From: Subject: Autocrine/Paracrine Platelet-Derived Growth Factor Regulates Proliferation of Neural Progenitor Cells -- Erlandsson et al. 66 (16): 8042 -- Cancer Research Date: Thu, 17 Jan 2008 17:53:33 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C85931.E0C31510" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.3198 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C85931.E0C31510 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://cancerres.aacrjournals.org/cgi/content/full/66/16/8042 Autocrine/Paracrine Platelet-Derived Growth Factor = Regulates Proliferation of Neural Progenitor Cells -- Erlandsson et al. = 66 (16): 8042 -- Cancer Research 3D"Cancer 3D"Ubiquitin 3D"
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[Cancer Research 66, 8042-8048, August 15, = 2006]
=A9 2006 American = Association=20 for Cancer Research

Cell, Tumor, and Stem Cell = Biology

Autocrine/Paracrine Platelet-Derived Growth Factor Regulates = Proliferation=20 of Neural Progenitor Cells

Anna=20 Erlandsson1, Karin = Br=E4nnvall1,=20 Sigrun Gustafsdottir2, Bengt=20 Westermark2 and Karin=20 Forsberg-Nilsson1

1 Department of Medical Biochemistry and=20 Microbiology, Uppsala University; and 2 Department of = Genetics and=20 Pathology, Uppsala University, Rudbeck Laboratory, Uppsala, Sweden =

Requests for reprints: Karin Forsberg-Nilsson, = Department of Medical Biochemistry and Microbiology, Uppsala University, = Box=20 582, 751 23 Uppsala, Sweden. Phone: 46-18-471-41-58; Fax: = 46-18-471-42-44;=20 E-mail: karin.nilsson{at}imbim.uu.se .


3D"   =20 Abstract
Top
3D"Abstract
Introduction
Materials and Methods
Results
Discussion
References
 
G= rowth=20 factors play an important role in regulating neural stem cell = proliferation and differentiation. This study shows that=20 platelet-derived growth factor (PDGF) induces a partial=20 differentiation of neural stem/progenitor cells (NSPCs) in = the=20 absence of other mitogens in vitro. NSPCs thus acquire = an=20 immature morphology and display markers for both neurons and = glia. In=20 addition, these cells do not readily mature in the absence of = further stimuli. When NSPC cultures treated with PDGF were = exposed=20 to additional differentiation factors, however, the=20 differentiation proceeded into neurons, astrocytes, and=20 oligodendrocytes. We find that NSPC cultures are endowed with = an=20 endogenous PDGF-BB production. The PDGF-BB expression peaks = during=20 early differentiation and is present both in cell lysates and = in=20 conditioned medium, allowing for autocrine as well as = paracrine=20 signaling. When the NSPC-derived PDGF was inhibited, = progenitor cell=20 numbers decreased, showing that PDGF is involved in NSPC=20 expansion. Addition of a PDGF receptor (PDGFR) inhibitor = resulted in=20 a more rapid differentiation. Neurons and oligodendrocytes=20 appeared earlier and had more elaborate processes than in = control=20 cultures where endogenous PDGFR signaling was not blocked. = Our=20 observations point to PDGF as an inducer of partial = differentiation=20 of NSPC that also sustains progenitor cell division. Such an=20 intermediate stage in stem cell differentiation is of = relevance for=20 the understanding of brain tumor development because = autocrine PDGF=20 stimulation is believed to drive malignant conversion of = central=20 nervous system progenitor cells. (Cancer Res 2006; 66(16):=20 8042-8)


3D"   =20 Introduction
Top
Abstract
3D"Introduction
Materials and Methods
Results
Discussion
References
 
D= uring=20 neural development, stem cell number and the subsequent = formation of=20 neurons, astrocytes, and oligodendrocytes must be tightly = controlled.=20 Failure in this regulation may lead to disorganization of the = central=20 nervous system (CNS) or the development of tumors. External = signals=20 are thought to play an important role for the regulation of = neural=20 stem/progenitor cell (NSPC) fate (1,=20 2).=20 In vitro, fibroblast growth factor-2 (FGF2) and = epidermal=20 growth factor are potent mitogens for NSPC (3),=20 whereas ciliary neurotrophic factor (CNTF) directs NSPC to an = astrocytic cell fate (4,=20 5)=20 and the thyroid hormone triiodothyronine (T3; ref. 4)=20 augments oligodendrocyte differentiation. To date, no = one-step=20 neuronal inducer has been described, although activation of = proneural=20 genes is clearly involved in lineage restriction and the = acquisition=20 of a neuronal fate (6).=20 Platelet-derived growth factor (PDGF) has been suggested to = support=20 neuronal differentiation (4,=20 7),=20 and we have previously reported that PDGF acts as a mitogen = for=20 immature neurons (8).=20

PDGF binds to and signal through the two cell surface receptor=20 tyrosine kinases, PDGF receptor (PDGFR) 3D{alpha}=20 and = PDGFR=DF=20 (9).=20 The PDGF ligands are present as homodimers (AA, BB, CC, and = DD) and=20 heterodimer (AB). Binding of the ligand causes dimerization = of the=20 PDGFR followed by an activation of intracellular signaling = cascade=20 (10,=20 11).=20 The PDGFRs have different binding capacities for the ligands. = The=20 =DF-receptor binds PDGF-B and PDGF-D, whereas the 3D{alpha}=20-receptor binds=20 PDGF-A, PDGF-B, and PDGF-C.

Both the PDGFRs and their ligands are expressed during CNS = development.=20 PDGF-A has been detected in the subventricular zone at = embryonic=20 day 11.5 (E11.5; ref. 12)=20 and in neurons in the spinal cord and dorsal root ganglia of = E12=20 mice. From E15 and onwards, PDGF-A is found in most neurons = (13).=20 Similar to PDGF-A, PDGF-B is expressed by neurons in the = adult brain,=20 and PDGF-B-positive cells were also found in the = subventricular zone=20 in E14 mice (14).=20 PDGFR3D{alpha} is predominantly expressed by oligodendrocyte = precursors=20 during neural development (15),=20 and loss-of-function studies further enhance the role for = PDGF in=20 oligodendrocyte development (9).=20

PDGF is a potent growth and differentiation factor during normal = development and has also been implied in tumor development.=20 Many types of brain tumors express the PDGFRs and their = ligands=20 (16),=20 and retroviruses encoding PDGF have been reported to induce = brain=20 tumors in mice (17=9619).=20 Several recent studies suggest that gliomas arise from = transformed=20 NSPCs and stem cells have also been isolated from brain = tumors (20,=20 21).=20 We therefore sought to define the effect of PDGF in NSPC=20 differentiation. In this study, we find that PDGF has=20 growth-promoting capacity for progenitors from the CNS in=20 vitro. Furthermore, autocrine/paracrine PDGF from NSPC = supports=20 early neurogenesis and gliogenesis. Autocrine PDGF signaling = may=20 therefore reflect a normal trait of progenitor cells that is = retained=20 in their transformed derivatives.


3D"   =20 Materials and Methods
Top
Abstract
Introduction
3D"Materials and = Methods
Results
Discussion
References
 
<= B>Cell=20 culture. NSPCs from embryonic cerebral cortex of timed = pregnant=20 Sprague-Dawley rats (B&K, Sollentuna, Sweden), E15, were = prepared=20 and maintained as described previously (8).=20 NSPCs were used for experiments 1 to 4 days after the first=20 passage.

For differentiation of NSPC, FGF2 was withdrawn and PDGF-AA = (10=20 ng/mL) or PDGF-BB (10 ng/mL) was added daily. After 4 days, = PDGF was=20 withdrawn and the cells were grown for an additional 4 days. = Control=20 cells were grown for 8 days in the absence of PDGF and FGF2. = B27=20 supplement (1:50), CNTF (10 ng/mL), and T3 (3 ng/mL) were = added daily=20 during the 4 last days after the withdrawal of PDGF-AA. =

To study the effect of endogenous PDGF on NSPC fate = determination,=20 NSPCs were initially placed on polyornithine/fibronectin = glass=20 coverslips (3,700 cells/cm2) in the presence of FGF2.=20 After 2 days, FGF2 was withdrawn and cells were treated with=20 either PDGF-BB (10 ng/mL), STI 571 (1 =B5mol/L), or vehicle = and=20 fixed in 4% paraformaldehyde in PBS (pH 7.4) after 1, 2, 4, = and=20 6 days of differentiation and processed for immunocytochemistry. = For=20 staining with antibodies to PDGF-BB, NSPCs grown in the = presence of=20 FGF2 were fixed and processed for immunocytochemistry.

Immunocytochemistry. NSPCs were grown on=20 polyornithine/fibronectin-coated glass coverslips and fixed = in=20 ice-cold acid ethanol (90% ethanol, 5% acetic acid) or in 4%=20 paraformaldehyde (O4 staining and STI 571 treatment of NSPC = during=20 differentiation). Cells were permeabilized with 0.2% Triton = X-100 in=20 PBS (except for O4 staining), washed in PBS, and incubated in = blocking solution [20% normal goat serum (DAKO A/S, Glostrup, = Denmark), 0.1% saponin (Sigma, St. Louis, MO), and 4 mg/mL = bovine=20 serum albumin (BSA; Sigma)] in PBS for 30 minutes at room=20 temperature. The cells were incubated with the following = primary=20 antibodies diluted in PBS with 0.1% saponin and 4 mg/mL BSA = for 1=20 hour at 37=B0C: mouse =DF-III tubulin (BabCO, 1:500; Nordic = Biosite AB,=20 T=E4by, Sweden), rabbit anti-GFAP (1:200; DAKO A/S), mouse = anti-CNPase=20 (1:500; Sigma), mouse anti-O4 (1:30; Chemicon, Temecula, CA), = mouse anti-PDGF-BB (007; 1:50; Biosciences Research=20 Laboratories, Mochida Pharmaceutical, Tokyo, Japan), rabbit=20 anti-PDGFR3D{alpha}=20 = (1:200;=20 Santa Cruz Biotechnology, Santa Cruz, CA), and mouse/rabbit=20 anti-nestin (1:200; Chemicon). NSPCs were washed 5 to 6 times = in PBS and incubated with Cy3-, AMCA-, or FITC-conjugated = goat=20 anti-mouse/rabbit secondary antibodies (Jackson = ImmunoResearch=20 Laboratories, West Grove, PA). Pictures were taken with a 20x objective on a Zeiss Axioplan = 2 (Carl=20 Zeiss, Oberkochen, Germany).

Cell counting. Following the immunostainings, positive = cells=20 were counted using a fluorescence microscope (Carl Zeiss). = For=20 duplicate samples, positively stained cells and the total = cell=20 numbers were counted in eight different fields. Cell counting = for the STI 571 inhibition studies was done using a Coulter = Z1=20 cell counter (Coulter Electronics, Herpendon, United Kingdom). = The=20 cells were removed from the dishes with a cell scraper, after = which=20 they were carefully dissociated and counted. Each time point = was=20 analyzed in triplicate.

Proximity ligation. Samples were analyzed for PDGF-BB by=20 homogenous proximity ligation using a polyclonal anti-PDGF-B = chain=20 antibody (PeproTech, Rocky Hill, NJ) as described before (22).=20 Before analysis, the samples were boiled for 10 minutes and=20 diluted 10 times in PBS containing 0.1% BSA. Each sample (1 = =B5L)=20 was analyzed in triplicates. Quantification of the total = protein=20 content was done using the NanoOrange kit (Molecular Probes,=20 Eugene, OR). Results of the analysis are expressed as = percentage=20 of maximal values (pg of PDGF-BB/mg of total amount of = protein=20 in lysates and pmol/L in the medium).

Statistical analysis. For statistical analysis, the ANOVA = method=20 was used followed by multiple comparison by Fisher's method = as=20 well as Student's t test. Values presented are the mean =B1=20 SEM.


3D"   =20 Results
Top
Abstract
Introduction
Materials and Methods
3D"Results
Discussion
References
 
<= B>PDGF=20 delays NSPC differentiation. NSPCs from the embryonic rat = cortex=20 were expanded in medium containing FGF2, which inhibits=20 differentiation. After FGF2 withdrawal, NSPC cultures were = treated=20 with PDGF during the first 4 days of differentiation. = Following=20 washout of PDGF, cultures were kept for an additional 4 days = in=20 the absence of growth factor, and the ability of PDGF-pretreated=20 cells to differentiate to mature neurons was studied. Control = cultures contained =DF-III tubulin-positive cells with = a mature=20 neuronal phenotype (Fig.=20 1A ). In contrast, in cultures that had been = pretreated=20 with PDGF-AA or PDGF-BB, =DF-III tubulin-positive cells were = immature:=20 had a rounded appearance, lacked processes, and tended to = grow in=20 clusters. A slightly more potent effect of PDGF-AA than = PDGF-BB was=20 noticed; particularly, the cell density was higher with = PDGF-AA and=20 the cell clusters contained more cells. Thus, NSPC pretreated = with=20 PDGF did not readily mature further when kept without = mitogen.=20 Instead, they maintained an immature phenotype, suggesting = that=20 exposure to PDGF delayed terminal differentiation. =



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  		Figure 1. PDGF alters = neuronal=20 maturation without increasing the proportion of neurons. = A,=20 after FGF2 withdrawal, NSPCs were grown either for 8 days in = medium=20 without mitogens (No add.) or for 4 days in the = presence of=20 PDGF-AA or PDGF-BB followed by 4 days in medium without = PDGF. Cells=20 were fixed and stained for the neuronal marker =DF-III = tubulin=20 (Tuj1). Depending on their morphology, the = Tuj1-positive=20 cells were divided into two subgroups. Cells extending long=20 processes were judged to be more mature neurons = (arrows),=20 whereas round cells bearing no processes were defined as = immature=20 neurons (arrowheads). B, quantification of = cells with=20 neuronal morphology. The proportion of Tuj1-positive cells = that=20 lacked processes (immature) was increased by a factor of 2 = in=20 PDGF-treated cell cultures (control, 19.0 =B1 5.5%; PDGF-AA, = 50.8 =B1=20 2.6%; PDGF-BB, 42.9 =B1 2.8%). Mature neurons (bearing = processes) were=20 almost exclusively found in cultures that had not been = exposed to=20 PDGF. (control, 35.4 =B1 6.7%; PDGF-AA, 1.3 =B1 0.6%; = PDGF-BB, 3.0 =B1=20 1.1%). The total number of TuJ1-positive cells remains = unchanged by=20 PDGF. =

 = ;
PDGF=20 alters neuronal maturation without increasing the proportion = of=20 neurons. The number of immature (no processes) and mature=20 (process bearing) =DF-III tubulin-positive cells was = counted and=20 displayed as the percentage of total cells (Fig.=20 1B). In control cultures, most neurons are mature, = 35.4=20 =B1 6.7% of the total cell number, compared with 19.0 =B1 = 5.5%=20 immature neurons. In contrast, the PDGF-treated cultures = contained=20 very few mature neurons but instead a large population of = immature=20 neurons. For the PDGF-AA-treated cultures, 1.3 =B1 0.6% = of the=20 neurons were mature and 50.8 =B1 2.6% were immature, whereas = the=20 corresponding numbers for PDGF-BB-treated cultures were 3.0 = =B1 1.1%=20 and 42.9 =B1 2.8%, respectively. We did not notice any = difference in=20 the total number of =DF-III tubulin-positive cells between = control and=20 PDGF-treated cultures, indicating that PDGF does not increase = the=20 amount of neurons.

Astrocyte cell number is not affected by PDGF. Next, we=20 investigated the effect of PDGF-AA and PDGF-BB on formation = of=20 astrocytes and oligodendrocytes. FGF2 was withdrawn, and = NSPCs were=20 treated with PDGF-AA or PDGF-BB or untreated for 4 days. This = was=20 followed by an additional 4 days without mitogens, after = which the=20 number of astrocytes and oligodendrocytes was counted. In = control=20 cultures, 34.6 =B1 3.3% of the cells were positive for GFAP, = and=20 this remained unchanged by PDGF treatment (36.4 =B1 1.4% = for=20 PDGF-AA and 33.6 =B1 3.1% for PDGF-BB). Although the amount = of=20 GFAP-positive cells was not affected by PDGF exposure, we = noted a=20 less developed morphology of astrocytes after PDGF treatment = (data=20 not shown). Oligodendrocytes derived from NSPC in culture = appear=20 later than neurons and astrocytes and are few in number = (2-10%). We=20 noticed a slight reduction in O4-positive cells in cultures = treated=20 with PDGF-AA (1.0 =B1 0.6%) or PDGF-BB (2.1 =B1 0.5%) = compared with=20 untreated cultures (6.4 =B1 0.9%).

The PDGF-induced maturation delay can be overcome by = additional=20 treatment. We next investigated if PDGF-treated cells that = were=20 unable to differentiate into mature neurons simply by = withdrawing=20 PDGF could be influenced to differentiate by additional = stimuli.=20 The medium supplement B27 used to enhance neuronal maturation = (23,=20 24),=20 CNTF, or the thyroid hormone T3 shown to facilitate = differentiation=20 into oligodendrocytes were used.

After PDGF-AA exposure in the absence of other mitogens, NSPCs=20 were grown for another 4 days in medium alone or in medium = containing=20 B27, CNTF, or T3. The cells were fixed and stained with = antibodies=20 against =DF-III tubulin, GFAP, or O4 (Fig.=20 2A ). In cultures that were kept in medium alone = for 4=20 days after PDGF-AA washout, both neurons and astrocytes = showed an=20 immature morphology as mentioned above and only a few=20 oligodendrocytes were detected. B27 treatment resulted in = more mature=20 cells of all three lineages, although the effect was most = pronounced=20 for neurons. In contrast to the round immature =DF-III = tubulin-positive=20 cells found in cultures treated with PDGF-AA, neurons in=20 B27-treated cultures extended long processes. Cells that were = treated=20 with CNTF following PDGF-AA treatment had almost exclusively=20 differentiated into astrocytes with a mature phenotype. Thus, = when=20 exchanging PDGF for CNTF, hardly any =DF-III tubulin-positive = cells or O4-positive cells could be detected, indicating that = CNTF could exert a similar effect on PDGF-AA-treated stem = cell=20 cultures as on naive NSPC. This further suggests that=20 PDGF-AA-treated cells were not committed to a neuronal cell = fate.=20 Treatment of the cells with T3 had a weakly enhancing effect = on the=20 number of O4-positive cells and caused a reduction in = =DF-III=20 tubulin-positive cells.



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  		Figure 2. The = PDGF-induced delay in=20 differentiation can be overcome by additional stimulation. = A,=20 NSPCs were first treated with PDGF-AA for 4 days and then = with CNTF,=20 B27, or T3 for 4 days. Cells were fixed and stained for GFAP = (green), Tuj1 (red), or O4 (green). = Control=20 cultures where PDGF was discontinued resulted in an immature = morphology after 4 days, whereas B27 increased the maturity = of all=20 three lineages. CNTF converted PDGF-pretreated cells into = almost=20 exclusively astrocytes. T3 treatment decreased the number of = neurons=20 formed and increased the number of O4-positive cells. = B to=20 D, quantification of cell lineages formed after PDGF = exposure=20 that was followed by addition of B27, CNTF, or T3. =

 = ;
The=20 proportion of neurons (Fig.=20 2B) was markedly reduced by CNTF and T3, 4.3 =B1 = 1.8% and=20 9.5 =B1 0.9%, respectively, compared with 52.8 =B1 3.9% in = the control=20 culture. In B27-treated cultures, neurons were more mature = but the=20 total neuronal number was not affected. The proportion of=20 astrocytes (Fig.=20 2C) was increased from 27 =B1 2.0% to 88.5 =B1 = 4.7% when the=20 PDGF-AA-exposed cells were treated with CNTF, whereas T3 and = B27 had=20 no obvious effect on the astrocyte number. A low proportion = (6.4 =B1=20 0.9%) of the PDGF-treated cells that received no further = stimulus=20 were positive for O4 (Fig.=20 2C), and this proportion was even lower in = CNTF-treated=20 cultures (0.4 =B1 0.3%). Both B27 and T3 increased the number = of=20 oligodendrocytes, resulting in 11 =B1 0.6% and 9. =B1 = 1.7%=20 O4-positive cells, respectively.

Progenitors for neurons and oligodendrocytes express PDGF3D{alpha}=20=20 receptors. We have previously shown that PDGFR3D{alpha}=20 is = expressed by=20 proliferating NSPC in culture (8),=20 whereas PDGFR=DF is not detectable until after several days = of=20 differentiation. It is not known which progenitor cell = population=20 continues to express PDGFR3D{alpha}=20 on=20 FGF2 withdrawal, and we therefore investigated if the distribution = of=20 PDGFR3D{alpha} differs between progenitor cell types. Immunostainings = done=20 after 3 days of differentiation show that PDGFR3D{alpha}=20-expressing=20 cells mainly displayed markers for neurons and = oligodendrocytes=20 and, to a lesser extent, for astrocytes (Supplementary Fig.=20 S1).

Autocrine/paracrine PDGF-BB from NSPC. The concept of=20 autocrine/paracrine signaling has been discussed in the = context of=20 PDGF-induced brain tumors as it enables cells to respond to = their own=20 growth factor or PDGF secreted by close neighboring cells. We = have noticed that NSPC density is a determining factor in = growth=20 arrest and that dense cultures continue to proliferate for a=20 few days after withdrawal of FGF2 presumably because of = supportive=20 factors from cells in close contact, whereas sparse cultures = do=20 not.3=20 We reasoned that PDGF might be one factor that signals in the = microenvironment of NSPC. To investigate if NSPCs produce = PDGF, we=20 used two independent methods: immunostaining and the more = recently=20 developed proximity ligation method. Figure=20 3A shows that NSPCs grown in the presence of FGF2 = are=20 immunopositive for PDGF-BB (95.7 =B1 1.5% of the = nestin-positive=20 cells also displayed PDGF-BB immunoreactivity). Small amounts = of=20 protein as well as changes in protein levels can accurately = be=20 detected by proximity ligation. PDGF-BB is produced (in the = range=20 of pg/mg protein) and is present in cell lysates of NSPC in=20 FGF2-containing medium (Fig.=20 3B). On withdrawal of FGF2, the levels increase = 10-fold at=20 2 days of differentiation, and by 6 days, the PDGF-BB = concentration=20 has returned to 15% of its maximal value. PDGF-BB is also = secreted=20 into the medium of NSPC cultures with the same peak at 2 days = of=20 differentiation (Fig.=20 3B). Thus, although some PDGF-BB remained cell = associated=20 or within the cell, some of the secreted protein was also = made=20 available to other cells pointing at possibilities for both = short-=20 and long-range effects.



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  		Figure 3. Endogenous = PDGF-BB=20 produced by NSPC. A, the vast majority (95.7 =B1 = 1.5%) of NSPCs=20 in culture are immunoreactive for PDGF-BB. B, levels = of=20 PDGF-BB were measured with the proximity ligation assay in = cell=20 lysates and in conditioned medium from NSPC during = conditions of=20 proliferation or differentiation. In the presence of FGF2, = PDGF-BB=20 can be detected at low levels. After FGF2 withdrawal, = PDGF-BB levels=20 in both lysate and conditioned medium increased by 10-fold = at day 2=20 of differentiation. By 4 days, the levels of PDGF-BB have = returned=20 to 20% of its maximal value. =

 = ;
NSPC-derived=20 PDGF-BB drives progenitor cell proliferation during early=20 differentiation to neurons and glia. To investigate if = the=20 PDGF-BB produced by NSPC during early differentiation plays a = role in=20 progenitor proliferation, we blocked signaling through = PDGFRs. Cells=20 were grown in the presence or absence of a receptor tyrosine = kinase=20 inhibitor, STI 571, and counted at the starting point of the=20 experiment and at different times after initiation of=20 differentiation. In the absence of FGF2, there was a 3-fold = increase=20 in the total cell number after 6 days (Fig.=20 4A ). Clearly, one or several endogenous factor/s = are able=20 to expand progenitor cells at this stage. In cultures treated = with=20 STI 571, further cell division was stopped beyond 2 days of=20 differentiation, which coincides with the peak of PDGF = expression. In=20 control cultures, STI 571 was added to cells growing in the=20 presence of FGF2 (Fig.=20 4B). After 12 days, the total cell number was 25 = times=20 higher than at the starting point of the experiment = regardless of=20 whether STI 571 was added or not. The inhibitor thus had no=20 unspecific toxic or growth-arresting effects on NSPC. =



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  		Figure 4. Endogenous PDGF = from NSPC=20 stimulates progenitor proliferation. A, after = withdrawal of=20 FGF2, NSPCs were treated daily with the tyrosine kinase = inhibitor=20 STI 571 or vehicle and counted at 1, 2, 4, and 6 days. STI = 571=20 addition abolished the increase in cell numbers beyond 2 = days of=20 differentiation. Bars, SEM. B, endogenous PDGF = had no=20 effect on cell number in the presence of FGF2. NSPCs were = treated=20 daily with STI 571 or vehicle in the presence of FGF2. After = 12 days=20 in culture, the total cell number had increased 25 times, = showing=20 lack of toxic effect by STI 571. Bars, SEM. =

 = ;
The=20 above result prompted us to find out how the inhibition of = PDGF=20 signaling affected different cell lineages deriving from = NSPC. During=20 differentiation induced by withdrawal of FGF2, the amount of=20 nestin-positive cells declines as a result of cell fate = choice and=20 markers of neurons and glia gradually appear. At 2 days of=20 differentiation, there were fewer nestin-positive cells in = STI=20 571-treated cultures compared with control cells, indicating = a more=20 rapid differentiation when PDGFR signaling was blocked (Fig.=20 5A ). In the presence of STI 571, neurons (Fig.=20 5B) and oligodendrocytes (Fig.=20 5C) were more abundant at an earlier stage than in = control=20 cultures. A similar trend, which could not be statistically=20 confirmed, was also seen for astrocytes (Fig.=20 5D).



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  		Figure 5. Inhibition of = endogenous=20 PDGF alters the differentiation kinetics of NSPC. NSPCs were = differentiated by withdrawal of FGF2 in the presence or = absence of=20 STI 571 for 0, 1, 2, 4, and 6 days. Cells were fixed and = processed=20 for immunocytochemistry. A, in STI 571-treated = cultures,=20 there are fewer nestin-positive cells at day 2 of = differentiation.=20 B, STI 571 results in an increase in the number of = neurons=20 expressing =DF-III tubulin at day 4 of differentiation. = C, the=20 number of oligodendrocytes increases when STI 571 is added.=20 D, STI 571 has no significant effect on the = generation of=20 astrocytes. =

 = ;
In=20 addition to an altered cell number, the morphology of neurons = and=20 oligodendrocytes was altered (Fig.=20 6 ). In cultures treated with STI 571 for 4 days, = elaborate=20 neurite extensions with long processes were seen on =DF-III=20 tubulin-positive cells. Oligodendrocytes were also = morphologically=20 more mature. Thus, the NSPC-derived PDGF expands the pool of = immature=20 progenitors during early differentiation and has most = profound=20 effects on neurons and oligodendrocytes.



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  		Figure 6. Blocking of = PDGFR=20 signaling increases the morphologic maturation of neurons = and=20 oligodendrocytes. STI 571 increases the maturity of =DF-III = tubulin=20 and CNPase-positive cells with regard to neurite outgrowth = and=20 process arborization, respectively. DAPI,=20 4',6-diamidino-2-phenylindole. =

 = ;

3D"   =20 Discussion
Top
Abstract
Introduction
Materials and Methods
Results
3D"Discussion
References
 
W= e here=20 describe that PDGF retains progenitors from NSPCs in an = immature=20 state in vitro, thus keeping them from terminal=20 differentiation. This is particularly shown for the neuronal=20 and oligodendrocyte lineages, although PDGF did not markedly=20 affect differentiation to astrocytes. Furthermore, we show = that=20 a peak of autocrine/paracrine PDGF-BB produced by NSPC during = initial differentiation acts to expand precursors of neurons=20 and oligodendrocytes.

In the context of neural stem cell culture, PDGF was first shown = to stimulate the formation of neurons from NSPC (4).=20 Subsequently, the potential by which PDGF acts as a = neurogenic=20 inducer has been a subject of some debate. Although there are = reports=20 that a short exposure to PDGF suffices to induce neuronal=20 differentiation (7),=20 others, including our own laboratory, find the effect of PDGF = mainly=20 to expand neural progenitors (8).=20 Data to support this conclusion come from the study by = Mohapel et al.=20 (25)=20 who detected an overall increase in striatal cell number but = not=20 in newly generated neurons after intracerebroventricular = infusion=20 of PDGF. Others suggest that PDGF instead reduces the number = of=20 neurons in human neural stem cell cultures (26).=20 We report that NSPC-derived neurons generated in the presence = of=20 PDGF were almost completely devoid of neurites, thus only=20 partially differentiated. However, the total number of = neurons did=20 not differ from control cultures and this shows that the = effect=20 of PDGF relates to suppression of maturation as shown by the=20 shift in neuronal morphology.

More mature cell types of all three lineages were induced by=20 addition of B27 after washout of PDGF. In contrast, cultures=20 that were not replenished with other supplements after = removal=20 of PDGF were slower to establish a differentiated phenotype.=20 This is indicative of a block of terminal differentiation = that=20 continued over several days. In accordance, addition of T3,=20 which is known to augment the differentiation of = oligodendrocytes,=20 led to a higher proportion of oligodendrocytes compared with=20 untreated cultures (4).=20 The instructive effect of CNTF on NSPC is well documented (4,=20 5).=20 We found that it was possible to direct PDGF-treated cells = toward the=20 astrocytic lineage using CNTF. This means that cultures, in = which 50%=20 of the cells express =DF-III tubulin, can be redirected to = 90%=20 GFAP-positive cells. The cytoskeletal protein =DF-III tubulin = is=20 commonly used as a neuronal marker (27,=20 28),=20 but transient =DF-III tubulin expression has also been = reported in=20 telencephalic subventricular zones comprising putative neural = and=20 glial precursors (reviewed in ref. 29).=20 We can therefore not conclude whether =DF-III = tubulin-positive cells=20 switching into GFAP expression reflect a transient marker = expression=20 of cells that can adopt a glial fate or whether this reflects = lineage=20 plasticity.

We have observed that NSPCs in culture continue to proliferate=20 even after FGF2 withdrawal in a density-dependent manner.3=20 This would indicate that diffusible factors and/or cell-cell=20 contacts inherent to the stem cell culture stimulate = proliferation=20 in the absence of exogenous FGF2. A previous study reported = that=20 conditioned medium from dense neural stem cells increased the = number of viable cells and clone size when stem cells were = seeded=20 at clonal density (30),=20 but the diffusible factor(s) secreted remained unidentified. = Given=20 our findings that PDGF can act as a mitogen for progenitors = from stem=20 cells and reduces apoptosis (8),=20 PDGF could be one such inherent factor. Using two independent=20 detection methods, we find that NSPC cultures produce = PDGF-BB.=20 The highly specific and sensitive proximity ligation assay (22,=20 31)=20 revealed a peak of PDGF production during early NSPC = differentiation.=20 This method of measuring protein concentrations, which = converts=20 the actual protein detection to the analysis of a DNA = sequence,=20 enabled detection of PDGF-BB both in cell lysates and in = conditioned=20 medium. PDGF-BB in the cell lysate may be located either=20 intracellularly or associated with proteoglycans of the = extracellular=20 matrix (32,=20 33),=20 owing to a retention signal in the PDGF-BB protein. This = could result=20 in a localized high concentration of the growth factor. The = proximity=20 ligation assay also detected PDGF in the conditioned medium. = Making=20 PDGF available to other cells allows for paracrine as well as = autocrine signaling and may account, at least in part, for=20 proliferative effect described above.

Because addition of the small-molecule inhibitor STI 571 reduces = cell number in progenitor cultures, we suggest that = endogenous=20 PDGF-BB stimulates progenitor proliferation. STI 571 blocks = the=20 activation of type III tyrosine kinase receptors, including = the=20 receptors for PDGF, stem cell factor (SCF), and the abl = tyrosine=20 kinase (34).=20 We have earlier reported that SCF has no effect on neural = stem cell=20 proliferation (35)=20 and its involvement is therefore less likely. The importance = of the=20 ableson nonreceptor kinase (abl) for proper axon outgrowth = and path=20 finding was shown for Drosphila (36),=20 and cytoskeletal rearrangements in developing mammalian = neurons=20 involve abl (reviewed in ref. 37).=20 A role for abl in neural stem cells has not been suggested = but=20 can at present not be ruled out.

When PDGFR signaling was blocked during differentiation of NSPC, = neurons and oligodendrocytes differentiated more rapidly. For = the astrocytic lineage, there was a similar trend, which = could=20 not be statistically confirmed. Neurons that form from NSPC = in=20 the presence of STI 571 have longer and more elaborate processes = than=20 those of control cultures. The same was true for oligodendrocytes=20 that exhibited an extensive arborization of their processes,=20 indicative of a mature morphology. Thus, PDGF inherent to = NSPC=20 cultures from the embryonic rat cortex expands pools of = immature=20 precursors during early differentiation.

Progenitors of neurons and glia with an autocrine PDGF signaling = have been suggested as an origin of malignant brain tumors (18).=20 PDGF and PDGFRs are frequently coexpressed in human glioma = cell=20 lines, and some glioma cell lines can be growth arrested by=20 PDGFR inhibitors (16).=20 In vivo, during normal development, the ligand is = generally=20 confined to one cell type and the receptor is expressed by an = adjacent cell layer (9).=20 The concomitant expression of both PDGF and its receptor in = cultures=20 of normal cells from the CNS has to our knowledge not been = reported=20 before. This study is based on in vitro cell culture;=20 nonetheless, the finding that cultures of primary NSPC = regulate their=20 own proliferation by autocrine/paracrine PDGF provides a new = link=20 between regulation of normal cell growth and the perturbed = growth=20 pathways of malignant brain tumors. Autocrine PDGFR signaling = in=20 glioma cells may reflect a natural trait of their normal=20 progenitor.


3D"   =20 Acknowledgments
 
The = costs of=20 publication of this article were defrayed in part by the = payment of=20 page charges. This article must therefore be hereby marked=20 advertisement in accordance with 18 U.S.C. Section = 1734 solely=20 to indicate this fact.


3D"   =20 Footnotes
 
Note: Supplementary data for this = article are=20 available at Cancer Research Online (http://cancerres.aacrjournals= .org/).=20

3 A. Erlandsson, = unpublished=20 observation.

Received 3/ 9/06. Revised 5/ 2/06. Accepted 6/ 1/06.


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

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quoted-printable Content-Location: http://cancerres.aacrjournals.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_01C85931.E0C31510 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://cancerres.aacrjournals.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 if (name =3D=3D "margin-right") {=0A= e.style.marginRight =3D value;=0A= }=0A= else if (name =3D=3D "margin-top") {=0A= e.style.marginTop =3D value;=0A= }=0A= else if (name =3D=3D "min-height") {=0A= e.style.minHeight =3D value;=0A= }=0A= else if (name =3D=3D "overflow") {=0A= e.style.overflow =3D value;=0A= }=0A= else if (name =3D=3D "overflow-x") {=0A= e.style.overflowX =3D value;=0A= }=0A= else if (name =3D=3D "overflow-y") {=0A= e.style.overflowY =3D value;=0A= }=0A= else if (name =3D=3D "padding") {=0A= e.style.padding =3D value;=0A= }=0A= else if (name =3D=3D "padding-bottom") {=0A= e.style.paddingBottom =3D value;=0A= }=0A= else if (name =3D=3D "padding-left") {=0A= e.style.paddingLeft =3D value;=0A= }=0A= else if (name =3D=3D "padding-right") {=0A= e.style.paddingRight =3D value;=0A= }=0A= else if (name =3D=3D "padding-top") {=0A= e.style.paddingTop =3D value;=0A= }=0A= else if (name =3D=3D "page-break-after") {=0A= e.style.pageBreakAfter =3D value;=0A= }=0A= else if (name =3D=3D "page-break-before") {=0A= e.style.pageBreakBefore =3D value;=0A= }=0A= else if (name =3D=3D "pixelBottom") {=0A= e.style.pixelBottom =3D value;=0A= }=0A= else if (name =3D=3D "pixelHeight") {=0A= e.style.pixelHeight =3D value;=0A= }=0A= else if (name =3D=3D "pixelLeft") {=0A= e.style.pixelLeft =3D value;=0A= }=0A= else if (name =3D=3D "pixelRight") {=0A= e.style.pixelRight =3D value;=0A= }=0A= else if (name =3D=3D "pixelTop") {=0A= e.style.pixelTop =3D value;=0A= }=0A= else if (name =3D=3D "pixelWidth") {=0A= e.style.pixelWidth =3D value;=0A= }=0A= else if (name =3D=3D "posBottom") {=0A= e.style.posBottom =3D value;=0A= }=0A= else if (name =3D=3D "posHeight") {=0A= e.style.posHeight =3D value;=0A= }=0A= else if (name =3D=3D "position") {=0A= e.style.position =3D value;=0A= }=0A= else if (name =3D=3D "posLeft") {=0A= e.style.posLeft =3D value;=0A= }=0A= else if (name =3D=3D "posRight") {=0A= e.style.posRight =3D value;=0A= }=0A= else if (name =3D=3D "posTop") {=0A= 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{=0A= e.style.scrollbarHighlightColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-shadow-color") {=0A= e.style.scrollbarShadowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-track-color") {=0A= e.style.scrollbarTrackColor =3D value;=0A= }=0A= else if (name =3D=3D "float") {=0A= e.style.styleFloat =3D value;=0A= }=0A= else if (name =3D=3D "table-layout") {=0A= e.style.tableLayout =3D value;=0A= }=0A= else if (name =3D=3D "text-align") {=0A= e.style.textAlign =3D value;=0A= }=0A= else if (name =3D=3D "text-align-last") {=0A= e.style.textAlignLast =3D value;=0A= }=0A= else if (name =3D=3D "text-autospace") {=0A= e.style.textAutospace =3D value;=0A= }=0A= else if (name =3D=3D "text-decoration") {=0A= e.style.textDecoration =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationBlink") {=0A= e.style.textDecorationBlink =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationLineThrough") {=0A= e.style.textDecorationLineThrough =3D value;=0A= }=0A= else if (name =3D=3D 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