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Copyright =C2=A9 1999=20 Cell Press. All rights reserved.

Article

Subventricular Zone Astrocytes Are Neural Stem Cells in the Adult = Mammalian=20 Brain

Fiona Doetsch1,=20 Isabelle Caill=C3=A91,=20 Daniel A. Lim1,=20 Jose Manuel Garc=C3=ADa-Verdugo2=20 and Arturo Alvarez-Buylla, , 1,=20 =E2=80=A1=20
1 The Rockefeller University, = New York,=20 New York 10021, USA
2 University of = Valencia,=20 Burjassot-46100, Valencia, Spain
Received 15 December 1998;  = Revised 10=20 May 1999.  Available online 28 June 2004.

Abstract

Neural stem cells reside in the subventricular zone (SVZ) of the = adult=20 mammalian brain. This germinal region, which continually generates new = neurons=20 destined for the olfactory bulb, is composed of four cell types: = migrating=20 neuroblasts, immature precursors, astrocytes, and ependymal cells. Here = we show=20 that SVZ astrocytes, and not ependymal cells, remain labeled with = proliferation=20 markers after long survivals in adult mice. After elimination of = immature=20 precursors and neuroblasts by an antimitotic treatment, SVZ astrocytes = divide to=20 generate immature precursors and neuroblasts. Furthermore, in untreated = mice,=20 SVZ astrocytes specifically infected with a retrovirus give rise to new = neurons=20 in the olfactory bulb. Finally, we show that SVZ astrocytes give rise to = cells=20 that grow into multipotent neurospheres in vitro. We conclude that SVZ=20 astrocytes act as neural stem cells in both the normal and regenerating = brain.=20


Article Outline

=E2=80=A2=20 Introduction
=E2=80=A2=20 Results
=E2=80=A2=20 Type B Cells Divide Early during Regeneration of the SVZ
=E2=80=A2=20 Dividing Type B Cells Give Rise to Type C Cells and Neuroblasts
=E2=80=A2=20 Type B Cells Are the Only Label-Retaining Cells in the Adult SVZ
=E2=80=A2=20 GFAP-Positive Cells in Untreated Mice Give Rise to New Neurons in = the=20 Olfactory Bulb
=E2=80=A2=20 Neurospheres Are Derived from SVZ Cells
=E2=80=A2=20 Neurospheres Are Derived from Type B Cells
=E2=80=A2=20 Discussion
=E2=80=A2=20 Experimental Procedures
=E2=80=A2=20 Ara-C Infusions
=E2=80=A2=20 Immunohistochemistry
=E2=80=A2=20 BrdU Administration and Detection
=E2=80=A2=20 [3H]Thymidine Autoradiography and EM Analysis
=E2=80=A2=20 Retroviral Injections for In Vivo Studies
=E2=80=A2=20 Whole-Mount Cultures
=E2=80=A2=20 Neurosphere Preparation and Differentiation
=E2=80=A2=20 Adenoviral Vectors
=E2=80=A2=20 Production of pAd5-GFAPp-GFP Adenoviral Vector
=E2=80=A2=20 Adenoviral Injections into the SVZ
=E2=80=A2=20 Labeling of the Ependymal Layer with Vital Dyes
=E2=80=A2=20 Fluorescent Tracers
=E2=80=A2=20 Adenoviral Tracers
=E2=80=A2=20 Scoring of Vitally Labeled Cells and Neurospheres
=E2=80=A2=20 Acknowledgements
=E2=80=A2=20 References

Display=20 Full Size version of this image (3K)

Figure 1. Cells Remaining in the SVZ after Ara-C Treatment Are GFAP=20 PositiveGFAP-postembedding immunostaining of the SVZ in 1.5 =CE=BCm = semithin (A) and=20 3D"not,70 nm=20 ultrathin sections (B and C).(A) SVZ cells immediately after Ara-C are = GFAP=20 positive. Asterisks show cell bodies of labeled cells.(B) = Silver-enhanced=20 immunogold labeling for GFAP immediately after Ara-C treatment. At the = EM, these=20 cells are characterized by dense bundles of intermediate filaments that = are=20 immunostained by GFAP antibodies.(C) Detail of immunostained = intermediate=20 filaments shown in (B) (rectangle). Scale bars: (A), 20 =CE=BCm; (B), 1 = =CE=BCm; (C), 0.25=20 =CE=BCm.Dividing cells reappear scattered throughout the SVZ 0.5 days = after=20 termination of Ara-C treatment (F. D. et al., unpublished data). To = identify=20 which cells divided after Ara-C treatment, mice were injected = systemically with=20 [3H]thymidine, a marker of DNA synthesis, 0.5 days after = Ara-C=20 treatment and killed 1 hr later. [3H]thymidine-labeled cells = in both=20 the injected and contralateral hemispheres were identified at the EM (Figure=20 2A and Table=20 1). Of 154 cells studied, 152 corresponded to type B cells and 2 to=20 microglia. None corresponded to ependymal cells. Some=20 [3H]thymidine-labeled cells were very close to the ependymal = layer=20 but were always separated from the ventricle by the cytoplasm of an = ependymal=20 cell.
Display=20 Full Size version of this image (3K)

Figure 2. Proliferating Type B Cells Give Rise to Type C Cells and=20 NeuroblastsTime lines illustrate the duration of Ara-C treatment and = time of=20 pulse labeling (double arrow) with [3H]thymidine (top panels) = or=20 retrovirus (bottom panels). Thin arrows indicate the time points at = which the=20 SVZ was analyzed with respect to the end of Ara-C treatment.(A) In=20 [3H]thymidine-injected mice killed 1 hr after pulse labeling, = type B=20 cells (b) were observed with the EM; upper inset shows the same cell = with silver=20 grains overlying its nucleus in a semithin section (see Table=20 1). Lower right inset shows a bundle of intermediate filaments,=20 characteristic of type B cells, in the [3H]thymidine-labeled = cell.(B)=20 Two days after pulse labeling proliferating astrocytes, type C cells (c) = were=20 observed in addition to type B cells. Inset shows silver grains = overlying the=20 nuclei of the two cells identified at the EM as type C cells.(C) = Alkaline=20 phosphatase=E2=80=93positive cells with a stellate morphology were = observed 1 day after=20 infection with DAP retrovirus.(D) At 4 days, small clusters of tightly=20 associated cells are present.(E) At 6 days, many cells with the = morphology of=20 migrating neuroblasts (type A cells) were observed.(F) Semithin section = of=20 =CE=B2-galactosidase retrovirally labeled cell (blue precipitate) 1.5 = days after=20 Ara-C.(G) This same cell analyzed at the EM corresponds to a type B = cell. Scale=20 bars: (A) and (B), upper inset, 10 =CE=BCm; photomicrograph, 2.5 = =CE=BCm; lower inset, 0.5=20 =CE=BCm; (C), 30 =CE=BCm; (D), 40 =CE=BCm; (E), 20 =CE=BCm; (F), 10 = =CE=BCm; (G), 2 =CE=BCm.
Display=20 Full Size version of this image (4K)

Figure 3. GFAP-Positive Cells Give Rise to Neuroblasts In Vitro(A)=20 Whole-mount dissection of the lateral wall of the lateral ventricle 1 = day after=20 infection with a retrovirus encoding GFP. The retrovirus was injected = 0.5 days=20 after the end of Ara-C treatment. Only cells with a branched morphology = were=20 present at this survival. These whole mounts were sectioned transversely = and=20 stained with an anti-GFAP mAb.(B) One micrometer confocal optical slice = showing=20 the GFP-labeled cells.(C) GFAP staining of this same section.(D) = GFP-positive=20 cells are GFAP positive. The ependymal layer in (B=E2=80=93D) came off = during=20 immunocytochemical processing. Ependymal cells were not labeled by the = reporter=20 gene.(E=E2=80=93K) Direct observation of the production of neuroblasts = by type B cells.=20 Proliferating type B cells were infected as in (A). One day after = infection,=20 whole mounts were dissected, embedded in a collagen gel, and cultured to = follow=20 the fate of GFP-labeled cells daily. (E) Two GFP cells with a branched=20 morphology after 1 day in culture. (F) The lower of the two branched = cells=20 divided and generated a tight cluster of globular cells (arrowhead). The = upper=20 cell also divided into branched (arrow) and unbranched cells. (G) By 3 = days, the=20 lower cluster began to disperse, and some cells with a leading process = and a=20 growth cone typical of type A cells (arrow) were present. By this time, = the=20 upper cell had also generated a tight cluster of globular cells = (arrowhead). (H)=20 One day later, several type A cells were present (arrows), and some had = moved=20 away from the clusters. (I=E2=80=93K) At this time, the cultures were = fixed and stained=20 for TuJ1, revealing that some GFP-positive cells (arrow) were TuJ1 = positive.=20 (E=E2=80=93H) are photomicrographs of live three-dimensional cultures. = (B=E2=80=93D) and (I=E2=80=93K)=20 are confocal 1 =CE=BCm optical sections through fixed cultures. Scale = bars: (A=E2=80=93D), 10=20 =CE=BCm; (E=E2=80=93K), 5 =CE=BCm.To follow the fate of individual live = GFP-labeled cells over=20 time, dividing cells were infected 0.5 days after Ara-C treatment in = vivo. One=20 day after infection, fragments of the SVZ were dissected and cultured in = a=20 collagen gel. Individual GFP-positive cells were marked and photographed = daily=20 to follow their progression (Figure=20 3E=E2=80=933K). Some of the branched cells divided to give rise to = tight clusters of=20 globular cells ( Figure=20 3F). The tight clusters dispersed after 4 days ( Figure=20 3G), and cells with a small round or elongated cell body and a = leading=20 process appeared ( Figure=20 3H). Some of these cells were TuJ1 positive, confirming their = neuronal=20 identity ( Figure=20 3I=E2=80=933K). These results directly show that dividing type B = cells give rise to=20 neurons after Ara-C treatment via an intermediate globular cell type.
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Figure 4. Type B Cells in the SVZ Are Labeled at Long Survivals after = BrdU=20 Administration or [3H]Thymidine InjectionsBrdU was = administered=20 continuously in the drinking water for 2 weeks, and sections were double = stained=20 for BrdU (green) and mCD24 (red).(A) One week after treatment, no = BrdU-positive=20 mCD24 ependymal cells were observed. While BrdU-labeled nuclei were = closely=20 associated with ependymal cells, these nuclei were not surrounded by = mCD24=20 staining. Ependymal nuclei (arrows) were surrounded on all sides by = mCD24.(B)=20 Upper inset shows silver grains overlying the nucleus (arrow) of a=20 3H-labeled cell in the SVZ after [3H]thymidine was = infused=20 into the lateral ventricles for 12 days. Labeled nuclei that at the = light=20 microscope appeared to be in the ependymal layer were studied at the EM. = These=20 cells did not correspond to ependymal cells but had the ultrastructural=20 characteristics of type B cells. Thin ependymal cell processes separate = most of=20 these cells from the ventricle.(C) Upper inset shows silver grains = overlying the=20 nucleus (arrow) of a 3H-labeled cell in the SVZ 30 days after = four=20 [3H]thymidine injections. Ultrastructural analysis of this = labeled=20 cell (asterisk) reveals that it is a type B cell (b) adjacent to a chain = of type=20 A cells (a). Scale bars: (A), 10 =CE=BCm; (B=E2=80=93C), inset, 10 = =CE=BCm; photomicrograph, 2 =CE=BCm.=20 LV, lateral ventricle.
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Figure 5. GFAP-Expressing Cells Generate Olfactory Bulb Neurons under = Normal=20 ConditionsAlkaline phosphatase=E2=80=93positive cells observed after the = targeted=20 infection of GFAP-positive cells using GFAP-Tva mice.(A) Section through = an=20 AP-positive branched cell in the SVZ 1 day after infection. At this = time, no=20 globular or migrating neuroblasts were encountered.(B) Three and a half = days=20 after infection, AP-positive migrating cells were observed in the RMS en = route=20 to the olfactory bulb (arrows).(C and D) Fourteen days after infection, = many=20 AP-labeled cells were found in the olfactory bulb. These cells had the = typical=20 morphology of granule (C) and periglomerular neurons (D).(E) In the SVZ = of these=20 mice, clusters containing branched globular and migratory cells were = found.=20 Scale bars: (A=E2=80=93E), 20 =CE=BCm.
Display=20 Full Size version of this image (4K)

Figure 6. Ependymal Cells Do Not Give Rise to Neurospheres(A) Frontal = section=20 showing the labeling of the ependymal layer in the contralateral = hemisphere of a=20 mouse that received an intraventricular injection of DiI.(B=E2=80=93E) = Phase contrast (B=20 and D) and epifluorescent (C and E) images of two neurospheres cultured = from the=20 lateral wall of the lateral ventricle of DiI-injected mice. Both the=20 contralateral (B and C) and the ipsilateral hemispheres (D and E) gave = rise to=20 neurospheres containing DiI-positive cells. However, we noticed that = these=20 cultures contained unincorporated DiI (arrow) and patchy DiI labeling of = neurospheres.(F) Section showing the labeling of the ependymal layer in = the=20 contralateral hemisphere of a mouse that received an intraventricular = injection=20 of rhodamine beads.(G) Confocal image showing the region marked by a = rectangle=20 in (F) at higher magnification. Rhodamine-positive cells are also = immunopositive=20 for mCD24 (green), which labels ependymal cells.(H=E2=80=93K) Phase = contrast (H and J)=20 and epifluorescent (I and K) images of two neurospheres cultured from = the=20 lateral wall of the lateral ventricle of rhodamine bead=E2=80=93injected = mice.=20 Neurospheres arising from the contralateral hemisphere did not show any=20 rhodamine labeling (H and I), whereas some neurospheres from the = ipsilateral=20 hemisphere were fluorescent (J and K).(L) Section showing the labeling = of the=20 ependymal layer in the contralateral hemisphere of a mouse that received = an=20 intraventricular injection of an adenovirus carrying LacZ.(M and N) = Nomarski=20 images of five DIV neurospheres from the lateral wall of the lateral = ventricle=20 of =CE=B2-gal adenovirus-infected mice. Neurospheres grown from the = contralateral=20 hemisphere (M) are X-gal negative, whereas some neurospheres from the=20 ipsilateral hemisphere are X-gal positive (N). These small neurospheres = were=20 allowed to attach to poly-L-ornithine for 5 hr for staining. Scale bars: = (A),=20 100 =CE=BCm; (B=E2=80=93E), 25 =CE=BCm; (F), 100 =CE=BCm; (G), 10 = =CE=BCm; (H=E2=80=93K), 50 =CE=BCm; (L), 100 =CE=BCm; (M and=20 N), 25 =CE=BCm. LV, lateral ventricle; CL, contralateral; IL, = ipsilateral.
Display=20 Full Size version of this image (3K)

Figure 7. GFAPp-GFP Adenovirus Selectively Labels AstrocytesAll = panels are 1=20 =CE=BCm confocal optical sections of frontal sections through the SVZ = (A=E2=80=93C and G=E2=80=93L)=20 and striatum (D=E2=80=93F) of mice infected with the GFAPp-GFP = adenovirus 48 hr earlier.=20 The rhodamine channel is shown in the left column and the FITC channel = (GFP) in=20 the middle column, and the two are merged in the right = column.(A=E2=80=93F) GFP-positive=20 cells in the SVZ (B) and striatum (E) are GFAP positive (A and D and C = and F).=20 Arrows in (A) indicate the GFAP-negative regions that correspond to = chains of=20 migrating neuroblasts in the SVZ.(G=E2=80=93I) CD24-positive cells in = the ependymal wall=20 (G) were not GFP positive (H and I). The GFP-positive cells correspond = to SVZ=20 cells.(J=E2=80=93L) PSA-NCAM-positive cells ([J], the migrating = neuroblasts, type A=20 cells) were not GFP positive (K and L). Instead, astrocytes around=20 PSA-NCAM-positive migrating neuroblasts were positive. Scale bars: 10 = =CE=BCm. LV,=20 lateral ventricle. The schematic at the bottom shows the GFAP promoter = reporter=20 construct used for the adenoviral vector.
Display=20 Full Size version of this image (5K)

Figure 8. GFAP-GFP-Positive Cells Give Rise to = Neurospheres(A=E2=80=93F) SVZ cells=20 were infected in vivo with the GFAPp-GFP adenovirus; 48 hr later, cells = were=20 dissociated and cultured in multiwell plates. Epifluorescent (A, C, and = E) and=20 phase contrast (B, D, and F) images of a GFP-positive cell monitored as = it=20 generated a neurosphere. The GFP fluorescence is progressively reduced = as the=20 neurosphere grows due to dilution of the adenovirus or to promoter = inactivation=20 (E). (A and B): 3DIV; (C and D): 5DIV, (E and F): 8DIV.(G) Double = staining with=20 Tuj1 (red) and GFAP (green) of a secondary neurosphere derived from the=20 neurosphere shown in (F) differentiated into neurons and glia upon = removal of=20 growth factor and plating onto an adhesive substrate. Scale bars: = (A=E2=80=93F), 25 =CE=BCm;=20 (G), 50 =CE=BCm. The table shows the percentage of GFP-positive cells = after=20 dissociation and the proportion of these cells that gave rise to = neurospheres in=20 four independent experiments.
Display=20 Full Size version of this image (4K)

Figure 9. Model of Neurogenesis in the SVZ of the Adult Mouse = BrainSVZ=20 astrocytes (type B cells, blue) are the primary neuronal precursors in = the SVZ.=20 These cells divide to give rise to clusters of type C cells (green), = which in=20 turn generate migrating neuroblasts (type A cells, red). Ependymal cells = (gray)=20 line the wall of the ventricles and in our experiments did not divide.=20


Table 2. Characterization of Cellular Composition of Retrovirally = Infected=20 Clusters after Ara-C Treatment
3D"Full

Table 1. Cell Types Labeled with [3H]thymidine after Pulse = Labeling at 12 Hr after Ara-C Treatment

[3H]thymidine-labeled cell types at different survivals = following=20 pulse labeling at 12 hr after Ara-C treatment. [3H]thymidine = was=20 injected at 0.5 days after Ara-C treatment and mice killed 1 hr or 1.5 = or 5.5=20 days later. Percentage of each cell type labeled with = [3H]thymidine=20 of total labeled cells analyzed at the EM (absolute numbers of cells in=20 brackets).


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Corresponding author. Correspondence: Arturo = Alvarez-Buylla,=20 212 327 7729 (phone), 212 327 8312 (fax)




Volume=20 97, Issue 6, 11 June 1999, Pages 703-716 		Result=20 list |  previous=20  < 9 of 17 >  next =20
 
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