We deeply thank Laure Bally-Cuif, Alice Davy and Elisa Gomez Perdiguero for critical and helpful comments on this review. We are very grateful to Mónica Folgueira for discussion about the zebrafish *tela choroidea* and telencephalic eversion.

## Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

## Publisher's note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

* Baccino-Calace, M., Prieto, D., Cantera, R., and Egger, B. (2020). Compartment and cell-type specific hypoxia responses in the developing Drosophila brain. *Biol. Open* 9, bio053629. doi:10.1242/bio.053629

* Bahrami, N., and Childs, S. J. (2018). Pericyte biology in zebrafish. *Adv. Exp. Med. Biol.* 1109, 33-51. doi:10.1007/978-3-030-02601-1_4

* Bailey, A. P., Koster, G., Guillermier, C., Hirst, E. M. A., MacRae, J. I., Lechene, C. P., et al. (2015). Antioxidant role for lipid droplets in a stem cell niche of Drosophila. *Cell* 163, 340-353. doi:10.1016/j.cell.2015.09.020

* Bainton, R. J., Tsai, L. T.-Y., Schwabe, T., DeSalvo, M., Gaul, U., and Heberlein, U. (2005). Moody encodes two GPCRs that regulate cocaine behaviors and blood-brain barrier permeability in Drosophila. *Cell* 123, 145-156. doi:10.1016/j.cell.2005.07.029

* Banach-Latapy, A., Rincheval, V., Briand, D., Guénal, I., and Spéder, P. (2023). Differential adhesion during development establishes individual neural stem cell niches and shapes adult behaviour in Drosophila. *PLoS Biol.* 21, e3002352. doi:10.1371/journal.pbio.3002352

* Barbosa, J. S., and Ninkovic, J. (2016). Adult neural stem cell behavior underlying constitutive and restorative neurogenesis in zebrafish. *Neurogenesis* 3, e1148101-e1148109. doi:10.1080/23262133.2016.1148101

* Barbosa, J. S., Sanchez-Gonzalez, R., Di Giaimo, R., Baumgart, E. V., Theis, F. J., Götz, M., et al. (2015). Neurodevelopment. Live imaging of adult neural stem cell behavior in the intact and injured zebrafish brain. *Science* 348, 789-793. doi:10.1126/SCIENCE.AAA2729

* Baumgart, E. V., Barbosa, J. S., Bally-cuif, L., Götz, M., and Ninkovic, J. (2012). Stab wound injury of the zebrafish telencephalon: a model for comparative analysis of reactive gliosis. *Glia* 60, 343-357. doi:10.1002/GLIA.22269

* Bayraktar, O. A., Fuentealba, L. C., Alvarez-Buylla, A., and Rowitch, D. H. (2014). Astrocyte development and heterogeneity. *Cold Spring Harb. Perspect. Biol.* 7, a020362. doi:10.1101/CSHPERSPECT.A020362

* Beckervordersandforth, R. M., Rickert, C., Altenhein, B., and Technau, G. M. (2008). Subtypes of glial cells in the Drosophila embryonic ventral nerve cord as related to lineage and gene expression. *Mech. Dev.* 125, 542-557. doi:10.1016/j.mod.2007.12.004

* Bello, B. C., Hirth, F., and Gould, A. P. (2003). A pulse of the Drosophila Hox protein Abdominal-A schedules the end of neural proliferation via neuroblast apoptosis. *Neuron* 37, 209-219. doi:10.1016/S0896-6273(02)01181-9

* Bello, B. C., Izergina, N., Caussinus, E., and Reichert, H. (2008). Amplification of neural stem cell proliferation by intermediate progenitor cells in Drosophila brain development. *Neural Dev.* 3, 5. doi:10.1186/1749-8104-3-5

* Benz, F., and Liebner, S. (2022). Structure and function of the blood-brain barrier (BBB). *Handb. Exp. Pharmacol.* 273, 3-31. doi:10.1007/164_2020_404

* Ben-Zvi, A., Lacoste, B., Kur, E., Andreone, B. J., Mayshar, Y., Yan, H., et al. (2014). Mfsd2a is critical for the formation and function of the blood-brain barrier. *Nat* 509, 507-511. doi:10.1038/nature13324
The answers to all these questions bid exciting times for stem cell biology, converging with other outstanding questions in the field touching their regulation under stress and their double-edged potential for curative (e.g., regeneration) and pathological (e.g., tumourigenesis) outcomes.

## Author contributions

GV: Writing-original draft, Writing-review and editing. PS: Funding acquisition, Writing-original draft, Writing-review and editing,

## Funding

The author(s) declare financial support was received for the research, authorship, and/or publication of this article. This work has been funded by a core funding from Institut Pasteur to PS. GV was supported by a post-doctoral fellowship from the LabEx Revive (ANR-10-LABX-0073).

## References

Ackerman, S. D., and Monk, K. R. (2016). The scales and tales of myelination: using zebrafish and mouse to study myelinating glia. *Brain Res.* 1641, 79-91. doi:10.1016/J.BRAINRES.2015.10.011

Adolf, B., Chapouton, P., Lam, C. S., Topp, S., Tannhäuser, B., Strähle, U., et al. (2006). Conserved and acquired features of adult neurogenesis in the zebrafish telencephalon. *Dev. Biol.* 295, 278-293. doi:10.1016/J.YDBIO.2006.03.023

Agirman, G., Broix, L., and Nguyen, L. (2017). Cerebral cortex development: an outside-in perspective. *FEBS Lett.* 591, 3978-3992. doi:10.1002/1873-3468.12924

Alliot, F., Godin, I., and Pessac, B. (1999). Microglia derive from progenitors, originating from the yolk sac, and which proliferate in the brain. *Dev. Brain Res.* 117, 145-152. doi:10.1016/S0165-3806(99)00113-3

Altenhein, B., Cattenzop, P. B., and Giangrande, A. (2015). The early life of a fly glial cell. *Wiley Interdiscip. Rev. Dev. Biol.* 5, 67-84. doi:10.1002/wdev.200

Altman, J., and Das, G. D. (1965). Autoradiographic and histological evidence of postnatal hippocampal neurogenesis in rats. *J. Comp. Neurol.* 124, 319-335. doi:10.1002/CNE.901240303

Alunni, A., and Bally-Cuif, L. (2016). A comparative view of regenerative neurogenesis in vertebrates. *Development* 143, 741-753. doi:10.1242/dev.122796

Araya, C., Ward, L. C., Girdler, G. C., and Miranda, M. (2016). Coordinating cell and tissue behavior during zebrafish neural tube morphogenesis. *Dev. Dyn.* 245, 197-208. doi:10.1002/DVDY.24304

Armulik, A., Genové, G., and Betsholtz, C. (2011). Pericytes: developmental, physiological, and pathological perspectives, problems, and promises. *Dev. Cell* 21, 193-215. doi:10.1016/J.DEVCEL.2011.07.001

Arnò, B., Grassivaro, F., Rossi, C., Bergamaschi, A., Castiglioni, V., Furlan, R., et al. (2014). Neural progenitor cells orchestrate microglia migration and positioning into the developing cortex. *Nat. Commun.* 5, 5611. doi:10.1038/NCOMMS6611

Avet-Rochex, A., Kaul, A. K., Gatt, A. P., McNeill, H., and Bateman, J. M. (2012). Concerted control of gliogenesis by InR/TOR and FGF signalling in the Drosophila post-embryonic brain. *Development* 139, 2763-2772. doi:10.1242/dev.074179

Awasaki, T., Lai, S.-L., Ito, K., and Lee, T. (2008). Organization and postembryonic development of glial cells in the adult central brain of Drosophila. *J. Neurosci.* 28, 13742-13753. doi:10.1523/JNEUROSCI.4844-08.2008

Atwatramanil, R., Soriano, P., Rodriguez, C., Mai, J. J., and Dymecki, S. M. (2003). Cryptic boundaries in roof plate and choroid plexus identified by intersectional gene activation. *Nat. Genet.* 35, 70-75. doi:10.1038/NG1228

Babatz, F., Faffin, E., and Klämbt, C. (2018). The Drosophila blood-brain barrier adapts to cell growth by unfolding of pre-existing septate junctions. *Dev. Cell* 47, 697-710. doi:10.1016/j.devcel.2018.10.002

## Acknowledgments

We deeply thank Laure Bally-Cuif, Alice Davy and Elisa Gomez Perdiguero for critical and helpful comments on this review. We are very grateful to Mónica Folgueira for discussion about the zebrafish *tela choroidea* and telencephalic eversion.

## Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

## Publisher's note

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.

Baccino-Calace, M., Prieto, D., Cantera, R., and Egger, B. (2020). Compartment and cell-type specific hypoxia responses in the developing Drosophila brain. *Biol. Open* 9, bio053629. doi:10.1242/bio.053629

Bahrami, N., and Childs, S. J. (2018). Pericyte biology in zebrafish. *Adv. Exp. Med. Biol.* 1109, 33-51. doi:10.1007/978-3-030-02601-1_4

Bailey, A. P., Koster, G., Guillermier, C., Hirst, E. M. A., MacRae, J. I., Lechene, C. P., et al. (2015). Antioxidant role for lipid droplets in a stem cell niche of Drosophila. *Cell* 163, 340-353. doi:10.1016/j.cell.2015.09.020

Bainton, R. J., Tsai, L. T.-Y., Schwabe, T., DeSalvo, M., Gaul, U., and Heberlein, U. (2005). Moody encodes two GPCRs that regulate cocaine behaviors and blood-brain barrier permeability in Drosophila. *Cell* 123, 145-156. doi:10.1016/j.cell.2005.07.029

Banach-Latapy, A., Rincheval, V., Briand, D., Guénal, I., and Spéder, P. (2023). Differential adhesion during development establishes individual neural stem cell niches and shapes adult behaviour in Drosophila. *PLoS Biol.* 21, e3002352. doi:10.1371/journal.pbio.3002352

Barbosa, J. S., and Ninkovic, J. (2016). Adult neural stem cell behavior underlying constitutive and restorative neurogenesis in zebrafish. *Neurogenesis* 3, e1148101-e1148109. doi:10.1080/23262133.2016.1148101

Barbosa, J. S., Sanchez-Gonzalez, R., Di Giaimo, R., Baumgart, E. V., Theis, F. J., Götz, M., et al. (2015). Neurodevelopment. Live imaging of adult neural stem cell behavior in the intact and injured zebrafish brain. *Science* 348, 789-793. doi:10.1126/SCIENCE.AAA2729

Baumgart, E. V., Barbosa, J. S., Bally-cuif, L., Götz, M., and Ninkovic, J. (2012). Stab wound injury of the zebrafish telencephalon: a model for comparative analysis of reactive gliosis. *Glia* 60, 343-357. doi:10.1002/GLIA.22269

Bayraktar, O. A., Fuentealba, L. C., Alvarez-Buylla, A., and Rowitch, D. H. (2014). Astrocyte development and heterogeneity. *Cold Spring Harb. Perspect. Biol.* 7, a020362. doi:10.1101/CSHPERSPECT.A020362

Beckervordersandforth, R. M., Rickert, C., Altenhein, B., and Technau, G. M. (2008). Subtypes of glial cells in the Drosophila embryonic ventral nerve cord as related to lineage and gene expression. *Mech. Dev.* 125, 542-557. doi:10.1016/j.mod.2007.12.004

Bello, B. C., Hirth, F., and Gould, A. P. (2003). A pulse of the Drosophila Hox protein Abdominal-A schedules the end of neural proliferation via neuroblast apoptosis. *Neuron* 37, 209-219. doi:10.1016/S0896-6273(02)01181-9

Bello, B. C., Izergina, N., Caussinus, E., and Reichert, H. (2008). Amplification of neural stem cell proliferation by intermediate progenitor cells in Drosophila brain development. *Neural Dev.* 3, 5. doi:10.1186/1749-8104-3-5

Benz, F., and Liebner, S. (2022). Structure and function of the blood-brain barrier (BBB). *Handb. Exp. Pharmacol.* 273, 3-31. doi:10.1007/164_2020_404

Ben-Zvi, A., Lacoste, B., Kur, E., Andreone, B. J., Mayshar, Y., Yan, H., et al. (2014). Mfsd2a is critical for the formation and function of the blood-brain barrier. *Nat* 509, 507-511. doi:10.1038/nature13324
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Bill, B. R., Balcuinas, D., McCarra, J. A., Young, E. D., Xiong, T., Spahn, A. M., et al. (2008). Development and Notch signaling requirements of the zebrafish choroid plexus. *PLoS One* 3, e3114. doi:10.1371/JOURNAL.PONE.0003114

Binder, M. D., Hirokawa, N., and Windhorst, U. (2009). Encyclopedia of neuroscience. Available at: [https://books.google.com/books/about/Encyclopedia_of_Neuroscience.html?id=mcvhAAAACAAJ](Accessed November 4, 2023).4393.

Biswas, S., Cottarelli, A., and Agalliu, D. (2020). Neuronal and glial regulation of CNS angiogenesis and barriergenesis. *Dev* 147, dev182279. doi:10.1242/dev.182279

Bjornson, C. R., Rietze, R. L., Reynolds, B. A., Magli, M. C., and Vescovi, A. L. (1999). Turning brain into blood: a hematopoietic fate adopted by adult neural stem cells *in vivo*. *Science* 283 (5401). New York, NY: JPEG, 534–537. doi:10.1126/science.283.5401.534

Bjornsson, C. S., Apostolopoulou, M., Tian, Y., and Temple, S. (2015). It takes a village: constructing the neurogenic niche. *Dev. Cell* 32, 435–446. doi:10.1016/j.devcel.2015.01.010

Blaschke, A. J., Weiner, J. A., and Chun, J. (1998). Programmed cell death is a universal feature of embryonic and postnatal neuroproliferative regions throughout the central nervous system. *J. Comp. Neurol.* 396, 39–50. doi:10.1002/(sici)1096-9861(19980622)396:1<39;aid-cne4>3.0.co;2-j(19980622)

Bond, A. M., Ming, G. L., and Song, H. (2015). Adult mammalian neural stem cells and neurogenesis: five decades later. *Cell Stem Cell* 17, 385–395. doi:10.1016/j.stem.2015.09.003

Bonvento, G., and Bolaños, J. P. (2021). Astrocyte-neuron metabolic cooperation shapes brain activity. *Cell Metab.* 33, 1546–1564. doi:10.1016/j.CMET.2021.07.006

Boone, J. Q., and Doe, C. Q. (2008). Identification of Drosophila type II neuroblast lineages containing transit amplifying ganglion mother cells. *Dev. Neurobiol.* 68, 1185–1195. doi:10.1002/dneu.20648

Bossing, T., Udolph, G., Doe, C. Q., and Technau, G. M. (1996). The embryonic central nervous system lineages of *Drosophila melanogaster*. *I. Neuroblast lineages* derived from the ventral half of the neuroectoderm. *Dev. Biol.* 179, 41–64. doi:10.1006/dbio.1996.0240

Bowman, S. K., Rolland, V., Betschinger, J., Kinsey, K. a., Emery, G., and Knoblich, J. a. (2008). The tumor suppressors Brat and Numb regulate transit-amplifying neuroblast lineages in Drosophila. *Dev. Cell* 14, 535–546. doi:10.1016/j.devcel.2008.03.004

Brankatschk, M., and Eaton, S. (2010). Lipoprotein particles cross the blood-brain barrier in Drosophila. *J. Neurosci.* 30, 10441–10447. doi:10.1523/JNEUROSCI.5943-09.2010

Breunig, J. J., Haydar, T. F., and Rakic, P. (2011). *Neural stem cells: historical perspective and future prospects age of rationalism: origin of neural stem cell research.* doi:10.1016/j.neuron.2011.05.005

Britton, J. S., and Edgar, B. A. (1998). Environmental control of the cell cycle in Drosophila: nutrition activates mitotic and endreplicative cells by distinct mechanisms. *Development* 125, 2149–2158. doi:10.1242/dev.125.11

Broom, E. R., Gilthorpe, J. D., Butts, T., Campo-Paysaa, F., and Wingate, R. J. T. (2012). The roof plate boundary is a bi-directional organiser of dorsal neural tube and choroid plexus development. *Development* 139, 4261–4270. doi:10.1242/DEV.082255

Byrd, C. A., and Brunjes, P. C. (2001). Neurogenesis in the olfactory bulb of adult zebrafish. *Neuroscience* 105, 793–801. doi:10.1016/S0306-4522(01)00215-9

Campos-Ortega, J. A. (1993). Mechanisms of early neurogenesis in *Drosophila melanogaster*. *J. Neurobiol.* 24, 1305–1327. doi:10.1002/NEU.480241005

Casano, A. M., Albert, M., and Peri, F. (2016). Developmental apoptosis mediates entry and positioning of microglia in the zebrafish brain. *Cell Rep.* 16, 897–906. doi:10.1016/J CELREP.2016.06.033

Chell, J. M., and Brand, A. H. (2010). Nutrition-responsive glia control exit of neural stem cells from quiescence. *Cell* 143, 1161–1173. doi:10.1016/j.cell.2010.12.007

Cheng, H., and Leblond, C. P. (1974). Origin, differentiation and renewal of the four main epithelial cell types in the mouse small intestine. V. Unitarian Theory of the origin of the four epithelial cell types. *Am. J. Anat.* 141, 537–561. doi:10.1002/AJA.1001410407

Cheng, L. Y., Bailey, A. P., Leevers, S. J., Ragan, T. J., Driscoll, P. C., and Gould, A. P. (2011). Anaplastic lymphoma kinase spares organ growth during nutrient restriction in Drosophila. *Cell* 146, 435–447. doi:10.1016/j.cell.2011.06.040

Clark, S. E., Williams, R. W., and Meyerowitz, E. M. (1997). The CLAVATA1 gene encodes a putative receptor kinase that controls shoot and floral meristem size in Arabidopsis. *Cell* 89, 575–585. doi:10.1016/S0092-8674(00)80239-1

Clavreul, S., Dumas, L., and Loulier, K. (2022). Astrocyte development in the cerebral cortex: complexity of their origin, genesis, and maturation. *Front. Neurosci.* 16, 916055. doi:10.3389/FNINS.2022.916055

Codega, P., Silva-Vargas, V., Paul, A., Maldonado-Soto, A. R., Deleo, A. M., Pastrana, E., et al. (2014). Prospective identification and purification of quiescent adult neural stem cells from their *in vivo* niche. *Neuron* 82, 545–559. doi:10.1016/j.neuron.2014.02.039

Colucci-D'Amato, L., Bonavita, V., and Porzio, U. (2006). The end of the central dogma of neurobiology: stem cells and neurogenesis in adult CNS. *Neurol. Sci.* 27, 266–270. doi:10.1007/S10072-006-0682-Z

Conover, J. C., and Todd, K. L. (2017). Neuronal stem cell niches of the brain. *Biol. Eng. Stem Cell Niches*, 75-91. doi:10.1016/B978-0-12-802734-9.00006-8

Contreras, E. G., Sierralta, J., and Oliva, C. (2019). Novel strategies for the generation of neuronal diversity: lessons from the fly visual system. *Front. Mol. Neurosci.* 12, 140. doi:10.3389/FNMOL.2019.00140

Corty, M. M., and Coutinho-Budd, J. (2023). Drosophila glia take shape to sculpt the nervous system. *Curr. Opin. Neurobiol.* 79, 102689. doi:10.1016/j.CONB.2023.102689

Costell, M., Gustafsson, E., Aszódi, A., Mörgelin, M., Bloch, W., Hunziker, E., et al. (1999). Perlecan maintains the integrity of cartilage and some basement membranes. *J. Cell Biol.* 147, 1109–1122. doi:10.1083/JCB.147.5.1109

Cotsarelis, G., Sun, T. T., and Lavker, R. M. (1990). Label-retaining cells reside in the bulge area of pilosebaceous unit: implications for follicular stem cells, hair cycle, and skin carcinogenesis. *Cell* 61, 1329–1337. doi:10.1016/0092-8674(90)90696-8

Coutinho-Budd, J. C., Sheehan, A. E., and Freeman, M. R. (2017). The secreted neurotrophin spätze 3 promotes glial morphogenesis and supports neuronal survival and function. *Genes Dev.* 31, 2023–2038. doi:10.1101/gad.305888.117

Cui, C., Wang, F., Zheng, Y., Wei, H., and Peng, J. (2023). From birth to death: the hardworking life of Paneth cell in the small intestine. *Front. Immunol.* 14, 1122258. doi:10.3389/FIMMU.2023.1122258

Daneman, R., Agalliu, D., Zhou, L., Kuhnert, F., Kuo, C. J., and Barres, B. A. (2009). Wnt/beta-catenin signaling is required for CNS, but not non-CNS, angiogenesis *Proc. Natl. Acad. Sci. U. S. A.* 106 (2), 641–646. doi:10.1073/pnas.0805165106

Daneman, R., and Prat, A. (2015). *The blood - brain barrier*, 1-24. doi:10.1002/ana.23648

Daneman, R., Zhou, L., Kebede, A. A., and Barres, B. A. (2010). Pericytes are required for blood-brain barrier integrity during embryogenesis. *Nat* 468, 562–566. doi:10.1038/nature09513

Datta, S. (1995). Control of proliferation activation in quiescent neuroblasts of the Drosophila central nervous system. *Development* 121, 1173–1182. doi:10.1242/dev.121.4.1173

Del Bigio, M. R. (2010). Ependymal cells: biology and pathology. *Acta Neuropathol.* 119, 55–73. doi:10.1007/s00401-009-0624-y

de Torres-Jurado, A., Manzanero-Ortiz, S., and Carmena, A. (2022). Glial-secreted Netrins regulate Robo1/Rac1-Cdc42 signaling threshold levels during Drosophila asymmetric neural stem/progenitor cell division. *Curr. Biol.* 32, 2174–2188.e3. doi:10.1016/j.CUB.2022.04.001

Dexter, T. M., Allen, T. D., and Lajtha, L. G. (1977). Conditions controlling the proliferation of haemopoietic stem cells *in vitro*. *J. Cell. Physiol.* 91, 335–344. doi:10.1002/JCP.1040910303

D'Gama, P. P., Qiu, T., Cosacak, M. I., Rayamajhi, D., Konac, A., Hansen, J. N., et al. (2021). Diversity and function of motile ciliated cell types within ependymal lineages of the zebrafish brain. *Cell Rep.* 37, 109775. doi:10.1016/j CELREP.2021.109775

Dias Moura Prazeres, P. H., Sena, I. F. G., Borges, I. da T., de Azevedo, P. O., Andreotti, J. P. P., de Paiva, A. E., et al. (2017). PERICYTES ARE HETEROGENEOUS IN THEIR ORIGIN WITHIN THE SAME TISSUE. *Dev. Biol.* 427, 6–11. doi:10.1016/J.YDBIO.2017.05.001

Di Marco, B., Crouch, E. E., Shah, B., Duman, C., Paredes, M. F., Ruiz de Almodovar, C., et al. (2020). Reciprocal interaction between vascular filopodia and neural stem cells shapes neurogenesis in the ventral telencephalon. *Cell Rep.* 33, 108256. doi:10.1016/J.CELREP.2020.108256

Ding, W. Y., Huang, J., and Wang, H. (2020). Waking up quiescent neural stem cells: molecular mechanisms and implications in neurodevelopmental disorders. *PLoS Genet.* 16, e1008653. doi:10.1371/journal.pgen.1008653

Diotel, N., Charlier, T. D., Lefebvre d'Hellencourt, C., Couret, D., Trudeau, V. L., Nicolau, J. C., et al. (2018). Steroid transport, local synthesis, and signaling within the brain: roles in neurogenesis, neuroprotection, and sexual behaviors. *Front. Neurosci.* 12, 84. doi:10.3389/FNINS.2018.00084
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Bill, B. R., Balcuinas, D., McCarra, J. A., Young, E. D., Xiong, T., Spahn, A. M., et al. (2008). Development and Notch signaling requirements of the zebrafish choroid plexus. *PLoS One* 3, e3114. doi:10.1371/JOURNAL.PONE.0003114

Binder, M. D., Hirokawa, N., and Windhorst, U. (2009). Encyclopedia of neuroscience. Available at: [https://books.google.com/books/about/Eencyclopedia_of_Neuroscience.html?id=mcvhAAAACAAJ](Accessed November 4, 2023).4393.

Biswas, S., Cottarelli, A., and Agalliu, D. (2020). Neuronal and glial regulation of CNS angiogenesis and barriergenesis. *Dev* 147, dev182279. doi:10.1242/dev.182279

Bjornson, C. R., Rietze, R. L., Reynolds, B. A., Magli, M. C., and Vescovi, A. L. (1999). Turning brain into blood: a hematopoietic fate adopted by adult neural stem cells *in vivo*. *Science* 283 (5401). New York, NY: JPEG, 534–537. doi:10.1126/science.283.5401.534

Bjornsson, C. S., Apostolopoulou, M., Tian, Y., and Temple, S. (2015). It takes a village: constructing the neurogenic niche. *Dev. Cell* 32, 435–446. doi:10.1016/j.devcel.2015.01.010

Blaschke, A. J., Weiner, J. A., and Chun, J. (1998). Programmed cell death is a universal feature of embryonic and postnatal neuroproliferative regions throughout the central nervous system. *J. Comp. Neurol.* 396, 39–50. doi:10.1002/(sici)1096-9861(19980622)396:1<39;aid-cne4>3.0.co;2-j(19980622)

Bond, A. M., Ming, G. L., and Song, H. (2015). Adult mammalian neural stem cells and neurogenesis: five decades later. *Cell Stem Cell* 17, 385–395. doi:10.1016/j.stem.2015.09.003

Bonvento, G., and Bolaños, J. P. (2021). Astrocyte-neuron metabolic cooperation shapes brain activity. *Cell Metab.* 33, 1546–1564. doi:10.1016/j.CMET.2021.07.006

Boone, J. Q., and Doe, C. Q. (2008). Identification of Drosophila type II neuroblast lineages containing transit amplifying ganglion mother cells. *Dev. Neurobiol.* 68, 1185–1195. doi:10.1002/dneu.20648

Bossing, T., Udolph, G., Doe, C. Q., and Technau, G. M. (1996). The embryonic central nervous system lineages of *Drosophila melanogaster*. *I. Neuroblast lineages* derived from the ventral half of the neuroectoderm. *Dev. Biol.* 179, 41–64. doi:10.1006/dbio.1996.0240

Bowman, S. K., Rolland, V., Betschinger, J., Kinsey, K. a., Emery, G., and Knoblich, J. a. (2008). The tumor suppressors Brat and Numb regulate transit-amplifying neuroblast lineages in Drosophila. *Dev. Cell* 14, 535–546. doi:10.1016/j.devcel.2008.03.004

Brankatschk, M., and Eaton, S. (2010). Lipoprotein particles cross the blood-brain barrier in Drosophila. *J. Neurosci.* 30, 10441–10447. doi:10.1523/JNEUROSCI.5943-09.2010

Breunig, J. J., Haydar, T. F., and Rakic, P. (2011). *Neural stem cells: historical perspective and future prospects age of rationalism: origin of neural stem cell research.* doi:10.1016/j.neuron.2011.05.005

Britton, J. S., and Edgar, B. A. (1998). Environmental control of the cell cycle in Drosophila: nutrition activates mitotic and endreplicative cells by distinct mechanisms. *Development* 125, 2149–2158. doi:10.1242/dev.125.11

Broom, E. R., Gilthorpe, J. D., Butts, T., Campo-Paysaa, F., and Wingate, R. J. T. (2012). The roof plate boundary is a bi-directional organiser of dorsal neural tube and choroid plexus development. *Development* 139, 4261–4270. doi:10.1242/DEV.082255

Byrd, C. A., and Brunjes, P. C. (2001). Neurogenesis in the olfactory bulb of adult zebrafish. *Neuroscience* 105, 793–801. doi:10.1016/S0306-4522(01)00215-9

Campos-Ortega, J. A. (1993). Mechanisms of early neurogenesis in *Drosophila melanogaster*. *J. Neurobiol.* 24, 1305–1327. doi:10.1002/NEU.480241005

Casano, A. M., Albert, M., and Peri, F. (2016). Developmental apoptosis mediates entry and positioning of microglia in the zebrafish brain. *Cell Rep.* 16, 897–906. doi:10.1016/J CELREP.2016.06.033

Chell, J. M., and Brand, A. H. (2010). Nutrition-responsive glia control exit of neural stem cells from quiescence. *Cell* 143, 1161–1173. doi:10.1016/j.cell.2010.12.007

Cheng, H., and Leblond, C. P. (1974). Origin, differentiation and renewal of the four main epithelial cell types in the mouse small intestine. V. Unitarian Theory of the origin of the four epithelial cell types. *Am. J. Anat.* 141, 537–561. doi:10.1002/AJA.1001410407

Cheng, L. Y., Bailey, A. P., Leevers, S. J., Ragan, T. J., Driscoll, P. C., and Gould, A. P. (2011). Anaplastic lymphoma kinase spares organ growth during nutrient restriction in Drosophila. *Cell* 146, 435–447. doi:10.1016/j.cell.2011.06.040

Clark, S. E., Williams, R. W., and Meyerowitz, E. M. (1997). The CLAVATA1 gene encodes a putative receptor kinase that controls shoot and floral meristem size in Arabidopsis. *Cell* 89, 575–585. doi:10.1016/S0092-8674(00)80239-1

Clavreul, S., Dumas, L., and Loulier, K. (2022). Astrocyte development in the cerebral cortex: complexity of their origin, genesis, and maturation. *Front. Neurosci.* 16, 916055. doi:10.3389/FNINS.2022.916055

Codega, P., Silva-Vargas, V., Paul, A., Maldonado-Soto, A. R., Deleo, A. M., Pastrana, E., et al. (2014). Prospective identification and purification of quiescent adult neural stem cells from their *in vivo* niche. *Neuron* 82, 545–559. doi:10.1016/j.neuron.2014.02.039

Colucci-D'Amato, L., Bonavita, V., and Porzio, U. (2006). The end of the central dogma of neurobiology: stem cells and neurogenesis in adult CNS. *Neurol. Sci.* 27, 266–270. doi:10.1007/S10072-006-0682-Z

Conover, J. C., and Todd, K. L. (2017). Neuronal stem cell niches of the brain. *Biol. Eng. Stem Cell Niches*, 75-91. doi:10.1016/B978-0-12-802734-9.00006-8

Contreras, E. G., Sierralta, J., and Oliva, C. (2019). Novel strategies for the generation of neuronal diversity: lessons from the fly visual system. *Front. Mol. Neurosci.* 12, 140. doi:10.3389/FNMOL.2019.00140

Corty, M. M., and Coutinho-Budd, J. (2023). Drosophila glia take shape to sculpt the nervous system. *Curr. Opin. Neurobiol.* 79, 102689. doi:10.1016/j.CONB.2023.102689

Costell, M., Gustafsson, E., Aszódi, A., Mörgelin, M., Bloch, W., Hunziker, E., et al. (1999). Perlecan maintains the integrity of cartilage and some basement membranes. *J. Cell Biol.* 147, 1109–1122. doi:10.1083/JCB.147.5.1109

Cotsarelis, G., Sun, T. T., and Lavker, R. M. (1990). Label-retaining cells reside in the bulge area of pilosebaceous unit: implications for follicular stem cells, hair cycle, and skin carcinogenesis. *Cell* 61, 1329–1337. doi:10.1016/0092-8674(90)90696-9

Coutinho-Budd, J. C., Sheehan, A. E., and Freeman, M. R. (2017). The secreted neurotrophin spätze 3 promotes glial morphogenesis and supports neuronal survival and function. *Genes Dev.* 31, 2023–2038. doi:10.1101/gad.305888.117

Cui, C., Wang, F., Zheng, Y., Wei, H., and Peng, J. (2023). From birth to death: the hardworking life of Paneth cell in the small intestine. *Front. Immunol.* 14, 1122258. doi:10.3389/FIMMU.2023.1122258

Daneman, R., Agalliu, D., Zhou, L., Kuhnert, F., Kuo, C. J., and Barres, B. A. (2009). Wnt/beta-catenin signaling is required for CNS, but not non-CNS, angiogenesis *Proc. Natl. Acad. Sci. U. S. A.* 106 (2), 641–646. doi:10.1073/pnas.0805165106

Daneman, R., and Prat, A. (2015). *The blood - brain barrier*, 1-24. doi:10.1002/ana.23648

Daneman, R., Zhou, L., Kebede, A. A., and Barres, B. A. (2010). Pericytes are required for blood-brain barrier integrity during embryogenesis. *Nat* 468, 562–566. doi:10.1038/nature09513

Datta, S. (1995). Control of proliferation activation in quiescent neuroblasts of the Drosophila central nervous system. *Development* 121, 1173–1182. doi:10.1242/dev.121.4.1173

Del Bigio, M. R. (2010). Ependymal cells: biology and pathology. *Acta Neuropathol.* 119, 55–73. doi:10.1007/s00401-009-0624-y

de Torres-Jurado, A., Manzanero-Ortiz, S., and Carmena, A. (2022). Glial-secreted Netrins regulate Robo1/Rac1-Cdc42 signaling threshold levels during Drosophila asymmetric neural stem/progenitor cell division. *Curr. Biol.* 32, 2174–2188.e3. doi:10.1016/j.CUB.2022.04.001

Dexter, T. M., Allen, T. D., and Lajtha, L. G. (1977). Conditions controlling the proliferation of haemopoietic stem cells *in vitro*. *J. Cell. Physiol.* 91, 335–344. doi:10.1002/JCP.1040910303

D'Gama, P. P., Qiu, T., Cosacak, M. I., Rayamajhi, D., Konac, A., Hansen, J. N., et al. (2021). Diversity and function of motile ciliated cell types within ependymal lineages of the zebrafish brain. *Cell Rep.* 37, 109775. doi:10.1016/J CELREP.2021.109775

Dias Moura Prazeres, P. H., Sena, I. F. G., Borges, I. da T., de Azevedo, P. O., Andreotti, J. P. P., de Paiva, A. E., et al. (2017). PERICYTES ARE HETEROGENEOUS IN THEIR ORIGIN WITHIN THE SAME TISSUE. *Dev. Biol.* 427, 6–11. doi:10.1016/J. YDBIO.2017.05.001

Di Marco, B., Crouch, E. E., Shah, B., Duman, C., Paredes, M. F., Ruiz de Almodovar, C., et al. (2020). Reciprocal interaction between vascular filopodia and neural stem cells shapes neurogenesis in the ventral telencephalon. *Cell Rep.* 33, 108256. doi:10.1016/J. CELREP.2020.108256

Ding, W. Y., Huang, J., and Wang, H. (2020). Waking up quiescent neural stem cells: molecular mechanisms and implications in neurodevelopmental disorders. *PLoS Genet.* 16, e1008653. doi:10.1371/journal.pgen.1008653

Diotel, N., Charlier, T. D., Lefebvre d'Hellencourt, C., Couret, D., Trudeau, V. L., Nicolau, J. C., et al. (2018). Steroid transport, local synthesis, and signaling within the brain: roles in neurogenesis, neuroprotection, and sexual behaviors. *Front. Neurosci.* 12, 84. doi:10.3389/FNINS.2018.00084
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Dong, Q., Zavortink, M., Froldi, F., Golenkina, S., Lam, T., and Cheng, L. Y. (2021). Glial Hedgehog signalling and lipid metabolism regulate neural stem cell proliferation in Drosophila. *EMBO Rep.* 22, e52130. doi:10.15252/embr. 202052130

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Engelhardt, B., and Liebner, S. (2014). Novel insights into the development and maintenance of the blood-brain barrier. *Cell Tissue Res.* 355 (3), 687-699. doi:10.1007/ s00441-014-1811-2

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Fame, R. M., Chang, J. T., Hong, A., Aponte-Santiago, N. A., and Sive, H. (2016). Directional cerebrospinal fluid movement between brain ventricles in larval zebrafish. *Fluids Barriers CNS* 13, 11. doi:10.1186/S12987-016-0036-Z

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Fernández-Hernández, I., Rhiner, C., and Moreno, E. (2013). Adult neurogenesis in Drosophila. *Cell Rep.* 3, 1857-1865. doi:10.1016/j.celrep.2013.05.034

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Ferrero, G., Mahony, C. B., Dupuis, E., Yvernogoe, L., Di Ruggiero, E., Miseroocchi, M., et al. (2018). Embryonic microglia derive from primitive macrophages and are replaced by cmyb-dependent definitive microglia in zebrafish. *Cell Rep.* 24, 130-141. doi:10.1016/j CELREP.2018.05.066

Ferrero, G., Miseroocchi, M., Di Ruggiero, E., and Wittamer, V. (2021). A c sfrrb mutation uncouples two waves of microglia development in zebrafish. *Development* 148, dev194241. doi:10.1242/DEV.194241

Foerster, P., Daclin, M., Asm, S., Faucourt, M., Boletta, A., Genovesio, A., et al. (2017). mTORC1 signaling and primary cilia are required for brain ventricle morphogenesis. *Development* 144, 201-210. doi:10.1242/DEV.138271

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Furlan, G., Cuccioli, V., Vuillemin, N., Dirian, L., Muntasell, A. J., Coolen, M., et al. (2017). Life-long neurogenic activity of individual neural stem cells and continuous growth establish an outside-in architecture in the teleost pallium. *Curr. Biol.* 27, 3288-3301. doi:10.1016/J.CUB.2017.09.052

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Ghersi-Egea, J. F., Strazielle, N., Catala, M., Silva-Vargas, V., Doetsch, F., and Engelhardt, B. (2018). Molecular anatomy and functions of the choroidal blood-cerebrospinal fluid barrier in health and disease. *Acta Neuropathol.* 135 (135), 337-361. doi:10.1007/S00401-018-1807-1

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Ginhoux, F., Greter, M., Leboeuf, M., Nandi, S., See, P., Gokhan, S., et al. (2010). Fate mapping analysis reveals that adult microglia derive from primitive macrophages. *Science* 330, 841-845. doi:10.1126/SCIENCE.1194637

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Gore, A. V., Monzo, K., Cha, Y. R., Pan, W., and Weinstein, B. M. (2012). Vascular development in the zebrafish. *Cold Spring Harb. Perspect. Med.* 2, a006684. doi:10.1101/ CSHPERSPECT.A006684

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Graus-Porta, D., Blaess, S., Senften, M., Littlewood-Evans, A., Damsky, C., Huang, Z., et al. (2001). Beta1-class integrins regulate the development of laminae and folia in the cerebral and cerebellar cortex. *Neuron* 31, 367-379. doi:10.1016/S0896-6273(01) 00374-9

Grimes, D. T., Boswell, C. W., Morante, N. F. C., Henkelman, R. M., Burdine, R. D., and Ciruna, B. (2016). Zebrafish models of idiopathic scoliosis link cerebrospinal fluid flow defects to spine curvature. *Science* 352, 1341-1344. doi:10.1126/SCIENCE.AAF6419

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Ferrero, G., Miseroocchi, M., Di Ruggiero, E., and Wittamer, V. (2021). A c sfrrb mutation uncouples two waves of microglia development in zebrafish. *Development* 148, dev194241. doi:10.1242/DEV.194241

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Grimes, D. T., Boswell, C. W., Morante, N. F. C., Henkelman, R. M., Burdine, R. D., and Ciruna, B. (2016). Zebrafish models of idiopathic scoliosis link cerebrospinal fluid flow defects to spine curvature. *Science* 352, 1341-1344. doi:10.1126/SCIENCE.AAF6419

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