From: Subject: Expression of oligodendrocyte progenitor cell antigens by gliomas: Implications for the histogenesis of brain tumors -- Shoshan et al. 96 (18): 10361 -- Proceedings of the National Academy of Sciences Date: Wed, 16 Jan 2008 19:20:49 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C85874.E6E50B00" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.3198 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C85874.E6E50B00 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.pnas.org/cgi/content/full/96/18/10361 Expression of oligodendrocyte progenitor cell = antigens by gliomas: Implications for the histogenesis of brain tumors = -- Shoshan et al. 96 (18): 10361 -- Proceedings of the National Academy = of Sciences
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Vol. 96, Issue 18, 10361-10366, August 31, = 1999

Medical = Sciences
Expression of=20 oligodendrocyte progenitor cell antigens by gliomas: Implications for = the=20 histogenesis of brain tumors

Yigal=20 Shoshan*,3D"dagger,, Akiko=20 Nishiyama*,=A7<= /SUP>,=20 Ansi Chang*, Sverre M=F6rk*, Gene H.=20 Barnett3D"dagger, John K. = Cowell*,, Bruce D. Trapp*, and Susan = M.=20 Staugaitis*,3D"dagger,=B6<= /SUP>=20

* Department of Neurosciences, The Lerner = Research=20 Institute, and 3D"dagger Department of Neurosurgery, Cleveland = Clinic=20 Foundation, Cleveland, OH 44195=20

Communicated by George R. Stark, Cleveland Clinic Foundation, = Cleveland,=20 OH, June 29, 1999 (received for review March 8, 1999) =


    ABSTRACT
TOP
ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
RESULTS
DISCUSSION
REFERENCES

The early events in neoplastic transformation can be understood only = by=20 comparison of the neoplastic cell with its nontransformed=20 counterpart. The most common central nervous system gliomas=20 traditionally are thought to arise from mature astrocytes and = oligodendrocytes. We examined the possibility that gliomas = arise from=20 a population of glia that has properties of oligodendrocyte=20 progenitors. These glial cells express the NG2 chondroitin = sulfate=20 proteoglycan and the 3D"alpha receptor of platelet-derived growth factor in = vivo.=20 We identified NG2 and the 3D"alpha receptor of platelet-derived growth factor = expression=20 in tissue from seven of seven oligodendrogliomas, three of = three=20 pilocytic astrocytomas, and one of five glioblastoma = multiforme.=20 These data provide evidence that glial tumors arise from = glial=20 progenitor cells. Molecules expressed by these progenitor = cells=20 should be considered as targets for novel therapeutics. =


    INTRODUCTION
TOP
ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
RESULTS
DISCUSSION
REFERENCES

In 1926, Bailey and Cushing classified glial neoplasms on the = basis of=20 the morphological similarities of the tumor cells to = nonneoplastic=20 cells (1).=20 Although the classification and nomenclature have evolved = over the=20 years, this general principle is still applied. The most = common=20 gliomas, astrocytoma [including glioblastoma multiforme = (GBM)] and=20 oligodendroglioma (OLIGO), are defined as being composed of=20 neoplastic astrocytes and oligodendrocytes, respectively (2, = 3). The = implication of such definitions is that these neoplasms = originate=20 from mature cell types. The alternative possibility, that = gliomas=20 arise from a dividing progenitor cell, has been raised on = numerous=20 occasions (4-12); = however,=20 the testing of this hypothesis has been limited by the lack = of=20 phenotypic markers that identify specific progenitor cell = populations=20 in tissue sections.

One progenitor that has attracted much interest is the O-2A = progenitor (13). = In=20 vitro studies have shown that this cell is responsive to = the=20 platelet-derived growth factor (14-16) = because of its expression of the 3D"alpha receptor (PDGF3D"alpha-R) (17). = O-2A=20 progenitors also express the NG2 chondroitin sulfate = proteoglycan (18-20).= =20 Both NG2 and PDGF3D"alpha-R can be=20 detected reliably in tissue sections (18). = Although=20 these antigens are expressed on a variety of = nonneuroectodermal cells=20 (21-23),= glia=20 that coexpress these antigens represent a unique cell = population.=20 These cells are abundant throughout the neuroaxis (18, 24, 25) = and show=20 evidence of DNA synthesis even in adulthood (24). = NG2 is not=20 expressed by mature oligodendrocytes, astrocytes, or = microglia (18, 26). = Some=20 NG2+ cells also express oligodendrocyte markers in a spatial = and=20 temporal pattern that closely precedes myelination, = indicating that=20 these cells are oligodendrocyte progenitors (24, 27); = however,=20 NG2+ cells may have additional functions (20).=20 NG2+ glia recently have been demonstrated in adult human = brain=20 tissue sections (see Results). In pathological = conditions such=20 as multiple sclerosis, individual cells are stained more = intensely by=20 NG2 and PDGF3D"alpha-R than=20 cells in normal adult brain (A.C. and B.D.T., = unpublished=20 observation).

The response of different gliomas to therapy shows a correlation with = cell=20 lineage. For example, patients with anaplastic = oligodendroglioma=20 (AOLIGO) have a higher response rate to chemotherapy (28) = than do=20 patients with tumors thought to be derived from astrocytes. = Most=20 recently, this chemoresponsiveness has been correlated with = specific=20 genetic changes (29). = These=20 observations warrant renewed investigation into the cellular = origin=20 of various glial tumors by using these more recently = characterized=20 cell lineage markers. In the present study we have used=20 immunohistochemistry and immunoblotting to detect NG2 and = PDGF-R in OLIGO and different types of=20 astrocytoma.


    MATERIALS AND = METHODS
TOP
ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
RESULTS
DISCUSSION
REFERENCES

Antibodies. Antibodies from the following sources = were=20 used: monoclonal anti-human melanoma-associated chondroitin sulfate=20 proteoglycan [mAb 9.2.27, 1:2,000; R. Reisfeld, = Scripps=20 Research Institute, La Jolla, CA, (30, 31)]; = rabbit=20 553 anti-rat NG2 antibody (1:2,000; = W. B. Stallcup,=20 Burnham Institute, La Jolla, CA); rabbit anti-human PDGF-R [R7, 1:2,000; C.-H. Heldin, Ludwig Institute,=20 Uppsala, Sweden (32)]; = rabbit=20 anti-glial fibrillary acidic protein (GFAP, 1:5,000, Dako);=20 monoclonal anti-GFAP (1:2,500, Dako); monoclonal anti-myelin = basic=20 protein (MBP, 1:2,000; Dako); and leukocyte common antigen = (LCA,=20 1:40; Dako).

Tissue Processing. Brain tumor samples were = obtained by=20 biopsy from 16 patients. These studies were approved by the = Institutional=20 Review Board of the Cleveland Clinic Foundation, and the = specimens=20 were obtained after representative sampling for diagnostic = purposes.=20 Original slides were reviewed by two neuropathologists = (S.M.S. and=20 S.M.). The diagnoses were: four OLIGO, three AOLIGO, three=20 pilocytic astrocytoma (PA), one fibrillary astrocytoma (A), = and five=20 GBM. The tissues collected were snap-frozen in liquid = nitrogen=20 and stored at 3D-=2070=B0C.=20 Portions of five of the tumors were fixed immediately in 4%=20 paraformaldehyde at 4=B0C overnight and cryoprotected in = 0.1 M=20 phosphate buffer, pH 7.6, containing 20% glycerol at 4=B0C = until=20 used for sectioning (27). =

Histochemical Studies. Unfixed, frozen samples = were thawed=20 in 4% paraformaldehyde overnight at 4=B0C and cryoprotected. Sections = (30 =B5m=20 thick) were obtained by using a sliding microtome (Leica SM = 2000R)=20 and maintained in cryostorage solution (27). = Sections=20 adjacent to those used for immunostaining were dried onto = slides at=20 50=B0C for 30 min, stained with hematoxylin/eosin, and = compared=20 with the sections used for diagnosis. Only those samples that = contained tumor were included in the study. One AOLIGO (no. = 10), one=20 GBM (no. 4), and the A (no. 24) consisted of gray matter = infiltrated=20 by variable numbers of neoplastic cells. One specimen from a = GBM (no.=20 57) showed low-grade histology only. The high-grade areas of = the=20 AOLIGO were focal in the paraffin sections and could not = readily be=20 identified in the specimens used in our studies. =

Immunohistochemistry. Sections were rinsed in PBS = and=20 treated with 10% Triton X-100 and 3% H2O2 for = 30 min=20 at room temperature. Sections were stained by the avidin-biotin complex=20 procedure as described previously (27). = Tissue=20 sections were incubated overnight at 4=B0C in primary antibody diluted = in PBS=20 containing 3% normal goat serum. Diluent without primary antibody served = as a=20 negative control. Sections were photographed by using an Axiophot = microscope=20 (Zeiss). Five OLIGO were analyzed further by = double-immunofluorescence=20 labeling. The antibody combinations examined were: NG2 and PDGF3D"alpha-R,=20 GFAP, or LCA and PDGF3D"alpha-R and GFAP.=20 Sections were processed as described above, and the primary=20 antibodies were applied to the sections simultaneously for = 1-5 days.=20 Sections were incubated with a mixture of biotinylated = anti-mouse=20 (1:500; Vector Laboratories) and fluorescein-conjugated = anti-rabbit=20 antibody (1:500; Jackson ImmunoResearch) for 2 hr at = room=20 temperature, followed by incubation in Texas red-conjugated = avidin D=20 (1:500; Vector Laboratories) for 1 hr at room = temperature. For=20 NG2 and LCA double-labeling, biotinylated anti-mouse IgG2a = and=20 fluorescein-conjugated anti-mouse IgG1 (Southern = Biotechnology=20 Associates) were used. Sections were rinsed, mounted in = Vectashield=20 (Vector Laboratories), and analyzed by confocal microscopy=20 (Aristoplan; Leica).

Immunoprecipitations and Western Blots. Cultured = cells and=20 frozen tissues were used for biochemical studies. MG63 human = osteosarcoma cells=20 and B49 rat glioma cells express NG2, whereas U251MG human = glioma=20 cells do not express NG2 (19). = To verify=20 the presence of tumor tissue, frozen samples (3D"approx1=20 cm3) were sectioned on a cryostat (Leica) along their = peripheral=20 surfaces and stained with hematoxylin/eosin. Extracts from = the=20 occipital cortex of an epileptic patient and normal gray and = white=20 matter from a multiple sclerosis patient were used as = nontumor=20 controls. Cells and tissues were lysed in 50 mM Tris=B7HCl, = pH=20 8/150 mM NaCl/1% NP40/1 mM PMSF/10 =B5g/ml leupeptin/2 = =B5g/ml=20 aprotinin for 30 min at room temperature. To remove=20 glycosaminoglycan chains from NG2, 0.1 unit/ml = chondroitinase=20 ABC (EC 4.2.2.4; ICN) was added during the extraction. For=20 immunoprecipitation, chondroitinase-treated extracts were=20 precipitated with mAb 9.2.27 by using protein A-Sepharose. = Total=20 extracts and immunoprecipitates were resolved on 4-12% = polyacrylamide=20 gradient gels under reducing conditions. Equal amounts of = protein=20 from tissue extracts, as determined by the Lowry method, were = loaded in each well. After electrophoresis, proteins were = transferred=20 to PVDF Immobilon membranes (Millipore, MA) in a buffer = containing=20 25 mM Tris=B7HCl, 192 mM glycine, and 10% methanol. The=20 membranes were blocked for 2 hr at room temperature in = 5% nonfat=20 dry milk in PBS and then incubated with the rabbit = 553 anti-rat=20 NG2 antibody (1:2,000) or R7 anti-human PDGF3D"alpha-R antibody=20 (1:1,000) diluted in blocking solution at 4=B0C overnight. = After four=20 washes, the membranes were incubated with horseradish=20 peroxidase-labeled goat anti-rabbit immunoglobulins = (1:10,000;=20 Bio-Rad). Labeled bands were visualized by using ECL = detection=20 reagents (Amersham).


    RESULTS
TOP
ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
RESULTS
DISCUSSION
REFERENCES

Characterization of NG2 Antibodies. The mAb=20 9.2.27 recognizes the core protein of a proteoglycan originally=20 characterized in malignant melanoma (30). = The=20 amino acid sequence of this proteoglycan has 84% identity = with=20 that of rat NG2 and is considered to be the human homologue = of=20 NG2 (33, 34). = Because mAb=20 9.2.27 does not recognize denatured NG2, the = 553 rabbit=20 anti-rat NG2 antibody was used for Western blots. To = demonstrate that=20 these two antibodies recognize the same molecule, the=20 553 antibody was used to probe a Western blot containing = immunoprecipitates from MG63 cells that had been formed with = mAb=20 9.2.27. As shown in Fig. 1a, lane=20 5, mAb 9.2.27 precipitated a 300-kDa species that = was=20 detected by the 553 antibody and had the same = electrophoretic=20 mobility as the NG2 core protein in total extracts from MG63 = (lane 4)=20 and B49 cells (lanes 2 and 3). The U251MG cell line, = which does=20 not express NG2, was used as a negative control.



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  		Fig. 1.   = (a)=20 mAb 9.2.27 and rabbit 553 anti-rat NG2 antibodies=20 recognize the same NG2 core protein. Extracts from human = U251MG,=20 MG63 and rat B49 cells lines before (3D-=20) and after (+) chondroitinase ABC (ch) = treatment=20 were subjected to Western blot by using the rabbit = 553 anti-rat=20 NG2 antibody. Lane 5 shows immunoprecipitates from MG63 = cells=20 that were formed with mAb 9.2.27. (b) mAb=20 9.2.27 recognizes complex process-bearing cells in = human=20 cerebral cortex resected for intractable epilepsy. = (c-f)=20 OLIGO express NG2 and PDGF3D"alpha-R. At low magnification, NG2 = immunostaining=20 (c) demarcates a hypercellular neoplastic area. High=20 magnification of hematoxylin/eosin-stained tissue (d) = demonstrates densely packed round cells typical of OLIGO. = The=20 majority of neoplastic cells show a membrane-labeling = pattern with=20 NG2 (e) and PDGF3D"alpha-R+ (f) antibodies. Tumor = 30. [Bar =3D 35 =B5m (b), = 120 =B5m=20 (c), and 50 =B5m (d-f).]

NG2 Proteoglycan and PDGF3D"alpha-R=20 Immunohistochemistry. NG2+ cells are not detectable in = normal=20 human brain under the conditions that routinely demonstrate NG2 = immunoreactivity=20 in gliomas. Prolonged incubations of normal tissues with increased = antibody=20 concentrations reveals an abundant population of process-bearing cells = in both=20 gray and white matter (Fig. 1b). The=20 cytoplasmic morphology of these cells is identical to that observed in = rodent=20 NG2+ cells (18), = and they=20 are morphologically and immunohistochemically distinct from human = astrocytes and=20 microglia (A.C. and B.D.T., unpublished results; see also Figs. 2 and = 3). All=20 OLIGO and PA demonstrated intense NG2 immunoreactivity on neoplastic = cells=20 (Table 1).=20 This is best illustrated in sections representing the = interface=20 between neoplastic foci and normal-appearing tissue (Fig. 1c). The=20 neoplastic cells of OLIGO are small and round by routine = histology=20 (Fig. 1d). Cells=20 with the same morphology show predominantly cell-surface = staining=20 with antibodies to NG2 and PDGF3D"alpha-R=20 (Fig. 1=20 e and f). In sections in which the tumor mass = was=20 adjacent to infiltrated brain parenchyma, neoplastic cells = showed a=20 few short, thin processes. In tumor 10, which contained = only=20 infiltrated gray matter, scattered NG2+ cells with short = processes=20 were observed. The morphology of these cells was distinct = from=20 resident NG2+ cells in normal brain (Fig. 1b) and=20 also from that observed in MS lesions and at the edge of = high-grade=20 neoplasms composed of NG23D-=20=20 cells (data not shown). In PA, intense membrane staining was = seen in=20 the microcystic areas of the tumor. More diffuse NG2 = immunoreactivity=20 was seen in the compact bipolar areas of the tumor. One GBM = (no.=20 54) showed numerous NG2+ tumor cells; the other GBM and = the A=20 showed no NG2 immunoreactivity. NG2 immunoreactivity was seen = in=20 capillaries of many of the tumors irrespective of diagnosis = (Table 1). = Capillary=20 staining was seen only in regions containing tumor. Tumors = with=20 NG2+ capillaries also contained areas with NG23D-=20=20 capillaries.



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  		Fig. 2.   = OLIGO=20 cells coexpress NG2 and PDGF3D"alpha-R but not GFAP or LCA. (a-c) = Sections=20 from an OLIGO double-labeled with antibodies to PDGF-R (a, green) and NG2 (b, = red). Merged=20 images demonstrate colocalization of NG2 and PDGF3D"alpha-R (c, yellow). (d-f)=20 Double-labeling with antibodies to PDGF3D"alpha-R (d, green) and GFAP (e, = red). No=20 colocalization is observed in the merged image (f).=20 (g-i) Double-labeling with antibodies to NG2 = (g, red)=20 and LCA (h, green). No colocalization is observed in = the=20 merged image (i). Tumor = 3. [Bar =3D 40 =B5m=20 (a-c), 30 =B5m (d-f), and = 20 =B5m=20 (g-i).]



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  		Fig. 3.   = OLIGO=20 contain reactive astrocytes, neoplastic GFAP+ = cells,=20 residual myelin, and activated microglia. GFAP+ = cells=20 were present in all OLIGO (a and b). Many had = the=20 appearance of reactive astrocytes (a, arrowhead). = Occasional=20 cells had one or two short GFAP+ processes = (b,=20 arrowheads) and may reflect neoplastic cells. (c) MBP = antibodies stained myelin in normal brain adjacent to OLIGO=20 (Right) and residual myelin within the tumor = (Left),=20 but not neoplastic cells. Arrowheads denote the border = between tumor=20 and adjacent normal brain. (d) = LCA+-activated=20 microglia were seen frequently within the neoplasms. = (a and=20 b) Tumor 3. (c and d) Tumor=20 30. [Bar =3D 70 =B5m (a and = b),=20 100 =B5m (c), and 60 =B5m (d).]


        &= nbsp;           &n= bsp;         
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  		Table=20 1.   NG2 and PDGF3D"alpha-R immunostain and Western blot analysis of = 16 brain tumors=20 =

Five of seven OLIGO, three of three PA, and one of five GBM = contained a=20 large number of cells that stained with anti-human PDGF-R antibody (Fig. 1f). The=20 two OLIGO that did not show immunoreactivity on tissue = sections=20 showed clear, strong bands on a Western blot (see below). The = lack of=20 staining in these cases may be due to overfixation of the = specimen or=20 other technical reasons. The morphology of the PDGF3D"alpha-R+ tumor cells was similar to that of = NG2+=20 tumor cells (Fig. 1 = e and=20 f).=20

Double-immunofluorescence labeling by using antibodies against = human NG2=20 and PDGF3D"alpha-R was performed on tumors=20 2, 3, 30, 39, and 44. Confocal=20 laser-scanning microscopy revealed extensive colocalization = of these=20 two antigens (Fig. 2 = a-c).=20

GFAP, MBP, and LCA Immunostaining. GFAP = reactivity=20 was detected in normal glia and tumor cells of most of the = OLIGO=20 examined in this study. Some of the = GFAP+ cells=20 in the OLIGO had long, ramified processes with large cell = bodies,=20 suggesting that they are reactive astrocytes (Fig. 3a). Some=20 OLIGO contained GFAP+ = cells with=20 one or two short processes and nuclei identical to those of neoplastic=20 GFAP3D-=20 cells (Fig. 3b). The=20 morphology is similar to the glial fibrillary oligodendrocytes described = previously in OLIGO (35).=20 Double-labeling of five OLIGO with antibodies to=20 GFAP and NG2 or GFAP and PDGF3D"alpha-R failed to=20 reveal colocalization in all tumors examined (Fig. 2=20 d-f). Immunohistochemistry for MBP was performed in = five=20 OLIGO. Brain adjacent to tumor showed immunoreactivity in myelinated = axons=20 (Fig. 3c).=20 Residual myelinated fibers within the tumor mass also were = stained=20 (Fig. 3c). MBP=20 antibodies did not label neoplastic cells in any of the = OLIGO=20 examined.

Antibodies to LCA were used to identify bone = marrow-derived=20 cells including microglia in three OLIGO. Variable numbers of=20 ramified and amoeboid microglia were present within the tumor = tissues (Fig. 3d).=20 Double-labeling by using antibodies to NG2 and LCA showed no=20 colocalization of these antigens (Fig. 2=20 g-i), indicating that these antibodies = recognize=20 nonoverlapping populations of cells.=20

Western Blot Analysis. Immunoblot analysis of = solubilized=20 extracts from brain tumors was carried out to confirm the presence of = PDGF-R and NG2. The R7 anti-human PDGF3D"alpha-R antibody=20 recognized a major component of approximately 180 kDa in extracts = from MG63=20 human osteosarcoma cells, the expected size for the mature PDGF3D"alpha-R protein,=20 and a faster migrating band at 140 kDa (Fig. 4 = Upper),=20 as described previously (19). = No clear=20 band was observed in the nonneoplastic human brain extracts (Fig. 4 = Upper).=20 Detectable levels of PDGF3D"alpha-R were=20 identified in all tumor samples. Intense bands were detected in four of = seven=20 OLIGO, two of two PA, one of five = GBM,=20 and one of one A (Table 1). = There was no=20 significant difference in the range of band intensities or the average = intensity=20 seen in the different tumors (Fig. 4=20 Upper). Blots also were probed with rabbit anti-rat NG2 = antibody.=20 A 300-kDa NG2-immunoreactive band was detected in = chondroitinase-treated=20 extracts from MG63 cells and two of seven OLIGO, two of two = PA,=20 and three of five GBM (Fig. 4 = Lower).=20 These data support the interpretation that the = immunoreactivity=20 observed in the tissue sections represents the NG2 core = glycoprotein.=20 We also observed strong bands migrating at 60-100 kDa. It is = possible=20 that these bands represent degradation products of NG2 = because of=20 delays in freezing of the tissue at the time of surgery. The = NG2=20 identified in Western blots of tissue extracts represents the = sum of=20 the NG2 produced by the tumor cells and by endothelial cells. = Therefore, the amount of NG2 on the blots is not directly=20 proportional to levels of tumor cell-derived NG2.



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  		Fig. 4.   = Western=20 blots detect PDGF3D"alpha-R and NG2 in gliomas. (Upper) = Although barely=20 visible in normal brain tissue, PDGF3D"alpha-R was detectable in extracts from all = tumors=20 examined. (Lower) A 300-kDa band corresponding to the = NG2=20 core glycoprotein was present in half of the tumors.=20 Lower-molecular-weight bands also were seen (see text). B, = normal=20 brain; M, human MG63 osteosarcoma cells.


    DISCUSSION
TOP
ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
RESULTS
DISCUSSION
REFERENCES

All OLIGO and PA examined in this study consisted of cells with = surface=20 antigens characteristic of oligodendrocyte progenitor cells. = There=20 are several possible interpretations of these results: these = tumors=20 arise from NG2+/PDGF3D"alpha-R+ cells, they arise from a less mature = progenitor=20 cell that acquires NG2 and PDGF3D"alpha-R=20 immunoreactivity, or they arise from mature cells that = acquire NG2=20 and PDGF3D"alpha-R immunoreactivity as they dedifferentiate. = We prefer=20 the first interpretation for the following reasons. In the = adult=20 mammalian brain, NG2+/PDGF3D"alpha-R+ progenitor cells are abundant, are = located=20 throughout the neuroaxis, and retain the ability to divide. = Cell=20 division is an important characteristic for tumorigenicity = because it=20 perpetuates alterations in growth-regulating genes. In = experimental=20 animals, for example, proliferating subventricular zone cells = are=20 more susceptible to chemically and virally induced = oncogenesis than=20 less proliferative brain regions (8, 9).=20 NG2+/PDGF3D"alpha-R+ cells, therefore, have a greater = predisposition to=20 oncogenesis than the postmitotic oligodendrocyte. = Collectively, these=20 observations support the hypothesis that = NG2+/PDGF-R+ cells are the precursors of these = gliomas. Whether=20 or not these neoplasms arise from a progenitor to the=20 NG2+/PDGF3D"alpha-R+ cell can be tested only when markers for = such=20 progenitors are discovered. Dedifferentation of = oligodendrocytes to=20 cells with progenitor phenotypes has not been described in = vitro or in vivo; therefore, it is unlikely that = OLIGO=20 arise from mature oligodendrocytes that lose their myelin = markers=20 while regaining characteristics of their precursors. Our = results=20 confirm previous reports (36-38)<= SUP>=20 that mature oligodendrocyte markers are not expressed by the=20 neoplastic cells in OLIGO. Recently, mRNA encoding=20 oligodendrocyte-specific proteins was reported in OLIGO (39), = however our=20 results and other studies (40) = indicate=20 that these myelin protein gene products are located within=20 preexisting myelinating oligodendrocytes.

A provocative finding of this study was the detection of high levels = of NG2=20 and PDGF3D"alpha-R in PA. Although OLIGO and PA are viewed = as distinct=20 clinical and pathological entities, they share several = interesting=20 properties (3). = Both are=20 typically slowly growing tumors and have better prognoses = than=20 fibrillary astrocytomas. Both PA and low-grade OLIGO are=20 cytologically benign but often histologically heterogeneous. = PA=20 characteristically have a biphasic histology composed of = compact=20 areas containing bipolar cells and microcystic areas composed = of=20 stellate cells. Some PA can even have areas histologically = identical=20 to OLIGO (3).=20 Our results show that antibodies to NG2 and PDGF3D"alpha-R labeled=20 the round cells of OLIGO and both the stellate cells and = bipolar=20 cells of PA. These observations suggest that NG2+ = cells=20 have the potential give rise to neoplasms with distinctive = clinical=20 and pathological phenotypes, perhaps as a result of different=20 environmental influences within the brain parenchyma or = different=20 acquired genetic alterations. In support of environmental = influences,=20 PA are most commonly located in the cerebellum, optic nerve, = and=20 hypothalamus, whereas OLIGO usually are located in the = cerebral=20 hemispheres. PA also typically occur at an earlier age than = OLIGO=20 (3).=20 Genetic changes are rarely described in PA, whereas loss of=20 heterozygosity in chromosomes 1p and 19q are common in OLIGO = (41-43).= These=20 genetic alterations may predispose OLIGO to histological = progression,=20 which is rare in PA.

NG2 immunoreactivity was observed in four of five GBM examined; = however, in=20 three of these cases, the staining was restricted to = capillary=20 endothelial cells. The expression of NG2 by endothelial cells = most=20 likely accounts for the NG2 seen in our Western blots of GBM. = Strong=20 staining of capillaries and weak staining of tumor cells with = antibody to NG2 has been reported previously (31).=20 In our series, one of five GBM showed strong immunoreactivity = for both NG2 and PDGF3D"alpha-R on the=20 majority of tumor cells. This GBM was histologically=20 indistinguishable from the others examined. High expression = of=20 PDGF3D"alpha-R was detected on Western blots of two = additional=20 astrocytomas (one low grade and one GBM). Although we did not = observe=20 PDGF3D"alpha-R immunoreactivity on tissue sections from = these=20 tumors, we believe this was a result of overfixation. High = expression=20 of PDGF3D"alpha-R mRNA in a variety of glial neoplasms is = well=20 documented in the literature (12, 44-48),= and the=20 possibility that endogenous PDGF3D"alpha-R+ cells are the precursors to gliomas has = been noted=20 previously (12, 48). = Detection=20 of both NG2 and PDGF3D"alpha-R on a=20 subset of GBM, in addition to OLIGO and PA, supports the = notion that=20 multiple types of glial neoplasms arise from the=20 NG2+/PDGF3D"alpha-R+ progenitor cell. Gliomas that do not = express NG2=20 and PDGF3D"alpha-R may have down-regulated these markers = because of=20 acquired genetic changes or environmental influences. = Alternatively,=20 they may have arisen from a different progenitor. =

At least three distinct glial progenitor cells may exist in the adult = central=20 nervous system. The NG2+/PDGF3D"alpha-R+ cells have been studied extensively = in=20 vitro, where they have been referred to as O-2A = progenitor cells=20 because they can differentiate into oligodendrocytes in = serum-free=20 medium and type 2 astrocytes in serum-containing medium = (13). = To date,=20 bipotentiality of NG2+/PDGF3D"alpha-R+ cells has not been demonstrated in the = normal=20 brain in vivo; however, as discussed above, neoplastic = NG2+/PDGF3D"alpha-R+ cells may have the potential to display = either=20 OLIGO or PA characteristics. A number of gliomas show a mixed = OLIGO/fibrillary astrocytoma phenotype (3). = Loss of=20 heterozygosity of chromosomes 1p and 19q occurs in both the = OLIGO and=20 the astrocytoma areas of the tumors, indicating that some = OLIGO and=20 fibrillary astrocytomas have a common origin (43). = In=20 vitro studies have characterized another astrocyte, = termed type=20 1, which does not arise from the O-2A progenitor cell = (13). = Studies on=20 the type 1 astrocyte progenitor have lagged, and = phenotypic=20 markers for this progenitor have not been reported. It is = possible=20 that this cell also is present in the adult central nervous = system=20 and is the source of tumors with more astrocytic = characteristics.=20 Other possible progenitors in adult brain include multipotent = glial=20 stem cells and multipotent neuroepithelial stem cells (49, 50). = Such cells=20 have the capacity to differentiate when injected into = neonatal rodent=20 brain (49,=20 51).=20 Further characterization of these progenitor cells in vitro=20 and in vivo is essential for testing their potential role = in=20 glial tumorigenesis. In addition, application of progenitor = cell=20 culture techniques to primary glioma specimens may provide = better=20 in vitro models for glioma research (for review see = ref. 11). =

Studying glial tumors from the viewpoint of progenitor cell biology = will help=20 identify markers that may facilitate diagnosis, predict = prognosis,=20 and direct therapy. This is particularly important for OLIGO = because=20 there is often disagreement among pathologists regarding the = criteria=20 for diagnosis of this glioma (52). = The=20 most common differential diagnosis is OLIGO vs. fibrillary=20 astrocytoma. The majority of patients with anaplastic OLIGO = show a=20 response to chemotherapy (28, 29), = but this=20 option might not be offered to a patient given a diagnosis of = astrocytoma. Our results also should stimulate research into = new=20 treatments. For example, cell surface molecules such as NG2 = and=20 PDGF3D"alpha-R could be targets for cytotoxic therapies. = From a=20 broader perspective, investigation of the pathways that = regulate the=20 proliferation, survival, or migration of normal progenitor = cells may=20 identify additional targets for therapy of = gliomas.=20


    ACKNOWLEDGEMENTS

We thank Drs. W. B. Stallcup, C.-H. Heldin, and = R. Reisfeld=20 for antibodies and Dr. Xinghua Yin for figure preparation. Dr. = M=F6rk=20 was on sabbatical leave from the Department of Pathology, = University=20 of Bergen, Norway. Dr. Nishiyama's current address is = Department of=20 Physiology and Neurobiology, University of Connecticut, = Storrs, CT.=20 This work was supported by National Institutes of Health = Grant=20 NS35058 (B.D.T.) and by the John Gagliarducci Fund (Y.S.). = This paper=20 is dedicated to the memory of Tim Gullickson.


    ABBREVIATIONS

PDGF3D"alpha-R, 3D"alpha = receptor of=20 platelet-derived growth factor; GBM, glioblastoma multiforme; OLIGO,=20 oligodendroglioma, AOLIGO, anaplastic oligodendroglioma; GFAP, glial = fibrillary=20 acidic protein; MBP, myelin basic protein; LCA, leukocyte common = antigen; PA,=20 pilocytic astrocytoma; A, fibrillary astrocytoma.


    FOOTNOTES

3D"Dagger Present address: Department of Neurosurgery, = Hadassah=20 University Hospital, Jerusalem, Israel 91120.=20

=A7 Present address: Department of = Physiology and=20 Neurobiology, University of Connecticut, Storrs, CT 06269.=20

=B6 To whom reprint requests should be = addressed=20 at: Department of Neurosciences #NC30, Cleveland Clinic Foundation,=20 9500 Euclid Avenue, Cleveland, OH 44195. E-mail: = staugas{at}ccf.org .=20


    REFERENCES
TOP
ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
RESULTS
DISCUSSION
REFERENCES =

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Copyright =A9=20 1999 by The National Academy of Sciences=20  0027-8424/99/9610361-6$2.00/0
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Opera = requires=0A= the Sun Java Runtime Environment .=0A= =0A= by Andrew Gregory=0A= http://www.scss.com.au/family/andrew/webdesign/xmlhttprequest/=0A= =0A= This work is licensed under the Creative Commons Attribution License. To = view a=0A= copy of this license, visit = http://creativecommons.org/licenses/by-sa/2.5/ or=0A= send a letter to Creative Commons, 559 Nathan Abbott Way, Stanford, = California=0A= 94305, USA.=0A= =0A= Attribution: Leave my name and web address in this script intact.=0A= =0A= Not Supported in Opera=0A= ----------------------=0A= * user/password authentication=0A= * responseXML data member=0A= =0A= Not Fully Supported in Opera=0A= ----------------------------=0A= * async requests=0A= * abort()=0A= * getAllResponseHeaders(), getAllResponseHeader(header)=0A= =0A= */=0A= // IE support=0A= if (window.ActiveXObject && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= var msxmls =3D new Array(=0A= 'Msxml2.XMLHTTP.5.0',=0A= 'Msxml2.XMLHTTP.4.0',=0A= 'Msxml2.XMLHTTP.3.0',=0A= 'Msxml2.XMLHTTP',=0A= 'Microsoft.XMLHTTP');=0A= for (var i =3D 0; i < msxmls.length; i++) {=0A= try {=0A= return new ActiveXObject(msxmls[i]);=0A= } catch (e) {=0A= }=0A= }=0A= return null;=0A= };=0A= }=0A= // Gecko support=0A= /* ;-) */=0A= // Opera support=0A= if (window.opera && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= this.readyState =3D 0; // = 0=3Duninitialized,1=3Dloading,2=3Dloaded,3=3Dinteractive,4=3Dcomplete=0A= this.status =3D 0; // HTTP status codes=0A= this.statusText =3D '';=0A= this._headers =3D [];=0A= this._aborted =3D false;=0A= this._async =3D true;=0A= this._defaultCharset =3D 'ISO-8859-1';=0A= this._getCharset =3D function() {=0A= var charset =3D _defaultCharset;=0A= var contentType =3D = this.getResponseHeader('Content-type').toUpperCase();=0A= val =3D contentType.indexOf('CHARSET=3D');=0A= if (val !=3D -1) {=0A= charset =3D contentType.substring(val);=0A= }=0A= val =3D charset.indexOf(';');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= val =3D charset.indexOf(',');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= return charset;=0A= };=0A= this.abort =3D function() {=0A= this._aborted =3D true;=0A= };=0A= this.getAllResponseHeaders =3D function() {=0A= return this.getAllResponseHeader('*');=0A= };=0A= this.getAllResponseHeader =3D function(header) {=0A= var ret =3D '';=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= if (header =3D=3D '*' || this._headers[i].h =3D=3D header) {=0A= ret +=3D this._headers[i].h + ': ' + this._headers[i].v + '\n';=0A= }=0A= }=0A= return ret;=0A= };=0A= this.getResponseHeader =3D function(header) {=0A= var ret =3D getAllResponseHeader(header);=0A= var i =3D ret.indexOf('\n');=0A= if (i !=3D -1) {=0A= ret =3D ret.substring(0, i);=0A= }=0A= return ret;=0A= };=0A= this.setRequestHeader =3D function(header, value) {=0A= this._headers[this._headers.length] =3D {h:header, v:value};=0A= };=0A= this.open =3D function(method, url, async, user, password) {=0A= this.method =3D method;=0A= this.url =3D url;=0A= this._async =3D true;=0A= this._aborted =3D false;=0A= this._headers =3D [];=0A= if (arguments.length >=3D 3) {=0A= this._async =3D async;=0A= }=0A= if (arguments.length > 3) {=0A= opera.postError('XMLHttpRequest.open() - user/password not = supported');=0A= }=0A= this.readyState =3D 1;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= };=0A= this.send =3D function(data) {=0A= if (!navigator.javaEnabled()) {=0A= alert("XMLHttpRequest.send() - Java must be installed and = enabled.");=0A= return;=0A= }=0A= if (this._async) {=0A= setTimeout(this._sendasync, 0, this, data);=0A= // this is not really asynchronous and won't execute until the = current=0A= // execution context ends=0A= } else {=0A= this._sendsync(data);=0A= }=0A= }=0A= this._sendasync =3D function(req, data) {=0A= if (!req._aborted) {=0A= req._sendsync(data);=0A= }=0A= };=0A= this._sendsync =3D function(data) {=0A= this.readyState =3D 2;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= // open connection=0A= var url =3D new java.net.URL(new = java.net.URL(window.location.href), this.url);=0A= var conn =3D url.openConnection();=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= conn.setRequestProperty(this._headers[i].h, this._headers[i].v);=0A= }=0A= this._headers =3D [];=0A= if (this.method =3D=3D 'POST') {=0A= // POST data=0A= conn.setDoOutput(true);=0A= var wr =3D new = java.io.OutputStreamWriter(conn.getOutputStream(), this._getCharset());=0A= wr.write(data);=0A= wr.flush();=0A= wr.close();=0A= }=0A= // read response headers=0A= // NOTE: the getHeaderField() methods always return nulls for me :(=0A= var gotContentEncoding =3D false;=0A= var gotContentLength =3D false;=0A= var gotContentType =3D false;=0A= var gotDate =3D false;=0A= var gotExpiration =3D false;=0A= var gotLastModified =3D false;=0A= for (var i =3D 0; ; i++) {=0A= var hdrName =3D conn.getHeaderFieldKey(i);=0A= var hdrValue =3D conn.getHeaderField(i);=0A= if (hdrName =3D=3D null && hdrValue =3D=3D null) {=0A= break;=0A= }=0A= if (hdrName !=3D null) {=0A= this._headers[this._headers.length] =3D {h:hdrName, = v:hdrValue};=0A= switch (hdrName.toLowerCase()) {=0A= case 'content-encoding': gotContentEncoding =3D true; break;=0A= case 'content-length' : gotContentLength =3D true; break;=0A= case 'content-type' : gotContentType =3D true; break;=0A= case 'date' : gotDate =3D true; break;=0A= case 'expires' : gotExpiration =3D true; break;=0A= case 'last-modified' : gotLastModified =3D true; break;=0A= }=0A= }=0A= }=0A= // try to fill in any missing header information=0A= var val;=0A= val =3D conn.getContentEncoding();=0A= if (val !=3D null && !gotContentEncoding) = this._headers[this._headers.length] =3D {h:'Content-encoding', v:val};=0A= val =3D conn.getContentLength();=0A= if (val !=3D -1 && !gotContentLength) = this._headers[this._headers.length] =3D {h:'Content-length', v:val};=0A= val =3D conn.getContentType();=0A= if (val !=3D null && !gotContentType) = this._headers[this._headers.length] =3D {h:'Content-type', v:val};=0A= val =3D conn.getDate();=0A= if (val !=3D 0 && !gotDate) this._headers[this._headers.length] = =3D {h:'Date', v:(new Date(val)).toUTCString()};=0A= val =3D conn.getExpiration();=0A= if (val !=3D 0 && !gotExpiration) = this._headers[this._headers.length] =3D {h:'Expires', v:(new = Date(val)).toUTCString()};=0A= val =3D conn.getLastModified();=0A= if (val !=3D 0 && !gotLastModified) = this._headers[this._headers.length] =3D {h:'Last-modified', v:(new = Date(val)).toUTCString()};=0A= // read response data=0A= var reqdata =3D '';=0A= var stream =3D conn.getInputStream();=0A= if (stream) {=0A= var reader =3D new java.io.BufferedReader(new = java.io.InputStreamReader(stream, this._getCharset()));=0A= var line;=0A= while ((line =3D reader.readLine()) !=3D null) {=0A= if (this.readyState =3D=3D 2) {=0A= this.readyState =3D 3;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= }=0A= reqdata +=3D line + '\n';=0A= }=0A= reader.close();=0A= this.status =3D 200;=0A= this.statusText =3D 'OK';=0A= this.responseText =3D reqdata;=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onload) {=0A= this.onload();=0A= }=0A= } else {=0A= // error=0A= this.status =3D 404;=0A= this.statusText =3D 'Not Found';=0A= this.responseText =3D '';=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onerror) {=0A= this.onerror();=0A= }=0A= }=0A= };=0A= };=0A= }=0A= // ActiveXObject emulation=0A= if (!window.ActiveXObject && window.XMLHttpRequest) {=0A= window.ActiveXObject =3D function(type) {=0A= switch (type.toLowerCase()) {=0A= case 'microsoft.xmlhttp':=0A= case 'msxml2.xmlhttp':=0A= case 'msxml2.xmlhttp.3.0':=0A= case 'msxml2.xmlhttp.4.0':=0A= case 'msxml2.xmlhttp.5.0':=0A= return new XMLHttpRequest();=0A= }=0A= return null;=0A= };=0A= }=0A= ------=_NextPart_000_0000_01C85874.E6E50B00 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.pnas.org/javascript/ajax/utility.js /************************************************************************= *****=0A= * javascript/ajax/utility.js=0A= *=0A= * Utility functions for working with XMLHttpRequest data.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Copy XML nodes into an HTMLElement. This effectively=0A= * clones XML markup which uses XHTML naming conventions=0A= * into an HTML DOM.=0A= */=0A= function copy_xml_to_html(src, dst) {=0A= if (src.nodeType =3D=3D 1) { /* Node.ELEMENT_NODE */=0A= var e =3D document.createElement(src.nodeName);=0A= for (var i =3D 0; i < src.childNodes.length; i++) {=0A= copy_xml_to_html(src.childNodes[i], e);=0A= }=0A= for (var i =3D 0; i < src.attributes.length; i++) {=0A= var n =3D src.attributes[i].name;=0A= var v =3D unescape_xml_string(src.attributes[i].value); =0A= e.setAttribute(n, v);=0A= if (n =3D=3D "class") {=0A= e.className =3D v;=0A= }=0A= else if (n =3D=3D "style") {=0A= set_css_style(v, e, "");=0A= }=0A= }=0A= dst.appendChild(e);=0A= }=0A= else if (src.nodeType =3D=3D 3) { /* Node.TEXT_NODE */=0A= dst.appendChild(document.createTextNode(src.nodeValue));=0A= }=0A= }=0A= =0A= /* =0A= * It is unclear that this is the right thing to be calling=0A= * from copy_xml_to_html, but it appears that Safari decides=0A= * to convert & to the NCR #, and then encodes that=0A= * NCR to &%26%2338;. So, I'm going to treat the DOM Attr=0A= * value as a plain string, and run our XML string input=0A= * through the decoding routine below.=0A= */=0A= function unescape_xml_string(s) {=0A= return s.replace(/'/g, "'")=0A= .replace(/'/g, "'")=0A= .replace(/"/g, "\"")=0A= .replace(/"/g, "\"")=0A= .replace(/>/g, ">")=0A= .replace(/>/g, ">")=0A= .replace(/</g, "<")=0A= .replace(/</g, "<")=0A= .replace(/&/g, "&")=0A= .replace(/&/g, "&");=0A= }=0A= =0A= /*=0A= * Parse set of CSS rules and apply them to an element.=0A= * This is quite horrifying, but I'm unable to determine=0A= * how else to handle this with IE 6. FireFox and other=0A= * sane browsers let you simply set the style attribute=0A= * or use e.style.setProperty(rule, value, priority),=0A= * IE 6 appears to have neither of these capabilities..=0A= */=0A= function set_css_style(css, e, priority) {=0A= var rules =3D css.split(";");=0A= for (var i =3D 0; i < rules.length; i++) {=0A= var nvpair =3D rules[i].split(":");=0A= if (nvpair.length =3D=3D 2) {=0A= try {=0A= var name =3D nvpair[0]; /* style attribute */=0A= var value =3D nvpair[1]; /* attribute value */=0A= =0A= /*=0A= * For each possible style attribute, set the=0A= * appropriate style property in the element.=0A= */=0A= if (name =3D=3D "background") {=0A= e.style.background =3D value;=0A= }=0A= else if (name =3D=3D "background-attachment") {=0A= e.style.backgroundAttachment =3D value;=0A= }=0A= else if (name =3D=3D "background-color") {=0A= e.style.backgroundColor =3D value;=0A= }=0A= else if (name =3D=3D "background-image") {=0A= e.style.backgroundImage =3D value;=0A= }=0A= else if (name =3D=3D "background-position") {=0A= e.style.backgroundPosition =3D value;=0A= }=0A= else if (name =3D=3D "background-position-x") {=0A= e.style.backgroundPositionX =3D value;=0A= }=0A= else if (name =3D=3D "background-position-y") {=0A= e.style.backgroundPositionY =3D value;=0A= }=0A= else if (name =3D=3D "background-repeat") {=0A= e.style.backgroundRepeat =3D value;=0A= }=0A= else if (name =3D=3D "behavior") {=0A= e.style.behavior =3D value;=0A= }=0A= else if (name =3D=3D "border") {=0A= e.style.border =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom") {=0A= e.style.borderBottom =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-color") {=0A= e.style.borderBottomColor =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-style") {=0A= e.style.borderBottomStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-width") {=0A= e.style.borderBottomWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-collapse") {=0A= e.style.borderCollapse =3D value;=0A= }=0A= else if (name =3D=3D "border-color") {=0A= e.style.borderColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left") {=0A= e.style.borderLeft =3D value;=0A= }=0A= else if (name =3D=3D "border-left-color") {=0A= e.style.borderLeftColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left-style") {=0A= e.style.borderLeftStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-left-width") {=0A= e.style.borderLeftWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-right") {=0A= e.style.borderRight =3D value;=0A= }=0A= else if (name =3D=3D "border-right-color") {=0A= e.style.borderRightColor =3D value;=0A= }=0A= else if (name =3D=3D "border-right-style") {=0A= e.style.borderRightStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-right-width") {=0A= e.style.borderRightWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-style") {=0A= e.style.borderStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top") {=0A= e.style.borderTop =3D value;=0A= }=0A= else if (name =3D=3D "border-top-color") {=0A= e.style.borderTopColor =3D value;=0A= }=0A= else if (name =3D=3D "border-top-style") {=0A= e.style.borderTopStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top-width") {=0A= e.style.borderTopWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-width") {=0A= e.style.borderWidth =3D value;=0A= }=0A= else if (name =3D=3D "bottom") {=0A= e.style.bottom =3D value;=0A= }=0A= else if (name =3D=3D "clear") {=0A= e.style.clear =3D value;=0A= }=0A= else if (name =3D=3D "clip") {=0A= e.style.clip =3D value;=0A= }=0A= else if (name =3D=3D "color") {=0A= e.style.color =3D value;=0A= }=0A= else if (name =3D=3D "cssText") {=0A= e.style.Sets =3D value;=0A= }=0A= else if (name =3D=3D "cursor") {=0A= e.style.cursor =3D value;=0A= }=0A= else if (name =3D=3D "direction") {=0A= e.style.direction =3D value;=0A= }=0A= else if (name =3D=3D "display") {=0A= e.style.display =3D value;=0A= }=0A= else if (name =3D=3D "font") {=0A= e.style.font =3D value;=0A= }=0A= else if (name =3D=3D "font-family") {=0A= e.style.fontFamily =3D value;=0A= }=0A= else if (name =3D=3D "font-size") {=0A= e.style.fontSize =3D value;=0A= }=0A= else if (name =3D=3D "font-style") {=0A= e.style.fontStyle =3D value;=0A= }=0A= else if (name =3D=3D "font-variant") {=0A= e.style.fontVariant =3D value;=0A= }=0A= else if (name =3D=3D "font-weight") {=0A= e.style.fontWeight =3D value;=0A= }=0A= else if (name =3D=3D "height") {=0A= e.style.height =3D value;=0A= }=0A= else if (name =3D=3D "ime-mode") {=0A= e.style.imeMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-flow") {=0A= e.style.layoutFlow =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid") {=0A= e.style.layoutGrid =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-char") {=0A= e.style.layoutGridChar =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-line") {=0A= e.style.layoutGridLine =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-mode") {=0A= e.style.layoutGridMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-type") {=0A= e.style.layoutGridType =3D value;=0A= }=0A= else if (name =3D=3D "left") {=0A= e.style.left =3D value;=0A= }=0A= else if (name =3D=3D "letter-spacing") {=0A= e.style.letterSpacing =3D value;=0A= }=0A= else if (name =3D=3D "line-break") {=0A= e.style.lineBreak =3D value;=0A= }=0A= else if (name =3D=3D "line-height") {=0A= e.style.lineHeight =3D value;=0A= }=0A= else if (name =3D=3D "list-style") {=0A= e.style.listStyle =3D value;=0A= }=0A= else if (name =3D=3D "list-style-image") {=0A= e.style.listStyleImage =3D value;=0A= }=0A= else if (name =3D=3D "list-style-position") {=0A= e.style.listStylePosition =3D value;=0A= }=0A= else if (name =3D=3D "list-style-type") {=0A= e.style.listStyleType =3D value;=0A= }=0A= else if (name =3D=3D "margin") {=0A= e.style.margin =3D value;=0A= }=0A= else if (name =3D=3D "margin-bottom") {=0A= e.style.marginBottom =3D value;=0A= }=0A= else if (name =3D=3D "margin-left") {=0A= e.style.marginLeft =3D value;=0A= }=0A= else 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