From: Subject: Glial Progenitors in Adult White Matter Are Driven to Form Malignant Gliomas by Platelet-Derived Growth Factor-Expressing Retroviruses -- Assanah et al. 26 (25): 6781 -- Journal of Neuroscience Date: Wed, 16 Jan 2008 18:59:51 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C85871.F932BDA0" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.3198 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C85871.F932BDA0 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.jneurosci.org/cgi/content/full/26/25/6781 Glial Progenitors in Adult White Matter Are Driven to = Form Malignant Gliomas by Platelet-Derived Growth Factor-Expressing = Retroviruses -- Assanah et al. 26 (25): 6781 -- Journal of = Neuroscience
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The Journal of = Neuroscience, June=20 21, 2006, 26(25):6781-6790; doi:10.1523/JNEUROSCI.0514-06.2006

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Neurobiology of=20 Disease
Glial Progenitors in = Adult=20 White Matter Are Driven to Form Malignant Gliomas by Platelet-Derived = Growth=20 Factor-Expressing Retroviruses

Marcela Assanah,1 = Richard=20 Lochhead,2 Alfred Ogden,2=20 Jeffrey Bruce,2 James=20 Goldman,1,3 and Peter=20 Canoll1

Departments of 1Pathology and 2Neurological = Surgery,=20 and 3Center for Neurobiology and Behavior, Columbia = University, New=20 York, New York 10032=20


3D"   =20 Abstract
Top
3D"Abstract
Introduction
Materials and Methods
Results
Discussion
References
 
T= o test=20 the gliomagenic potential of adult glial progenitors, we = infected=20 adult rat white matter with a retrovirus that expresses high = levels=20 of PDGF and green fluorescent protein (GFP). Tumors that = closely=20 resembled human glioblastomas formed in 100% of the animals = by 14 d=20 postinfection. Surprisingly, the tumors were composed of a=20 heterogeneous population of cells, <20% of which expressed = the=20 retroviral reporter gene (GFP). The vast majority of both = GFP+ and=20 GFP=96 tumor cells expressed markers of glial progenitors. = Thus, the=20 tumors arose from the massive expansion of both infected and=20 uninfected glial progenitors, suggesting that PDGF was = driving tumor=20 formation via autocrine and paracrine stimulation of glial = progenitor=20 cells. To explore this possibility further, we coinjected a=20 retrovirus expressing PDGF-IRES-DsRed with a control = retrovirus=20 expressing only GFP. The resulting tumors contained a mixture = of red=20 cells (PDGF-expressing/tumor-initiating cells) and green = cells=20 (recruited progenitors). Both populations were highly = proliferative=20 and infiltrative. In contrast, when the control GFP = retrovirus was=20 injected alone, the animals never formed tumors and the = majority of=20 infected cells differentiated along the oligodendrocyte = lineage.=20 Together, these results reveal that adult white matter = progenitors=20 not only have the capacity to give rise to gliomas, but = resident=20 progenitors are recruited to proliferate within the mitogenic = environment of the tumor and in this way contribute = significantly to=20 the heterogeneous mass of cells that compose a malignant = glioma.=20

Key words: platelet-derived growth factor; tumor = environment;=20 oligodendrocyte; glioblastoma multiforme; animal model; tumor = clonality


3D"   =20 Introduction
Top
Abstract
3D"Introduction
Materials and Methods
Results
Discussion
References
 
I= dentifying=20 the cells that give rise to human gliomas has long been a = major goal=20 of neuro-oncology research. Recent studies showing that many = human=20 gliomas contain a subpopulation of cells with stem cell-like=20 properties have focused attention on the role of neural stem = cells or=20 progenitors as possible "cells of origin" (Reya et al., = 20013DGo;=20 Ignatova et al., 20023DGo;=20 Singh et al., 20033DGo).=20 This idea is supported by studies that have shown that = infecting=20 progenitor cells in the neonatal rodent brain with = PDGF-expressing=20 retroviruses induces the formation of tumors that closely = resemble=20 human gliomas (Uhrbom et al., 19983DGo;=20 Dai et al., 20013DGo;=20 Shih et al., 20043DGo).=20 However, no previous studies have tested the effects of = delivering a=20 PDGF-expressing retrovirus to the adult brain, and it is not = yet=20 known whether adult progenitors possess the proliferative and = self-renewing capacity to form tumors. This question is of = great=20 clinical relevance, because most malignant gliomas occur in=20 adults.

The subcortical white matter contains one of the largest = populations=20 of cycling cells in the adult brain. It is estimated that = glial=20 progenitors account for 3D~=202% of the adult = rat white=20 matter and as many as 4% of the adult human white matter = (Hommes and=20 Leblond, 19673DGo;=20 Roy et al., 19993DGo;=20 Gensert and Goldman, 20013DGo;=20 Dawson et al., 20033DGo).=20 Stereotactic injections of replication incompetent retrovirus = into=20 adult rat white matter selectively infects the resident = population of=20 cycling glial progenitors, most of which belong to the=20 oligodendrocyte lineage (Gensert and Goldman, 19963DGo).=20 We have chosen to test the effects of PDGF overexpression on = glial=20 progenitors in the adult white matter for several reasons. = (1)=20 Subcortical white matter is the most common location in which = malignant gliomas arise. (2) Many human gliomas express = markers also=20 expressed by glial progenitors, including olig2, NG2, and = PDGF=20 receptor 3D{alpha} (PDGFR3D{alpha}), suggesting a close relationship (Shoshan et = al., 19993DGo;=20 Chekenya and Pilkington, 20023DGo;=20 Bouvier et al., 20033DGo;=20 Ligon et al., 20043DGo).=20 (3) PDGF is a powerful mitogen for both glioma cells and = adult glial=20 progenitors (Wolswijk et al., 19913DGo;=20 Wolswijk and Noble, 19923DGo;=20 Vassbotn et al., 19943DGo). (4) Human gliomas often express both PDGF = and its=20 receptor, suggesting that autocrine and paracrine PDGF = signaling=20 plays a role in glioma growth and progression (Hermanson et = al.,=20 1992; Westermark et al., 19953DGo;=20 Di Rocco et al., 19983DGo).=20 This also raises the possibility that nontransformed glial=20 progenitors entrapped within an infiltrating glioma will be = driven to=20 proliferate by paracrine PDGF signaling.

In this study, we show that infecting adult white matter = progenitors=20 with a PDGF-B-expressing retrovirus induces rapid and = consistent=20 formation of tumors that closely resemble glioblastomas. The=20 retroviruses also express high levels of fluorescent reporter = genes, allowing us to identify infected cells and their = progeny.=20 This analysis has led to the finding that the majority of = cells=20 comprising the tumors are uninfected progenitor cells, = suggesting=20 that adult glial progenitors not only have the proliferative=20 and self-renewing capacity to form tumors when infected with=20 the PDGF retrovirus but also that uninfected progenitor cells = can be recruited and induced to proliferate via paracrine = signaling.=20


3D"   =20 Materials and Methods
Top
Abstract
Introduction
3D"Materials and Methods
Results
Discussion
References
 
<= FONT=20 size=3D-1>Retrovirus production and injections. A 0.8 kb = fragment=20 encoding PDGF-B-hemagglutinin (HA) (Shih et al., 20043DGo)=20 was ligated into the MCS1 region of the retroviral vectors=20 pQ-MCS1-IRES-eGFP (pQ-GFP) and pQ-MCS1-IRES-DsRed. = Replication-deficient=20 viruses with vsv-G coats were generated from these constructs = as well as from pNIT-GFP as described previously (Kakita and=20 Goldman, 19993DGo).=20 Viral titers were determined in colony-forming units (CFU) by = incubating C6 glioma cells with serial dilutions of = retrovirus in=20 10-fold steps. At 48 h postinfection the number of GFP+ (or = DsRed+)=20 cell clusters were counted. The CFU were calculated by = multiplying=20 the number of GFP+ (or DsRed+) cell clusters by the dilution = factor.=20 Viruses were injected into the forceps minor of the corpus = callosum=20 (stereotactic coordinates relative to bregma: 2 mm lateral, = 2.5 mm=20 rostral, 3.5 mm deep) of adult Sprague Dawley rats using a = Hamilton=20 syringe with a 33 gauge needle (5 =B5l at 0.2 =B5l/min) (see = Fig. = 1).=20 The injection of retrovirally infected progenitor cells (see=20 below) was performed using the same procedure including the=20 same volume, flow rate, and stereotactic coordinates. All = animal=20 work was performed according to Institutional Animal Care and = Use Committee (IACUC) guidelines of Columbia University. =

Survival study. Adult Sprague Dawley = rats were=20 injected with either pQ-GFP (six animals) or PDGF-IRES-GFP = (six=20 animals) as described above. Animals were monitored daily for = signs=20 of morbidity, such as weight loss, seizures, posturing, and = nasal=20 and/or periorbital hemorrhage. In addition to this survival = study, 26=20 animals were injected with the PDGF-IRES-GFP retrovirus and = followed=20 until they showed signs of tumor morbidity (between 14 and 20 = dpi). In accordance with IACUC guidelines, animals were = killed=20 at the first sign of morbidity. All animals were perfused = with=20 4% paraformaldehyde, and the brains were examined = histologically=20 for the presence of tumor.

Magnetic resonance imaging study. = Adult Sprague=20 Dawley rats were injected with PDGF-IRES-GFP (three animals) = as=20 described above. At 5, 10, and 17 d postinfection (dpi), the = animals=20 were anesthetized and 0.1 ml of gadolinium was given=20 intraperitoneally. Animals were immobilized in a Plexiglas = frame and=20 T1 and T2 images were collected from a magnetic resonance = imaging=20 (MRI) unit with a 1.5 T magnet.

Immunohistochemistry, microscopy, and cell=20 counting. Brains were fixed at 3, 14, or 17 dpi by cardiac = perfusion=20 with 4% paraformaldehyde. Hematoxylin and eosin stains were=20 performed on 5 =B5m sections from paraffin-embedded blocks.=20 Immunofluorescence analysis was performed on 10 =B5m = cryosections as=20 described previously (Marshall et al., 20053DGo)=20 using the following antibodies: anti-Olig2 (kind gift from = Dr. Thomas=20 Jessel, Columbia University, New York, NY), mouse anti-Nestin = (1:200;=20 Chemicon, Temecula, CA), rabbit anti-GFAP (1:500; = DakoCytomation,=20 Carpinteria, CA), mouse anti-NeuN (1:500; Chemicon), rabbit = anti-GFP=20 (1:500; Invitrogen, Eugene, OR), rabbit anti-DsRed (1:100; BD = Biosciences, Palo Alto, CA), mouse anti-HA (1:100; Covance, = Berkeley,=20 CA), rabbit anti-Ki67 (1:1000; NovoCastra, New Castle, UK), = goat=20 anti-PDGFR3D{alpha} (1:80; NeoMarkers, Fremont, CA), anti-NG2 (kind = gift from=20 Dr. William Stallcup, Burnham Institute, La Jolla, CA), mouse = anti-smooth muscle actin (SMA) (1:200; DakoCytomation), and = mouse=20 anti-CC1 (1:50; Calbiochem, La Jolla, CA). Stained sections = were=20 examined and photographed using a Zeiss (Oberkochen, Germany) = Axiophot 200 fluorescent microscope equipped with an Axiocam=20 (Zeiss) and OpenLab imaging software (Improvision, Lexington, = MA).=20 Multiple sections from three to six brains from each = condition were=20 photographed, and the number of cells staining positive for = each=20 marker was manually counted. Statistical analysis was = performed using=20 InStat, version 3.0, program.

Cell culture. White matter = progenitors were=20 isolated from adult Sprague Dawley rats as described = previously=20 (Gensert and Goldman, 20013DGo;=20 Mason and Goldman, 20023DGo).=20 The isolated cells were cultured in B104 conditioned media, = as=20 described previously (Canoll et al., 19963DGo) for 5 d, and then infected with either = pNIT-GFP or=20 PDGF-IRES-DsRed retrovirus. Cells were injected into animals = (10,000=20 cells in 5 =B5l) at 2 dpi, as described above. In some=20 experiments, infected cells were kept in culture with basal = media=20 (DMEM, N2, T3, 0.5% FBS, and=20 penicillin/streptomycin/amphotericin) for 10 dpi, and then = fixed in=20 4% paraformaldehyde, and analyzed by fluorescence = microscopy.=20

Flow cytometry and fluorescence activated cell=20 sorting. Tumors were dissected between 17 and 19 dpi and = cells were=20 isolated using the same method of isolation of adult white = matter=20 progenitors (Gensert and Goldman, 20013DGo;=20 Mason and Goldman, 20023DGo)=20 and then stained at 4=B0C with A2B5 antibody (American Type=20 Culture Collection, Manassas, VA) for 1 h followed by = anti-IgM Cy5=20 secondary for 30 min (Jackson ImmunoResearch, West Grove, = PA). Cells=20 were resuspended in PBS/10% FBS and FACS-sorted in a BD FACS=20 ARIA system (BD Bioscience). The FlowJo program was used for=20 flow cytometry data analysis. Cells isolated from the tumors=20 that formed from the coinjection of pNIT-GFP and = PDGF-IRES-DsRed=20 were sorted on the basis of endogenous GFP and DsRed.

ELISA. Confluent cultures of human = glioma cell=20 lines (U87, U251, U343, and U373) or primary cultures of = PDGF-induced=20 tumor cells (isolated from tumors at 17=9619 dpi as described = above)=20 were rinsed with PBS and incubated for 24 h with basal = medium.=20 Quantikine kits from R&D Systems (Minneapolis, MN) were = used for=20 quantification of PDGF-AA or PDGF-BB of cell culture media = according=20 to the manufacturer=92s instructions. Optical densities were=20 obtained with a Bio-Rad (Hercules, CA) 680 plate reader, and=20 concentrations were determined with the Microplate Manager, = version=20 II, program.


3D"   =20 Results
Top
Abstract
Introduction
Materials and Methods
3D"Results
Discussion
References
 
<= STRONG>PDGF-expressing=20 retrovirus introduced into adult white matter progenitors induces tumors = that=20 closely resemble human glioblastomas
To determine whether = PDGF=20 overexpression could induce adult white matter progenitors to = give=20 rise to gliomas, we generated a Moloney-based retrovirus that = coexpresses PDGF-B-HA and GFP, separated by an internal = ribosomal=20 entry site (PDGF-IRES-GFP). The infected cells expressed GFP = at high=20 levels, making it easy to identify the infected cells and = their=20 progeny in tissue sections by fluorescence microscopy and to = isolate=20 and analyze the cells by FACS. We stereotactically injected 5 = =B5l of=20 the retrovirus (at titers of 3D~=20105 = CFU/ml) into=20 the rostral subcortical white matter (the forceps minor of = the corpus=20 callosum) of adult rats (Fig. = 1).=20 Brain tumors developed in 100% of the animals injected (86 of = 86).=20 Survival studies showed that all animals developed signs of=20 tumor-induced morbidity between 14 and 20 dpi (Fig. = 2).=20 We were also able to follow tumor progression through MRI = scanning=20 at 5, 10, and 17 dpi. PDGF retrovirus-induced tumors were = visible=20 by 10 dpi with some edema around the small tumor. A large = tumor=20 was visible at 17 dpi with marked edema throughout the = ipsilateral=20 hemisphere and extending into the contralateral hemisphere (Fig. = 3).=20



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  		Figure 1. PDGF=20 overexpression induces the formation of malignant gliomas.=20 Hematoxylin and eosin stains were performed on coronal = sections of=20 adult rat brains 14 dpi with pQ-GFP (A,=20 C) or PDGF-IRES-GFP (B, = D,=20 E). A, No tumors formed in = brains=20 injected with the control retrovirus, but a small area of = reactive=20 gliosis is seen around the needle track (arrow). = C,=20 Higher-magnification micrograph showing the injection site.=20 B, Large infiltrative tumors with the = histological=20 features of human glioblastoma formed by 14 dpi with = PDGF-IRES-GFP=20 retrovirus. This section, 3 mm caudal from injection site, = shows the=20 tumor extending across the corpus callosum into the = contralateral=20 hemisphere. D, Higher-magnification micrograph = showing=20 an area of pseudopalisading necrosis (N), a hallmark of = glioma=20 malignancy seen in human glioblastomas. E, = Tumor cells=20 crossing the corpus callosum (CC) and infiltrating the = cortex (CX).=20 Scale bar: (in B) A, = B, 2=20 mm. Insets in A are coronal (a') = and=20 sagittal (a'') schematic diagrams of the = injection=20 site (arrow) at the level of the forceps minor corpus = callosum. The=20 dotted line in a'' shows the level of the = coronal=20 section shown in B. =

 = ;



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  		Figure 2. PDGF = retrovirus=20 causes rapid tumor formation and morbidity. Kaplan=96Meier = survival=20 curve showing the rapid onset of tumor-induced morbidity. = Equal=20 volumes and titers of pQ-GFP or PDGF-IRES-GFP were injected = into the=20 subcortical white matter of adult rats (6 rats in each = group). All=20 rats injected with the PDGF-IRES-GFP retrovirus showed signs = of=20 tumor-induced morbidity between 14 and 19 dpi. None of the = rats=20 injected with pQ-GFP showed any signs of tumor-induced = morbidity at=20 35 dpi. The graph shows the results of one representative=20 experiment. In total, we have performed survival studies on = 32 rats=20 injected with the PDGF-IRES-GFP retrovirus. All have = developed=20 tumor-induced morbidity between 14 and 20 dpi. In all cases, = the=20 presence of a malignant glioma was confirmed by histologic = analysis. =

 = ;



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  		Figure 3. = Serial MRI=20 studies show tumor progression. MRI scans were performed on = adult=20 rats at 5, 10, and, 17 dpi with the PDGF-IRES-GFP = retrovirus. Here,=20 we show a representative series of coronal images from one = animal.=20 No tumor is visible at 5 dpi. By 10 dpi, a small tumor is = seen at=20 the injection site on postcontrast T1 images. Tumor-induced = edema is=20 visible on T2 images. At 17 dpi, a large tumor is visible on = postcontrast T1 image, and edema involves the entire = hemisphere and=20 part of the contralateral hemisphere as seen in the T2 = image. gad,=20 Gadolinium. =

 = ;
The=20 tumors had all of the histologic features of glioblastoma = multiforme,=20 including marked vascular proliferation as seen with = immunostain for=20 SMA (Fig.=20 4C) and pseudopalisading necrosis (Figs.=20 1D, 4= B).=20 Fluorescence microscopy revealed that numerous GFP+ cells had = infiltrated the surrounding brain tissue and migrated across = the=20 corpus callosum into the contralateral hemisphere (Fig.=20 4A). In contrast, none of the animals injected with = pQ-GFP=20 or pNIT-GFP control retroviruses developed tumors (0 of 72) = (Figs.=20 1A,C, 7= A;=20 supplemental Fig. 2, available at http://www.jneurosci.org/ = as supplem= ental=20 material).



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  		Figure 4. GFP = expression=20 reveals the distribution of retrovirally infected cells.=20 Immunofluorescence analysis for GFP and SMA was performed on = sections of tumor at 17 dpi with the PDGF-IRES-GFP = retrovirus.=20 A, Micrograph shows GFP+ cells (green) = crossing the=20 corpus callosum (CC) and invading the contralateral = hemisphere.=20 Hoechst stain (blue) shows increased cellular density in and = around=20 the main tumor mass. Note that only a subset of the cells is = GFP+.=20 B, GFP+ and GFP=96 cells are seen intermingled = throughout the tumor, including in areas of pseudopalisading = necrosis (N). C, SMA immunofluorescence (red) = shows=20 marked vascular proliferation with recruitment of = perivascular=20 smooth muscle cells. Scale bar, 1 mm. =

 = ;
Retroviral=20 delivery to the adult white matter predominantly infects NG2+/GFAP=96 = glial=20 progenitors
To characterize the identity of the cells that = were=20 infected with the retroviruses and gave rise to the = PDGF-driven=20 tumors, we performed immunohistochemistry of = retrovirus-infected=20 cells at 3 dpi with PDGF-IRES-GFP or a control retrovirus = that=20 expresses only GFP (pNIT-GFP). At 3 dpi, the infected (GFP+)=20 populations were very similar in number, distribution, = morphology,=20 and immunophenotype for each retrovirus. Our results showed = that the=20 large majority of GFP+ cells were NG2+ (87.8 =B1 1.7% for = pNIT-GFP=20 and 91.0 =B1 1.1% for PDGF-IRES-GFP) (supplemental Fig.=20 1A,C, available at http://www.jneurosci.org/ as supplem= ental=20 material). Less than 1% of the GFP+ cells expressed = detectable=20 levels of GFAP (0.5 =B1 0.2% for pNIT-GFP and 0.4 =B1 0.4% = for=20 PDGF-IRES-GFP) (supplemental Fig. 1B,D, = available at http://www.jneurosci.org/ as supplem= ental=20 material). The majority of cells was located in the = subcortical=20 white matter near the injection site. A few GFP+ cells were = also seen=20 in the overlying cortex adjacent to the injection tract and=20 these cells were also NG2+/GFAP=96. These results are = consistent=20 with previous BrdU-labeling studies showing that NG2+ glial=20 progenitors account for the majority of cycling cells in the=20 adult white matter and cortex (Dawson et al., 20033DGo).=20 By 14 dpi with the control retroviruses, the majority of the = GFP+=20 cells had begun to differentiate along the oligodendrocyte=20 lineage: 83.9 =B1 2.5% expressed the oligodendrocyte marker = CC1,=20 and 1.3 =B1 0.7% expressed GFAP (supplemental Fig. 2, = available=20 at http://www.jneurosci.org/ as supplem= ental=20 material). It is important to note that, although a = gliotic=20 response formed around the injection site, this did not lead = to the=20 retroviral labeling of reactive astrocytes. Previous studies = have=20 shown that there is a 3=964 d delay before GFAP+ astrocytes = proliferate=20 in response to a cortical stab wound (Latov et al., 19793DGo;=20 Takamiya et al., 19883DGo).=20 The retroviral titers decay rapidly and in vitro = analysis has=20 shown that these retroviruses have a half-life of 3D~=204 h at 37=B0C = (Gensert and=20 Goldman, 19973DGo).=20 Therefore, the effective viral titer has dropped to near zero = by the=20 time astrocytes begin to proliferate. These results are = consistent=20 with previous results showing that retroviral injection into = the=20 adult white matter predominantly labels glial progenitors, = most of=20 which belong to the oligodendrocyte lineage and does not = label=20 GFAP+ astrocytes (Gensert and Goldman, 19963DGo).

The tumors are composed of infected and uninfected glial=20 progenitors
To characterize the cellular composition of the = tumors,=20 we performed immunohistochemical analysis for retrovirally = encoded=20 genes, GFP and PDGF-HA, and a variety of neuronal and glial=20 markers. Remarkably, only a subset of the cells in the tumor=20 expressed detectable levels of GFP or PDGF-HA, suggesting = that=20 tumors were composed of both infected and uninfected cells = (Figs. = 5,=20 6). = Furthermore, double immunofluorescence showed that GFP and HA = colocalized to the same subset of cells (Fig.=20 5A), confirming that GFP is a reliable marker for = cells=20 that are expressing both retroviral genes. The vast majority = of tumor=20 cells expressed markers seen in immature glia, including = olig2, NG2,=20 nestin, and PDGFR3D{alpha}=20, and=20 double-immunofluorescence analysis showed that the majority = of both=20 GFP+ and GFP=96 cells were positive for these markers (Figs. = 5, 6). = Numerous=20 GFAP+/GFP=96 reactive astrocytes were scattered throughout = the tumor=20 (Fig.=20 5E=96E''), but <3% of the GFP+ cells were GFAP+ = (data not=20 shown). None of the GFP+ cells expressed the neuronal marker, = NeuN=20 (data not shown).



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  		Figure 5. GFP+ = and GFP=96=20 tumor cells express markers of glial progenitor cells.=20 Double-immunofluorescence analysis of tumors, 17 dpi with = the=20 PDGF-IRES-GFP retrovirus, shows that GFP (green) is = expressed in=20 only a subset of tumor cells. A=96A'', PDGF-HA = expression (red) is seen in the same subset of cells that = express=20 GFP. However, PDGFR3D{alpha}=20=20 (B', B''), NG2 = (C',=20 C''), and nestin (D',=20 D''), each stained red, are expressed in the = vast=20 majority of GFP+ and GFP=96 tumor cells. E',=20 E'', GFAP+/GFP=96 reactive astrocytes (red) = are seen=20 scattered throughout the tumor. Rare GFAP+/GFP+ cells were = seen=20 (<3%). =

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  		Figure 6. = Quantitative=20 analysis shows that the majority of cells in the PDGF-driven = tumors=20 are GFP=96 progenitor cells. A, Double=20 immunofluorescence of tumors, 17 dpi with PDGF-IRES-GFP, = shows that=20 the vast majority of tumor cells are olig2+ (red), but only = a subset=20 of olig2+ cells are GFP+ (green). B, Double=20 immunofluorescence for GFP (green) and Ki67 (red), a marker = for=20 cycling cells, shows that tumor cells are highly = proliferative, but=20 only a subset of Ki67+ cells are GFP+. The triptych at the = bottom of=20 A and B show green, red, and = green/red=20 overlay. C, Bar graph showing the number of = total=20 cells (identified by Hoechst nuclear staining), olig2+, = GFP+, and=20 GFP+/olig2+, cells per high-powered field. D, = Bar=20 graph showing the number of total, Ki67+, GFP+, and = GFP+/Ki67+ cells=20 per high-powered field. Data represent the mean =B1 SEM of = multiple=20 high-powered fields from three separate brain tumors.=20 E, F, Cells isolated from = freshly=20 dissected tumors (17 dpi with PDGF-IRES-GFP retrovirus) were = immunostained with the progenitor cell marker A2B5 and = analyzed by=20 flow cytometry. E, Scatter plot from a single=20 representative experiment showing the relative abundance of = four=20 populations of tumor cells: GFP+/A2B5+, GFP=96/A2B5+, = GFP+/A2B5=96, and=20 GFP/A2B5=96. F, Bar graph showing the = percentage of=20 cells in each population. Data represent the mean =B1 SEM = from three=20 separate experiments. =

 = ;
To=20 determine the relative abundance and proliferation rates of = GFP+ and=20 GFP=96 cells in the tumors, we performed quantitative = analysis after=20 double immunofluorescence for GFP and olig2, or GFP and the=20 proliferation marker Ki67 on sections of tumors 17 dpi with = the=20 PDGF-IRES-GFP retrovirus, and counted the number of cells = positive=20 for each marker. The majority of cells in the tumor (77.6 =B1 = 4.6%)=20 expressed olig2. However, only 16.8 =B1 2.0% of the total = cells=20 expressed GFP, almost all of which (96.5 =B1 0.5%) were = olig2+ (Fig.=20 6A,C). Similarly, a large fraction of tumor = cells=20 (34.0 =B1 4.1%) were Ki67+, but only 18.2 =B1 1.8% of the = Ki67+ cells=20 were GFP+ (Fig.=20 6B,D). Thus, the majority of olig2+ and = Ki67+ cells=20 in the tumor were GFP negative, suggesting that most of the=20 proliferating cells in the tumor were uninfected glial = progenitors=20 that had been recruited via paracrine signaling.

As a second approach to quantitate the relative abundance of = the=20 different cell types in the tumor, we performed flow cytometry = on=20 cells isolated from tumors (17=9619 dpi) using the A2B5 = monoclonal=20 antibody, which is a well established marker for a subset of=20 progenitor cells found in the adult white matter (Mason and = Goldman,=20 2002; Nunes et al., 20033DGo).=20 Flow cytometry showed that 13.2 =B1 1.1% of the isolated = cells=20 expressed GFP (consistent with our immunohistochemical = analysis=20 noted above). Of the total cells, 51.2 =B1 1.9% were A2B5+, = but=20 only 9.0 =B1 2.6% were A2B5+/GFP+ (Fig.=20 6E,F). From a typical experiment, we = isolated 5 x 106 A2B5+ cells per tumor (of = which the majority was GFP=96). In contrast, using the same=20 procedure, we typically isolate on the order of = 104 A2B5+=20 cells from a normal adult brain (data not shown). Thus, the = PDGF=20 retrovirus caused a massive expansion of the A2B5+ = population, most=20 of which are GFP=96 (uninfected) cells.

Adult white matter progenitors infected with the = PDGF-expressing=20 retrovirus in vivo and in vitro are able to recruit and = activate=20 normal progenitors
To test the hypothesis that = PDGF-expressing cells=20 were recruiting other glial progenitors to proliferate, we = coinjected=20 a GFP-expressing retrovirus (pNIT-GFP) and the = PDGF-IRES-DsRed=20 retrovirus into adult white matter. All of the rats = coinjected with=20 pNIT-GFP and PDGF-IRES-DsRed retrovirus formed malignant = gliomas by=20 17 dpi (35 of 35) (Fig.=20 7B). Immunofluorescence analysis and FACS for GFP = and=20 DsRed, showed that tumors were composed of four populations = of cells:=20 GFP=96/DsRed=96, GFP+/DsRed=96, GFP=96/DsRed+, and a small = subpopulation was=20 GFP+/DsRed+, demonstrating that coinfection was a rare event = (Fig.=20 7D). The red cells = (PDGF-expressing/tumor-initiating=20 cells) and green cells (recruited progenitors) were = intermingled=20 within the tumor, including regions of pseudopalisading = necrosis (Fig.=20 7C). The large majority of both populations had an = immature morphology and were nestin+, NG2+, and olig2+ (data = not=20 shown). Both red and green cells diffusely infiltrated the = brain,=20 crossing the corpus callosum and invading the contralateral=20 hemisphere (Fig.=20 7B).



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  		Figure 7. = Coinjection=20 studies show that PDGF-expressing cells recruit other = progenitors to=20 proliferate within the tumor. A, Low-power = micrograph=20 showing a small collection of GFP+ cells (green) along the = injection=20 track (white arrow) at 17 dpi with pNIT-GFP. = B, In=20 contrast, a large tumor containing many GFP+ cells (green) = and=20 DsRed+ cells (red) are seen at 17 dpi with pNIT-GFP and=20 PDGF-IRES-DsRed. Note that this section, 3 mm caudal of the=20 injection site, shows that many of the GFP+ cells are seen = crossing=20 the corpus callosum (CC) into the contralateral hemisphere. = Scale=20 bars: A, B, 2 mm: = A,=20 B, C, Higher-magnification = micrograph=20 showing red and green cells intermingled throughout the = tumor,=20 including areas of pseudopalisading necrosis (N). = D,=20 Cells isolated from tumors (17=9619 dpi with pNIT-GFP and=20 PDGF-IRES-DsRed) were FACS-sorted into four populations:=20 GFP=96/DsRed=96, GFP+/DsRed=96, DsRed+/GFP=96, and = DsRed+/GFP+. Totals of=20 1.5 x 106 = GFP+/DsRed=96=20 cells and 2.8 x = 105=20 DsRed+/GFP=96 cells were sorted from two pooled tumors in = this=20 representative experiment (the experiment was repeated five = times).=20 E=96E'', In brains injected with pNIT-GFP (17 = dpi), the=20 GFP+ cells have a highly branched morphology and are = negative for=20 the proliferation marker Ki67 (labeling index of <5%).=20 F=96F'', In tumors generated by coinfection of = pNIT-GFP=20 and PDGF-IRES-DsRed, the GFP+ cells have an immature = morphology and=20 many are Ki67+ (labeling index of 26%). Insets in = A=20 and B are schematic diagrams illustrating the=20 distribution and number of cells (green and red dots) in = coronal=20 sections of adult brain at their corresponding levels. =

 = ;
To=20 further characterize the normal fate and behavior of the = retrovirally=20 infected cells, we injected the pNIT-GFP virus alone, in the = same=20 volume and at the same titer (106 CFU/ml). As = anticipated,=20 none of these animals developed tumors (0 of 19). By 17 dpi, = the=20 pNIT-GFP cells remained few (3D~=20100 cells = per=20 brain) and all of the pNIT-GFP cells were distributed within=20 500 =B5m of the injection track (Fig.=20 7A). The majority of pNIT-GFP cells had acquired a = complex=20 morphology with numerous thin processes characteristic of=20 differentiating oligodendrocytes (Fig.=20 7E=96E''). Immunofluorescence analysis showed that = 77% of=20 the pNIT-GFP+ cells stained positive for the oligodendrocyte = marker=20 CC1 (61 of 79 cells counted), and <1% of the pNIT-GFP = cells=20 expressed GFAP (consistent with the result of our analysis at = 14 dpi=20 described above). Less than 1% of the pNIT-GFP cells were = Ki67+ in=20 brains injected with the control virus (Fig.=20 7E=96E''). In contrast, 26% of the = pNIT-GFP/DsRed=96 cells=20 were Ki67+ in tumors formed by the coinjection of the two = viruses (Fig.=20 7F=96F''). Together, these results show that, in = normal=20 adult white matter, the majority of retrovirally infected = cells=20 differentiated along the oligodendrocyte lineage. However, = within the=20 mitogenic rich environment of the tumor, the = pNIT-GFP-infected cells=20 were induced to proliferate and migrate, and thus mimic the = behavior=20 of malignant glioma cells.

As another test of the recruitment hypothesis, and to eliminate=20 the possibility of any cells being coinfected with both = viruses,=20 we isolated progenitors from adult white matter, expanded = them=20 in vitro for 5 d in B104 conditioned media, and infected = them=20 in vitro. Equal numbers of progenitors were separately=20 infected with either pNIT-GFP or PDGF-IRES-DsRed. In some=20 experiments, the infected progenitor cells were grown in = culture with=20 basal media for 10 dpi (Fig.=20 8A,B). Cells infected with pNIT-GFP stopped=20 proliferating and acquired a highly branched morphology=20 characteristic of maturing oligodendrocytes (Fig.=20 8A). In contrast, cells infected with = PDGF-IRES-DsRed=20 remained immature and highly proliferative, forming large = clusters of=20 cells by 10 dpi (Fig.=20 8B). In a separate set of experiments, the in=20 vitro-infected progenitor cells were transplanted into = adult rat=20 subcortical white matter (same coordinates as shown in Fig. = 1) 2 d=20 after infection, by either coinjecting pNIT-GFP and = PDGF-IRES-DsRed=20 cells (2000 of each cell type), or pNIT-GFP+ cells alone = (2000).=20 Coinjected cells formed infiltrative tumors by 20 dpi (Fig.=20 8D). The tumors were composed of three = populations:=20 DsRed+/GFP=96, DsRed=96/GFP+, and DsRed=96/GFP=96. No = double-positive cells=20 were observed. Both the PDGF-IRES-DsRed and pNIT-GFP cells in = the=20 coinjections remained immature, infiltrative, and highly=20 proliferative (Fig.=20 8D). In contrast, pNIT-GFP cells injected alone = remained=20 few, had branched morphologies characteristic of = differentiating=20 glia, and did not develop into tumors (Fig.=20 8C). All of the pNIT-GFP cells were located within = a few=20 hundred micrometers of the injection site, demonstrating = that,=20 when injected alone, the transplanted adult progenitors do = not=20 accumulate or migrate far from the injection site (Fig.=20 8C).



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  		Figure 8. Adult = white=20 matter progenitors infected with PDGF-IRES-DsRed in = vitro=20 form malignant gliomas through autocrine and paracrine = signaling.=20 Normal adult white matter progenitors were isolated and = expanded for=20 5 d in vitro with B104-containing media. = Proliferating=20 progenitors were then infected with pNIT-GFP or = PDGF-IRES-DsRed.=20 A, B, Adult white matter = progenitors=20 grown in culture for 10 dpi in basal media. A, = pNIT-GFP-infected cells stopped proliferating and acquired a = highly=20 branched morphology. B, = PDGF-IRES-DsRed-infected cells=20 retained a simple morphology and remained highly = proliferative,=20 forming large clusters of cells. C, = D,=20 At 2 d postinfection, the cells were implanted into adult = brains.=20 Brains were analyzed at 20 dpi. C, = pNIT-GFP-infected=20 cells injected alone remained close to the injection site = and no=20 tumors formed. D, When pNIT-GFP-infected cells = were=20 coinjected with PDGF-IRES-DsRed-infected cells, a tumor = formed by 20=20 dpi, and this tumor is composed of a mixture of red and = green cells.=20 Equal numbers of pNIT-GFP cells were injected in = C and=20 D. Note the marked proliferation and = dispersion of=20 pNIT-GFP cells in the presence of PDGF-IRES-DsRed cells=20 (D). Insets in C and = D are=20 schematic diagrams illustrating the distribution and number = of cells=20 (green and red dots) in coronal sections of adult brain at = the level=20 of the injection site. =

 = ;
Secretion=20 of PDGF-A and PDGF-B into glioma cell conditioned medium
The = above=20 results strongly suggest that retrovirus-infected cells = secrete PDGF=20 at high enough levels to drive proliferation of progenitors = cells in=20 their local environment. How do the levels of PDGF produced = and=20 secreted by the retrovirus-induced tumor cells compare with = those=20 produced by human glioma cells? Do human glioma cells secrete = high=20 enough levels of PDGF to drive glial progenitor = proliferation? To=20 address these questions, we collected conditioned medium from = confluent cultures of the retrovirus-induced tumors (17 dpi) = and=20 several human glioma cell lines (U87, U251, U343, and U373). = The=20 levels of PDGF-A and PDGF-B in conditioned medium were = measured by=20 ELISA (Table=20 1). These results show that the primary cultures from the = retrovirus-induced tumors secreted detectable levels of both = PDGF-A=20 (1.4 =B1 0.1 ng/ml) and high levels of PDGF-B (29.2 =B1 7.4 = ng/ml).=20 The human glioma cell lines we tested did not secrete = detectable=20 levels of PDGF-B but did secrete significant levels of = PDGF-A.=20 In fact, one of the cell lines (U343) expressed very high = levels=20 of PDGF-A (52.1 =B1 1.6 ng/ml). Previous studies have shown = that=20 many glioma cells express PDGF-A, PDGF-B, or both, but most = glioma=20 cell lines secrete predominantly PDGF-A (Betsholtz et al., = 19863DGo;=20 Nister et al., 19863DGo,=20 1991). The levels of PDGF measured in these = previous studies=20 (20=9630 ng/ml) are very similar to our results. It is = important to=20 note that these levels are well within the range of PDGF=20 concentrations needed to drive glial progenitor = proliferation. In=20 vitro studies have shown that both PDGF-A and PDGF-B are = potent=20 mitogens for glial progenitors with PDGF-A having = half-maximal=20 effects at 2 ng/ml and PDGF-B having half-maximal effects at = 10 ng/ml=20 (Pringle et al., 19893DGo). Together, these results suggest that some = human=20 glioma cells produce high enough levels of PDGF to drive the=20 proliferation of glial progenitors that are in and around the = tumor.


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  		Table = 1.=20 Secretion of PDGF-A and PDGF-B into culture medium =

 

3D"   =20 Discussion
Top
Abstract
Introduction
Materials and Methods
Results
3D"Discussion
References
 
W= e have=20 shown that infecting glial progenitors in adult rat white = matter with=20 a retrovirus that expresses high levels of PDGF-B induces the = rapid=20 and consistent formation of tumors that closely resemble = human=20 glioblastomas. The tumors are composed of infected and = uninfected=20 progenitors, suggesting that tumor formation was being driven = by both=20 autocrine and paracrine growth factor stimulation. This = conclusion=20 was further supported by our coinjection experiments that = allowed us=20 to compare the behavior (and fate) of GFP-tagged glial = progenitors=20 (and their progeny) in the presence or absence of=20 PDGF-IRES-DsRed-expressing cells. When the GFP-expressing = virus was=20 injected alone, tumors never formed and the majority of = GFP-tagged=20 cells differentiated along the oligodendrocyte lineage by 14 = dpi. In=20 contrast, when both viruses were injected, the animals formed = tumors=20 that were composed of a mixture of red and green cells. Both=20 populations were highly proliferative and migratory. We also = infected=20 adult glial progenitors in vitro with one or the other = of=20 these retroviruses and then injected the pNIT-GFP cells alone = or with=20 PDGF-IRES-DsRed cells. This approach again showed that the = GFP-tagged=20 progenitors were recruited to proliferate within the tumor = and=20 effectively ruled out the possibility that any of the green = cells had=20 been infected with the PDGF-expressing retrovirus. =

The tumor formation seen with this model was remarkably rapid = and=20 consistent. Although this makes the model very convenient and = attractive for the use of controlled studies, such as preclinical=20 trials for new therapies, one may question the degree to = which=20 this resembles the growth of human tumors. It is not known = how=20 long human gliomas have been growing before they manifest = clinically;=20 however, by the time of diagnosis, human glioblastomas can = grow=20 remarkably fast. Serial imagining studies have shown that = humans=20 gliomas can grow with volume doubling times as short as 1 = week=20 (Blankenberg et al., 19953DGo;=20 Swanson et al., 20033DGo).=20 Furthermore, human glioblastomas often have Ki67 labeling = indices as=20 high or higher than that seen in the PDGF glioma model. The=20 major difference in survival times (<3 weeks for the rats = vs=20 <1 year for treated human glioblastomas) has more to do = with=20 the size of the skull and the room available to accommodate = the=20 tumor than it has to do with growth rate of the tumor per se. =

Adult glial progenitors as a cell of origin for malignant=20 gliomas
Several different types of brain cells have been=20 implicated in the formation of human gliomas including mature = astrocytes, progenitors, and neural stem cells (Bachoo et = al., 20023DGo;=20 Singh et al., 20043DGo;=20 Fomchenko and Holland, 20053DGo;=20 Sanai et al., 20053DGo). Using the RCAS system, Holland and = coworkers have=20 shown that, in the neonatal brain, GFAP+ astrocytes and = nestin+=20 progenitors have the capacity to form tumors when infected = with a=20 PDGFB-HA-expressing retrovirus (Dai et al., 20013DGo).=20 In these studies, the frequency of tumor formation and the = histologic=20 features of the tumor (grade and tumor type) depended on the = genetic=20 background of the mice used and the population of cells = targeted by=20 the retrovirus (Uhrbom et al., 19983DGo;=20 Dai et al., 20013DGo;=20 Hesselager et al., 20033DGo). Shih et al. (2004)3DGo=20 compared the tumorigenic effects of retroviruses expressing = different=20 levels of PDGFB-HA. The PDGFB-HA retrovirus expressing higher = levels=20 of growth factor caused more malignant tumors with a shorter = latency=20 and in a higher percentage of animals.

In this study, we used the same high expressing PDGFB-HA = construct=20 (cloned into an amphoteric retrovirus) to selectively infect=20 progenitors in the adult brain. Our results show that adult=20 white matter progenitors have a remarkable capacity to form=20 tumors. This is perhaps surprising considering that these = cells=20 are normally nonmigratory and slowly proliferating (Noble et=20 al., 19923DGo;=20 Wren et al., 19923DGo;=20 Gensert and Goldman, 19963DGo;=20 Roy et al., 19993DGo;=20 Dawson et al., 20033DGo).=20 In contrast, neonatal progenitors are highly migratory and=20 proliferative and resemble glioma cells in their capacity to=20 infiltrate long distances through brain tissue (Goldman et = al.,=20 1997; Kakita and Goldman, 19993DGo;=20 Suzuki and Goldman, 20033DGo).=20 However, in vitro studies have shown that adult glial=20 progenitors can be induced to become more migratory and = proliferative=20 when treated with growth factors including PDGF, basic = fibroblast=20 growth factor, and glial growth factor (Wolswijk et al., = 19913DGo;=20 Wolswijk and Noble, 19923DGo;=20 Shi et al., 19983DGo).=20 Our results provide the first in vivo evidence that PDGF = can=20 drive adult glial progenitors to acquire a more migratory and = proliferative behavior. When this behavior proceeds unchecked, = it=20 leads to the formation of malignant tumors with all of the = histologic=20 features of glioblastoma. In particular, the patterns of = infiltration=20 seen in this model, with a predilection for fiber tracks, = closely=20 resemble those seen in human gliomas. This also resembles the = migration of glial progenitors that occurs during brain = development=20 (Kakita and Goldman, 19993DGo;=20 Kakita et al., 20033DGo;=20 Suzuki and Goldman, 20033DGo),=20 suggesting that the infiltrative capacity of glioma cells may = be a=20 re-expression of a neonatal glial progenitor phenotype. =

Constitutive overexpression of PDGF is sufficient to drive = malignant=20 behavior
Molecular and cytogenetic analysis has shown that = human=20 glioblastomas often contain multiple genetic lesions (Ohgaki = and=20 Kleihues, 20053DGo).=20 These findings have led to the assumption that malignant = behavior=20 requires the accumulation of multiple genetic lesions. = However,=20 studies using molecular and cytogenetic analysis of human = tumors are=20 correlative in nature, and it is not known how many genetic = lesions=20 are actually required for malignant transformation. One very=20 important implication to our study is that the histologic = hallmarks=20 of malignancy can be driven by overexpression of a single = growth=20 factor. By 14 dpi, 100% of the PDGF-driven tumors have shown = features=20 of glioblastoma, including marked vascular proliferation and=20 pseudopalisading necrosis. Furthermore, the areas of = pseudopalisading=20 necrosis contained a mixture of infected and uninfected = cells,=20 suggesting that even genetically normal cells can be driven = to=20 display a malignant phenotype. These findings point to an=20 important concept; the progression to malignancy does not = merely=20 reflect the autonomous misbehavior of genetically deranged = cells,=20 but rather results from complex interactions between tumor = cells=20 and their environment. We propose that PDGF-expressing = retroviruses=20 drive the formation of malignant tumors so quickly because = they=20 not only transform the infected cells but they also transform = the environment.

Although PDGF overexpression clearly initiates tumor formation = in=20 this model, it is possible that other genetic or epigenetic=20 alterations are at play. Other growth factors may be = upregulated=20 (either by the progenitor cells or by other cells within the=20 tumor) and cooperate with PDGF in the paracrine signaling, as = has been shown to occur with neonatal and adult glial = progenitors=20 in vitro (Bogler et al., 19903DGo;=20 Wolswijk and Noble, 19923DGo;=20 Shi et al., 19983DGo).=20 It is also possible that the retrovirally infected cells = acquire=20 additional genetic lesions as a result of insertional = mutagenesis.=20 Previous studies using a Moloney-based retrovirus expressing = PDGF=20 have shown evidence that insertional mutagenesis may = cooperate with=20 PDGF in the development of gliomas (Johansson et al., 20043DGo,=20 2005). However, the speed and consistency with = which tumors=20 formed in our model (100% of animals formed tumors by 14 = dpi), and=20 the fact that tumors never formed when animals were injected = with=20 control virus alone, make it unlikely that insertional = mutagenesis=20 and subsequent clonal selection played an essential role in = tumor=20 formation. Rather, the tumors are primarily driven by = autocrine and=20 paracrine growth factor stimulation. Still, the acquisition = of=20 additional genetic lesions (either by insertion mutagenesis = or some=20 other mechanism) may confer a selective advantage to a = subpopulation=20 of tumor cells (infected or recruited).

Recruitment as a mechanism of glioma growth
Human = gliomas=20 are composed of a heterogeneous mass of tumor cells = intermingled with=20 entrapped and reactive brain cells. Several studies have = provided=20 evidence that paracrine growth factor signaling and/or = recruitment=20 may play an important role in the rapid growth and = progression of=20 gliomas (Hermanson et al., 19923DGo;=20 Westermark et al., 19953DGo;=20 van der Valk et al., 19973DGo; Shih and Holland, 20053DGo).=20 Vascular proliferation, which is one of the most reliable = markers of=20 malignant progression of gliomas, is well known to involve=20 recruitment of endothelial cells, pericytes, and smooth = muscle cells.=20 This recruitment is mediated, in part, by paracrine signaling = of=20 multiple growth factors, including PDGF (Dunn et al., 20003DGo;=20 Guo et al., 20033DGo;=20 Lamszus et al., 20043DGo). Recent studies have shown that gliomas can = also=20 recruit GFAP+ astrocytes and nestin+ neural stem = cells/progenitors=20 from the subventricular zone (Duntsch et al., 20053DGo;=20 Glass et al., 20053DGo). In this study, we show, for the first time, = that=20 local recruitment of glial progenitors that reside in the = adult white=20 matter can contribute significantly to the proliferative = mass.=20 Immunohistochemical analysis showed that the majority of the=20 recruited cells were of the oligodendroglial lineage = (PDGFR+, NG2+,=20 olig2+, GFAP=96) (Figs. = 5, 6). = PDGFR+/NG2+ glial=20 progenitors are abundant and widely distributed in the = subcortical=20 white matter of adult humans and rodents (Gogate et al., = 19943DGo;=20 Nishiyama et al., 19963DGo; Scolding et al., 19983DGo;=20 Dawson et al., 20003DGo).=20 As with our model, human gliomas grow by diffusely = infiltrating the=20 brain and the tumor cells intermingle with normal brain = cells. As a=20 result, glial progenitors entrapped within the infiltrative=20 margins of the tumor are subject to paracrine growth factor=20 stimulation. Therefore, in the case of human tumors that = express a=20 high level of PDGF, one would expect recruited glial = progenitors to=20 contribute to tumor growth regardless of the cell of origin. = However,=20 because of the remarkable phenotypic similarities between = glial=20 progenitors and glioma cells, it may be difficult to = distinguish a=20 recruited glial progenitor from a "true glioma cell." =

How can we define the difference between a true glioma cell = and a=20 recruited glial progenitor? In human gliomas, the "true tumor = cells"=20 most likely arise from the clonal expansion of a genetically=20 transformed cell of origin. In principle, these cells would = have the=20 capacity to initiate the formation of new tumors if they were = isolated and reinjected into the brain of a nude rodent. The=20 recruited glial progenitors would be proliferating cells that = are not=20 clonally derived from the genetically transformed cell of = origin.=20 Some of the recruited cells may acquire genetic lesions as a = result=20 of their abnormally driven proliferation, but many may be = genetically=20 normal and their proliferation would be dependent on the = mitogenic=20 environment of the tumor. Such cells would not have the = capacity to=20 form new tumors if isolated and transplanted into a nude = rodent.=20

An exceptional advantage of the PDGF tumor model is that the=20 retroviral reporters enable us to identify the infected cells = and their progeny, and thus to distinguish the cells that = arise=20 from clonal expansion of the "tumor-initiating cells" from = cells=20 that have been recruited. We must consider the possibility = that=20 transcriptional silencing of retroviral genes could cause us = to=20 underestimate the number of infected cells. The established=20 mechanisms of retroviral silencing (including methylation of=20 promotor regions and histone acetylation) inhibit gene = expression=20 at the level of transcription. The retrovirus encodes PDGF = and=20 GFP on a single bicistronic message; therefore, = transcriptional=20 silencing, if it were to occur, should block expression of = both=20 viral genes. The immunohistochemical data showing a = colocalization=20 of PDGF-HA and GFP (Fig.=20 5A=96A'') provides direct evidence that GFP is a = reliable=20 reporter of retroviral gene expression. Our coinjection = studies also=20 address this issue, allowing us to tag a separate population = of=20 progenitors with a control retrovirus and track the fate of = their=20 progeny. The massive expansion of progenitors infected with = the=20 pNIT-GFP control retrovirus (which were never infected with=20 PDGF-expressing virus) cannot be explained by retroviral = silencing=20 and provides direct evidence of the recruitment = phenomenon.=20

Progenitor recruitment is not inconsistent with the clonal = character=20 of human gliomas
Clonal analysis has shown that most (but = not all)=20 human gliomas contain a prominent (or at least detectable) = clonal=20 population (Berkman et al., 19923DGo;=20 Morse et al., 19943DGo;=20 Kattar et al., 19973DGo). These studies have used X chromosome = inactivation=20 analyses, such as the HUMARA assay, which can detect a clonal = population even if it accounts for only 15% of the total = cells in a=20 polyclonal background (Willman, 19943DGo).=20 Thus, a clonal character does not indicate that human gliomas = are=20 solid tumors composed exclusively of a monoclonal population = of=20 cells. They are not. Because of their highly infiltrative = nature,=20 glioma cells intermingle with the surrounding brain tissue. = As a=20 result, human gliomas are composed of a mixture of neoplastic = and=20 non-neoplastic cells, including entrapped neurons, = astrocytes,=20 microglia, blood vessels, and progenitor cells. The relative=20 abundance of these different cell types varies from one = region to the=20 next. In some regions, the tumor may be composed = predominantly of a=20 clonal population of glioma cells, but in the more = infiltrative=20 regions (which, for many gliomas accounts for the vast = majority of=20 the tumor) the true tumor cells are a minority. Not = surprisingly,=20 clonal analysis of these more infiltrative regions of gliomas = has=20 shown that the clonal population accounts for a minority of = the=20 total cells, and in some cases the clonal population cannot = be=20 detected (Kattar et al., 19973DGo).=20 It is within these regions in which glioma cells are = intermingling=20 with and interacting with non-neoplastic cells, including = many=20 PDGF-responsive glial progenitors, that recruitment is likely = to be=20 most robust. Thus, the finding that most human gliomas = contain a=20 clonal population is not inconsistent with the possibility = that=20 extensive recruitment of glial progenitors is occurring = (supplemental=20 Fig. 3, available at http://www.jneurosci.org/ = as supplem= ental=20 material).

These findings shed new light on the relationship between = gliomas=20 and adult glial progenitors and suggest a novel mechanism of=20 tumor growth by progenitor recruitment, which may play a role = in the rapid progression and heterogeneity that characterize=20 malignant gliomas. In addition to the novel implications = regarding=20 the relationship between progenitors and glioma cells, the = utility=20 of this model as a tool for additional studies of gliomas = should=20 not be understated. The ability to generate tumors that = closely=20 resemble human glioblastomas in a rapid and highly = reproducible=20 manner makes this an ideal model for preclinical animal = studies=20 involving new treatment strategies.


3D"   =20 Footnotes
 
Received Feb. = 3, 2006;=20 revised May 8, 2006; accepted May 13, 2006.

This work was supported by = National=20 Institutes of Health Grants NS17125 and NS045070. We thank = Drs. Eric=20 Holland, Alan Shih, and Chankai Dai for their PDGF-HA = construct; Drs.=20 Phyllis Faust and Carol Mason for sharing their microscope = systems=20 with us; Dr. Truman Brown and Jaime Cruz for their expert = help with=20 the magnetic resonance imaging; Kristie Gordon for her expert = help with the FACS; and Dr. Fred Gage for the pNIT retrovirus = construct.

Correspondence should be addressed to = Dr. Peter=20 Canoll, Department of Pathology, Columbia University, 630 West 168th = Street, New=20 York, NY 10032. Email: pc561{at}columbia.edu

DOI:10.1523/JNEUROSCI.0514-06.2006

Copyright =A9 2006 Society for Neuroscience = 0270-6474/06/266781-10$15.00/0=20


3D"   =20 References
Top
Abstract
Introduction
Materials and Methods
Results
Discussion
3D"References
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Bachoo RM, Maher EA, Ligon KL, Sharpless NE, Chan SS, You MJ, Tang Y, = DeFrances J, Stover E, Weissleder R, Rowitch DH, Louis DN, DePinho RA = (2002)=20 Epidermal growth factor receptor and Ink4a/Arf: convergent mechanisms = governing=20 terminal differentiation and transformation along the neural stem cell = to=20 astrocyte axis. Cancer Cell 1:269=96277.[CrossRef][ISI][Medline]

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Blankenberg FG, Teplitz RL, Ellis W, Salamat MS, Min BH, Hall L, = Boothroyd=20 DB, Johnstone IM, Enzmann DR (1995) The influence of volumetric tumor = doubling=20 time, DNA ploidy, and histologic grade on the survival of patients with=20 intracranial astrocytomas. AJNR Am J Neuroradiol 16:1001=961012.[Abstract]

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R0lGODlhMQAQAJECAAAAANYTNP///wAAACH5BAEAAAIALAAAAAAxABAAQAJQlI+py80BwwG0AkPn FbZj64TiSJbmiaYOlGTfC1fa5jYsUrs5TcebCgwKh0SgpPXbyWCaD0gRwSV5TY7sWXvcmLrp60pd KXYzstS5LKrXqAIAOw== ------=_NextPart_000_0000_01C85871.F932BDA0 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.jneurosci.org/javascript/ajax/xmlhttprequest.js /*=0A= =0A= Cross-Browser XMLHttpRequest v1.2=0A= =3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D= =3D=3D=3D=3D=3D=3D=3D=3D=0A= =0A= Emulate Gecko 'XMLHttpRequest()' functionality in IE and Opera. 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_0000_01C85871.F932BDA0 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 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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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// set the web path to the __utm.gif file=0A= var _utsp=3D"|"; // transaction field separator=0A= var _uflash=3D1; // set flash version detect option (1=3Don|0=3Doff)=0A= var _utitle=3D1; // set the document title detect option = (1=3Don|0=3Doff)=0A= var _ulink=3D0; // enable linker functionality (1=3Don|0=3Doff)=0A= var _uanchor=3D0; // enable use of anchors for campaign = (1=3Don|0=3Doff)=0A= var _utcp=3D"/"; // the cookie path for tracking=0A= var _usample=3D100; // The sampling % of visitors to track (1-100).=0A= =0A= //-- UTM Campaign Tracking Settings=0A= var _uctm=3D1; // set campaign tracking module (1=3Don|0=3Doff)=0A= var _ucto=3D"15768000"; // set timeout in seconds (6 month default)=0A= var _uccn=3D"utm_campaign"; // name=0A= var _ucmd=3D"utm_medium"; // medium (cpc|cpm|link|email|organic)=0A= var _ucsr=3D"utm_source"; // source=0A= var _uctr=3D"utm_term"; // term/keyword=0A= var _ucct=3D"utm_content"; // content=0A= var _ucid=3D"utm_id"; // id number=0A= var _ucno=3D"utm_nooverride"; 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path=3D"+_utcp+";"+nx+_udo;=0A= _ubd.cookie=3D"__utmb=3D"+_udh+"; path=3D"+_utcp+";"+x+_udo;=0A= _ubd.cookie=3D"__utmc=3D"+_udh+"; path=3D"+_utcp+";"+_udo;=0A= _ufns=3D1;=0A= } else if (a>=3D0 && b>=3D0 && c>=3D0) {=0A= _ubd.cookie=3D"__utmb=3D"+_udh+"; path=3D"+_utcp+";"+x+_udo;=0A= } else {=0A= if (a>=3D0) a=3D_uFixA(_ubd.cookie,";",_ust);=0A= else a=3D_udh+"."+_uu+"."+_ust+"."+_ust+"."+_ust+".1";=0A= _ubd.cookie=3D"__utma=3D"+a+"; path=3D"+_utcp+";"+nx+_udo;=0A= _ubd.cookie=3D"__utmb=3D"+_udh+"; path=3D"+_utcp+";"+x+_udo;=0A= _ubd.cookie=3D"__utmc=3D"+_udh+"; path=3D"+_utcp+";"+_udo;=0A= _ufns=3D1;=0A= }=0A= if (_ulink && xx && xx!=3D"" && xx!=3D"-") {=0A= xx=3D_uUES(xx);=0A= if (xx.indexOf(";")=3D=3D-1) _ubd.cookie=3D"__utmx=3D"+xx+"; = path=3D"+_utcp+";"+nx+_udo;=0A= }=0A= if (_ulink && v && v!=3D"" && v!=3D"-") {=0A= v=3D_uUES(v);=0A= if (v.indexOf(";")=3D=3D-1) _ubd.cookie=3D"__utmv=3D"+v+"; = path=3D"+_utcp+";"+nx+_udo;=0A= }=0A= _uInfo(page);=0A= _ufns=3D0;=0A= _ufno=3D0;=0A= if (!page || page=3D=3D"") _uff=3D1;=0A= }=0A= function _uInfo(page) {=0A= var p,s=3D"",dm=3D"",pg=3D_udl.pathname+_udl.search;=0A= if (page && page!=3D"") pg=3D_uES(page,1);=0A= _ur=3D_ubd.referrer;=0A= if (!_ur || _ur=3D=3D"") { _ur=3D"-"; 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