From: Subject: Olig2 Directs Astrocyte and Oligodendrocyte Formation in Postnatal Subventricular Zone Cells -- Marshall et al. 25 (32): 7289 -- Journal of Neuroscience Date: Thu, 16 Nov 2006 18:44:00 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0066_01C709AF.2E622820" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.2962 This is a multi-part message in MIME format. ------=_NextPart_000_0066_01C709AF.2E622820 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.jneurosci.org/cgi/content/full/25/32/7289?ijkey=88958bd79eba18cf7ca01fbbf30cd5269877dcff Olig2 Directs Astrocyte and Oligodendrocyte Formation = in Postnatal Subventricular Zone Cells -- Marshall et al. 25 (32): 7289 = -- Journal of Neuroscience
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The Journal of = Neuroscience, August=20 10, 2005, 25(32):7289-7298; doi:10.1523/JNEUROSCI.1924-05.2005

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Development/Plasticity/Repair
Olig2 Directs Astrocyte and Oligodendrocyte Formation in = Postnatal=20 Subventricular Zone Cells

Christine A. G. Marshall,1=20 Bennett G. Novitch,1,2,4 = and=20 James E. Goldman1,3

1Center for Neurobiology and Behavior, = 2Department of=20 Biochemistry and Molecular Biophysics, and 3Department of = Pathology,=20 Columbia University, College of Physicians and Surgeons, New York, New = York=20 10032, and 4Department of Cell and Developmental Biology, = University=20 of Michigan Medical School, Ann Arbor, Michigan 48109=20


3D"   =20 Abstract
Top
3D"Abstract
Introduction
Materials and Methods
Results
Discussion
References
 
T= he=20 subventricular zone (SVZ) in the neonatal mammalian forebrain=20 simultaneously generates olfactory interneurons, astrocytes,=20 and oligodendrocytes. The molecular cues that enable SVZ = progenitors=20 to generate three distinct cell lineages without a temporal=20 switching mechanism are not known. Here, we demonstrate that=20 the basic helix-loop-helix transcription factor Olig2 plays = a=20 central role in this process. Olig2 is specifically expressed = in=20 gliogenic progenitors in the postnatal SVZ and by all glial = lineages=20 derived from this structure. By expressing normal and=20 dominant-interfering forms of Olig2 in vivo, we show that=20 Olig2 repressor function is both sufficient and necessary to=20 prevent neuronal differentiation and to direct SVZ = progenitors=20 toward astrocytic and oligodendrocytic fates. Although Olig2=20 activity has been associated previously with motor neuron and = oligodendrocyte development, our findings establish a = previously=20 unappreciated role for Olig2 in the development of = astrocytes.=20 Furthermore, these results indicate that Olig2 serves a = critical role=20 in pan-glial versus neuronal fate decisions in SVZ = progenitors,=20 making it the first intrinsic fate determinant shown to = operate=20 in the early postnatal SVZ.

Key words: SVZ; olig2; = neurogenesis;=20 gliogenesis; development; forebrain


3D"   =20 Introduction
Top
Abstract
3D"Introduction
Materials and Methods
Results
Discussion
References
 
I= n the=20 developing telencephalon, the neuroepithelium gives rise to = the=20 subventricular zone (SVZ), a secondary proliferative layer of = progenitors that simultaneously generates both neurons and = glia (for=20 review, see Marshall et al., 2003). The contemporaneous production of neurons = and glia=20 makes the SVZ a unique system in which to study the molecular = mechanisms directing cell fate decisions. During the = perinatal=20 period, the SVZ expands to include an angular prominence = between the=20 developing striatum and subcortical white matter (see Fig.=20 1A,B) and forms a continuous structure with the = rostral=20 migratory stream (RMS) (see Fig.=20 1A,C). SVZ progenitors specified to a neuronal = lineage=20 migrate rostrally to the olfactory bulb and differentiate = into=20 granular and periglomerular interneurons (Altman, 1969; Luskin, 1993; Lois and Alvarez-Buylla, 1994). In contrast, cells specified to become glia migrate=20 radially from the SVZ into the subcortical white matter, = corpus=20 callosum, striatum, and medial, dorsal, and lateral areas of = the=20 cerebral cortex, where they differentiate into astrocytes and = oligodendrocytes (see Fig.=20 1C) (Marshall et al., 2003).

The point at which SVZ progenitors commit to neuronal versus = glial=20 lineages is not known. Studies of SVZ cell migration suggest = that=20 distinct glial and neuronal cell lineages are established = within the=20 SVZ before long-range migrations into the forebrain (Suzuki = and=20 Goldman, 2003). However, a delineation of committed = neuronal and=20 glial populations within the SVZ has not been possible thus = far, and=20 molecules directing SVZ cell fate decisions (see Fig.=20 1D) have not been identified.

Comprehensive work on neuronal and glial specification in the=20 embryonic CNS has provided insight into molecular pathways = involved=20 in fate decisions. Basic helix-loop-helix (bHLH) = transcription=20 factors are important regulators of neuronal and glial cell=20 fates in the embryo (for review, see Bertrand et al., 2002), but their involvement in postnatal fate = decisions,=20 while anticipated, has not been well described. One family of = bHLH=20 proteins, the Oligs, has received a great deal of attention = for their=20 essential involvement in both motor neuron and = oligodendrocyte=20 development in the embryonic spinal cord. Both Olig1 and = Olig2 are=20 expressed in the embryonic and postnatal forebrain, but their = function in the development of this structure remains poorly=20 understood. During embryogenesis, constitutive expression of = Olig1 in=20 forebrain progenitors has been shown to promote the formation = of=20 oligodendrocytes (Lu et al., 2001). The function of Olig genes in the postnatal=20 forebrain, however, has not been evaluated, and progress in=20 this area has been hampered by the failure of Olig mutant = mice=20 to survive embryonic development. Thus, other methods must be = used to study the postnatal functions of Olig genes in = vivo.=20

In this study, we use retroviral-mediated gene transduction = to=20 express normal and dominant-interfering forms of Olig2 in SVZ = cells.=20 We show that Olig2 repressor function is both sufficient and=20 necessary to prevent neuronal differentiation and to direct = SVZ=20 progenitors toward astrocytic and oligodendrocytic fates. = Together,=20 our findings demonstrate a previously unappreciated role for = Olig2 in=20 astrocyte development and establish that Olig2 directs the = primary=20 specification of pan-glial versus neuronal lineages within = SVZ=20 progenitors in the neonatal forebrain.


3D"   =20 Materials and Methods
Top
Abstract
Introduction
3D"Materials and Methods
Results
Discussion
References
 
<= I>Retroviral=20 vectors and virus preparation. All fragments encoding = Olig2 or=20 mutant Olig2 and enhanced green fluorescent protein (eGFP) = cDNAs were=20 subcloned into the bicistronic replication-incompetent viral = vector=20 pQCXIX (Clontech, Cambridge, UK) 5' and 3' relative to an = internal=20 ribosomal entry site (IRES), respectively. The pQCXIX vector = uses a=20 cytomegalovirus immediate early promoter downstream of a 5' = long=20 terminal repeat (LTR) to drive expression of both = transcripts. Coding=20 fragments (Novitch et al., 2001) were as follows: mOlig2 (wild-type = mouse Olig2,=20 EcoRI fragment from pCS2+6MTmOlig2), cOlig2bHLH = (chick=20 Olig2 bHLH domains; amino acids 92-180; ClaI fragment = from=20 pCS2+MTcOlig2bHLH-VP16), MTcOlig2bHLH-VP16 (chick = Olig2 bHLH=20 domains fused to the herpes simplex VP16 transactivation = domain;=20 ClaI fragment from pCS2+MTcOlig2bHLH-VP16), and = eGFP=20 [EcoRV fragment provided by Satoshi Suzuki (Kyushu = University,=20 Fukuoka, Japan)]. To generate virus, retroviral vectors were=20 cotransfected with the vesicular stomatitis virus G (VSV-G) = viral=20 envelope into gp293 cells using Lipofectamine 2000 = (Invitrogen, San=20 Diego, CA). Growth medium was added to cells after 6 h, and=20 supernatant was harvested 48 h posttransfection. Viral stocks = were=20 concentrated and titered to 1-3 x=20 105 cfu/ml. The pNIT vector (provided by T. Palmer = and S.=20 Suhr, The Salk Institute, La Jolla, CA) is a bicistronic=20 replication-incompetent retrovirus that uses a 5' LTR to = drive=20 expression of the tetracycline activator and the tetracycline = operon=20 enhancer/promoter to drive the expression of eGFP. pNIT virus = was=20 prepared as described previously (Kakita and Goldman, 1999).

Retroviral injections. Sprague Dawley rat pups at = postnatal=20 day 2 (P2) to P3 were deeply anesthetized by immersion in ice = water and positioned in a stereotactic apparatus, and 0.5 = =B5l of=20 retrovirus solution was injected unilaterally at a rate of = 0.2 =B5l/min=20 using a 10 =B5l Hamilton (Reno, NV) syringe. Coordinates of = the=20 injection site used for all experiments are 1 mm anterior and = 1.5 mm=20 lateral to bregma at a depth of 2.0 mm.

Animals, histology, immunohistochemistry, and terminal=20 deoxynucleotidyl transferase-mediated biotinylated UTP nick = end=20 labeling assay. Human glial fibrillary acidic protein = (hGFAP)-GFP=20 mice (The Jackson Laboratory, Bar Harbor, ME), C57BL/6J mice, = and=20 Sprague Dawley rats (Charles River, Quebec, Canada) were=20 anesthetized by intraperitoneal injection of ketamine (75 = mg/kg;=20 Aveco, Fort Dodge, IA) and xylazine (5 mg/kg; Mobay, Shawnee, = KS) and=20 were transcardially perfused with 4% paraformaldehyde in PBS, = pH=20 7.4. Mouse brains were postfixed after <6 h, rat brains = were=20 postfixed overnight, and all brains were cryoprotected with = 30%=20 sucrose/PBS. Olig2-GFP mice (B.G. Novitch and T. M. Jessell,=20 unpublished observation) were generated by disruption of the=20 coding region of Olig2 with an Ires-EGFP cassette and = processed=20 as above. Serial coronal or parasagittal sections (see Fig.=20 1 A) were cut (16-30 =B5m) using a cryostat, = air-dried,=20 and stored at -80=B0C until use. Immunofluorescence was = performed=20 using the following antibodies: rabbit anti-GFP (1:500; = Molecular=20 Probes, Eugene, OR), mouse anti-GFP (1:200; Molecular=20 Probes), rabbit anti-Dlx2 (1:200; provided by J. = Rubinstein,=20 University of California, San Francisco, San Francisco, CA), = guinea=20 pig anti-olig2 [1:20,000 (Wichterle et al., 2002); antibody specificity was confirmed by a = complete lack=20 of labeling in olig2 mutants (supplemental Fig. 1, available = at http://www.jneurosci.org/ as supplem= ental=20 material), an expression pattern similar to in = situ=20 hybridizations for olig2 transcript and a nonoverlap with = olig1=20 immunoreactivity], mouse anti-CNPase (1:200; Sigma, St. = Louis, MO),=20 mouse anti-ZebrinII (1:100; provided by R. Hawkes, University = of=20 Calgary, Calgary, Alberta, Canada), rabbit anti-GFAP (1:1000; = Dako,=20 High Wycombe, UK), mouse anti-vimentin (1:100; Dako), NG2 = (1:400;=20 provided by W. Stallcup, The Burnham Institute, La Jolla, = CA),=20 mouse anti-CC1 (1:20; Oncogene Sciences, Uniondale, NY), = mouse=20 anti-neuronal-specific nuclear protein (NeuN; 1:250; = Chemicon,=20 Temecula, CA), mouse anti-Map2 (1:1000; Sternberger = Monoclonals,=20 Exeter, UK), mouse anti-calretinin (1:100; Chemicon), and = rabbit=20 anti-calbindin (1:1000; Swant, Bellinzona, Switzerland). = Biotinylated=20 secondary antibodies with avidin-conjugated fluorescein = (1:300;=20 Vector Laboratories, Burlingame, CA) were used to detect GFP = in=20 tissue sections from injected brains. Antigen retrieval was=20 performed before immunolabeling with antibodies to Zebrin II = and=20 GFAP as described previously (Marshall and Goldman, 2002). For terminal deoxynucleotidyl = transferase-mediated=20 biotinylated UTP nick end labeling (TUNEL) assay, an in = situ=20 cell death detection kit (Roche Products, Hertfordshire, UK) = was=20 used.

SVZ cell cultures and immunocytochemistry. SVZs from P3 = mice=20 were dissected, dissociated with a 0.025%Trypsin-EDTA/0.25 = mg/ml=20 DNase solution, and filtered through a 35 =B5m Nitex mesh.=20 Dissociated cells were grown in neurobasal medium with 0.1%=20 fetal bovine serum (FBS) for 3 d. Cells were then cultured = for=20 one additional day in neurobasal medium with 3% FBS to = generate=20 cultures enriched for immature astrocytes and an additional = 5 d=20 for cultures enriched for mature astrocytes. Cell cultures = were=20 labeled using the following antibodies: mouse anti-vimentin = (1:400;=20 Dako), mouse anti-GFAP (1:600; Dako), and guinea pig = anti-Olig2=20 (1:20,000) (Wichterle et al., 2002).


3D"   =20 Results
Top
Abstract
Introduction
Materials and Methods
3D"Results
Discussion
References
 
<= STRONG>Olig2=20 expression in the SVZ and RMS
We first examined the = expression of=20 Olig2 in the neonatal rodent forebrain SVZ and RMS using = antibodies=20 to Zebrin II (aldolase C) and the homeodomain transcription = factor=20 Dlx2 to define two major SVZ cell subpopulations. Zebrin II = is=20 specifically expressed by radial glia and astrocytes within = the=20 perinatal forebrain and is a useful marker for these cells=20 (Staugaitis et al., 2001; Marshall and Goldman, 2002). Zebrin II expression in astrocytic "border = cells"=20 identifies the limits of the perinatal SVZ, whereas a = reciprocal,=20 Zebrin II negative, "central" cell population is defined by = the=20 expression of Dlx2 (Marshall and Goldman, 2002). Immunohistochemical analysis of mouse forebrain = tissue=20 at P0-P7 revealed that Olig2 was expressed in both SVZ border = and central cells. Olig2 expression in Zebrin II+ border = cells,=20 which line the ventricles as well as the dorsolateral edges = of=20 the SVZ adjacent to the subcortical white matter and striatum, = is=20 shown in coronal sections (Fig.=20 1E-G). Olig2 expression in Dlx2+ central SVZ = cells, which=20 migrate radially into forebrain areas overlying the SVZ as = well=20 rostrally into the RMS, is best viewed in parasagittal = sections (Fig.=20 1H-J). We found that only a minority of the Dlx+ = cells=20 also express olig2. The RMS was strikingly devoid of Olig2 = expression=20 (Fig.=20 1K), suggesting that Olig2 might be expressed = specifically=20 by glia or glial progenitors but not by interneurons or = migratory=20 neuronal progenitors in the perinatal forebrain. We also = examined=20 cells from acutely dissociated P1 SVZs and found that = expression of=20 neuronal 3D{beta}-tubulin (TuJ1) and Olig2 do not overlap but, in = sum, can=20 account for the large majority of cells (data not shown). If = TuJ1 is=20 a marker for immature cells of the neuronal lineage, then = Olig2 may=20 be expressed by glioblasts and also perhaps by multipotent=20 cells, if such cells exist in the neonatal SVZ.

Developing astrocytes and oligodendrocytes express=20 Olig2
We next used immunohistochemical labeling to examine = the=20 expression of Olig2 throughout the rest of the perinatal = forebrain=20 [embryonic day 18 (E18) to P10], with particular attention to = developing glia. Olig2 was expressed in astrocytes as well as = oligodendrocytes throughout the cortex, subcortical white = matter, and=20 striatum. Because astrocytes arise from at least two sources = within=20 the perinatal forebrain, ZebrinII+ radial glia, which = transform=20 directly into astrocytes, and Dlx2+ migratory SVZ = progenitors,=20 which colonize the forebrain and complete their = differentiation=20 in the white matter and gray matter, we examined Olig2 = expression=20 in both radial glia and immature astrocytes. Radial glia = throughout=20 the perinatal dorsal forebrain, including those with an = intermediate=20 morphology suggestive of an ongoing transformation into a = mature=20 astrocyte phenotype (Fig.=20 2A, arrowheads) (Voigt, 1989), expressed Zebrin II and Olig2. Immature = astrocytes,=20 defined by their relatively simple morphology (in contrast to = the=20 complex, bushy appearance of mature astrocytes) and their = expression=20 of Zebrin II but not GFAP, also expressed Olig2+ (Fig.=20 2B, arrowheads). In addition, astrocytes that were = more=20 differentiated and expressed detectable GFAP in their = perinuclear=20 region (Fig.=20 2B-D, asterisk) as well as astrocytes in late = stages of=20 maturation, which expressed GFAP in the perinuclear region as = well as=20 throughout their processes (Fig.=20 2E-G), were also Olig2+. Notably, the GFAP+/Olig2+ = astrocytes=20 that we identified were distributed widely throughout the = forebrain=20 and in the cerebral cortex, white matter, and striatum. =

Although most, if not all, GFAP+ astrocytes in the neonatal=20 forebrain expressed Olig2, mature and fully arborized = astrocytes=20 expressing the highest amounts of GFAP (Fig.=20 2E, arrowheads) consistently appeared to express = slightly=20 lower levels of Olig2, as judged by Olig2 immunoreactivity, = than=20 astrocytes with a more immature morphology and expressing = less GFAP=20 (Fig.=20 2E, asterisk), suggesting a downregulation of = Olig2 in=20 mature astrocytes. Immunohistochemical analysis of forebrain=20 astrocytes at various developmental time points, from = neonatal=20 (P0-P7) to adult (P30), confirmed a gradual downregulation of = Olig2=20 expression (R. Ventura, unpublished observations). Thus, = Olig2=20 appears to be expressed transiently in forebrain astrocytes. = Olig2 is=20 expressed strongly as astrocytes become specified and = differentiated,=20 but it is subsequently downregulated after their development = is=20 mostly complete. In contrast, oligodendrocytes throughout the = forebrain, defined here by their expression of CNPase (Fig.=20 2H,I), expressed Olig2 consistently from early=20 developmental stages into adulthood (data not shown). =



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  		Figure = 1. Olig2=20 expression in the SVZ. A, Diagram of coronal = and=20 parasagittal planes highlighting the SVZ (B) = and RMS=20 (C) (both green), respectively. SVZ-derived = glial=20 precursors migrate radially within a coronal plane into = adjacent=20 forebrain areas, whereas SVZ-derived neuronal progenitors = migrate=20 rostrally in a parasagittal plane to the olfactory bulb.=20 D, Transcriptional regulators directing the = formation=20 of neuronal and glial lineages in SVZ cells are not known.=20 E-G, Olig2 expression in Zebrin II+ (border) = and=20 Zebrin II- (middle) SVZ cells, as indicated by = immunostaining of P6=20 mouse forebrain. The inset in E depicts the = area of=20 photomicrographs shown in E and = F. The=20 boxed area in F is magnified in = G.=20 H-J, Olig2 is expressed in Dlx2+ (central) SVZ = progenitors. The inset in H depicts the area = of the=20 photomicrograph shown in H and = I. The=20 boxed area in I is magnified in = J.=20 K, Cells within the RMS do not express Olig2. = lv,=20 Lateral ventricle; wm, white matter. =

 = ;
Because=20 Olig2 expression in forebrain astrocytes has not yet been = documented,=20 we examined this expression further using three independent = methods.=20 First, we analyzed neonatal forebrains from hGFAP-eGFP = transgenic mice. The human GFAP promoter drives expression of = GFAP at=20 very early stages of astrocyte development relative to the = mouse GFAP=20 promoter. Hence, in these mice, the hGFAP promoter drives = expression=20 of GFP in immature astrocytes (Malatesta et al., 2000). Many developing astrocytes in the perinatal = (P6)=20 telencephalon of hGFAP-eGFP transgenics coexpressed = GFP and=20 Olig2 (Fig.=20 2J,K). Second, we performed a short-term lineage = tracing=20 analysis of cells that had expressed Olig2 by examining = forebrain=20 sections taken from Olig2-GFP knock-in mice. Olig2-GFP+ = progenitors=20 appeared to give rise to all glial lineages, including many = GFAP+=20 astrocytes, in the white matter and cerebral cortex (Fig.=20 2L-N). Third, we analyzed the expression pattern = of Olig2=20 at different stages of astrocyte development in vitro.=20 Astrocytes express a predictable, sequential pattern of = intermediate=20 filaments during their differentiation. Immature astrocytes=20 express vimentin (Dahl et al., 1981), whereas the additional expression of GFAP = (Chiu et=20 al., 1981) is often used as an indicator of maturation. = SVZ=20 progenitors, harvested at P3, were cultured for 4 d in=20 astrocyte-supporting conditions to generate immature = astrocytes. Most=20 cells coexpressed vimentin and Olig2 after 4 d in = vitro (div)=20 (Fig.=20 2O). These cells did not express immunodetectable = levels=20 of GFAP. In contrast, older astrocyte cultures at 8 div were=20 comprised of more mature, GFAP+ cells. However, few of these=20 astrocytes coexpressed Olig2 strongly (Fig.=20 2P), pointing again to a downregulation of Olig2 = in mature=20 astrocytes. These findings support results from previous = studies in=20 which Olig2 expression was positively correlated with an = immature=20 astrocyte phenotype but negatively correlated with a mature=20 astrocyte phenotype in neural progenitor cell cultures = (Fukuda et=20 al., 2003; Gabay et al., 2003). A summary of Olig2 expression with respect = to glial=20 cell lineages in the postnatal forebrain is depicted in Figure=20 2Q.



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  		Figure = 2.=20 Developing astrocytes and oligodendrocytes express Olig2.=20 A, Cortical radial glia with an intermediate=20 morphology (arrowheads), indicative of a transformation into = astrocytes, express Zebrin II and Olig2 (P3, coronal section = through=20 murine cerebral cortex). B-D, Olig2 and Zebrin = II are=20 expressed robustly in developing astrocytes (P6, murine = cerebral=20 cortex) at early stages of differentiation before GFAP = expression=20 (B, arrowheads) and at later stages as GFAP is = upregulated (B, asterisk). The cell marked = with an=20 asterisk in B is magnified to show perinuclear = GFAP=20 (C) and Zebrin II (D) = immunolabeling=20 with respect to nuclear Olig2 expression. E-G, = Mature=20 and fully arborized astrocytes (P6, murine cerebral cortex)=20 expressing high amounts of GFAP and Zebrin II = (E,=20 arrowheads) consistently appeared to express lower levels of = Olig2=20 than astrocytes with immature morphologies that express less = GFAP=20 (E, asterisk). H, = I,=20 CNPase+/Olig2+ oligodendrocytes in parasagittal sections of = P6 mouse=20 forebrain. J, K, Developing = astrocytes=20 in the hGFAP-GFP mouse (P7) forebrain coexpress GFP = and=20 Olig2. L, M, Astrocytes in the=20 mOlig2-I-eGFP mouse forebrain (P6, cortex) coexpress = GFAP,=20 GFP, and Olig2. O, Cell cultures generated = from P3=20 mouse SVZ contained many vimentin+/Olig2+, immature = astrocytes after=20 4 div. P, After 8 div, cultures were mostly = comprised=20 of mature, GFAP+ astrocytes. Olig2 immunoreactivity in these = mature=20 astrocytes was markedly reduced. Q, Summary of = Olig2=20 expression with respect to glial cell lineages in the = postnatal=20 forebrain. =

 = ;
Olig2=20 is expressed exclusively by SVZ-derived glia
SVZ progenitors = generate interneurons, astrocytes, and oligodendrocytes = during the=20 perinatal period. Those cells specified to become glia = migrate=20 radially into the cerebral cortex, subcortical white matter, = and=20 striatum, whereas those specified to become interneurons = migrate=20 rostrally, via the RMS, to the olfactory bulb. As noted = above,=20 astrocytes and oligodendrocytes in the forebrain expressed = Olig2,=20 whereas cells in the RMS did not. Furthermore, a short-term = lineage=20 analysis using Olig2-GFP knock-in mice revealed GFP = expression in=20 postnatal forebrain glia but not olfactory interneurons. To = determine=20 definitively which SVZ-derived cells express Olig2, we = transduced SVZ=20 cells in vivo with GFP using a replication-incompetent = retrovirus encoding GFP (X-ires-eGFP) and = analyzed=20 infected cells at several time points. A virus was injected=20 unilaterally into the SVZ of neonatal rat pups (P2/P3) at the = rostral=20 aspect of the SVZ, in which the RMS diverges from the SVZ (Fig.=20 3F, arrow), using a stereotax. Infected cells were = examined at 2 and 4 d postinjection (dpi) in coronal (Fig.=20 3A-C) and parasagittal (Fig.=20 3D-I) sections. The distribution and phenotypes of = GFP+=20 cells in the olfactory bulb, RMS, SVZ, white matter, and = cortex (Fig.=20 3F) were similar to those observed previously = (Suzuki and=20 Goldman, 2003). At 2 dpi, most infected cells were found = within the=20 SVZ, whereas only a few had migrated into the overlying white = matter=20 (Fig.=20 2A). By 4 dpi, infected cells colonized the white = matter,=20 cortex, striatum, RMS, and olfactory bulb. Many of the = infected=20 SVZ cells that had migrated radially into the white matter,=20 striatum, and cortex were GFP+/Olig2+ and exhibited = morphologic=20 characteristics of oligodendrocytes (Fig.=20 3B) as well as immature astrocytes (Fig.=20 3C; the asterisk marks a blood vessel associated with = the=20 astrocyte). In contrast, GFP+ cells migrating in the RMS (Fig.=20 3D-F) or colonizing the olfactory bulb (Fig.=20 3G-I) did not express Olig2. Interestingly, Olig2+ = cells=20 lining the RMS (Fig.=20 1K) are closely adjacent to Olig2- migratory neuroblasts = (Fig.=20 3E) and perhaps comprise part of the astrocyte = population=20 that encapsulates neuroblasts within the RMS (Doetsch et al., = 1997).



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  		Figure = 3. Olig2 is=20 expressed exclusively by SVZ-derived glia. A, = GFP=20 expression 2 d after stereotactic injection of = replication-deficient=20 retrovirus (pNIT-eGFP) into P2/P3 rat SVZ. See=20 F for the site of injection and distribution = of=20 infected cells at 4 dpi. B, C, = At 4 dpi,=20 GFP+/Olig2+ cells with oligodendrocyte (B) and = astrocyte (C) morphologies were observed in = the=20 parenchyma (the asterisk in C indicates a = blood vessel=20 enwrapped by an astrocyte). D, = E,=20 SVZ-derived neuroblasts do not express Olig2 as they migrate = in the=20 RMS, which is outlined for clarity. The boxed area is = magnified in=20 E. F, Distribution of SVZ cells=20 transduced with GFP. The red box indicates the area of the=20 photomicrograph in D. G-I, = SVZ-derived=20 interneuronal precursors remain Olig2-within the olfactory = bulb. The=20 boxed area in G is magnified in = H.=20 I, The red box indicates the area of the=20 photomicrograph in G. lv, Lateral ventricle; = wm, white=20 matter; str, striatum. =

 = ;
In=20 summary, SVZ cells that became specified to a gliogenic lineage = and=20 migrated radially into the parenchyma expressed Olig2. In = contrast,=20 SVZ cells specified to an olfactory interneuron lineage that = migrated=20 rostrally via the RMS and differentiated in the olfactory = bulb did=20 not express Olig2. These findings demonstrated an intriguing=20 correlation between Olig2 expression and glial fates, raising = the=20 possibility that Olig2 transcriptional regulatory function = may direct=20 the specification of glial versus neuronal lineages in SVZ=20 progenitors.

SVZ progenitors transduced with Olig2 differentiate = exclusively into=20 glia
To examine the potential functions of Olig2 in = directing=20 SVZ progenitors to glial lineages, we transduced SVZ cells = with=20 a bicistronic retrovirus encoding Olig2 and eGFP=20 (Olig2-ires-eGFP) or a control virus encoding = eGFP=20 alone (X-ires-eGFP). All experiments were performed = using the=20 same retroviral vector (pQCXIX), protein coat (VSV-G), and=20 stereotaxic coordinates.

Retrovirus was injected into neonatal rats as described above, = and=20 brains (n =3D 8) were harvested at 4 and 8 dpi. Olig2 = expression=20 in SVZ cells appeared to prevent the migration and = differentiation=20 of olfactory interneurons, because no infected cells were = observed=20 in the distal RMS or olfactory bulb (Fig.=20 4C). Immunohistochemistry using horseradish = peroxidase and=20 diaminobenzidine precipitation was performed to ensure that = cells=20 expressing low levels of GFP, and thus undetectable by=20 immunofluorescence, were evaluated. This more sensitive = analysis did=20 not reveal any labeled cells in the RMS or olfactory bulb = (data not=20 shown). Instead, the large majority (3D~=2088%) of cells = misexpressing=20 Olig2 migrated radially from the SVZ into the white matter = and=20 cerebral cortex, areas typically colonized by glia (Fig.=20 4C). Others (3D~12%) differentiated within the SVZ/proximal RMS = (Fig.=20 4A-D) before their migration. We did not observe=20 differentiated cells within the SVZ or RMS in brains injected = with=20 our control virus; this premature differentiation is not a = behavior=20 typical of SVZ cells (Suzuki and Goldman, 2003).

To quantitate the relative distributions of infected cells, = a=20 sampling of three comparable parasagittal sections was taken = from=20 five brains misexpressing Olig2 and three brains expressing = the=20 control vector at 4 dpi. Selected sections contained the = injection=20 site and the entire RMS. Cells infected with the X-ires-eGFP = control virus (Fig.=20 5A) migrated radially into the white matter, = striatum, and=20 all layers of the cerebral cortex as well as rostrally into = the RMS=20 and olfactory bulb (Fig.=20 5B). In these control parasagittal sections, 51% = of the=20 cells were found in the olfactory bulb, and 27% were found in = the=20 RMS. In contrast, the majority of cells misexpressing=20 Olig2-ires-eGFP (Fig.=20 5C) migrated into the white matter (51%) and all = layers of=20 the cerebral cortex (37%); 12% were found in the proximal = RMS, but no=20 cells were identified in the distal RMS or olfactory bulb (Fig.=20 5D). Infected brains were cut along a parasagittal = plane,=20 enabling us to analyze the rostral versus radial migration of = SVZ=20 cells within each section. Our quantitative distributions (Fig.=20 5B,D) represent cells solely within this = parasagittal=20 plane, however, and do not predict the distribution of = labeled cells=20 throughout the entire brain. Because many SVZ-derived glia = migrate to=20 dorsolateral areas of the forebrain, these cells are not = represented=20 in our quantitations.



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  		Figure = 4. SVZ=20 progenitors transduced with Olig2 differentiate exclusively = into=20 astrocytes and oligodendrocytes. A-C, = Parasagittal=20 section of rat forebrain at 4 dpi, showing cells transduced = with=20 Olig2-ires-eGFP. The boxed area in A is = magnified in B, highlighting differentiated = glial=20 cells within the SVZ/RMS. C, Distribution of = cells=20 expressing Olig2-ires-eGFP at 4 dpi. Thered box = indicates the=20 area of SVZ and proximal RMS shown in A.=20 D, Infected cells differentiated ectopically = in the=20 SVZ, adjacent to the lateral ventricles. E-K, = More=20 than 70% of infected cells exhibited the morphology of=20 oligodendrocytes (E) and expressed the=20 oligodendrocyte-specific markers NG2 (F-H) and = CC1=20 (I-K). L-N, Approximately 20% of = infected cells exhibited astrocyte morphologies and = expressed GFAP=20 (asterisk). lv, Lateral ventricle; wm, white matter; str, = striatum. =

 = ;



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  		Figure = 5.=20 Distribution of SVZ cells expressing X-ires-eGFP and=20 Olig2-ires-eGFP. A-D, Retroviral = constructs=20 used to transduce SVZ cells with GFP (A) or = Olig2-GFP=20 (C) and quantified distributions of cells=20 misexpressing GFP (B) or Olig2-ires-GFP=20 (D), presented as a percentage of infected = cells per=20 brain located in each area (mean =B1 SEM). OB, Olfactory = bulb; WM,=20 white matter; CTX, cerebral cortex. =

 = ;
The=20 skewed, radial distribution of cells misexpressing Olig2 = suggests a=20 redirection of cells migrating rostrally from the SVZ/RMS to = areas=20 normally colonized by SVZ-derived glia. It is unlikely that a = difference in viral titer or apoptosis is responsible for the = cellular distributions or phenotypes we observed. All viruses = were=20 titered to 1-3 x 105 = cfu/ml.=20 Furthermore, the numbers of infected cells were consistent = across=20 experiments. In brains harvested at 4 dpi, an average of 66.3 = cells=20 infected with Olig2-ires-eGFP and 77 cells infected = with the=20 X-ires-eGFP control were observed per section. = Furthermore,=20 our findings do not appear to be the result of apoptosis. Few = infected cells contained TUNEL+ nuclei in brains injected = with either=20 the X-ires-eGFP control (<5%) (Fig.=20 5B) or Olig2-ires-eGFP (<10%) (Fig.=20 5D); therefore, Olig2 misexpression did not selectively = kill all SVZ cells destined to become olfactory interneurons. = Rather, the neuroblasts, glioblasts, and progenitors that = were=20 infected with Olig2-ires-GFP appeared to differentiate = unilaterally along a glial lineage. Over 70% of infected = cells=20 exhibited morphologies typical of lacy (Fig.=20 4E) or myelinating oligodendrocytes and expressed = NG2 (Fig.=20 4F-H) or CC1 (Fig.=20 4I-K), whereas 20% had the morphology of = astrocytes and=20 expressed GFAP (Fig.=20 4L-N). Approximately 10% of the infected cells = appeared to=20 exhibit glial morphologies but did not express = oligodendrocytic=20 (NG2, CC1), astrocytic (GFAP), or neuronal [NeuN,=20 microtubule-associated protein 2 (MAP2)] markers. In control=20 experiments, the same relative percentage of oligodendrocytes = (70%)=20 and astrocytes (30%) were generated by perinatal SVZ cells = that=20 emigrated into the white matter and cortex (Levison and = Goldman,=20 1993). Notably, Olig2-ires-eGFP+ cells that = expressed=20 GFAP (Fig.=20 4L-N) exhibited an immature astrocytic morphology = with=20 limited arborization, suggesting that constitutive Olig2 = expression=20 inhibited complete astrocyte maturation. In contrast, brains = infected=20 with control virus contained many GFP+/GFAP+ fully mature = astrocytes=20 and olfactory interneurons, as described previously (Levison = et=20 al., 1993; Luskin et al., 1993; Suzuki and Goldman, 2003).



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  		Figure = 6. A=20 dominant-interfering form of Olig2 blocks glial = differentiation.=20 A, B, SVZ cells misexpressing = Olig2=20 bHLH exhibited normal migration to the olfactory bulb = via the=20 RMS (A), which is outlined for clarity. The=20 distribution of total infected cells at 4 dpi is shown in=20 B. The red box indicates the area of RMS = pictured=20 (A). Infected cells also colonized the white = matter=20 and cerebral cortex, but glial differentiation was blocked = in=20 >90% of cells (data not shown). C-E, = However, 25% of=20 infected cells in the white matter and cortex ectopically = expressed=20 the neuronal marker NeuN, suggesting a potential = respecification=20 along a neuronal lineage. wm, White matter. =

 = ;
In=20 summary, the expression of Olig2 in neonatal SVZ cells is = sufficient=20 to direct glial versus neuronal identities, perhaps = prematurely in=20 some cells. Because no infected cells expressed either the = neuronal=20 marker NeuN or MAP2, our findings suggest that constitutive=20 expression of Olig2 directs SVZ progenitors to migrate = radially, away=20 from the SVZ/RMS, and differentiate into glia at the expense = of=20 interneurons. Apoptotic indices do not appear to account for = the=20 differences in distributions of infected cells. However, = rapid cell=20 death of neuroblasts misexpressing Olig2 might not have been=20 observed.

A dominant-interfering form of Olig2 blocks glial=20 differentiation
Because Olig2 expression appeared to specify = SVZ=20 cells toward glial lineages, we examined the consequences of = a loss=20 of Olig2 activity. Olig2 influences cell identity through its = ability to bind DNA and repress target gene expression = (Novitch et=20 al., 2001; Zhou et al., 2002). We reasoned that misexpression = of=20 the Olig2 bHLH DNA binding domain alone might interfere with=20 the repressor function of endogenous Olig2. SVZ cells were = therefore=20 transduced with an Olig2 construct containing the bHLH domain = alone (Olig2bHLH-ires-eGFP) (Fig.=20 6), as described above.

In all brains examined (4 and 8 dpi; n =3D 8), cells = expressing=20 Olig2bHLH-ires-eGFP migrated into the RMS, olfactory bulb,=20 white matter, and cortex (Fig.=20 6B). However, glial differentiation was profoundly = disrupted. Infected cells that migrated radially exhibited a = normal=20 but immature, unipolar or bipolar morphology. Few = differentiated=20 GFAP+ astrocytes (<5% per brain) and CC1+ oligodendrocytes = (<5%) were observed, and those that were seen were found=20 exclusively in the white matter (data not shown). = Furthermore, 5% of = infected cells=20 in the white matter and cortex retained Dlx2 expression, = which was=20 downregulated completely by SVZ-derived glia at the same time = points=20 in control experiments. In contrast, interneuron migration = and=20 differentiation did not appear to be affected by expression = of the=20 Olig2bHLH-ires-eGFP construct. Many infected cells = (40%)=20 migrated rostrally into the RMS (Fig.=20 6A,B) and olfactory bulb, where they differentiated=20 normally (data not shown). We infer from these results that=20 Olig2 serves a required function in both astrocyte as well as = oligodendrocyte formation but is not necessary for = interneuron=20 development in SVZ cells.

Although Olig2bHLH-ires-eGFP+ cells colonized the white = matter=20 and cerebral cortex, 3D~=2025% of these = infected cells=20 ectopically expressed the neuronal marker NeuN (Fig.=20 6C-E). This was never observed in brains injected = with the=20 X-ires-eGFP control virus and is not a behavior = typical of=20 SVZ-derived cells (Suzuki and Goldman, 2003), suggesting that disruption of Olig2 function = may=20 cause cells in these normally gliogenic regions to adopt = neuronal=20 fates. We surmise that the appearance of cells with neuronal=20 characteristics in the white matter and cerebral cortex most = likely=20 results from SVZ progenitors in which the onset of Olig2 bHLH = domain=20 expression had occurred at the time of or shortly after these = progenitors had begun their radial migration, and thus too = late for=20 these cells to enter the RMS. In addition to these aberrantly = positioned neurons, we found that 3D~=2030% of=20 Olig2bHLH-ires-eGFP infected cells in each brain lacked=20 processes and had TUNEL+ nuclei. A likely possibility is that = cells=20 that become respecified to a neuronal fate ectopically may = not=20 receive appropriate environmental trophic support, leading to = their premature death.

In summary, ectopic expression of a competitor form of Olig2=20 prevented both astrocyte and oligodendrocyte differentiation = in=20 >90% of infected SVZ cells. However, expression of the = truncated=20 Olig2 did not interfere with the specification of neuroblasts = or=20 their migration to the olfactory bulb. Cells that migrated = radially=20 into the developing forebrain either remained = undifferentiated or=20 began to differentiate ectopically along a neuronal lineage.=20 Together, these results strongly support a requirement for = Olig2=20 during SVZ progenitor specification to a glial fate. =

An antimorphic activator form of Olig2 promotes neurogenesis = in SVZ=20 cells
To confirm that Olig2 repressor activity is required = for=20 SVZ progenitor specification to glial fates, we used a=20 dominant-activator form of Olig2 to reverse endogenous Olig2=20 repressor function. A retrovirus encoding the Olig2 bHLH = subdomains=20 fused to a herpes simplex VP16 activator domain was generated = (Olig2bHLH/VP16-ires-eGFP) (Fig.=20 7), and injections were performed as described.

Infected cells from brains harvested at 4 and 8 dpi (n =3D = 9)=20 were examined. Infected cells either remained within the SVZ = or=20 migrated a short distance into the overlying white matter, = layer 6,=20 or proximal RMS (Fig.=20 7A-C). No infected cells exhibited glial = morphologies or=20 expressed the glial markers vimentin, GFAP, CC1, or NG2. = Rather,=20 cells expressing the antimorph appeared to differentiate = prematurely=20 into interneurons. Strikingly, >75% of infected cells = positioned=20 within the SVZ, proximal descending arm of the RMS, white = matter, and=20 cortex, expressed the neuronal markers MAP2 (Fig.=20 7D-F) or NeuN (Fig.=20 7G-I). Furthermore, 3D~=2030% of infected = cells also=20 expressed markers specific to interneurons, such as calbindin = (Fig.=20 7J-L) or calretinin (Fig.=20 7M-O) but exhibited relatively immature neuronal=20 morphologies. Thus, expression of the dominant-activator form = of=20 Olig2 appeared to elicit an aberrant and premature program of = neuronal differentiation. As in the case of the bHLH = domain-alone=20 construct, misexpression of the activator form of Olig2 led = to a=20 significant amount of apoptosis, 3D~=2050% of the = infected cell=20 population. Together, these findings provide evidence that = Olig2=20 repressor function is required to prevent neuronal = differentiation=20 and to direct both astrocytic and oligodendrocytic fates in = postnatal=20 SVZ progenitors.



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  		Figure = 7. An=20 antimorphic activator form of Olig2 promotes neurogenesis in = SVZ=20 cells. A-C, Expression of an = Olig2bHLH-VP16=20 activator fusion construct limited the migration of SVZ = cells. Cells=20 migrated short distances into the proximal RMS, white = matter, and=20 layer 6 of the cerebral cortex. The boxed area in = A,=20 containing infected SVZ cells, is magnified in = B. The=20 distribution of infected cells is shown in C.=20 D-I, More than 75% of infected cells expressed = the=20 neuronal markers Map2 (D-F) and NeuN=20 (G-I). J-O, Approximately 30% of = cells=20 in the white matter and SVZ/RMS expressed = interneuron-specific=20 markers such as calbindin (J-L) and calretinin = (M-O). WM/VI, White matter/layer 6 border; LV, = lateral=20 ventricle. =

 = ;

3D"   =20 Discussion
Top
Abstract
Introduction
Materials and Methods
Results
3D"Discussion
References
 
W= e provide=20 evidence that Olig2 specifies SVZ cells to pan-glial versus = neuronal=20 lineages. By repressing a neuronal phenotype in SVZ = progenitors,=20 Olig2 expression permits the production of astrocyte and=20 oligodendrocyte precursors. Our findings establish a = previously=20 unappreciated role for Olig2 in astrocyte development and = suggest=20 that Olig2 directs the primary specification of glial versus = neuronal=20 lineages in SVZ progenitors, making it the first intrinsic = fate=20 determinant shown to operate within the postnatal SVZ. =

A role for Olig2 in astrocyte = differentiation
Olig2=20 expression has not been reported previously in forebrain = astrocytes.=20 It is not surprising, however, that this is the case. GFAP is = the=20 most common molecular marker used for the identification of=20 astrocytes, and it is expressed at relatively late stages of=20 astrocyte development in rodents. Still, we observed many = astrocytes=20 throughout the forebrain that coexpressed Olig2 and GFAP. One = possible explanation for the discrepancy between our findings = and=20 those of other studies is the method of detecting GFAP in = tissue=20 sections. Using basic immunofluorescence techniques, GFAP = expression=20 is readily detectable in a subset of the astrocytes = populating the=20 SVZ, white matter, and subpial area. However, other = astrocytes,=20 located in gray matter, express lower levels of GFAP, which = is less=20 easily detectable. To visualize all forebrain astrocytes with = GFAP=20 immunofluorescence, we used an antigen retrieval technique. = Many of=20 these "unmasked" GFAP-expressing astrocytes were positive for = Olig2.=20 Thus, this difference in GFAP detection might account for = differences=20 between our data and those reporting no Olig2 expression in = forebrain=20 astrocytes (Zhou et al., 2002).

In addition, Olig2 expression is downregulated in mature = astrocytes.=20 Astrocytes in the adult forebrain express little, if any,=20 immunodetectable Olig2 (R. Ventura, unpublished = observations). Thus,=20 studies focused on Olig2 expression in mature glia would not = have=20 seen expression in astrocytes. A transient expression in=20 developing astrocytes supports in vitro studies in = which Olig2=20 expression was positively correlated with an immature = astrocyte=20 phenotype but negatively correlated with a fully mature = phenotype=20 (Gabay et al., 2003) and partially inhibited GFAP transcription (Fukuda = et=20 al., 2003) in neural progenitor cultures. However, in these=20 experiments, a significant proportion of cells misexpressing=20 Olig2 remained GFAP+. Thus, Olig2 may not be sufficient to = regulate=20 GFAP expression. Together with our findings in vivo, these=20 results point to two potential roles for Olig2 in astrocyte=20 development (supplemental Fig. 2, available at http://www.jneurosci.org/ as supplem= ental=20 material). First, Olig2 function is required for = astrocyte=20 formation in SVZ cells. Second, Olig2 may also serve to limit = GFAP=20 expression.

Olig1/2 mutants suffer a complete loss of oligodendrocytes, = but no=20 astrocyte phenotype was observed (Zhou et al., 2002). Several = potential interpretations of the Olig1/2 mutants can be made = with=20 respect to the forebrain. First, there are at least two = sources of=20 astrocytes in the postnatal forebrain. One population arises = directly=20 from radial glia that transform during the perinatal period = into=20 astrocytes (Voigt, 1989), whereas another population arises from = migratory=20 progenitors within the SVZ (Levison and Goldman, 1993; Luskin et al., 1993). Therefore, even a complete deficit in one = class of=20 astrocyte may not be noticeable on a gross level of = examination. Our=20 findings only demonstrate a requirement for Olig2 in the = development=20 of SVZ-derived astrocytes. Importantly, our retroviral = labeling=20 technique does not infect postnatal radial glia, which = proliferate=20 infrequently at this time relative to SVZ progenitors. Thus, = it is=20 possible that astrocytes observed in the Olig1/2 mutants = could be=20 derived exclusively from radial glial origins but not from=20 SVZ-derived astrocytes, which require Olig2 expression for = their=20 formation.

Could Olig1 compensate for Olig2? Olig1 and Olig2 expression = is=20 similar if not identical in the postnatal SVZ (C. A. G. Marshall,=20 unpublished observations), and Olig1 misexpression increases=20 astrocytes and oligodendrocytes in the developing forebrain = (Lu=20 et al., 2001); the two factors may work together in repressing=20 neurogenesis and/or promoting gliogenesis. However, if Olig1=20 compensated fully for Olig2 in the Olig2 blocking = experiments,=20 we would have expected to see some gliogenesis in SVZ cells=20 expressing the truncated Olig2 construct, but we did not. It = is=20 unlikely that Olig1 and Olig2 bind to identical DNA sequences, = given=20 the low homology these genes share in their bHLH domains. The = Olig2=20 constructs used for functional experiments are specific = (Novitch et=20 al., 2001; Zhou and Anderson, 2002; Gabay et al., 2003).

Olig2 expression in radial glia
Although it is = not clear=20 whether Olig2 is required for the production of astrocytes = from=20 postmitotic radial glia, we found that cortical radial glia = did=20 express Olig2 during the perinatal period (we examined = P0-P10).=20 Unlike the spinal cord, the cerebral cortex does not express = Olig2=20 during earlier, neurogenic stages of development (Nery et = al., 2001; our unpublished data). Rather, Olig2 is = expressed in=20 radial glia later, at a time when they have finished = generating=20 projection neurons and have "switched" from a primarily = neurogenic=20 state to a primarily gliogenic state (Anthony et al., 2004). Therefore, another potential role for Olig2 = in=20 neonatal forebrain development is to repress neurogenesis in = cortical=20 radial glia. Radial glia express astrocytic molecules such as = glutamate transporters and intermediate filaments and may be=20 committed to their astrocytic fate, however, before expression = of=20 Olig2. Experiments aimed at understanding the function of = Olig2 in=20 these dynamic cells will be required to understand any part = it might=20 play in determining their fates or the fates of their = progeny.=20

SVZ cell phenotype and migration
SVZ cells = specified to=20 become interneurons migrate rostrally via the RMS into the = olfactory=20 bulb (Altman, 1969; Luskin, 1993; Lois and Alvarez-Buylla, 1994), whereas SVZ cells specified to become glia = migrate=20 radially into the developing parenchyma (Fig.=20 1A) (Altman, 1966; Paterson et al., 1973; Imamoto et al., 1978; Levison and Goldman, 1993; Luskin et al., 1993; Suzuki and Goldman, 2003). The synchronization of SVZ cell phenotype = and=20 migration behavior is not understood. Do SVZ progenitors=20 stochastically choose a rostral versus radial migration = pathway=20 before becoming committed to neuronal and glial lineages? Or, = does the phenotype of a cell dictate the direction of its = migration?=20 One model that would explain this tight correlation between=20 cell fate and migration behavior is one in which progenitors=20 become specified to a glial lineage before their emigration=20 from the SVZ. Our findings strongly support such a model. We=20 demonstrate that SVZ cell fate decisions are likely to depend = critically on the expression of transcriptional regulators = such=20 as Olig2. Olig2-directed glial specification may subsequently = confer on these cells an ability to migrate across SVZ = borders=20 through transcriptional regulation of specific receptors or=20 adhesion molecules involved in accessing the radial glial = scaffold=20 and migrating through the perinatal brain. Because Olig2 = functions=20 as a transcriptional repressor, it might serve to = downregulate=20 genes that otherwise prevent emigration from the SVZ. Thus, = a=20 knowledge of downstream targets for Olig2 may contribute to = an=20 understanding of phenotypic specification as well as cell=20 distribution within the postnatal brain.

SVZ cell potential
Our model (supplemental Fig. = 2,=20 available at http://www.jneurosci.org/ = as supplem= ental=20 material) reconciles data from in vitro studies = (Noble et=20 al., 2004), which point to bipotent and tripotent cells = in the=20 developing forebrain (Levison and Goldman, 1997) and spinal cord (Gabay et al., 2003), with in vivo work (Levison and = Goldman, 1993; Luskin et al., 1993; Parnavelas, 1999; Liu and Rao, 2004; Zerlin et al., 2004), in which a smaller population of cells = demonstrate=20 such broad potential. What intrinsic and extrinsic cues = direct=20 progenitors that are multipotent in vitro to a limited = range=20 of fates in vivo? Studies of progenitor specification = have=20 been performed predominantly in embryonic systems, such as = the spinal=20 cord, in which a sequential assignment of neuronal and glial=20 phenotypes has been revealed and important molecular fate = regulators=20 have been identified (Lee and Jessell, 1999; Briscoe and Ericson, 2001). Although recent studies have indicated = functions for=20 some of these molecular regulators in forebrain development = (Woodruff=20 et al., 2001; Bertrand et al., 2002), the dynamics of neuronal and glial cell = production in=20 the subventricular zone are distinct from those in the = embryonic cord=20 and are less clear.

The SVZ generates interneurons, astrocytes, and oligodendrocytes = simultaneously during the perinatal period, but its output = changes=20 over time, so that the adult SVZ primarily generates neurons=20 (Doetsch et al., 1999, 2002) and only limited numbers of glia (Hack et = al., 2005). The molecular mechanisms regulating this = switch to=20 neurogenesis are not known, and a lack of temporospatial = segregation=20 among neuronal and glial progenitors has made this switch = difficult=20 to analyze. Even in the adult SVZ, however, constitutive = expression=20 of Olig2 via retrovirus leads to a substantial increase in=20 gliogenesis (Hack et al., 2005), suggesting that perinatal mechanisms may = persist into=20 adulthood. Our data showing the sufficiency of Olig2 in = directing=20 primary, pan-glial versus neuronal fate decisions in the = perinatal=20 SVZ is a fundamental step toward understanding the molecular=20 mechanisms that regulate SVZ cell fates. In the future, a = detailed=20 knowledge of SVZ cell fate regulation under normal conditions = will be=20 paramount for studying the responses of such cells to = pathologic=20 cues, as well as accessing this endogenous source of = progenitors=20 for potential therapeutic interventions in the diseased = brain.=20


3D"   =20 Footnotes
 
Received May = 13, 2005;=20 revised June 28, 2005; accepted June 29, 2005.

C.A.G.M. and J.E.G. were supported by = National=20 Institute of Neurological Disorders and Stroke Grant = NS-17125. B.G.N.=20 was a Marion Abbe Fellow of the Damon Runyon Cancer Research=20 Fund (DRG-1569). We thank Tom Jessell and Fiona Doetsch for=20 insightful discussions and comments on this manuscript. We = are=20 grateful to Carol Mason for graciously sharing imaging = equipment,=20 Rachel Ventura for help with Olig2 expression analysis, and=20 Michael Hack and Magdalena Gotz for sharing unpublished work. = Finally, we thank Satoshi Suzuki and Ana Milosevic for = providing=20 expert assistance with retroviral production and stereotaxic=20 injections.

Correspondence should be addressed to = Dr. James=20 E. Goldman, Division of Neuropathology, Department of = Pathology,=20 Columbia University, College of Physicians and Surgeons, 630 = West=20 168th Street, New York, NY 10032. E-mail: Jeg5{at}columbia.edu .

Copyright =A9 2005 Society for = Neuroscience=20 0270-6474/05/257289-10$15.00/0


3D"   =20 References
Top
Abstract
Introduction
Materials and Methods
Results
Discussion
3D"References
&nb= sp;

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Opera = requires=0A= the Sun Java Runtime Environment .=0A= =0A= by Andrew Gregory=0A= http://www.scss.com.au/family/andrew/webdesign/xmlhttprequest/=0A= =0A= This work is licensed under the Creative Commons Attribution License. To = view a=0A= copy of this license, visit = http://creativecommons.org/licenses/by-sa/2.5/ or=0A= send a letter to Creative Commons, 559 Nathan Abbott Way, Stanford, = California=0A= 94305, USA.=0A= =0A= Attribution: Leave my name and web address in this script intact.=0A= =0A= Not Supported in Opera=0A= ----------------------=0A= * user/password authentication=0A= * responseXML data member=0A= =0A= Not Fully Supported in Opera=0A= ----------------------------=0A= * async requests=0A= * abort()=0A= * getAllResponseHeaders(), getAllResponseHeader(header)=0A= =0A= */=0A= // IE support=0A= if (window.ActiveXObject && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= var msxmls =3D new Array(=0A= 'Msxml2.XMLHTTP.5.0',=0A= 'Msxml2.XMLHTTP.4.0',=0A= 'Msxml2.XMLHTTP.3.0',=0A= 'Msxml2.XMLHTTP',=0A= 'Microsoft.XMLHTTP');=0A= for (var i =3D 0; i < msxmls.length; i++) {=0A= try {=0A= return new ActiveXObject(msxmls[i]);=0A= } catch (e) {=0A= }=0A= }=0A= return null;=0A= };=0A= }=0A= // Gecko support=0A= /* ;-) */=0A= // Opera support=0A= if (window.opera && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= this.readyState =3D 0; // = 0=3Duninitialized,1=3Dloading,2=3Dloaded,3=3Dinteractive,4=3Dcomplete=0A= this.status =3D 0; // HTTP status codes=0A= this.statusText =3D '';=0A= this._headers =3D [];=0A= this._aborted =3D false;=0A= this._async =3D true;=0A= this._defaultCharset =3D 'ISO-8859-1';=0A= this._getCharset =3D function() {=0A= var charset =3D _defaultCharset;=0A= var contentType =3D = this.getResponseHeader('Content-type').toUpperCase();=0A= val =3D contentType.indexOf('CHARSET=3D');=0A= if (val !=3D -1) {=0A= charset =3D contentType.substring(val);=0A= }=0A= val =3D charset.indexOf(';');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= val =3D charset.indexOf(',');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= return charset;=0A= };=0A= this.abort =3D function() {=0A= this._aborted =3D true;=0A= };=0A= this.getAllResponseHeaders =3D function() {=0A= return this.getAllResponseHeader('*');=0A= };=0A= this.getAllResponseHeader =3D function(header) {=0A= var ret =3D '';=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= if (header =3D=3D '*' || this._headers[i].h =3D=3D header) {=0A= ret +=3D this._headers[i].h + ': ' + this._headers[i].v + '\n';=0A= }=0A= }=0A= return ret;=0A= };=0A= this.getResponseHeader =3D function(header) {=0A= var ret =3D getAllResponseHeader(header);=0A= var i =3D ret.indexOf('\n');=0A= if (i !=3D -1) {=0A= ret =3D ret.substring(0, i);=0A= }=0A= return ret;=0A= };=0A= this.setRequestHeader =3D function(header, value) {=0A= this._headers[this._headers.length] =3D {h:header, v:value};=0A= };=0A= this.open =3D function(method, url, async, user, password) {=0A= this.method =3D method;=0A= this.url =3D url;=0A= this._async =3D true;=0A= this._aborted =3D false;=0A= this._headers =3D [];=0A= if (arguments.length >=3D 3) {=0A= this._async =3D async;=0A= }=0A= if (arguments.length > 3) {=0A= opera.postError('XMLHttpRequest.open() - user/password not = supported');=0A= }=0A= this.readyState =3D 1;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= };=0A= this.send =3D function(data) {=0A= if (!navigator.javaEnabled()) {=0A= alert("XMLHttpRequest.send() - Java must be installed and = enabled.");=0A= return;=0A= }=0A= if (this._async) {=0A= setTimeout(this._sendasync, 0, this, data);=0A= // this is not really asynchronous and won't execute until the = current=0A= // execution context ends=0A= } else {=0A= this._sendsync(data);=0A= }=0A= }=0A= this._sendasync =3D function(req, data) {=0A= if (!req._aborted) {=0A= req._sendsync(data);=0A= }=0A= };=0A= this._sendsync =3D function(data) {=0A= this.readyState =3D 2;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= // open connection=0A= var url =3D new java.net.URL(new = java.net.URL(window.location.href), this.url);=0A= var conn =3D url.openConnection();=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= conn.setRequestProperty(this._headers[i].h, this._headers[i].v);=0A= }=0A= this._headers =3D [];=0A= if (this.method =3D=3D 'POST') {=0A= // POST data=0A= conn.setDoOutput(true);=0A= var wr =3D new = java.io.OutputStreamWriter(conn.getOutputStream(), this._getCharset());=0A= wr.write(data);=0A= wr.flush();=0A= wr.close();=0A= }=0A= // read response headers=0A= // NOTE: the getHeaderField() methods always return nulls for me :(=0A= var gotContentEncoding =3D false;=0A= var gotContentLength =3D false;=0A= var gotContentType =3D false;=0A= var gotDate =3D false;=0A= var gotExpiration =3D false;=0A= var gotLastModified =3D false;=0A= for (var i =3D 0; ; i++) {=0A= var hdrName =3D conn.getHeaderFieldKey(i);=0A= var hdrValue =3D conn.getHeaderField(i);=0A= if (hdrName =3D=3D null && hdrValue =3D=3D null) {=0A= break;=0A= }=0A= if (hdrName !=3D null) {=0A= this._headers[this._headers.length] =3D {h:hdrName, = v:hdrValue};=0A= switch (hdrName.toLowerCase()) {=0A= case 'content-encoding': gotContentEncoding =3D true; break;=0A= case 'content-length' : gotContentLength =3D true; break;=0A= case 'content-type' : gotContentType =3D true; break;=0A= case 'date' : gotDate =3D true; break;=0A= case 'expires' : gotExpiration =3D true; break;=0A= case 'last-modified' : gotLastModified =3D true; break;=0A= }=0A= }=0A= }=0A= // try to fill in any missing header information=0A= var val;=0A= val =3D conn.getContentEncoding();=0A= if (val !=3D null && !gotContentEncoding) = this._headers[this._headers.length] =3D {h:'Content-encoding', v:val};=0A= val =3D conn.getContentLength();=0A= if (val !=3D -1 && !gotContentLength) = this._headers[this._headers.length] =3D {h:'Content-length', v:val};=0A= val =3D conn.getContentType();=0A= if (val !=3D null && !gotContentType) = this._headers[this._headers.length] =3D {h:'Content-type', v:val};=0A= val =3D conn.getDate();=0A= if (val !=3D 0 && !gotDate) this._headers[this._headers.length] = =3D {h:'Date', v:(new Date(val)).toUTCString()};=0A= val =3D conn.getExpiration();=0A= if (val !=3D 0 && !gotExpiration) = this._headers[this._headers.length] =3D {h:'Expires', v:(new = Date(val)).toUTCString()};=0A= val =3D conn.getLastModified();=0A= if (val !=3D 0 && !gotLastModified) = this._headers[this._headers.length] =3D {h:'Last-modified', v:(new = Date(val)).toUTCString()};=0A= // read response data=0A= var reqdata =3D '';=0A= var stream =3D conn.getInputStream();=0A= if (stream) {=0A= var reader =3D new java.io.BufferedReader(new = java.io.InputStreamReader(stream, this._getCharset()));=0A= var line;=0A= while ((line =3D reader.readLine()) !=3D null) {=0A= if (this.readyState =3D=3D 2) {=0A= this.readyState =3D 3;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= }=0A= reqdata +=3D line + '\n';=0A= }=0A= reader.close();=0A= this.status =3D 200;=0A= this.statusText =3D 'OK';=0A= this.responseText =3D reqdata;=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onload) {=0A= this.onload();=0A= }=0A= } else {=0A= // error=0A= this.status =3D 404;=0A= this.statusText =3D 'Not Found';=0A= this.responseText =3D '';=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onerror) {=0A= this.onerror();=0A= }=0A= }=0A= };=0A= };=0A= }=0A= // ActiveXObject emulation=0A= if (!window.ActiveXObject && window.XMLHttpRequest) {=0A= window.ActiveXObject =3D function(type) {=0A= switch (type.toLowerCase()) {=0A= case 'microsoft.xmlhttp':=0A= case 'msxml2.xmlhttp':=0A= case 'msxml2.xmlhttp.3.0':=0A= case 'msxml2.xmlhttp.4.0':=0A= case 'msxml2.xmlhttp.5.0':=0A= return new XMLHttpRequest();=0A= }=0A= return null;=0A= };=0A= }=0A= ------=_NextPart_000_0066_01C709AF.2E622820 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.jneurosci.org/javascript/ajax/utility.js /************************************************************************= *****=0A= * javascript/ajax/utility.js=0A= *=0A= * Utility functions for working with XMLHttpRequest data.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Copy XML nodes into an HTMLElement. This effectively=0A= * clones XML markup which uses XHTML naming conventions=0A= * into an HTML DOM.=0A= */=0A= function copy_xml_to_html(src, dst) {=0A= if (src.nodeType =3D=3D 1) { /* Node.ELEMENT_NODE */=0A= var e =3D document.createElement(src.nodeName);=0A= for (var i =3D 0; i < src.childNodes.length; i++) {=0A= copy_xml_to_html(src.childNodes[i], e);=0A= }=0A= for (var i =3D 0; i < src.attributes.length; i++) {=0A= var n =3D src.attributes[i].name;=0A= var v =3D unescape_xml_string(src.attributes[i].value); =0A= e.setAttribute(n, v);=0A= if (n =3D=3D "class") {=0A= e.className =3D v;=0A= }=0A= else if (n =3D=3D "style") {=0A= set_css_style(v, e, "");=0A= }=0A= }=0A= dst.appendChild(e);=0A= }=0A= else if (src.nodeType =3D=3D 3) { /* Node.TEXT_NODE */=0A= dst.appendChild(document.createTextNode(src.nodeValue));=0A= }=0A= }=0A= =0A= /* =0A= * It is unclear that this is the right thing to be calling=0A= * from copy_xml_to_html, but it appears that Safari decides=0A= * to convert & to the NCR #, and then encodes that=0A= * NCR to &%26%2338;. So, I'm going to treat the DOM Attr=0A= * value as a plain string, and run our XML string input=0A= * through the decoding routine below.=0A= */=0A= function unescape_xml_string(s) {=0A= return s.replace(/'/g, "'")=0A= .replace(/'/g, "'")=0A= .replace(/"/g, "\"")=0A= .replace(/"/g, "\"")=0A= .replace(/>/g, ">")=0A= .replace(/>/g, ">")=0A= .replace(/</g, "<")=0A= .replace(/</g, "<")=0A= .replace(/&/g, "&")=0A= .replace(/&/g, "&");=0A= }=0A= =0A= /*=0A= * Parse set of CSS rules and apply them to an element.=0A= * This is quite horrifying, but I'm unable to determine=0A= * how else to handle this with IE 6. FireFox and other=0A= * sane browsers let you simply set the style attribute=0A= * or use e.style.setProperty(rule, value, priority),=0A= * IE 6 appears to have neither of these capabilities..=0A= */=0A= function set_css_style(css, e, priority) {=0A= var rules =3D css.split(";");=0A= for (var i =3D 0; i < rules.length; i++) {=0A= var nvpair =3D rules[i].split(":");=0A= if (nvpair.length =3D=3D 2) {=0A= try {=0A= var name =3D nvpair[0]; /* style attribute */=0A= var value =3D nvpair[1]; /* attribute value */=0A= =0A= /*=0A= * For each possible style attribute, set the=0A= * appropriate style property in the element.=0A= */=0A= if (name =3D=3D "background") {=0A= e.style.background =3D value;=0A= }=0A= else if (name =3D=3D "background-attachment") {=0A= e.style.backgroundAttachment =3D value;=0A= }=0A= else if (name =3D=3D "background-color") {=0A= e.style.backgroundColor =3D value;=0A= }=0A= else if (name =3D=3D "background-image") {=0A= e.style.backgroundImage =3D value;=0A= }=0A= else if (name =3D=3D "background-position") {=0A= e.style.backgroundPosition =3D value;=0A= }=0A= else if (name =3D=3D "background-position-x") {=0A= e.style.backgroundPositionX =3D value;=0A= }=0A= else if (name =3D=3D "background-position-y") {=0A= e.style.backgroundPositionY =3D value;=0A= }=0A= else if (name =3D=3D "background-repeat") {=0A= e.style.backgroundRepeat =3D value;=0A= }=0A= else if (name =3D=3D "behavior") {=0A= e.style.behavior =3D value;=0A= }=0A= else if (name =3D=3D "border") {=0A= e.style.border =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom") {=0A= e.style.borderBottom =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-color") {=0A= e.style.borderBottomColor =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-style") {=0A= e.style.borderBottomStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-width") {=0A= e.style.borderBottomWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-collapse") {=0A= e.style.borderCollapse =3D value;=0A= }=0A= else if (name =3D=3D "border-color") {=0A= e.style.borderColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left") {=0A= e.style.borderLeft =3D value;=0A= }=0A= else if (name =3D=3D "border-left-color") {=0A= e.style.borderLeftColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left-style") {=0A= e.style.borderLeftStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-left-width") {=0A= e.style.borderLeftWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-right") {=0A= e.style.borderRight =3D value;=0A= }=0A= else if (name =3D=3D "border-right-color") {=0A= e.style.borderRightColor =3D value;=0A= }=0A= else if (name =3D=3D "border-right-style") {=0A= e.style.borderRightStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-right-width") {=0A= e.style.borderRightWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-style") {=0A= e.style.borderStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top") {=0A= e.style.borderTop =3D value;=0A= }=0A= else if (name =3D=3D "border-top-color") {=0A= e.style.borderTopColor =3D value;=0A= }=0A= else if (name =3D=3D "border-top-style") {=0A= e.style.borderTopStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top-width") {=0A= e.style.borderTopWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-width") {=0A= e.style.borderWidth =3D value;=0A= }=0A= else if (name =3D=3D "bottom") {=0A= e.style.bottom =3D value;=0A= }=0A= else if (name =3D=3D "clear") {=0A= e.style.clear =3D value;=0A= }=0A= else if (name =3D=3D "clip") {=0A= e.style.clip =3D value;=0A= }=0A= else if (name =3D=3D "color") {=0A= e.style.color =3D value;=0A= }=0A= else if (name =3D=3D "cssText") {=0A= e.style.Sets =3D value;=0A= }=0A= else if (name =3D=3D "cursor") {=0A= e.style.cursor =3D value;=0A= }=0A= else if (name =3D=3D "direction") {=0A= e.style.direction =3D value;=0A= }=0A= else if (name =3D=3D "display") {=0A= e.style.display =3D value;=0A= }=0A= else if (name =3D=3D "font") {=0A= e.style.font =3D value;=0A= }=0A= else if (name =3D=3D "font-family") {=0A= e.style.fontFamily =3D value;=0A= }=0A= else if (name =3D=3D "font-size") {=0A= e.style.fontSize =3D value;=0A= }=0A= else if (name =3D=3D "font-style") {=0A= e.style.fontStyle =3D value;=0A= }=0A= else if (name =3D=3D "font-variant") {=0A= e.style.fontVariant =3D value;=0A= }=0A= else if (name =3D=3D "font-weight") {=0A= e.style.fontWeight =3D value;=0A= }=0A= else if (name =3D=3D "height") {=0A= e.style.height =3D value;=0A= }=0A= else if (name =3D=3D "ime-mode") {=0A= e.style.imeMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-flow") {=0A= e.style.layoutFlow =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid") {=0A= e.style.layoutGrid =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-char") {=0A= e.style.layoutGridChar =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-line") {=0A= e.style.layoutGridLine =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-mode") {=0A= e.style.layoutGridMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-type") {=0A= e.style.layoutGridType =3D value;=0A= }=0A= else if (name =3D=3D "left") {=0A= e.style.left =3D value;=0A= }=0A= else if (name =3D=3D "letter-spacing") {=0A= e.style.letterSpacing =3D value;=0A= }=0A= else if (name =3D=3D "line-break") {=0A= e.style.lineBreak =3D value;=0A= }=0A= else if (name =3D=3D "line-height") {=0A= e.style.lineHeight =3D value;=0A= }=0A= else if (name =3D=3D "list-style") {=0A= e.style.listStyle =3D value;=0A= }=0A= else if (name =3D=3D "list-style-image") {=0A= e.style.listStyleImage =3D value;=0A= }=0A= else if (name =3D=3D "list-style-position") {=0A= e.style.listStylePosition =3D value;=0A= }=0A= else if (name =3D=3D "list-style-type") {=0A= e.style.listStyleType =3D value;=0A= }=0A= else if (name =3D=3D "margin") {=0A= e.style.margin =3D value;=0A= }=0A= else if (name =3D=3D "margin-bottom") {=0A= e.style.marginBottom =3D value;=0A= }=0A= else if (name =3D=3D "margin-left") {=0A= e.style.marginLeft =3D value;=0A= }=0A= else 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quoted-printable Content-Location: http://www.jneurosci.org/javascript/entrez/callback.js /************************************************************************= *****=0A= * javascript/entrez/callback.js=0A= *=0A= * Entrez Linking callback to populate content box.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Execute callback to fill content box with Entrez Linking information.=0A= */=0A= function entrez_callback(pmid, callback_url) {=0A= /*=0A= * MSIE 5.5 and below have issues with the JavaScript=0A= * used for Entrez Linking. 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