From: Subject: Isolation and Characterization of Tumorigenic, Stem-like Neural Precursors from Human Glioblastoma -- Galli et al. 64 (19): 7011 -- Cancer Research Date: Tue, 15 Jan 2008 15:12:21 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0041_01C85789.06931E20" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.3198 This is a multi-part message in MIME format. ------=_NextPart_000_0041_01C85789.06931E20 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://intl-cancerres.aacrjournals.org/cgi/content/full/64/19/7011 Isolation and Characterization of Tumorigenic, = Stem-like Neural Precursors from Human Glioblastoma -- Galli et al. 64 = (19): 7011 -- Cancer Research 3D"Cancer 3D"
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[Cancer Research 64, 7011-7021, October 1, = 2004]
=A9 2004 American= =20 Association for Cancer Research

Regular Articles

Isolation and Characterization of Tumorigenic, Stem-like Neural = Precursors=20 from Human Glioblastoma

Rossella=20 Galli1, Elena Binda1, = Ugo=20 Orfanelli1, Barbara = Cipelletti1,=20 Angela Gritti1, Simona De=20 Vitis2, Roberta Fiocco1, = Chiara=20 Foroni1, Francesco=20 Dimeco3,4 and Angelo=20 Vescovi1,5

1 Stem Cell Research Institute and = 2=20 Laboratory of Molecular Diagnostics, H. S. Raffaele, Milan, Italy; = 3=20 National Neurological Institute "C. Besta," Milan, Italy; 4=20 Department of Neurological Surgery, Johns Hopkins Medical School, = Baltimore,=20 Maryland; and 5 Department of Biological Sciences and = Biotechnology,=20 University of Milano-Bicocca, Milan, Italy


3D"   =20 ABSTRACT
Top
3D"ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
RESULTS
DISCUSSION
REFERENCES
 
T= ransformed=20 stem cells have been isolated from some human cancers. We = report=20 that, unlike other brain cancers, the lethal glioblastoma = multiforme=20 contains neural precursors endowed with all of the critical = features=20 expected from neural stem cells. Similar, yet not identical, = to their=20 normal neural stem cell counterpart, these precursors emerge = as=20 unipotent (astroglial) in vivo and multipotent=20 (neuronal-astroglial-oligodendroglial) in culture. More = importantly,=20 these cells can act as tumor-founding cells down to the = clonal level=20 and can establish tumors that closely resemble the main = histologic,=20 cytologic, and architectural features of the human disease, = even when=20 challenged through serial transplantation. Thus, cells = possessing all=20 of the characteristics expected from tumor neural stem cells = seem to=20 be involved in the growth and recurrence of adult human = glioblastomas=20 multiforme.


3D"   =20 INTRODUCTION
Top
ABSTRACT
3D"INTRODUCTION
MATERIALS AND METHODS
RESULTS
DISCUSSION
REFERENCES
 
C= ancer=20 arises from a series of mutations occurring in few or single = founder=20 cells. From a therapeutic standpoint, a critical issue = regards the=20 elucidation of the identity and physiology of the cell(s) = responsible=20 for tumor formation, progression, and recurrence (1)=20 . With some exceptions (2)=20 , the identity of tumorigenic cells within many cancers = remains=20 unclear, but recent observations point to the involvement of = somatic=20 stem cells in oncogenesis (3)=20 . Embryonic and fetal stem cells may accumulate mutations = during=20 their expansion/growth phase throughout development and may = form=20 full-blown tumors in adulthood (3)=20 , whereas adult stem cells and their progeny may represent = the=20 direct target of oncogenic transformation in mature tissues = (1)=20 . Accordingly, the involvement of cells displaying a stem = cell=20 phenotype in cancer formation and progression (1)=20 has been confirmed in acute myeloid leukemia (4,=20 5,=20 6)=20 and breast cancer (2)=20 .

If stem cells and/or their progeny represent the main target = of=20 neoplastic transformation, it is then reasonable to infer = that the=20 main foci of malignant transformation ought to coincide with = those=20 tissue regions that embody somatic stem cells. Indeed, the = main areas=20 of distribution of certain brain tumors overlap specific = regions of=20 the central nervous system (CNS; refs. 7=20 and 8=20 ) that comprise glia-like neural stem cells and their progeny = (9=20 ,=20 10) . Accordingly, deregulation of specific proto-oncogenes = such=20 as Ink4A/Arf, epidermal growth factor (EGF) receptor, and = c-myc (11,=20 12,=20 13)=20 or expression of SV40 T antigen (14)=20 in glial fibrillary acidic protein (GFAP)- or = nestin-expressing=20 neural cells induces high-grade gliomas in brain areas that=20 contain neural stem cells.

Recently, transformed, stem-like neural progenitors have been=20 isolated and cultured from human brain tumor tissues (15,=20 16,=20 17)=20 , which raises fundamental issues concerning their actual = ability to=20 drive cancer growth and recurrence (18)=20 .

Here, we report the initial finding that glioblastoma = multiforme,=20 the most malignant among the adult human CNS tumors, = comprises=20 transformed precursors that bear the full complement of = functional=20 characteristics expected from stem cells (19)=20 , including the capacity for tumor generation. In fact, these = cells=20 can establish tumors with all of the classical features of = human=20 glioblastomas multiforme, even upon serial transplantation = and,=20 therefore, can be identified as tumor neural stem cells (20)=20 .


3D"   =20 MATERIALS AND METHODS
Top
ABSTRACT
INTRODUCTION
3D"MATERIALS AND = METHODS
RESULTS
DISCUSSION
REFERENCES
 
<= STRONG>Sample=20 Classification.
All of the samples were classified according = to=20 World Health Organization guidelines. Lines 0627, 0821, 0913, = 1022,=20 and 1205 were obtained from patients with diagnosis of = primary=20 glioblastoma; line 1030 was derived from a patient suffering = from=20 secondary glioblastoma. Patients from whom lines 0627, 0821, = 0913,=20 and 1022 were established underwent recurrence after 6 to 12=20 months post-surgery.

Primary Culture, Culture Propagation, Population Analysis, = and=20 Cloning.
Tumor samples, either derived from patients or from = orthotopic transplantation of glioblastoma = multiforme=96derived=20 cell lines, were processed as by Gritti et al. (21)=20 . Primary cells were plated in 25 cm2 tissue = culture=20 flasks plated at clonal density (2500=965000 = cells/cm2) in=20 Dulbecco=92s modified Eagle=92s medium/F-12 medium containing = 20 ng/mL of=20 both EGF and fibroblast growth factor (FGF2; Peprotech, Rocky = Hill, NY). As reference cell lines, normal human fetal neural = stem cells (22)=20 and the glioblastoma cell line U87MG were used in all of the = assays.=20 Population and serial subclonogenic analysis were performed = as by=20 Galli et al. (23)=20 .

Differentiation of Stem Cell Progeny and=20 Immunocytochemistry.
To assess for multipotency, cells were = plated=20 at a density of 2.5 x=20 104 cells/cm2 onto Matrigel-coated glass = coverslips=20 (12 mm diameter) in the presence of leukemia inhibitory = factor=20 (10 ng/mL; ref. 24=20 ) for 7 to 10 days. Multiple immunofluorescence assays for = neural=20 antigens were performed as described by Gritti et al. = (21)=20 and Galli et al. (23)=20 .

Evaluation of Tumorigenicity by Subcutaneous Injection and by = Orthotopic Injection.
Tumorigenicity was determined by = injecting=20 glioblastoma multiforme=96derived neural stem cells, U87, and = human=20 fetal neural stem cells, either subcutaneously (s.c.) or=20 orthotopically. Cells (3 x = 106)=20 in 100 =B5L of PBS were s.c. injected into the right = flank=20 of Scid/bg mice while 2 =B5L of a 1 x 108 cells/mL cell = suspension=20 in PBS were delivered into the right striatum (0.2 =B5L/min) = by=20 stereotactic injection through a glass electrode connected to = a=20 Hamilton syringe. The following coordinates were used:=20 antero-posterior =3D 0; medio-lateral =3D +2.5 mm; dorso-ventral = =3D =963.5=20 mm. Mice were sacrificed at different times between 1 and 10 = weeks=20 post-injection, according to the cell line originally = injected.=20 Hematoxylin and eosin staining and immunohistochemistry were=20 performed on 15-=B5m-thick cryostat sections. Sections were = processed=20 as by Vescovi et al. (22)=20 . Antibodies/antisera used were as follows: mouse = antivimentin=20 (1:200; Chemicon, Temecula, CA), mouse antihuman mitochondria = (1:20;=20 Chemicon), mouse antihuman nuclei (1:20; Chemicon), mouse=20 anti-Galactocerebroside C (Gal C; 1:20; Chemicon), mouse = antineuronal=20 class III =DF-tubulin (Tuj1; 1:500; Babco, Richmond, CA), = rabbit=20 anti-GFAP (1:1000; Dako Corporation, Carpinteria, CA), rabbit = antilaminin (1:200; Sigma, St. Louis, MO), and rabbit = anti-Ki67=20 (1:1000; NovoCastra, Newcastle, UK).

Chromosome Analysis.
Cells were treated with = medium=20 containing 10 =B5g/mL Colcemid (Irvine Scientific, Santa Ana, = CA) for 1=20 to 2 hours and resuspended in hypotonic 1% sodium citrate at = room=20 temperature for 30 minutes. The cells were then washed in=20 methanol-acetic acid (3:1, v/v) fixative solution for 30 = minutes and=20 spread onto clean dry slides. Q-banding staining was then = performed,=20 and 10 metaphases were analyzed for each sample.

Telomeric Repeat Amplification Protocol and Telomere Length=20 Assessment.
Telomerase activity was detected using the = PCR-based=20 TRAPeze Telomerase Detection kit (Intergen, Purchase, NY),=20 according to the manufacturer=92s instructions. Total = proteins=20 (150 ng) were loaded into each lane. For the telomere length=20 assay, the length of the telomeres was determined using = TeloTAGGG=20 Telomere Length Assay (Roche, Gipf-Oberfrick, Switzerland)=20 according to the manufacturer=92s instructions. Two =B5g of = genomic=20 DNA of each sample were digested with a HinfI/RsaI = mixture=20 for 2 hours at 37=B0C and then loaded into a 0.8% agarose = gel.=20

Molecular Analysis.
One =B5g of total RNA from = tumor neural=20 stem cells, U87, and human fetal neural stem cells, extracted = using=20 the RNeasy Mini kit (Qiagen, Chatsworth, CA), was primed with = oligo(dT) for cDNA synthesis and reverse-transcribed by using = Superscript RNase H=96 Reverse Transcriptase (Life = Technologies, Rockville, MD). All cDNAs used as templates = were=20 previously normalized throughout a =DF-Actin reverse = transcriptase-PCR.=20 Sequence of primers and PCR conditions are available on = request.=20


3D"   =20 RESULTS
Top
ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
3D"RESULTS
DISCUSSION
REFERENCES
 
<= STRONG>Selection=20 and Expansion of Neural Precursors from Adult Human Brain = Tumors.
To=20 assess for the presence of neural precursors within human = brain=20 tumors, cells from post-surgery specimens of low-grade = gliomas,=20 glioblastoma multiformes, or high-grade medulloblastomas were = plated=20 at clonal density (21)=20 in serum-free medium, containing EGF and FGF2. This provides = a=20 stringent, low-density system (<5 x 103 cells/cm2), = which=20 selects away differentiated/differentiating cells in primary = CNS=20 cultures, leaving neural stem cells free to proliferate and = expand=20 exponentially (25=20 ,=20 26) .

Twenty to 40 days after plating, phase-bright clones resembling=20 the classical neurospheres formed in vitro by normal = neural=20 stem cells (ref. 27=20 ; Fig. 1A and B ) were detected in all of the cultures = established from=20 high-grade glioblastoma multiformes and medulloblastomas. = Clonal=20 frequency was between 0.5 and 31% of the total cells plated = for=20 glioblastomas multiforme (n =3D 12) and 50 and 80% for = medulloblastomas (n =3D 4). Conversely, clone = formation was=20 never observed in cultures from either World Health = Organization=20 grade I gliomas or World Health Organization grade II=20 oligodendrogliomas (n =3D 5; up to 90 days in vitro). = These data confirm and extend previous findings (15,=20 16,=20 17)=20 , which show that EGF/FGF2-responsive precursors are=20 specifically found in glioblastomas multiforme and=20 medulloblastomas.



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  		Fig. 1. Analysis of long-term=20 proliferation in glioblastoma multiforme=96derived tumor = neural stem=20 cells. Phase-bright microphotographs showing examples of=20 glioblastoma multiforme=96derived (A) and=20 medulloblastoma-derived (B) neurospheres; x20 objective. C. = Intrinsic=20 differences in the growth rate were observed among cluster = A,=20 consisting of faster-growing tumor neural stem cells such as = 0627,=20 0913, and 1022 (black lines), and cluster B, = comprising=20 slower-expanding glioblastoma multiforme (GBM) cell = lines=20 0821, 1030, and 1205 (red lines). Although = medulloblastoma=20 (MDB) tissues displayed the highest content of = clonogenic=20 cells (up to 80%), they never established tumor neural stem = cells=20 lines (green lines). D. Differently from = normal human=20 fetal neural stem cells (22)=20 , glioblastoma multiforme=96derived tumor neural stem cells = displayed=20 different rates of amplification at low- and = high-subculturing=20 stages, as shown for cell line 0627. E. The exposure = of cell=20 line 0627 to EGF or FGF2 influenced the proliferation rate, = which=20 was higher in the presence of both mitogens and lower when = using=20 only one factor, as shown previously for normal human fetal = neural=20 stem cells (22)=20 . =

 = ;
Glioblastomas=20 Multiforme Contain Multipotent Precursors Endowed with Long-term=20 Self-renewal.
The generation of clones from disaggregated = primary=20 cancers provides an index of the cancer clonogenicity (20)=20 . However, the sole formation of neurospheres in cultured CNS = tumors=20 does not provide, per se, the demonstration of the = presence of=20 tumor neural stem cells therein. Transiently amplifying=20 precursors are also known to produce neurospheres in this = system, and=20 they can even undergo a limited number of passages in culture = (20=20 ,=20 28) . In addition, neurosphere formation may also result=20 from spontaneous cell aggregation, usually as the consequence = of=20 excessive plating density (29)=20 .

Hence, we determined whether the clonogenic cells in our primary = glioblastoma multiforme and medulloblastoma cultures = represented=20 transient clonogens as generated by short-term proliferating, = transit amplifying cells or possessed the cardinal properties = expected from cultured tumor neural stem cells (20)=20 . To this end, we assessed their capacity for long-term=20 proliferation, self-renewal, multipotency (the ability to = generate=20 the three major neural cell types, i.e., neurons, = astroglia,=20 and oligodendroglia) and their tumorigenicity (27=20 ,=20 30, 31,=20 32)=20 , by combining cloning, subcloning, and population analysis = followed=20 by orthotopic, heterotopic, and serial transplantation = assays.=20

One of the primary consequences of dealing with candidate stem=20 cells endowed with extensive self-renewal capacity is the=20 establishment of steadily expanding, stable lines (19=20 ,=20 28) . Surprisingly, although medulloblastoma cultures = displayed=20 the highest clonogenic capacity, they failed in establishing=20 long-term cell lines (four of four specimens) under the = conditions=20 normally used for neural stem cells expansion. In agreement = with=20 previous findings (16=20 ,=20 17) , medulloblastoma cultures could be propagated for a = few=20 passages (Fig. 1C) , although in our hands, they could only = generate=20 Tuj1-immunoreactive cells when triggered to differentiate = (data not=20 shown). Conversely, under identical culture conditions, = one-half of=20 the 12 cultures established from glioblastomas multiforme = comprised=20 multipotent precursors, which established long-term expanding = cultures. These six different glioblastoma multiforme=96derived = lines=20 were expanded for more than 80 passages, with an average = doubling=20 time of 3 to 4 days (i.e., 180=96280 days in vitro), = and=20 were characterized as tumor neural stem cells through the = comparison=20 with normal human fetal neural stem cells (22)=20 and with the well-characterized human glioblastoma cell line=20 U87MG.

Human glioblastoma multiforme cell lines were named after the = date=20 of surgery: 0627, 0821, 0913, 1022, 1030, and 1205. When = their growth=20 rate was assessed, significant differences between specific = lines=20 emerged (Fig. 1C) . Nonetheless, despite the fact that the = kinetics of=20 expansion was characteristic for each tumor cell line and = clearly=20 depended upon the specimen of origin, all of the lines = exponentially=20 expanded in number, similarly to normal human fetal neural = stem cells=20 and U87 cells and as opposed to their medulloblastoma = counterpart=20 (Fig. 1C) . Based on growth properties, two distinct = clusters=20 emerged: cluster A, comprising faster-growing lines 0627, = 0913, and=20 1022; and cluster B, consisting of slower-expanding lines = 0821,=20 1030, and 1205 (Fig. 1C) . Of note, whereas normal human fetal neural = stem cells=20 displayed steady growth capacity over long-term culturing (22=20 ,=20 33) , the proliferative activity of glioblastoma = multiforme=96derived=20 cell lines increased at very late subculturing stages = (120=96150=20 days in vitro) compared with earlier passages (15=9635 = days=20 in vitro; Fig. 1D , in reference to line 0627). This may reflect = their=20 progressive enrichment in cells endowed with aberrant = proliferative=20 advantage, the tumor cell adaptation to the culture = conditions, or=20 the intrinsic, genetic fluctuation expected from tumor cells. = Nonetheless, this phenomenon was the only hint as to a = possible,=20 latent instability of glioblastoma multiforme=96derived cell = lines,=20 which displayed an otherwise remarkably stable cellular and = molecular=20 phenotype and karyotype (see next sections).

Additional proof that tumor neural stem cells were the likely=20 founders of glioblastoma multiforme=96derived cell lines = came=20 from the observation that single glioblastoma multiforme = cells could=20 give rise to clonal cell lines, which behaved identically to = their=20 parental (i.e., bulk culture) cell lines (not shown). = These=20 clonal tumor neural stem cells were used in all of the = experiments=20 described from now on.

We continued by analyzing the response of our candidate tumor=20 neural stem cells to mitogens that regulate proliferation in=20 normal neural stem cells (22=20 ,=20 34 ,=20 35) . As described for adult rodent neural stem cells (34)=20 , glioblastoma multiforme=96derived cell lines expanded = slowly in the=20 presence of the sole FGF2, switched to a faster growth mode = when=20 exposed to EGF alone, and expanded even faster when exposed = to both=20 mitogens simultaneously (Fig. 1E , cell line 0627). Thus, glioblastoma multiforme = cells=20 respond to the same mitogenic cues that control the fate of=20 normal neural stem cells.

We then assessed tumor neural stem cell multipotency by = determining=20 the ability of clonal glioblastoma multiforme=96derived = cells to=20 generate neurons, astrocytes, and oligodendrocytes. Early = passage=20 (passages 10=9615) clonally derived glioblastoma multiforme = cells=20 underwent terminal differentiation upon removal of mitogens = (34=20 ,=20 36) and the subsequent addition of leukemia inhibitory = factor to=20 the medium, as shown previously for human fetal neural stem = cells (24)=20 . Similar to normal neural stem cells (34)=20 , all of the glioblastoma multiforme clonal lines = differentiated into=20 GFAP-positive astrocyte-like cells (Fig. 2B and = F) but also into neuron-like cells, which were = immunoreactive=20 for Tuj1 (Fig. 2A and E) , MAP5 (Fig. 2I) , MAP2 (Fig. 2J) , neurofilament Mr 200,000 = (NF200;=20 Fig. 2K ), glutamate (Fig. 2L) , and 3D{gamma}=20-aminobutyric=20 acid (Fig. 2M) and into GalC-immunoreactive = oligodendrocyte-like cells=20 (Fig. 2D and H) . Conversely, U87 cells never gave rise to = cells=20 immunoreactive for any of the above pan-neural antigens, thus = demonstrating that multipotency is a unique property of = glioblastoma=20 multiforme=96derived cells. Such multipotency was maintained = unaltered=20 even after extensive culturing, up to passage 80 (not shown). = Notably, glioblastoma multiforme=96derived, differentiated=20 cultures comprised a fraction of cells, which colabeled,=20 promiscuously, with neuronal and glial markers (Fig. = 2C and=20 G) . This abnormal type of cells was never = observed in=20 normal human fetal neural stem cells and likely reflects the=20 activation of an aberrant differentiation program in tumor = neural=20 stem cells, as expected from developmentally deregulated = tumor cells.=20 Intriguingly, this same fraction of promiscuously = double-labeled=20 cells varied between each cell line and correlated with the=20 proliferation capacity index, with cluster A glioblastoma = multiforme=20 cells (fast-growing), showing the highest content of abnormal = end cells, and cluster B lines (slow-growing), displaying a=20 smaller proportion and a behavior somewhat reminiscent of = normal=20 human fetal neural stem cells (Fig. 2 , table).



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  		Fig. 2. Differentiation = potential of fast-=20 and slow-expanding tumor neural stem cells. Analysis of the=20 multipotency of the fast-growing line 0627 (representative = of=20 cluster A, passage 13; A=96D) and slow-growing = line=20 0821, (representative of cluster B, passage 15; = E=96H)=20 by immunofluorescence for neuronal (Tuj1; red = in=20 A, C, E, and G), glial = (GFAP;=20 green in B, C, F, and G) = and=20 oligodendroglial (GalC; red in D and = H)=20 markers. Additional markers of neuronal differentiation were = also=20 detected [line 0821, MAP5, MAP2, and=20 NF200, red in I, J, and = K;=20 glutamate and 3D{gamma}=20-aminobutyric acid (GABA), green in = L=20 and M]. Cells that colabeled promiscuously with both = neuronal=20 and glial markers (arrows in C and G) = could be=20 observed in all of the cell lines established; = magnification, x20. The table shows a = quantitative=20 analysis of the differentiation capacity of the various = glioblastoma=20 multiforme=96derived tumor neural stem cell lines, normal = human fetal=20 neural stem cells, and U87 cells with respect to their = ability to=20 generate cells of the three major neural lineages and to = cells that=20 display aberrant, promiscuous neuronal/astroglial = colabeling. =

 = ;
Ultimately,=20 the self-renewal capacity of glioblastoma multiforme cells = was=20 assessed directly, using subcloning experiments by which the = number=20 of stem cells in a given clonal neurosphere can be determined = by=20 assessing the number of secondary neurospheres generated = after=20 dissociation of the clone itself (23=20 ,=20 28 ,=20 34) . This measure also provides both an estimate of the=20 relative frequencies between symmetric proliferative = divisions and=20 symmetric differentiative divisions and an indirect index of = the=20 capacity of the stem cell population for size expansion (23=20 ,=20 28 ,=20 37) . The generation of multiple secondary clones could = be=20 observed in all of the lines tested (line 0627, 68.0 =B1 5.0; = line=20 0913, 55.3 =B1 4.0; line 1022, 85.0 =B1 5.0; line 0821, = 28.0 =B1 8.0;=20 line 1030, 26.1 =B1 2.2; line 1205, 63.8 =B1 4.0; human fetal = neural stem=20 cells, 37.9 =B1 6.0; U87, 90.0 =B1 5.0; means =B1 SEM, = n=20 =3D 4, Student=92s t test, * =3D P < = 0.001). When=20 the six different cell lines were compared with each other, a = significantly lower frequency of proliferative symmetric = divisions=20 was detected in some of the slow-growing cell lines (0821 and = 1030)=20 compared with the other lines. This is indicative of an = inherent=20 difference in the self-renewal capacity between the different = glioblastoma multiforme=96derived tumor neural stem cell = lines,=20 which strictly correlates with their clonogenic ability and=20 their growth profile.

These findings show that glioblastoma multiformes contain = multipotent,=20 long-term self-renewing, population-expanding cells that = satisfy=20 all of the defining criteria expected from tumor neural stem=20 cells in vitro. Nonetheless, because the most critical=20 attribute of tumor stem cells is their capacity to generate = and=20 perpetuate their tumor of origin (20)=20 , we sought to verify whether our glioblastoma = multiforme=96derived=20 tumor neural stem cells could function as tumor-founding = cells by=20 assessing their actual ability to give rise to tumors in=20 vivo.

The In vivo Tumorigenic Potential of Tumor = Neural Stem=20 Cells.
When injected into immunosuppressed animals, both = s.c. and=20 intracranially (i.c.), all of the six tumor neural stem cell = lines=20 reproducibly established tumors, with a take efficiency of = 50% s.c=20 and 100% orthotopically, whereas normal human fetal neural = stem=20 cells always failed. When compared with classical U87-derived = tumors, tumor neural stem cell=96derived gliomas developed at = a=20 much slower rate. The first appearance of well-defined tumor=20 masses could be observed after 1 week s.c. and 3 weeks i.c. = in=20 U87-treated animals, whereas it took at least 6 weeks s.c. = and 7=20 weeks i.c. to observe the same extent of tumorigenesis in=20 glioblastoma multiforme=96derived, tumor neural stem = cell=96transplanted=20 animals.

Histopathologic analysis of s.c. generated tumor neural stem=20 cell=96derived tumors demonstrated an unprecedented, striking = glioblastoma-like tissue pattern. This was characterized by = the=20 presence of (1) several areas of necrosis surrounded by = typical=20 pseudo-palisade structures (Fig. 3A) , (2) elevated vascular proliferation as = demonstrated=20 by laminin staining (Fig. 3B) , (3) GFAP immunoreactivity (Fig. 3C) , and (4) many mitotic figures, as measured = by Ki67=20 (not shown), which were significantly higher than U87-derived = tumors=20 (3.68 =B1 0.58% for line 0627-derived tumor versus = 1.38 =B1 0.13%=20 for U87-derived tumor; n =3D 4, P < 0.01, = Student=92s=20 t test). Notably, tumors derived from U87 implantation = never=20 displayed any glioblastoma-specific feature. Rather, they = were=20 characterized by the presence of cells with globoid = cytoplasm, which=20 lacked fibrillary structures and hyperchromatic nuclei, the = typical=20 hallmarks of highly malignant gliomas (Fig. 3D) .



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  		Fig. 3. Histopathologic features = of tumor=20 neural stem cell=96derived s.c. and i.c. tumors. A. = Large areas=20 of necrosis surrounded by highly packed tumor cells=20 ("pseudopalisade") were identified within the tumors = generated by=20 s.c. transplantation of tumor neural stem cells into = scid/bg=20 mice. Extensive neovascularization of the tumor tissue by = laminin=20 staining (B; x10 = objective)=20 as well as widespread and intense GFAP immunoreactivity = (C;=20 x20 objective) could be = retrieved.=20 D. U87-derived tumors did not display any of the = peculiar=20 histomorphologic features, typical of highly malignant = gliomas.=20 Tumor neural stem cell=96generated i.c. tumors were = characterized by a=20 peculiar morphology (E; x4=20 objective), and nest-like formations, reminiscent of = mitotically=20 active anomalous glial cells, could be identified in the = core of the=20 lesion (inset in E), as opposed to U87-derived = tumors=20 (F), which consisted of highly packed = undifferentiated small=20 rounded single mitotic elements (inset in F). =

 = ;
Strikingly,=20 when transplanted orthotopically, tumor neural stem cells = were able=20 to form i.c. tumors, which displayed an elevated extent of=20 proliferation and an exacerbated migratory and infiltration=20 capability, distinctive of human glioblastomas multiforme. As = shown by hematoxylin and eosin staining, tumor neural stem = cells=20 generated highly anaplastic tumors, characterized by marked=20 nuclear atypia and high mitotic index (hematoxylin and eosin = in=20 Fig. 3E ). Conversely, unlike any typical glioblastoma, U87 = cells gave rise to enormous but well-delimitated tumor = masses,=20 which developed strictly within the area of injection = (hematoxylin=20 and eosin in Fig. 3F ). On the same line, the i.c. tumors derived = from tumor=20 neural stem cells displayed a very peculiar histomorphology. = Six=20 weeks after implantation, peculiar nest-like structures, = composed of=20 highly mitotic polymorphic cell nuclei, identified as = aberrant glial=20 elements, could be retrieved especially in the center of the = lesion=20 (Fig. 3E , inset). Due to the physical = constraints=20 imposed by the walls of the skull, the maximum level of = growth=20 reached by i.c. tumor neural stem cell=96derived tumors did = not allow=20 the formation of clear areas of necrosis (pseudo-palisades), = as=20 opposed to s.c. tumors, which were allowed to grow up to 5=20 cm3. In agreement with findings on s.c. tumors, = also=20 orthotopic U87-derived lesions did not show any cyto-architectural=20 similarity with the actual human disease (Fig. 3F , inset).

Most importantly, by staining with human-specific = antimitochondria,=20 tumor neural stem cell=96derived tumor cells were shown = to escape=20 the site of transplantation, infiltrating both the adjacent=20 parenchyma and the corpus callosum, even extending into the=20 contralateral hemisphere, as bipolar migrating cells (Fig.=20 4A) . Consistent with our in vitro findings and with = the=20 study of Hemmati et al. (17)=20 , when passage seventh medulloblastoma-derived cells were=20 transplanted orthotopically, after 6 weeks from transplantation, = no=20 sign of tumor formation could be observed, and the transplanted = cells=20 did not migrate significantly from the graft (Fig. 4B) . The same noninfiltrative behavior could be = observed=20 in U87-derived tumors (Fig. 4C) .



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  		Fig. 4. Tumor neural stem cells=20 xenografted i.c. generate tumors, which reproduce the=20 intraparenchymal invasion pattern, typical of human = glioblastoma=20 multiformes. Six weeks after orthotopic implantation, tumor = neural=20 stem cell=96derived tumors (A; human-specific = antimitochondria=20 staining in green) were characterized by an expanding = area of=20 growth, located in the center of the tumor mass and by a = surrounding=20 "penumbra," from which highly migratory cells departed, = infiltrating=20 the adjacent parenchyma and white matter (A, = arrow;=20 x4 objective). At the = same time=20 point, medulloblastoma=96derived grafts (B; = human-specific=20 antinuclei staining in green; x10 objective) exhibited very = poor=20 capacity for migration, spreading, and invasion, if any. = Notably,=20 U87-derived tumors (C; human-specific = antimitochondria=20 staining in green; x4=20 objective) were quite large in size, but cells did not = exhibit any=20 migratory capacity, so that these tumors consisted of a=20 progressively enlarging, well-defined cell mass, which = remained=20 confined to the site of injection. An elevated mitotic index = was=20 exhibited by i.c. tumor neural stem cell tumors (D; = Ki67 in=20 red and human nuclei in green; x40 objective; arrows = point to=20 examples of double-labeled cells), whereas very few cells = within the=20 medulloblastoma-derived grafts were indeed proliferating = (E;=20 Ki67 in red and human nuclei in green; x40 objective; arrow = pointing to=20 a double-labeled cell). In medulloblastoma grafts, we could = never=20 find cells that expressed astroglial markers [F; GFAP = in=20 red and human mitochondria in green; x20 objective; note that GFAP=20 immunoreactivity (red) is confined outside the graft = mass=20 (green)]. Conversely, some cells in glioblastoma=20 multiforme=96derived tumor neural stem cell=96generated = tumors were=20 found to label with the anti-GFAP antibody (G, human=20 mitochondria in green; H, anti-GFAP in = red;=20 merged confocal image in I; x20 objective). =

 = ;
Notably,=20 when i.c. tumors generated by glioblastoma multiforme=96derived = tumor=20 neural stem cells were characterized for the expression of = specific=20 antigens, they displayed an intense mitotic activity as shown = by Ki67=20 staining (Fig. 4D , double-labeled cells in yellow), = whereas the=20 graft established by medulloblastoma cells contained fewer=20 proliferating cells (Fig. 4E) . Moreover, the tumors established by = glioblastoma=20 multiforme=96derived tumor neural stem cells were = immunoreactive for=20 the astroglial-specific marker GFAP (Fig. = 4G--I) and vimentin (not shown) and negative for = neuronal=20 (Tuj1) and oligodendroglial antigens (GalC), just like = typical human=20 glioblastomas multiforme (not shown). Similar to grafted U87 = cells,=20 medulloblastoma implants did not display immunoreactivity for = astroglial (Fig. 4F) , neuronal, or oligodendroglial antigens (not = shown).

Serial Transplantation of Tumor Neural Stem = Cells.
To=20 conclusively demonstrate the stemness of glioblastoma = multiforme=96derived=20 tumor neural stem cells, we performed sequential = transplantation=20 experiments. This was done in analogy with the classical = hemopoietic=20 serial repopulation paradigm used to identify true = hemopoietic=20 stem cells (38)=20 . Cultured tumor neural stem cells (primary tumor neural stem = cells)=20 were transplanted into the mouse CNS to establish a tumor = from which=20 secondary tumor neural stem cells were recultured and then=20 transplanted into new recipients. The successful, sequential=20 generation of brain tumors provided evidence that we had = isolated=20 bona fide tumor neural stem cells.

Tumor neural stem cells, isolated from post-surgery specimens,=20 were expanded, cloned, and injected orthotopically, giving = rise=20 to brain tumors. At the first sign of neurologic impairment=20 (8=9610 weeks after transplantation), as produced by the=20 expanding tumor mass (Fig. 5A and E) , animals were sacrificed. The tumor was = explanted by=20 careful dissection, which excluded the periventricular region = of the=20 forebrain, highly enriched for stem cells and progenitors = (ref. 9=20 ; Fig. 5B=96D ), and the cells were cultured under the same = conditions used to establish the original tumor neural stem = cell=20 cultures. Clonal neurospheres were readily detected in these=20 secondary cultures, which exclusively contained cells that = labeled=20 positive with human-specific markers and that were named=20 post-transplantation=96tumor neural stem cells (Fig. = 5F) . Generation of primary spheres from tumor = neural stem=20 cell=96induced tumors occurred much faster than parental, = primary=20 spheres, established from the original patient=92s specimen = (3 days for=20 post-transplantation=96tumor neural stem cells compared with = 20=9640 days=20 for primary, parental tumor neural stem cell lines). Cell = lines=20 established from the post-transplantation=96tumor neural stem = cells also expanded at a faster rate than their parental = tumor=20 neural stem cells (line glioblastoma multiforme 1022 as=20 reference; Fig. 5G ). This suggests that the malignancy of serially = transplanted=20 post-transplantation=96tumor neural stem cells might have = been=20 exacerbated by the exposure to the in vivo environment = or that=20 particularly aggressive cells had been selected upon = transplantation,=20 tumor growth, and reculturing.



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  		Fig. 5. Tumor neural stem cells = self-renew=20 in vivo, upon serial transplantation. The tumor mass=20 established from the transplanted human tumor neural stem = cells was=20 dissected out at the first sign of neurologic impairment, = taking=20 particular care of not including the stem cell=96enriched=20 periventricular region (arrows in = A=96D). This=20 procedure was facilitated by the large size of the = in-growing tumor=20 mass, which produced a macroscopic enlargement of the = injected=20 cerebral hemisphere (E, arrows). The tumor = cells were=20 then dissociated and placed in culture under conditions = identical to=20 those used to establish tumor neural stem cell lines from = the=20 original human specimen. This resulted in the establishment = of=20 secondary tumor neural stem cell (PT-TNSCs) lines, = which=20 contained exclusively cells of human origin, with no sign of = rodent=20 cell contamination (F; human-specific mitochondria = staining=20 in red and nuclear 4',6-diamidino-2-phenylindole = staining in=20 blue; x20 = objective).=20 G, Post-transplantation=96tumor neural stem cells = were able to=20 long-term expand in culture, even faster than the original, = parental=20 tumor neural stem cell line. GBM, glioblastoma = multiforme. =

 = ;
When=20 post-transplantation=96tumor neural stem cell lines were = differentiated=20 under the same conditions of their parental ones, all of the = three=20 cell types could be detected. Thus, post-transplantation=96tumor = neural=20 stem cells retained multipotency after transplantation and=20 reculturing (20 =B1 2.1% of Tuj1-IR cells; 52 =B1 5.4% of = GFAP-IR cells;=20 24 =B1 0.7% of Tuj1/GFAP cells, n =3D 10, two = different secondary=20 glioblastoma multiforme 1022 cell lines analyzed). = Furthermore,=20 post-transplantation=96tumor neural stem cells maintained the = same=20 karyotypic features (not shown) and molecular signature of = their=20 parental lines (see Fig. 6D , line 1022 and 1022PT), and when retransplanted=20 orthotopically, they were still capable of giving rise to new = glioblastoma multiforme=96like tumors, with characteristics = identical=20 to those generated by primary tumor neural stem cells (not = shown). In=20 conclusion, tumor neural stem cells satisfy all of the = critical=20 criteria to be defined as multipotent tumor neural stem cell = lines,=20 both in vitro and in vivo (20)=20 .



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  		Fig. 6. Karyotypic imbalance, = telomerase=20 reactivation, and molecular fingerprint characterize all=20 glioblastoma multiforme=96derived tumor neural stem cells. = A,=20 summary of the chromosomal aberrations found in all of the = different=20 tumor neural stem cell lines. HFNSC, human fetal = neural stem=20 cell. B, the enzymatic activity of the RNA catalytic = subunit=20 of telomerase as assessed by telomeric repeat amplification = protocol=20 assay in the different cell lines, normal human neural stem = cells,=20 and U87 glioma cells; C, telomere length, as measured = by=20 Southern blotting in glioblastoma multiforme cell lines, = normal=20 human neural stem cells, and U87 glioma cells. M, = marker;=20 LW, low-weight standard; HW, high-weight = standard.=20 D, a preliminary gene expression profile on tumor = neural stem=20 cells and control cell lines by semiquantitative reverse=20 transcription-PCR. Tumor neural stem cell lines are = displayed as=20 clustered into fast-growing (cluster A) or = slow-growing=20 (cluster B) cell lines. =

 = ;
Tumor=20 Neural Stem Cells Display Tumor-Specific Properties.
In = addition to=20 the exacerbated growth rate displayed by most of the = glioblastoma=20 multiforme=96derived lines when compared with normal human = fetal neural=20 stem cells (Fig. 1C) , tumor neural stem cells were also expected = to bear=20 tumor-specific features, such as highly unbalanced karyotypes = and=20 telomerase reactivation.

Karyotypic analysis was performed on all of the six lines, on = U87=20 cells and on human fetal neural stem cells (Fig. 6A) . Many different numerical and structural = chromosomal=20 aberrations could be detected in tumor neural stem cells and = in U87=20 but not in human fetal neural stem cells, which revealed a = normal=20 complement of chromosomes and no genetic abnormalities. = Intriguingly,=20 aneuploidy of specific autosomes and sex chromosomes = (monosomy 1,=20 trisomy 7 and 22, and disomy X) was retrieved most frequently = in=20 the three tumor neural stem cell lines with the highest=20 expansion potential (cluster A), suggesting that peculiar=20 aberrations may confer a certain degree of proliferation = advantage to=20 tumor neural stem cells. Conversely, slow-growing lines = (cluster=20 B) showed line-specific, chromosomal aberrations. Notably,=20 long-term culturing did not affect the karyotypic signature = of tumor=20 neural stem cells, which remained constant even at high = passages=20 in culture (not shown).

Malignant transformation entails the acquisition of an immortal=20 phenotype, which requires active telomere-maintenance = mechanisms=20 (39)=20 . We tested whether the catalytic subunit of the human=20 ribonucleoprotein enzyme telomerase reverse transcriptase, = which=20 provides tumor cells with unlimited proliferation capacity by = telomere lengthening, was at work in glioblastoma = multiforme=96derived=20 cell lines. By means of the telomeric repeat amplification = protocol=20 assay, high human telomerase reverse transcriptase activity = was=20 retrieved in all of the glioblastoma multiforme=96derived = cell lines=20 and in the corresponding original tumor specimens (not = shown),=20 whereas normal human fetal neural stem cells were telomerase = negative=20 (Fig. 6B) , as shown previously (40)=20 . We also assessed telomere length and detected significant=20 differences among the various tumor neural stem cell lines, = with=20 lines 0821 and 1030 displaying very short telomeres, even = shorter=20 than human fetal neural stem cells (Fig. 6C) . Our observations suggest that tumor neural = stem=20 cells, isolated from glioblastoma multiformes, although = displaying=20 typical stem cell features, possess line-intrinsic, = tumor-related=20 functional properties.

Molecular Phenotyping of Tumor Neural Stem = Cells.
To=20 assess whether the heterogeneity detected at the cellular and = biochemical level in various tumor neural stem cells could = also be=20 retrieved at the molecular level (Fig. 6D) , the gene expression profiles of the six = different=20 lines were analyzed by semiquantitative reverse = transcription-PCR.=20 The candidate genes for reverse transcription-PCR were = selected and=20 clustered taking into account their restricted pattern of = expression=20 in (1)=20 normal neural progenitors; (2)=20 brain tumors; (3)=20 CNS development. Normal human fetal neural stem cells, the=20 glioblastoma cell line U87MG, and human fetal whole brain = tissue were=20 used as specimens of reference for the above gene clusters 1, = 2,=20 and 3, respectively.

As expected, based on their cellular phenotype, the genetic=20 profiles of the different tumor neural stem cells could = readily=20 be clustered based on the expression of specific gene sets. = The=20 fast-growing cell lines 0627, 0913, and 1022 were characterized = by=20 similar gene expression patterns, whereas each one of the=20 slow-growing lines 0821, 1030, and 1205 displayed a unique = molecular=20 signature. Nonetheless, fast-growing lines (0627, 0913, and=20 1022) also showed cell-line=96specific variations in their = gene=20 expression profile, confirming that despite the evidence for=20 overlapping cellular and molecular characteristics, each = tumor neural=20 stem cell line was ultimately, inherently unique and = phenotypically=20 different from all of the others.

Intriguingly, the widest body of heterogeneity in terms of gene=20 expression occurred at the level of developmentally regulated = genes, which displayed line-specific, modulation patterns. = For=20 instance, the homeobox gene Emx2, a regulator of = self-renewal=20 in neural stem cells (23)=20 , also implicated in cortical development (41=20 ,=20 42) , and its repressor HOXA10(43)=20 were transcriptionally regulated in a reciprocal fashion, = with lines=20 0627, 0913, and 1022 (cluster A) displaying high levels of = Emx2=20 transcript and no detectable HOXA10mRNA and lines = 0821, 1030,=20 and 1205 (cluster B) showing the opposite trend. Similarly, = genes=20 such as Patched and Pax6 demonstrated a = scattered=20 distribution across the different tumor neural stem cell = lines,=20 although not correlated to any clear functional clustering.=20 Conversely, a higher degree of homogeneity between lines was=20 retrieved when analyzing tumor-related genes such as = PTEN,=20 p21, p27, p53, and MDR1. In = perspective,=20 the line-specific molecular signature and the identification = of=20 genes that play a key role in a given type of tumor neural = stem cell=20 may be exploited to develop custom-fit therapies, by taking = into=20 account the unique genetic background of the specific tumor = cells=20 found in the glioblastoma multiforme of a specific = patient.=20


3D"   =20 DISCUSSION
Top
ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
RESULTS
3D"DISCUSSION
REFERENCES
 
T= his work=20 reports on the isolation and identification of tumor neural = stem=20 cells from human adult glioblastoma multiforme, which possess = the=20 capacity to establish, sustain, and expand these tumors, even = under=20 the challenging settings posed by serial transplantation = experiments.=20 These data identify tumor neural stem cells as one of the = cell=20 lineages involved in establishing the malignant, aggressive = profile=20 of this lethal brain tumor.

Glioblastoma multiforme represents one of the most frequent = brain=20 cancers and either may develop from lower grade astrocytic = tumors=20 (secondary or progressive glioblastoma multiforme) or may = arise very=20 rapidly de novo (primary glioblastoma multiforme; ref. = 44=20 ). In both cases, neither chemotherapy nor radiotherapy has = been=20 effective in managing these cancers (45)=20 . Glioblastomas multiforme display a rather heterogeneous = cellular=20 composition, as indicated by the term "multiforme," with some = of the=20 tumor cells bearing significant migratory capacity. This = results=20 in invasive spread, in the formation of multiple secondary = foci,=20 and in frequent, recurrent growth. Therefore, the = identification=20 of the cell type(s) involved in some or all of these = phenomena=20 is critical from both a scientific and therapeutic = standpoint.

To date, three studies have indicated the presence of = undifferentiated=20 neural precursors within pediatric or adult human brain = tumors.=20 The initial study by Ignatova et al. (15)=20 described the isolation of neurosphere-forming, bipotent=20 (neuronal/astroglial) precursors from glioblastomas = multiforme and=20 was subsequently extended by Singh et al. (16)=20 , who also demonstrated that bipotent progenitors from CNS = tumors=20 (mostly medulloblastomas) displayed short-term self-renewal = (three=20 passages in culture). The latter work also proposed that=20 oligodendrocytes might have been generated from tumor cells, = but the=20 use of a promiscuous marker such as platelet-derived growth = factor=20 receptor 3D{alpha}=20, = which=20 simultaneously labels both oligodendrocytes and neurons in = culture (46)=20 , precluded any conclusion in this direction. Similar = findings were=20 later reported by Hemmati et al. (17)=20 , who described the absence of oligodendrocytes and the lack = of=20 tumorigenic ability after intracerebral transplantation of=20 tumor-derived cells.

Collectively, these studies show that cells endowed with some = of=20 the characteristics expected from stem cells can be found = within=20 brain tumors, thus raising some intriguing issues. First and=20 foremost, can aberrant neural precursors be involved in the=20 establishment, expansion, and recurrence of CNS cancers = (i.e.,=20 are they tumorigenic?) or, rather, do they emerge as the = byproduct of=20 the uncontrolled proliferation of the actual tumorigenic = cells, which=20 are developmentally deranged? Second, if these precursors are = tumorigenic, do they possess all of the features to qualify = as tumor=20 neural stem cells (20)=20 ?

The data presented here provide answers to these questions. = They=20 describe the isolation and identification of human glioblastoma=20 multiforme cells, which possess all of the defining features = of=20 somatic stem cells, including ex vivo multipotency = (i.e.,=20 the potential to simultaneously generate cells displaying = antigenic=20 reactivity for neuronal, astroglial, and oligodendroglial = markers=20 in culture; refs. 47,=20 48,=20 49=20 ) and the ability to establish and expand glioblastoma=20 multiforme=96like tumors. The latter contain exclusively = cells=20 expressing astroglial markers, as in typical human = glioblastomas=20 multiforme. Notably, these cells can perpetuate glioblastoma=20 multiforme=96like tumors even under the critical settings = imposed by=20 serial transplantation, thus satisfying the most rigorous = criteria to=20 be classified as tumor neural stem cells (20)=20 .

These findings were highly reproducible when analyzing tumor=20 neural stem cells established from different specimens, = regardless=20 of significant functional, karyotypic, and molecular = differences=20 between the various tumor neural stem cells. This may account = for the presence within different glioblastoma multiforme = specimens=20 of tumor neural stem cells with common general stem cell = properties=20 that, however, are also characterized by unique, = specimen-related,=20 specific functional and molecular attributes.

Our data seem to suggest that tumor neural stem cells are found=20 mainly within glioblastomas multiforme, although such a = conclusion=20 should be taken with caution. In fact, although our findings=20 are consistent with those from Hemmati et al. (17)=20 and show that cells from medulloblastomas expand poorly in = vitro and do not establish tumors upon intracerebral = grafting,=20 they should not be construed as evidence that tumor neural = stem cells=20 are not present in medulloblastomas. It is possible that=20 medulloblastomas contain cells that elicit their full stem = cell=20 potential under different settings than those used to date. = Thus, it=20 is intriguing that Kondo et al. (50)=20 have reported on the presence of tumorigenic stem-like cells = in a=20 rodent glioma cell line (C6) that require different growth = factors=20 than those used in previous studies to expand in = culture.

The knowledge that cells with stem cell characteristics may = be=20 involved in glioblastoma multiforme tumorigenesis is of paramount=20 importance for our understanding of how these tumors form and = expand and may provide an indication as to the key cellular = and=20 molecular mechanism to be investigated and tackled for diagnostic = and=20 therapeutic purposes. Unfortunately, it remains unclear = whether tumor=20 neural stem cells in glioblastomas multiforme derive from the = transformation of normal human neural stem cells or whether = their=20 presence therein reflects the result of the transformation = and=20 de-differentiation of a more mature brain cell, which brought = about=20 otherwise silent/latent stem cell properties (5)=20 . Nonetheless, the demonstration of the existence within=20 glioblastomas multiforme of tumor neural stem cells with the = capacity=20 of establishing this type of tumor, together with previous = reports=20 showing the existence within similar cancers of bipotent, = stem-like=20 cells (15,=20 16,=20 17)=20 , lends to the hypothesis that a hierarchical model of cell = genesis=20 may be at work within malignant brain cancers.

To date, only a few glioblastoma multiforme cell lines have = been=20 established (51,=20 52,=20 53)=20 and used for studies on basic glioma biology or to perfect = therapy=20 for glioblastoma multiforme (54,=20 55,=20 56,=20 57)=20 . The tumor neural stem cells described here can be used to = routinely=20 establish adult human glioblastoma multiforme cell lines in a = quick=20 and reproducible fashion. Of note, these cells reproduce the = overall=20 behavior of glioblastomas multiforme closer than any other = previously=20 available cell line. For example, when transplanted into=20 immunosuppressed mice, the most widely used human = glioblastoma=20 multiforme cell line, the U87MG, gave rise to tumors that did = not=20 mimic the typical glioblastoma multiforme characteristics.=20 Conversely, upon orthotopic transplant, all of the = glioblastoma=20 multiforme=96derived, tumor neural stem cell lines formed = tumors=20 bearing all of the primary traits of human glioblastomas = multiforme.=20 Most importantly, cells in tumor neural stem cell=96founded = tumors=20 displayed, in vivo, the typical migratory capacity = expected=20 from glioblastoma multiforme malignant cells, which was not = observed=20 when using U87, and, most importantly, retained their = distinctive,=20 line-specific proliferation and differentiation attributes. = Thus, the=20 identification of tumor neural stem cells also makes = available cell=20 lines of glioblastoma multiforme=96initiating cells, which = may=20 retain specimen-specific and, thus, patient-specific traits. = This=20 may open new avenues to identify novel tumor cell markers for = therapeutic and diagnostic purposes and to develop = patient-tailored=20 pharmacologic approaches for the cure of glioblastomas = multiforme,=20 aiming to target what may very well be the most aggressive = cell=20 type found in glioblastomas multiforme, to date, which is the = tumor neural stem cell.


3D"   =20 ACKNOWLEDGMENTS
 
We = thank Luigi=20 Cornaghi for his expert technical assistance with karyotyping = and Dr.=20 Cesare Covino for his help with confocal imaging.


3D"   =20 FOOTNOTES
 
Grant support: Compagnia S. = Paolo-Programma=20 Oncologia (R. Galli) and the Fondazione Agarini and BMW Italy = (A.=20 Vescovi).

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

Requests for reprints: Angelo = Vescovi,=20 Stem Cell Research Institute, DIBIT, H. S. Raffaele, Via = Olgettina=20 58, Milan, Italy. E-mail address: vescovi.angelo{at}hsr.it

Received 4/29/04. Revised 7/10/04. Accepted 7/26/04.


3D"   =20 REFERENCES
Top
ABSTRACT
INTRODUCTION
MATERIALS AND METHODS
RESULTS
DISCUSSION
3D"REFERENCES
&nb= sp;

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Genetic = alterations=20 associated with acquired temozolomide resistance in SNB-19, a = human glioma=20 cell line.
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Notch Pathway Inhibition Depletes Stem-like = Cells=20 and Blocks Engraftment in Embryonal Brain = Tumors.
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Consensus document on European = brain=20 research
J. Neurol. Neurosurg. Psychiatry,=20 August 1, 2006; 77(suppl_1): i1 - i49.
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Stem cell aging and = cancer.
Sci.=20 Aging Knowl. Environ., May 15, 2006; 2006(9): pe12 - = pe12.=20
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Recent Advances in the Treatment of Malignant=20 Astrocytoma
J. Clin. Oncol., March 10, 2006; = 24(8):=20 1253 - 1265.
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PBK/TOPK, a Proliferating Neural=20 Progenitor-Specific Mitogen-Activated Protein Kinase = Kinase
J.=20 Neurosci., November 16, 2005; 25(46): 10773 - 10785. =
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Emx2 Regulates = Mammalian=20 Reproduction by Altering Endometrial Cell = Proliferation
Mol.=20 Endocrinol., November 1, 2005; 19(11): 2839 - 2846. =
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D. Fang, T. K. Nguyen, K. Leishear, R. Finko, A. N. = Kulp, S. Hotz,=20 P. A. Van Belle, X. Xu, D. E. Elder, and M. Herlyn
A=20 Tumorigenic Subpopulation with Stem Cell Properties in=20 Melanomas
Cancer Res., October 15, 2005; = 65(20):=20 9328 - 9337.
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N. Sanai, A. Alvarez-Buylla, and M. S. = Berger
Neural=20 Stem Cells and the Origin of Gliomas
N. Engl. J. Med., = August 25, 2005; 353(8): 811 - 822.
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S.-K. Kim, T. G. Cargioli, M. Machluf, W. Yang, Y. Sun, = R.=20 Al-Hashem, S. U. Kim, P. M. Black, and R. S.=20 Carroll
PEX-Producing Human Neural Stem Cells Inhibit = Tumor=20 Growth in a Mouse Glioma Model
Clin. Cancer Res.,=20 August 15, 2005; 11(16): 5965 - 5970.
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L. Patrawala, T. Calhoun, R. Schneider-Broussard, J. = Zhou, K.=20 Claypool, and D. G. Tang
Side Population Is Enriched in = Tumorigenic, Stem-Like Cancer Cells, whereas ABCG2+ and ABCG2- = Cancer=20 Cells Are Similarly Tumorigenic
Cancer Res.,=20 July 15, 2005; 65(14): 6207 - 6219.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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D. Ponti, A. Costa, N. Zaffaroni, G. Pratesi, G. = Petrangolini, D.=20 Coradini, S. Pilotti, M. A. Pierotti, and M. G.=20 Daidone
Isolation and In vitro Propagation of = Tumorigenic=20 Breast Cancer Cells with Stem/Progenitor Cell=20 Properties
Cancer Res., July 1, 2005; = 65(13): 5506 -=20 5511.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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B. Dasgupta and D. H. Gutmann
Neurofibromin = Regulates=20 Neural Stem Cell Proliferation, Survival, and Astroglial = Differentiation=20 In Vitro and In Vivo
J. Neurosci., = June 8, 2005;=20 25(23): 5584 - 5594.
[Abstra= ct]=20 [Full=20 Text] [PDF] =
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S. A. Bapat, A. M. Mali, C. B. Koppikar, and N. K.=20 Kurrey
Stem and Progenitor-Like Cells Contribute to the = Aggressive Behavior of Human Epithelial Ovarian = Cancer
Cancer=20 Res., April 15, 2005; 65(8): 3025 - 3029.
[Abstract]=20 [Fu= ll=20 Text] [PDF]=20
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R. Glass, M. Synowitz, G. Kronenberg, J.-H. Walzlein, D. = S.=20 Markovic, L.-P. Wang, D. Gast, J. Kiwit, G. Kempermann, and H.=20 Kettenmann
Glioblastoma-Induced Attraction of = Endogenous Neural=20 Precursor Cells Is Associated with Improved = Survival
J.=20 Neurosci., March 9, 2005; 25(10): 2637 - 2646.
[Abstra= ct]=20 [Full=20 Text] [PDF] =
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HOME HELP FEEDBACK SUBSCRIPTIONS ARCHIVE SEARCH TABLE OF=20 CONTENTS
Cancer = Research Clinical Cancer=20 Research
Cancer=20 Epidemiology Biomarkers & Prevention Molecular=20 Cancer Therapeutics
Molecular=20 Cancer Research Cell Growth=20 & Differentiation
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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_0041_01C85789.06931E20 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://intl-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 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quoted-printable Content-Location: http://intl-cancerres.aacrjournals.org/javascript/entrez/callback.js /************************************************************************= *****=0A= * javascript/entrez/callback.js=0A= *=0A= * Entrez Linking callback to populate content box.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Execute callback to fill content box with Entrez Linking information.=0A= */=0A= function entrez_callback(pmid, callback_url) {=0A= /*=0A= * MSIE 5.5 and below have issues with the JavaScript=0A= * used for Entrez Linking. 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path=3D"+_utcp+";"+nx+_udo;=0A= _ubd.cookie=3D"__utmb=3D"+_udh+"; path=3D"+_utcp+";"+x+_udo;=0A= _ubd.cookie=3D"__utmc=3D"+_udh+"; path=3D"+_utcp+";"+_udo;=0A= _ufns=3D1;=0A= } else if (a>=3D0 && b>=3D0 && c>=3D0) {=0A= _ubd.cookie=3D"__utmb=3D"+_udh+"; path=3D"+_utcp+";"+x+_udo;=0A= } else {=0A= if (a>=3D0) a=3D_uFixA(_ubd.cookie,";",_ust);=0A= else a=3D_udh+"."+_uu+"."+_ust+"."+_ust+"."+_ust+".1";=0A= _ubd.cookie=3D"__utma=3D"+a+"; path=3D"+_utcp+";"+nx+_udo;=0A= _ubd.cookie=3D"__utmb=3D"+_udh+"; path=3D"+_utcp+";"+x+_udo;=0A= _ubd.cookie=3D"__utmc=3D"+_udh+"; path=3D"+_utcp+";"+_udo;=0A= _ufns=3D1;=0A= }=0A= if (_ulink && xx && xx!=3D"" && xx!=3D"-") {=0A= xx=3D_uUES(xx);=0A= if (xx.indexOf(";")=3D=3D-1) _ubd.cookie=3D"__utmx=3D"+xx+"; = path=3D"+_utcp+";"+nx+_udo;=0A= }=0A= if (_ulink && v && v!=3D"" && v!=3D"-") {=0A= v=3D_uUES(v);=0A= if (v.indexOf(";")=3D=3D-1) _ubd.cookie=3D"__utmv=3D"+v+"; = path=3D"+_utcp+";"+nx+_udo;=0A= }=0A= _uInfo(page);=0A= _ufns=3D0;=0A= _ufno=3D0;=0A= if (!page || page=3D=3D"") _uff=3D1;=0A= }=0A= function _uInfo(page) {=0A= var p,s=3D"",dm=3D"",pg=3D_udl.pathname+_udl.search;=0A= if (page && page!=3D"") pg=3D_uES(page,1);=0A= _ur=3D_ubd.referrer;=0A= if (!_ur || _ur=3D=3D"") { _ur=3D"-"; }=0A= else {=0A= dm=3D_ubd.domain;=0A= if(_utcp && _utcp!=3D"/") dm+=3D_utcp;=0A= p=3D_ur.indexOf(dm);=0A= if ((p>=3D0) && (p<=3D8)) { _ur=3D"0"; }=0A= if (_ur.indexOf("[")=3D=3D0 && = _ur.lastIndexOf("]")=3D=3D(_ur.length-1)) { _ur=3D"-"; }=0A= }=0A= s+=3D"&utmn=3D"+_uu;=0A= if (_ufsc) s+=3D_uBInfo();=0A= if (_uctm) s+=3D_uCInfo();=0A= if (_utitle && _ubd.title && _ubd.title!=3D"") = s+=3D"&utmdt=3D"+_uES(_ubd.title);=0A= if (_udl.hostname && _udl.hostname!=3D"") = s+=3D"&utmhn=3D"+_uES(_udl.hostname);=0A= s+=3D"&utmr=3D"+_ur;=0A= s+=3D"&utmp=3D"+pg;=0A= if ((_userv=3D=3D0 || _userv=3D=3D2) && _uSP()) {=0A= var i=3Dnew Image(1,1);=0A= i.src=3D_ugifpath+"?"+"utmwv=3D"+_uwv+s;=0A= i.onload=3Dfunction() {_uVoid();}=0A= }=0A= if ((_userv=3D=3D1 || _userv=3D=3D2) && _uSP()) {=0A= var i2=3Dnew Image(1,1);=0A= = i2.src=3D_ugifpath2+"?"+"utmwv=3D"+_uwv+s+"&utmac=3D"+_uacct+"&utmcc=3D"+= _uGCS();=0A= i2.onload=3Dfunction() { _uVoid(); }=0A= }=0A= return;=0A= }=0A= function _uVoid() { return; }=0A= function _uCInfo() {=0A= if (!_ucto || _ucto=3D=3D"") { _ucto=3D"15768000"; 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