From: Subject: From the Cover: Neural stem cells display extensive tropism for pathology in adult brain: Evidence from intracranial gliomas -- Aboody et al. 97 (23): 12846 -- Proceedings of the National Academy of Sciences Date: Wed, 16 Jan 2008 17:08:12 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C85862.60200690" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.3198 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C85862.60200690 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.pnas.org/cgi/content/full/97/23/12846 From the Cover: Neural stem cells display extensive = tropism for pathology in adult brain: Evidence from intracranial gliomas = -- Aboody et al. 97 (23): 12846 -- Proceedings of the National Academy = of Sciences
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PNAS | November 7, 2000 | vol. 97 | no. 23 |=20 12846-12851

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From the = Cover
Neurobiology
Neural=20 stem cells display extensive tropism for pathology in adult brain: = Evidence from=20 intracranial gliomas

Karen S.=20 Aboody*,3D"dagger,3D"Dagger, Alice Brown3D"dagger, Nikolai G. Rainov3D"dagger, Kate A. Bower*, Shaoxiong=20 Liu*, Wendy Yang*, Juan E.=20 Small*, Ulrich Herrlinger3D"dagger, Vaclav Ourednik*, Peter = McL.=20 Black=A7, Xandra O. Breakefield3D"dagger,=20 and Evan Y. Snyder*,=B6<= /SUP>=20

* Departments of Neurology, Pediatrics, and = Neurosurgery,=20 Children's Hospital; 3D"dagger Molecular Neurogenetics Unit, Department of = Neurology, Massachusetts General Hospital; and =A7 Brain = Tumor=20 Service, Department of Neurosurgery, Brigham and Women's Hospital, = Harvard=20 Medical School, Boston, MA 02115; and 3D"Dagger Layton Bioscience, Sunnyvale, CA 94086=20

Communicated by Richard L. Sidman, Harvard Medical School, = Southborough,=20 MA, July 24, 2000 (received for review May 19, 2000) =


    Abstract
Top
Abstract
Introduction
Experimental Methods
Results
Discussion
References

One of the impediments to the treatment of brain tumors (e.g., = gliomas) has=20 been the degree to which they expand, infiltrate surrounding = tissue,=20 and migrate widely into normal brain, usually rendering them=20 "elusive" to effective resection, irradiation, chemotherapy, = or gene=20 therapy. We demonstrate that neural stem cells (NSCs), when = implanted=20 into experimental intracranial gliomas in vivo in = adult=20 rodents, distribute themselves quickly and extensively = throughout the=20 tumor bed and migrate uniquely in juxtaposition to widely = expanding=20 and aggressively advancing tumor cells, while continuing to = stably=20 express a foreign gene. The NSCs "surround" the invading = tumor border=20 while "chasing down" infiltrating tumor cells. When implanted = intracranially at distant sites from the tumor (e.g., into = normal=20 tissue, into the contralateral hemisphere, or into the = cerebral=20 ventricles), the donor cells migrate through normal tissue = targeting=20 the tumor cells (including human glioblastomas). When = implanted=20 outside the CNS intravascularly, NSCs will target an = intracranial=20 tumor. NSCs can deliver a therapeutically relevant = moleculecytosine deaminase3D---=20such that quantifiable reduction in tumor burden = results.=20 These data suggest the adjunctive use of inherently migratory = NSCs as=20 a delivery vehicle for targeting therapeutic genes and = vectors to=20 refractory, migratory, invasive brain tumors. More broadly, = they=20 suggest that NSC migration can be extensive, even in the = adult brain=20 and along nonstereotypical routes, if pathology (as modeled = here by=20 tumor) is present.

gene therapy | transplantation | migration | brain = tumors |=20 vascular

    Introduction
Top
Abstract
Introduction
Experimental Methods
Results
Discussion
References

Malignant brain tumors, e.g., glioblastoma multiforme, remain = virtually=20 untreatable and inevitably lethal despite extensive surgical = excision=20 and adjuvant radio- and chemotherapy (1).= =20 Their treatment resistance is related to their exceptional = migratory=20 nature and ability to insinuate themselves seamlessly and = extensively=20 into normal neural tissue, often migrating great distances = from=20 the main tumor mass. These cells are responsible for the = recurrent=20 tumor growth near the borders of the resection cavity (1).= It=20 is this behavior that has also limited their accessibility to = otherwise promising gene therapeutic vectors and interventions = (2). = Intriguingly,=20 one of the cardinal features of neural stem cells (NSCs) is = their=20 exceptional migratory ability (3-10).=20 Indeed, it is their migratory capacity that has made them so = useful=20 in therapeutic paradigms demanding brainwide gene and cell=20 replacement in various animal models of neurodegeneration, = albeit=20 usually in the newborn (8-10). = We=20 hypothesized that pathology promotes NSC migration to an = extent not=20 assumed possible based on knowledge drawn from the normal = adult brain=20 and that, therefore, an approach for targeting gene therapy = to the=20 most migratory tumor cells in the adult central nervous = system (CNS)=20 might be the use of inherently migratory NSCs to deliver = therapeutic=20 genes and/or their products.


    Experimental = Methods
Top
Abstract
Introduction
Experimental Methods
Results
Discussion
References

In Vitro Migration Studies. CNS-1 is a = virulent,=20 invasive rat-derived glioblastoma cell line. Cells engineered to express = green=20 fluorescent protein (GFP) as previously described (11, 12) = were plated=20 to 60-70% confluence onto 100-mm culture dishes around a = central 5-mm=20 metal cylinder that was sealed and, therefore, remained = cell-free.=20 The plate was incubated overnight, by which time the glioma = cells had=20 attached. A suspension of 4 =D7 104 = dissociated=20 fibroblasts (in control dishes) or murine NSCs (in = experimental=20 dishes) were seeded into the central cylinder (i.e., no = direct=20 contact with CNS-1 cells) (Fig. 1, = arrowheads). A=20 similar number of fibroblasts or NSCs, respectively, were = placed=20 into a 5-mm cylinder placed directly on top of the adherent = CNS-1=20 monolayer and cultured as before (at extreme right edge of = plates)=20 (Fig. 1, = arrows). The=20 fibroblasts (clone TR-10) were derived from 3T3 cells = infected with a=20 retroviral vector encoding lacZ. The murine NSCs were = derived=20 from the prototypical constitutively lacZ-expressing = helper=20 virus-free murine NSC clonal line C17.2 (8-10, = 13), = which,=20 because of its well-documented ability to integrate into most = CNS structures and in a number of normal and abnormal animal = models,=20 has been useful for delineating the range of therapeutic=20 possibilities for NSCs (8-10, = 13). = (Although=20 self-renewing, NSCs become contact inhibited; never grow in = soft=20 agar; are nontumorigenic in nude mice; fail to incorporate = BrdUrd=20 after 48 h in vivo; and respond to normal cues = for cell=20 cycle withdrawal, differentiation, and interaction with host = cells.)=20 The day after plating of fibroblasts and NSCs, the cylinders = were=20 removed, and the dishes were rinsed and incubated for an = additional=20 5 days. Dishes were stained for the lacZ gene = product=20 Escherichia coli 3D"beta-galactosidase (3D"beta-gal)=20 by 5-bromo-4-chloro-3-indolyl 3D"beta-D-galactoside (X-Gal) histochemistry (8).



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  		Fig. 1.   = Migratory=20 capacity of NSCs in culture. CNS-1 glioblastoma cells were = plated=20 around a central cylinder (i.e., free of CNS-1 cells). = Fibroblasts=20 (A) or NSCs (B) were seeded into the center = cylinder=20 (i.e., no direct contact with CNS-1 cells) (arrowheads) or = into=20 cylinders placed directly on top of adherent tumor cells (at = the=20 extreme right edge of plates; arrows). After removal of the=20 cylinders and 5 additional days of incubation, there = was wide=20 distribution of blue X-Gal+ NSCs (B), = compared=20 with fibroblasts (A), which remained localized to = their area=20 of initial seeding.

Animal Studies in Vivo. For some studies, = not only=20 were murine NSCs (C17.2) used but also some of human derivation were = used (5). To=20 establish intracranial tumors, either the CNS-1 rat = glioblastoma line=20 (11,=20 12) or = the HGL21 human glioblastoma line (14) = was=20 implanted into the brains of adult female nude mice, or the = D74=20 glioma line was implanted into the brains of adult female = Fisher rats=20 by using procedures previously described (15). = Briefly,=20 animals received stereotactically guided injections over 3-5 = min into=20 the forebrain (2 mm lateral and 1 mm anterior to = bregma;=20 depth 2-4 mm from dura) of tumor cells (of a number specified = below)=20 suspended in 1 =B5l of PBS. Animals receiving a second = implant at=20 a later date of NSCs or fibroblasts [suspended in PBS at=20 2-4 =D7 104 cells per =B5l as detailed elsewhere = (8-10, = 13)] = were=20 injected stereotactically with cells in a quantity and = location to=20 yield the various paradigms described below. On the days = specified=20 below, the brains were processed as detailed below (e.g., for = -gal, GFP, BrdUrd, human markers, and/or = cell=20 type-specific antigen expression). CD-1 mice, when used, = received=20 daily cyclosporin injections (10 =B5g/g).

Specific Protocols for in Vivo = Experiments. NSC=20 implantation directly into tumor bed (Fig. 2, = Paradigm=20 1). On day 0, recipients received injections of tumor cells=20 (3-4 =D7 104 in 1 =B5l of PBS) into the right = frontal=20 lobe. On day 4-6, NSCs or fibroblasts=20 (4-10 =D7 104 in 1.5 =B5l of PBS) were = injected directly=20 into the tumor bed, using identical coordinates. Recipients = were=20 killed on days 6-9, 10-12, 14-16, and 21 after tumor=20 implantation.



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  		Fig. 2.   = NSCs=20 migrate extensively throughout a brain tumor mass in = vivo and=20 "trail" advancing tumor cells. Paradigm 1 is = illustrated=20 schematically. Section of brain under low (A) and = high=20 (B) power from an adult rat killed 48 h after = NSC=20 injection into an established D74 glioma, processed with = X-Gal to=20 detect blue-staining 3D"beta-gal-producing NSCs and counterstained with = neutral=20 red to show dark red tumor cells; arrowheads demarcate = approximate=20 edges of the tumor mass where it interfaces with normal = tissue.=20 Donor X-Gal+ blue NSCs (arrows) can be seen = extensively=20 distributed throughout the mass, interspersed among the red = tumor=20 cells. (C) Tumor, 10 days after NSC injection,=20 illustrating that, although NSCs (arrows) have infiltrated = the mass,=20 they largely stop at the junction between tumor and normal = tissue=20 (arrowheads) except where a tumor cell (dark red, elongated) = is=20 entering normal tissue; then NSCs appear to "follow" the = invading=20 tumor cell into surrounding tissue (upper right arrow). This = phenomenon becomes more dramatic when examining NSC behavior = in a=20 more virulent and aggressively invasive tumor, the CNS-1=20 glioblastoma in the adult nude mouse, pictured in D. = This=20 section illustrates extensive migration and distribution of = blue=20 NSCs (arrows) throughout the infiltrating glioblastoma up to = and=20 along the infiltrating tumor edge (red arrowheads) and into=20 surrounding tissue in juxtaposition to many dark = red+=20 tumor cells invading normal tissue. The "tracking" of = individual=20 glioblastoma cells is examined in greater detail in = E-L,=20 where CNS-1 cells have been labeled ex vivo by = transduction=20 with GFP cDNA. (E and F) Sister sections = showing a low=20 power view of transgene-expressing NSCs distributed = throughout the=20 main tumor mass to the tumor edge (outlined by arrowheads). = Sections=20 were either costained with X-Gal (NSCs, blue) and neutral = red (tumor=20 cells, dark red and elongated) (E) or processed for = double=20 immunofluorescence using an anti-3D"beta-gal antibody (NSCs, red) and an = FITC-conjugated=20 anti-GFP antibody (glioblastoma cells, green) (F). = Low=20 (G) and high (H) power views of tumor edge=20 (arrowheads) with blue NSCs (blue arrow) in immediate = proximity to=20 and intermixed with an invading tumor "island" (dark red=20 spindle-shaped cells) (red arrow). (I and J) = Low and=20 high power views, respectively (boxed area in I is = magnified=20 in J), of a blue NSC in direct juxtaposition to a = single=20 migrating neutral red+, spindle-shaped tumor cell = (arrow), the NSC "riding" the glioma cell in "piggy-back" = fashion.=20 (K and L) Low and high power views, = respectively,=20 under fluorescence microscopy, of single migrating = GFP+=20 tumor cells (green) in juxtaposition to 3D"beta-gal+ NSCs (red). Region = indicated by=20 white arrow in K and magnified in L = illustrates NSCs=20 apposed to tumor cells migrating away from the main tumor = bed.=20 (Scale bars: A, 40 =B5m, 30 =B5m in = B; C,=20 30 =B5m, 25 =B5m in D; E, = 90 =B5m,=20 100 =B5m in F; H, 15 =B5m, = 60 =B5m in=20 G; J, 30 =B5m, 60 =B5m in I, = 70 =B5m=20 in K, 35 =B5m in L.)

NSC implantation at distant intracranial site from tumor bed: = In same=20 hemisphere (Fig. 3, = Paradigm=20 2). On day 0, recipients received injections of = glioblastoma cells=20 (3 =D7 104 in 1 =B5l of PBS) into the right = frontal lobe.=20 On day 6, NSCs (4 =D7 104 in = 1.5 =B5l of=20 PBS) were injected also into the right frontoparietal lobe at = the=20 following coordinates: 3 mm lateral and 4 mm caudal = to=20 bregma; depth 3 mm from dura3D---=20i.e., 1 mm lateral and 4 mm behind = the=20 tumor. Animals were killed on days 12 and 21. 



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  		Fig. 3.   = NSCs=20 implanted at various intracranial sites distant from main = tumor bed=20 migrate through normal adult tissue toward glioma cells. = (A=20 and B) Same hemisphere but behind tumor = (Paradigm 2).=20 Shown here is a section through the tumor from an adult nude = mouse=20 6 days after NSC implantation caudal to tumor. In = A (as=20 per the schematic, a coned down view of a tumor populated as = pictured under low power in Figs. 2A=20 and 3 A and B), note X-Gal+ = blue NSCs=20 interspersed among dark neutral red+ tumor cells. = (B) High power view of NSCs in juxtaposition to = islands of=20 tumor cells. (C-H) Contralateral hemisphere (Paradigm = 3).=20 (C-E) As indicated on the schematic, these panels are = views=20 through the corpus callosum ("c") where 3D"beta-gal+ NSCs (red cells, arrows) = are seen=20 migrating from their site of implantation on one side of the = brain=20 toward tumor on the other. Two representative NSCs indicated = by=20 arrows in C are viewed at higher magnification in = D=20 and E, respectively, to visualize the classic = elongated=20 morphology and leading process of a migrating neural = progenitor=20 oriented toward its target. In F, 3D"beta-gal+ NSCs (red) are "homing in" = on the=20 GFP+ tumor (green) having migrated from the other = hemisphere. In G, and magnified further in H, = the=20 X-Gal+ blue NSCs (arrows) have now actually = entered the=20 neutral red+ tumor (arrowheads) from the opposite = hemisphere. (I and J) Intraventricular = (Paradigm 4).=20 Shown here is a section through the brain tumor of an adult = nude=20 mouse 6 days following NSC injection into the = contralateral=20 cerebral ventricle. In I, as per the schematic, blue=20 X-Gal+ NSCs are distributed within the neutral=20 red+ main tumor bed (edge delineated by = arrowheads). At=20 higher power in J, the NSCs are in juxtaposition to = migrating=20 islands of red glioblastoma cells. Fibroblast control cells = never=20 migrated from their injection site in any paradigm. All=20 X-Gal-positivity was corroborated by anti-3D"beta-gal immunoreactivity. (Scale bar: = A,=20 20 =B5m, and applies to C; B, = 8 =B5m, 14 =B5m=20 in D and E, 30 =B5m in F and = G,=20 15 =B5m in H, 20 =B5m in I, and = 15 =B5m in=20 J.)

In contralateral hemisphere (Fig. 3, = Paradigm=20 3). On day 0, recipients received injections of = glioblastoma cells=20 (3-5 =D7 104 in 1 =B5l of PBS) into the right = frontal lobe=20 (2.5 mm lateral and 2 mm caudal to bregma; depth = 3 mm=20 from dura). On day 6, NSCs = (8 =D7 104 in=20 2 =B5l of PBS) were injected into the left frontal lobe at = the=20 following coordinates: 2 mm lateral and 2 mm caudal to=20 bregma; depth 3 mm from dura. Animals were killed on = days=20 12 and 21.=20

Into ventricles (Fig. 3, = Paradigm=20 4). On day 0, recipients received injections of glioblastoma = cells=20 (5-8 =D7 104 in 1 =B5l of PBS) into the right = frontal lobe=20 (2 mm lateral to bregma on the coronal suture; depth = 3 mm=20 from dura). On day 6, NSCs = (8 =D7 104 in=20 2 =B5l of PBS) were injected into the contralateral or = ipsilateral=20 cerebral ventricle at the following coordinates on the = respective=20 side: 1 mm lateral and 3 mm caudal to bregma; depth = 2 mm=20 from dura. Animals were killed on days = 8, 12, and/or=20 21.=20

NSC implantation into a peripheral, intravascular site = (Fig. 4, = Paradigm=20 5). On day 0, adult nude mice received injections of CNS-1=20 glioblastoma cells (1 =D7 105 in 2 =B5l of = PBS) into the=20 right frontal lobe. On day 7, murine NSCs=20 (2 =D7 106 in 200 =B5l of PBS) were injected = into the tail=20 vein. Animals were killed on day 12. 



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  		Fig. 4.   = NSCs=20 injected into tail vein "target" intracerebral gliomas. = Paradigm=20 5 is illustrated. (A-C) Progressively higher = power views=20 of representative 10-=B5m sections through the brain = 4 days after=20 NSC injection, processed with X-Gal histochemistry = (A) and=20 anti-3D"beta-gal immunocytochemistry (B and = C) to=20 identify donor NSCs and counterstained with neutral red to = delineate=20 the tumor border. (The 3D"beta-gal immunoproduct, in addition to = providing=20 independent identity confirmation, typically fills cells and = processes much better than X-Gal.) At low power (A),=20 X-Gal+ NSCs (representative X-Gal precipitate = enlarged in=20 Inset) are distributed throughout the tumor but not = in=20 surrounding normal tissue. Sister sections, reacted with an=20 anti-3D"beta-gal antibody and visualized at higher = power in=20 B and further magnified in C confirm the = presence of=20 donor-derived cells (arrow) within the tumor. (Scale bar: = A,=20 25 =B5m, 20 =B5m in B, and 12 =B5m in = C.)

Retroviral Transduction of NSCs with Cytosine = Deaminase=20 (CD). A plasmid using the retroviral pBabePuro backbone (16) = was=20 constructed to include the E. coli CD cDNA (1.5-kb = fragment)=20 transcribed from the long terminal repeat. Vectors were = packaged=20 by cotransduction of the CDpuro plasmid with the MV12 = envelope=20 coding plasmid cDNA (17) = into 293T/17=20 cells (18).=20 CDpuro retroviral supernatant was used for multiple = infections of the=20 murine NSCs. Transduced NSCs ("CD-NSCs") were placed under = puromycin=20 selection for 3D"approx2=20 wk.

Oncolysis Assays of CD Bioactivity. For in = vitro=20 assays, CNS-1 cells (2 =D7 105) were plated onto = 10-cm dishes=20 (day 1). On day 2, murine or human CD-NSCs=20 (5-10 =D7 104) were added. On day=20 3, 5-fluorocytosine (5-FC, 500 =B5g/ml) was added. = Control=20 dishes included (i) cocultures with no 5-FC and = (ii)=20 tumor cells alone with 5-FC. On day 6, plates were stained = by=20 means of X-Gal histochemistry to visualize NSCs and with neutral = red=20 to visualize tumor cells (Fig. 6 = A and=20 B). The number of tumor cells was extrapolated from = the=20 average of 20 random high power fields per plate. For = in=20 vivo assays, animals bearing CNS-1 cells=20 (7 =D7 104) alone or CNS-1 cells interspersed = with=20 CD-NSCs (3.5 =D7 104) received 10 i.p. = injections of=20 5-FC (900 mg/kg) over 10 days. Control = tumor-bearing=20 animals received no 5-FC, 5-FC without NSCs, or NSCs without = 5-FC.=20 One day after the last 5-FC dose, the brains were = cryosectioned and=20 stained with X-Gal and neutral red, and measurements of the = tumor=20 were made from camera lucida drawings of the mass from = interval=20 sections through the tumor from which relative surface areas = were=20 then calculated by image analysis (Fig. 7). =

BrdUrd Uptake by Engrafted NSCs. Selected animals = received=20 three i.p. injections of BrdUrd (1 ml/100 g body weight of a = 20 =B5M=20 stock solution) over 24 h before sacrifice.

Histopathological and Immunocytochemical = Analysis. On the=20 days specified in the paradigms above, animals were killed, and = 10- to=20 15-=B5m serial coronal cryosections from 4% paraformaldehyde-fixed=20 brains were processed for light microscopy with X-Gal = histochemistry=20 to identify lacZ-expressing blue donor cells (8-10, = 13) = and=20 counterstained with neutral red to detect distinctively dark = red,=20 elongated tumor cells (2). = Sister=20 sections were prepared for dual-filter immunofluorescence = where an=20 anti-3D"beta-gal antibody was revealed with a Texas = Red-conjugated=20 secondary antibody (1:1,000; Vectastain) (to identify donor = cells as=20 red) (8-10, = 13), = and=20 an anti-GFP antibody (1:500; CLONTECH) was revealed with an=20 FITC-conjugated secondary antibody (1:1,000; Vectastain) (to=20 recognize CNS-1 tumor cells as green).=20 BrdUrd+-intercalated cells were identified by an anti-BrdUrd=20 antibody (5). = Sections=20 containing human NSCs were additionally stained with multiple = human-specific antibodies, including against to ribonuclear = protein=20 (Chemicon, 1:20), against NuMA (Chemicon, 1:40 and = Calbiochem,=20 1:400), and against the human EGF receptor (Upstate = Biotechnology,=20 1:100), revealed by a biotinylated goat anti-mouse IgM = (1:250,=20 Vectastain) followed by the standard ABC and diaminobenzidine = (DAB)=20 reaction (Vectastain).


    Results
Top
Abstract
Introduction
Experimental Methods
Results
Discussion
References

This study's focus was documenting the migratory behavior of NSCs=20 [specifically those reported to be effective delivery vehicles = for=20 genes (5, 8) and = viral=20 vectors (10)]=20 in relation to aggressively invasive experimental = intracranial tumors=20 in adult brain and then to arm some of these cells with a = bioactive,=20 therapeutic gene requiring relative proximity to tumor cells=20 (allowing oncolysis to be a measure of the efficiency and = specificity=20 of gene expression).

Migratory Capacity of NSCs in Culture. To = visualize the=20 migratory properties of NSCs when confronted with a tumor, in = vitro=20 studies first assessed the relative migratory capacity of = NSCs=20 compared with fibroblasts cocultured with glioma cells. In = contrast=20 to fibroblasts, which remained localized to the area of = initial=20 seeding (Fig. 1A), NSCs=20 migrated rapidly and interspersed throughout the glioma = monolayer,=20 far from their initial site of seeding, robustly expressing=20 lacZ (Fig. 1B). These=20 patterns were observed whether the cells were plated directly = on top=20 of (i.e., in direct contact with) the glioma cells (arrows) = or=20 merely within the same culture medium and environment without = direct contact (arrowheads). The migrating NSCs then became=20 contact-inhibited and quiescent (do not incorporate BrdUrd; = ref. 5). =

NSCs Migrate Throughout and "Surround" Tumor in=20 Vivo. To determine the behavior of NSCs introduced to brain = tumors in=20 vivo, adult rats first received an implant of syngeneic D74 = rat=20 glioma cells (19) = into the=20 right frontal lobe. Four days later, lacZ-expressing = murine=20 NSCs were injected directly into the tumor bed (Paradigm = 1, Fig.=20 2). = Animals=20 were killed at 2- to 4-day intervals after intratumoral=20 injection and brain sections processed to detect 3D"beta-gal-producing NSCs. As early as 2 days after = injection, X-Gal+ blue donor NSCs were found = distributed=20 extensively throughout the darkly neutral red-stained tumor = mass=20 (Fig. 2A).=20 Although the transgene-expressing NSCs remained stably = intermixed=20 throughout the tumor (Fig. 2B,=20 arrows), up to and along the infiltrating tumor edge, they = largely=20 stopped at the border of the tumor where it interfaced with = normal=20 tissue (Fig. 2C,=20 arrowheads) as if "surrounding" the advancing neoplasm. = Normal adult=20 parenchyma presented a less permissive migratory environment = to NSCs,=20 except under one circumstance: where tumor cells began to = infiltrate=20 normal brain. In those instances, the NSCs migrated slightly = beyond=20 the tumor edge in conjunction with3D---=20 as=20 if following or "trailing"3D---=20individual infiltrating tumor cells (Fig. 2C,=20 arrow). This phenomenon was most dramatic when examined in = the=20 context of the more virulent and aggressively infiltrative = CNS-1=20 glioblastoma cell line (11) = (Fig. 2D),=20 which, in adult nude mice, demonstrates single cell invasive=20 characteristics analogous to those of human glioblastomas. = After=20 implantation as per Paradigm 1, extensive migration and=20 distribution of 3D"beta-gal+ donor cells was again noted = throughout=20 the darkly red-stained infiltrating tumor mass, up to, and = along the=20 encroaching tumor edge (red arrowheads), with further = migration into=20 the surrounding tissue in concert with and in virtual = juxtaposition=20 to aggressively invading tumor cells (Fig. 2D,=20 black arrows).

NSCs "Track" Infiltrating Tumor Cells. To better=20 distinguish single tumor cells migrating away from the main tumor mass, = CNS-1=20 glioblastoma cells were labeled ex vivo by retroviral=20 transduction of GFP cDNA before implantation (12). = After=20 implantation (as per Paradigm 1) of lacZ-expressing = NSCs into=20 the GFP-expressing CNS-1 tumor bed (Fig. 2 = E-L),=20 NSCs could not only be seen to distribute themselves = throughout=20 the tumor to its invading edge (Fig. 2 = E and=20 F), but could even more clearly be seen to "trail" = islands of=20 tumor cells migrating away from the main tumor mass (Fig. 2 = G and=20 H) as well as individual aggressive, dark red, or=20 GFP+, elongated infiltrating tumor cells (Fig. 2 = I-L). Of=20 note is the frequently observed apposition of = transgene-expressing=20 NSCs to invasive tumor cells (Fig. 2 = J-L,=20 arrows). The NSCs themselves never became tumorigenic. BrdUrd = pulsing=20 of animals before sacrifice confirmed prior observations that = donor=20 NSCs were quiescent in normal tissue (5), = nonmitotic=20 (i.e., BrdUrd3D-=20)=20 in the heart of the tumor, and with an occasional NSC that could=20 still incorporate BrdUrd at the advancing edge, a situation = optimal=20 for targeting therapy toward invading tumors. The vast = majority=20 of NSCs remained not only quiescent but undifferentiated, = expressing=20 only nestin.

NSCs Implanted Intracranially at Distant Sites = Migrate Toward=20 Tumor. To determine whether NSCs have the capacity to migrate = specifically=20 toward the tumor, NSCs were injected into uninvolved intracranial=20 sites distant from the main tumor mass in three separate = paradigms=20 (Fig. 3). In = each case,=20 donor NSCs migrating through normal adult tissue "targeted" = the=20 tumor. In Paradigm 2, NSCs were injected behind the=20 glioblastoma. NSCs were always found distributed within the = main=20 tumor bed, as well as in apposition to migrating tumor cells = in=20 surrounding tissue (Fig. 3 = A and=20 B), with very few NSCs in other locations. In Paradigm = 3, murine NSCs were injected into the contralateral = hemisphere.=20 NSCs (fluorescent red or X-Gal+ blue) were seen = migrating=20 across the corpus callosum and central commissures (Fig. 3 = C-E,=20 arrows) toward the tumor (fluorescent green or dark neutral=20 red+ masses delineated by arrowheads in Fig. 3F),=20 ultimately entering and populating (Fig. 3 = G and=20 H, arrows) the tumor on the opposite side of the = brain. In=20 Paradigm 4, NSCs were injected into the ipsilateral or=20 contralateral cerebral ventricle. NSCs (blue) again were seen = within=20 the main tumor bed (Fig. 3I), as=20 well as in juxtaposition to migrating "islands" of tumor = cells (dark=20 red) (Fig. 3J).=20 The only source of blue cells in these paradigms was the=20 "distant" NSC implant. Very few NSC-derived cells were found = in=20 normal brain tissue beyond the injection site, except when = tracking=20 toward the main tumor mass or near infiltrating tumor cells,=20 suggesting that, whereas NSCs migrated freely within the = tumor, the=20 normal adult brain parenchyma presented a less permissive=20 environment for migration. The NSCs themselves never became=20 tumorigenic. The tumors in transplant recipients were never = larger=20 than those in non-transplant recipients. NSC-derived cells = continued=20 to express their lacZ transgene exuberantly, often in = direct=20 contact with tumor cells. These NSC behaviors appeared to be=20 independent of the size or location of the tumor; findings = were=20 similar for large tumors, small tumor foci, and even single = scattered=20 tumor cells surrounding the main tumor mass. Fibroblasts = grafted as=20 controls never showed this dispersion or tropism, consistent = with=20 previous reports (15). =

NSCs Injected into Systemic Circulation "Target" = Intracerebral=20 Gliomas. Murine NSCs were injected into the tail vein of adult nude = mice in=20 which a CNS-1 glioblastoma had been established 1 wk before = in=20 the frontal lobe. Four days after i.v. NSC injection, albeit = with low=20 efficiency, anti-3D"beta-gal+ NSCs were distributed = throughout the=20 intracerebral tumor mass, but were not found in surrounding=20 normal-appearing brain tissue, elsewhere in the brain, or in = the=20 brains of control animals (NSC-injected mice without = intracerebral=20 gliomas or tumor-bearing mice in the absence of = NSC-injection).

Human NSCs and NSCs Expressing a Therapeutic Gene = Migrate to=20 Tumors. Because of the clinical implications of these migratory = phenomena,=20 we asked two further questions: (i) did these migratory=20 properties extend to human NSCs? and (ii) could relevant=20 bioactive genes be expressed?

In answer to the = first,=20 human NSCs (brown nuclei, arrows in Fig. 5 = A and=20 B) transplanted into the hemisphere contralateral = to a=20 CNS-1 glioma (Paradigm 3) indeed migrated across the corpus = callosum=20 and infiltrated and distributed themselves throughout the = targeted=20 tumor (arrowheads) as previously observed. That human NSCs = could=20 similarly target a true human glioblastoma is suggested in = Fig. 5=20 C-E in which Paradigm 3 was repeated employing = human=20 NSCs implanted contralateral to an HGL-21-derived tumor = established=20 in the nude mouse cerebrum; again, human NSCs migrated from = one=20 hemisphere to the other to infiltrate the glioblastoma.



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  		Fig. 5.   = Human NSCs=20 (hNSCs) possess tumor tracking characteristics. (A = and=20 B) Rodent CNS-1 glioblastoma cells and human NSCs = were=20 implanted as per Paradigm 3 into opposite hemispheres = of an=20 adult mouse. Pictured 7 days later at low power = (A) and=20 high power (B) is a section through the neutral = red-stained=20 tumor (outlined by arrowheads) intermixed with human NSCs=20 (identified by their brown nuclei following reaction with an = anti-human nuclear antibody) (arrows) that migrated from the = contralateral side. (C-E) Human HGL21 = glioblastoma=20 cells and hNSCs were similarly implanted into opposite = hemispheres.=20 Pictured at progressively higher power are sections through = that=20 neutral red-stained tumor intermixed with human NSCs=20 (X-Gal+ blue) that migrated from the = contralateral side.=20 (Scale bars: A, 20 =B5m, 15 =B5m in = B; C,=20 60 =B5m, 30 =B5m in D, and 15 =B5m in = E.)

To address the second question, NSCs were stably transduced with = a=20 transgene encoding the enzyme CD. CD can convert the nontoxic=20 "prodrug" 5-FC to the oncolytic drug 5-fluorouracil, a=20 chemotherapeutic agent that readily diffuses into tumor cells = and has=20 selective toxicity to rapidly dividing cells (18, 19). = The CD gene=20 provided an opportunity to examine a prototypical bioactive = gene with=20 a relevant, specific, quantifiable read-out of functionality=20 (oncolysis) that might be enhanced by tumor proximity. = CD-bearing=20 NSCs retained their extensive migratory, tumor-tracking = properties.=20 To determine quantitatively whether a gene such as CD within = an NSC=20 retains its bioactivity3D---=20as=20 assayed in this case by its anti-tumor effect3D---=20CD-bearing NSCs were first cocultured with glioma = cells and=20 then, when nearly confluent (Fig. 6A),=20 exposed to 5-FC. Death of surrounding tumor cells was induced = (Fig.=20 6B),=20 even when the ratio of NSCs-to-tumor cells was as low as 1:4. = NSCs=20 that were mitotic at the time of 5-FC exposure = self-eliminated.=20 Control plates of tumor alone were not significantly killed = by the=20 same dose of 5-FC. To determine whether this bioactivity was = retained=20 in vivo, we used CD-transduced NSCs to express this = gene=20 within an intracranial glioma established in an adult nude = mouse=20 (3.5 =D7 104 NSCs to 7 =D7 104 = CNS-1 tumor=20 cells in a 1:2 ratio). After systemic treatment with 5-FC, = there was=20 dramatic (3D"approx80%) reduction in the resultant tumor mass = at=20 2 wk postimplantation as compared with that in untreated = animals=20 (Fig. 7),=20 indicative of CD bioactivity.



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  		Fig. 6.   = Bioactive=20 transgene (CD) remains functional (as assayed by in = vitro=20 oncolysis) when expressed within NSCs. CNS-1 glioblastoma = cells=20 (red) were cocultured with CD-transduced murine NSCs = (A and=20 B) (blue). Cocultures unexposed to 5-FC grew = healthily and=20 confluent (A), whereas plates exposed to 5-FC showed = dramatic=20 loss of tumor cells (B), represented quantitatively = by the=20 histograms (*, P < 0.001). The oncolytic = effect=20 was identical whether 1 =D7 105 CD-NSCs = or half=20 that number were cocultured with a constant number of tumor = cells.=20 (In this paradigm, subconfluent NSCs were still mitotic at = the time=20 of 5-FC exposure and thus also subject to self-elimination = by the=20 generated 5-fluorouracil and its toxic metabolites.)



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  		Fig. 7.   = Expression=20 of a bioactive transgene (CD) delivered by NSCs is retained = in=20 vivo as assayed by reduction in tumor mass. The size of = an=20 intracranial glioblastoma populated with CD-NSCs in an adult = nude=20 mouse treated with 5-FC was compared with that of tumor = treated with=20 5-FC but lacking CD-NSCs. These data, standardized against = and=20 expressed as a percentage of a control tumor populated with = CD-NSCs=20 receiving no treatment, are presented in the histograms in = A.=20 These measurements were derived from measuring the surface = area of=20 tumors (like those in Figs. 2-5),=20 representative camera lucidas of which are presented in = B-D.=20 Note the large areas of a control non-5-FC-treated tumor = containing=20 CD-NSCs (B) and a control 5-FC-treated tumor lacking = CD-NSCs=20 (C) as compared with the dramatically smaller tumor = areas of=20 the 5-FC-treated animal who also received CD-NSCs (D) = (80% reduction as per the histogram in = A; *,=20 P < 0.001), suggesting both activity and = specificity of the transgene. The lack of effect of 5-FC on = tumor=20 mass when no CD-bearing NSCs were within the tumor = (C) was=20 identical to the effect of CD-NSCs in the tumor without the = gene=20 being used (B).


    Discussion
Top
Abstract
Introduction
Experimental Methods
Results
Discussion
References

Transplanted NSCs have recently been recognized for their remarkable = ability=20 to migrate throughout the CNS, become normal constituents of = the host=20 cytoarchitecture, and disseminate bioactive molecules and = retroviral=20 vectors (3-10).=20 Intriguingly, this migratory ability emulates the invasive = spread of=20 some brain tumors, e.g., gliomas. Here, we show that an = unanticipated=20 benefit of the directed, migratory capacity of = transgene-expressing=20 NSCs, including of human origin, may be to target invasive = primary=20 brain tumors (also including of human origin) that have = proven=20 refractory to current treatments (1, 2). = Most gene=20 therapy strategies employ viral vectors to deliver genes = directly to=20 tumor cells in vivo; however, the distribution of = genes to the=20 extensive regions and large numbers of cells in need of = attack has=20 been limited. The present study demonstrates the ability of = NSCs to=20 migrate expeditiously throughout a tumor mass and, presumably = drawn=20 by the degenerative or inflammatory environment created at = the=20 infiltrating tumor edge, to "surround" the invading tumor = border, all=20 while continuing to express a bioactively relevant transgene. = Moreover, the foreign gene-expressing NSCs seem to follow or=20 "track"3D---virtually ride "piggy-back" upon3D---=20those aggressively infiltrating tumor cells = escaping into=20 normal tissue. Although the NSCs migrate freely through the = tumor,=20 the normal adult brain parenchyma seems to present a less = permissive=20 environment for their migration, except when NSCs (even at = sites=20 distant from the tumor) travel in a directed fashion through = normal=20 adult CNS tissue to target the main tumor mass as well as = individual=20 infiltrating, tumor cells. The practical implication is that = NSCs=20 might actually "seek out" tumor foci that may have migrated=20 undetected far away from the main tumor mass, not an uncommon = occurrence with glioblastoma. Such behaviors are not = displayed by=20 cells of nonneural origin (15). = Indeed,=20 targeting may be so powerful that even NSCs injected into the = systemic circulation may preferentially populate intracerebral=20 gliomas.

Hence, NSCs evince extensive tropism for the tumor itself or for the=20 degenerating CNS it engenders. That the tumor itself = elaborates at=20 least some of the tropic cues is suggested by our in = vitro=20 studies in which glioma cells in culture, isolated from = surrounding=20 brain, prompted NSCs to migrate, by both contact and=20 non-contact-mediated factors. Alternatively, or in addition, = tropic=20 cytokines may be released by extensively damaged normal = tissue.=20 That minor CNS destruction alone could not prompt the = dramatic=20 migration seen in vivo was suggested by NSC transplants = into=20 "mock" tumor bearing animals, i.e., animals in which a needle = was=20 inserted to emulate the tissue damage of establishing an = experimental=20 tumor bed but without the actual implantation of glioblastoma = cells; NSCs did not migrate toward the site in this = circumstance.=20 Nevertheless, in other previously reported experimental = situations in=20 which significant neuronal death was rendered (13), = NSC=20 differentiation was altered by apparent trophic influences.=20 Therefore, the signals to which the NSCs are responding are = most=20 likely complex, from multiple sources, and representing a = "mixture"=20 of attractants, adhesion and substrate molecules, chemokines, = etc. Of=20 broader biological significance, these findings suggest that=20 migration can be unexpectedly extensive, even in adult brain = and=20 along nonstereotypical routes, if pathology (as modeled here = by=20 tumor) is present.

Having documented this powerful tropic interaction between NSCs and=20 intracranial pathology, we believe that exogenous NSCs, = genetically=20 engineered ex vivo and strategically implanted, may = provide a=20 "platform" for the dissemination of therapeutic genes and/or = gene=20 products to previously inaccessible infiltrating tumor cells. = As=20 suggested in our CD/5-FC prodrug paradigm, NSCs were able to = express=20 a bioactive transgene in vivo to effect a significant=20 biologically relevant read-out (dramatic reduction in tumor = burden).=20 Cytotoxic 5-fluorouracil and its toxic metabolites can = readily=20 diffuse into surrounding tumor cells giving CD an impressive=20 "bystander" effect; as little as 2% of the tumor mass = containing=20 CD-expressing cells may generate significant oncolysis (22). = Indeed,=20 NSCs engineered to express CD are attractive as molecular = pumps=20 because they can generate agents that kill tumor cells yet = undergo=20 self-elimination should the NSCs themselves become mitotic. = This=20 prototypical genetic strategy represents one of many = potential=20 approaches to treating brain tumors with migratory = genetically=20 engineered NSCs. Other candidates include genes encoding: = proteins=20 that induce differentiation of neoplastic cells and/or their=20 signal-transduction mediators; cell cycle modulators;=20 apoptosis-promoting agents; anti-angiogenesis factors;=20 immune-enhancing agents (23); = fusion=20 agents; and oncolytic factors (24). = That these=20 same engraftable migratory NSCs have been demonstrated to = serve as=20 intracerebral viral vector producer cells (10) = may allow=20 "extended" delivery of lethal viral-mediated genes to larger = numbers=20 of tumor cells in broader regions of brain. Instilled into = the=20 resection or biopsy cavity, or applied intermittently into or = near=20 the tumor mass or suspicious tissue or into cerebral = ventricles,=20 engineered NSCs could be used in conjunction with other=20 interventions. An NSC-based strategy3D---=20responding to the altered biology of the abnormal = adult=20 brain and by virtue of their unique inherent biology3D---=20may=20 both optimize present approaches and make feasible new ones = for more=20 effectively, selectively, and safely targeting genes and = vectors to=20 refractory, migratory, invasive brain tumors. =


    Acknowledgements

We thank M. Sena-Esteves for help with generating retrovirus = vectors.=20 This work is supported by National Institutes of Health = Grants=20 HD07466 and CA86768 (to K.S.A.) and CA69246 (to X.O.B.), the = Toennies=20 Stiftung (to N.G.R.), the Deutsche Forschungsgemeinschaft (to = U.H.),=20 National Institutes of Health Grants NS33852 and NS34247, and = the=20 Brain Tumor Society (to E.Y.S.). Portions of this work were = presented=20 at the Second Conference on Cellular and Molecular Treatments = of=20 Neurological Diseases, Cambridge, MA, October=20 10, 1998. This work is dedicated to the memory of = Dr. James=20 A. Galambos.


    Abbreviations

NSC, neural stem cell; CNS, central nervous system; GFP, green = fluorescent=20 protein; 3D"beta-gal, 3D"beta-galactosidase; X-Gal, 5-bromo-4-chloro-3-indolyl -D-galactoside; CD, cytosine = deaminase;=20 5-FC, 5-fluorocytosine; CD-NSC, CD-transduced NSC.


    Footnotes

=B6 To whom reprint requests should be = addressed.=20 E-mail: Snyder{at}A1.TCH.Harvard.edu .=20

See commentary on page 12393.=20


    References
Top
Abstract
Introduction
Experimental Methods
Results
Discussion
References =

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23.=20 Benedetti, S. , Pirola, B. , Pollo, B. , Gagrassi, = L. ,=20 Briuzzone, M. G. , Rigamonti, D. , Galli, R. , Selleri, S. , = DiMeco, R. ,=20 DeFraja, C. , et al. (2000) Nat. Med. 6, = 447-450[CrossRef][ISI][Medline]=20 .
24.=20 Herrlinger, U. , Woiciechowski, C. , Aboody, K. S. = ,=20 Jacobs, A. H. , Rainov, N. G. , Snyder, E. Y. & Breakefield, = X. O.=20 (2000) Mol. Ther. 1, 347-357[CrossRef][ISI][Medline]=20 .

=20 CiteULike   =20 Complore   =20 Connotea   =20 Del.icio.us    Digg    What's this? =

Related=20 Commentary in PNAS:=20

Can neural stem cells be used to track down and destroy = migratory=20 brain tumor cells while also providing a means of repairing = tumor-associated=20 damage?=20
Mark Noble
PNAS 2000 97: 12393-12395. [Extract] [Full = Text]=20  =20



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 3D""=20 3D"Stem3D"Home
F.-J.=20 Mueller, N. Serobyan, I. U. Schraufstatter, R. DiScipio, D. = Wakeman, J. F.=20 Loring, E. Y. Snyder, and S. K. Khaldoyanidi
Adhesive=20 Interactions Between Human Neural Stem Cells and Inflamed Human = Vascular=20 Endothelium Are Mediated by Integrins
Stem Cells,=20 November 1, 2006; 24(11): 2367 - 2372.
[Abstract]=20 [= Full=20 Text] [PDF]<= /A>=20
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P. O.=20 Sakariassen, L. Prestegarden, J. Wang, K.-O. Skaftnesmo, R. = Mahesparan, C.=20 Molthoff, P. Sminia, E. Sundlisaeter, A. Misra, B. B. Tysnes, = et=20 al.
Angiogenesis-independent tumor growth mediated = by=20 stem-like cancer cells
PNAS, = October 31, 2006;=20 103(44): 16466 - 16471.
[Abstract]= =20 [Full=20 Text] [PDF]=20
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 3D""=20 3D"Clin.3D"Home
S.-K.=20 Kim, S. U. Kim, I. H. Park, J. H. Bang, K. S. Aboody, K.-C. Wang, = B.-K.=20 Cho, M. Kim, L. G. Menon, P. M. Black, et = al.
Human=20 neural stem cells target experimental intracranial medulloblastoma = and=20 deliver a therapeutic gene leading to tumor = regression.
Clin.=20 Cancer Res., September 15, 2006; 12(18): 5550 - 5556. =
[Abstract]=20 [Full=20 Text] [PD= F]=20
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 3D""=20 3D"Neuro3D"Home
K. S.=20 Aboody, J. Najbauer, N. O. Schmidt, W. Yang, J. K. Wu, Y. Zhuge, = W.=20 Przylecki, R. Carroll, P. M. Black, and G. = Perides
Targeting of=20 melanoma brain metastases using engineered neural stem/progenitor=20 cells
Neuro-oncol, April 1, 2006; 8(2): 119 = - 126.=20
[Abstract]=20 = [Full=20 Text] [PDF]= =20
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 3D""=20 3D"Home
A.=20 Belmadani, P. B. Tran, D. Ren, and R. J. = Miller
Chemokines=20 regulate the migration of neural progenitors to sites of=20 neuroinflammation.
J. Neurosci., = March 22, 2006;=20 26(12): 3182 - 3191.
[Abstra= ct]=20 [Full=20 Text] [PDF] =
=20 		3D""=20
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 3D""=20 3D"Cancer3D"Home
X.=20 Yuan, J. Hu, M. L. Belladonna, K. L. Black, and J. S.=20 Yu
Interleukin-23-Expressing Bone Marrow-Derived Neural = Stem-Like Cells Exhibit Antitumor Activity against Intracranial=20 Glioma.
Cancer Res., March 1, 2006; 66(5): = 2630 -=20 2638.
[Abstract]=20 [Fu= ll=20 Text] [PDF]=20
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 3D""=20 3D"Stem3D"Home
J. S.=20 Meyer, M. L. Katz, J. A. Maruniak, and M. D. = Kirk
Embryonic=20 Stem Cell-Derived Neural Progenitors Incorporate into Degenerating = Retina=20 and Enhance Survival of Host Photoreceptors
Stem = Cells,=20 February 1, 2006; 24(2): 274 - 283.
[Abstract]=20 [Fu= ll=20 Text] [PDF]=20
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 3D""=20 3D"Clin.3D"Home
S.-C.=20 Hung, W.-P. Deng, W. K. Yang, R.-S. Liu, C.-C. Lee, T.-C. Su, = R.-J. Lin,=20 D.-M. Yang, C.-W. Chang, W.-H. Chen, et=20 al.
Mesenchymal Stem Cell Targeting of Microscopic = Tumors=20 and Tumor Stroma Development Monitored by Noninvasive In vivo = Positron=20 Emission Tomography Imaging
Clin. Cancer Res.,=20 November 1, 2005; 11(21): 7749 - 7756.
[Abstract]=20 [Full=20 Text] [PD= F]=20
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 3D""=20 3D"Neuro3D"Home
O.=20 Heese, A. Disko, D. Zirkel, M. Westphal, and K. = Lamszus
Neural=20 stem cell migration toward gliomas in = vitro
Neuro-oncol,=20 October 1, 2005; 7(4): 476 - 484.
[Abstract]=20 [PDF]= =20
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 3D""=20 3D"Arch3D"Home
K.=20 Warfvinge, J. F. Kiilgaard, E. B. Lavik, E. Scherfig, R. Langer, = H. J.=20 Klassen, and M. J. Young
Retinal Progenitor Cell = Xenografts to=20 the Pig Retina: Morphologic Integration and Cytochemical=20 Differentiation
Arch Ophthalmol, = October 1, 2005;=20 123(10): 1385 - 1393.
[A= bstract]=20 [Full = Text] [PDF]=20
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G.=20 Tabatabai, O. Bahr, R. Mohle, I. Y. Eyupoglu, A. M. Boehmler, J.=20 Wischhusen, J. Rieger, I. Blumcke, M. Weller, and W.=20 Wick
Lessons from the bone marrow: how malignant glioma = cells=20 attract adult haematopoietic progenitor cells
Brain,=20 September 1, 2005; 128(9): 2200 - 2211.
= [Abstract]=20 [Ful= l=20 Text] [PDF]= =20
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N.=20 Sanai, A. Alvarez-Buylla, and M. S. Berger
Neural Stem = Cells=20 and the Origin of Gliomas
N. Engl. J. Med.,=20 August 25, 2005; 353(8): 811 - 822.
[Full = Text]=20 [PDF] =
=20 		3D""=20
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 3D""=20 3D"Clin.3D"Home
S.-K.=20 Kim, T. G. Cargioli, M. Machluf, W. Yang, Y. Sun, R. Al-Hashem, S. = U. Kim,=20 P. M. Black, and R. S. Carroll
PEX-Producing Human = Neural Stem=20 Cells Inhibit Tumor Growth in a Mouse Glioma = Model
Clin.=20 Cancer Res., August 15, 2005; 11(16): 5965 - 5970. =
[Abstract]=20 [Full=20 Text] [PD= F]=20
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 3D""=20 3D"Cancer3D"Home
A.=20 Nakamizo, F. Marini, T. Amano, A. Khan, M. Studeny, J. Gumin, J. = Chen, S.=20 Hentschel, G. Vecil, J. Dembinski, et = al.
Human Bone=20 Marrow-Derived Mesenchymal Stem Cells in the Treatment of=20 Gliomas
Cancer Res., April 15, 2005; 65(8): = 3307 -=20 3318.
[Abstract]=20 [Fu= ll=20 Text] [PDF]=20
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R.=20 Glass, M. Synowitz, G. Kronenberg, J.-H. Walzlein, D. S. Markovic, = L.-P.=20 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] =
=20 		3D""=20
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 3D""=20 3D"Home
J.=20 Imitola, K. Raddassi, K. I. Park, F.-J. Mueller, M. Nieto, Y. D. = Teng, D.=20 Frenkel, J. Li, R. L. Sidman, C. A. Walsh, et=20 al.
Directed migration of neural stem cells to = sites of=20 CNS injury by the stromal cell-derived factor 1{alpha}/CXC = chemokine=20 receptor 4 pathway
PNAS, December 28, 2004; = 101(52):=20 18117 - 18122.
[Abstract]= =20 [Full=20 Text] [PDF]=20
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S.=20 Kelly, T. M. Bliss, A. K. Shah, G. H. Sun, M. Ma, W. C. Foo, J. = Masel, M.=20 A. Yenari, I. L. Weissman, N. Uchida, et=20 al.
Transplanted human fetal neural stem cells = survive,=20 migrate, and differentiate in ischemic rat cerebral=20 cortex
PNAS, August 10, 2004; 101(32): 11839 = -=20 11844.
[Abstract]= =20 [Full=20 Text] [PDF]=20
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K.=20 Staflin, G. Honeth, S. Kalliomaki, C. Kjellman, K. Edvardsen, and = M.=20 Lindvall
Neural Progenitor Cell Lines Inhibit Rat Tumor = Growth=20 in Vivo
Cancer Res., August 1, 2004; 64(15): = 5347 -=20 5354.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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C. I.=20 FERNANDEZ, E. ALBERTI, Y. MENDOZA, L. MARTINEZ, J. COLLAZO, J. C. = ROSILLO,=20 and J. Y. BAUZA
Motor and Cognitive Recovery Induced by = Bone=20 Marrow Stem Cells Grafted to Striatum and Hippocampus of Impaired = Aged=20 Rats: Functional and Therapeutic Considerations
Ann. = N.Y.=20 Acad. Sci., June 1, 2004; 1019(1): 48 - 52.
[Abstra= ct]=20 [Full=20 Text] [PDF] =
=20 		3D""=20
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K.=20 Shah, C.-H. Tung, K. Yang, R. Weissleder, and X. O.=20 Breakefield
Inducible Release of TRAIL Fusion Proteins = from a=20 Proapoptotic Form for Tumor Therapy
Cancer Res.,=20 May 1, 2004; 64(9): 3236 - 3242.
[Abstract]=20 [Fu= ll=20 Text] [PDF]=20
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D.-E.=20 Kim, D. Schellingerhout, K. Ishii, K. Shah, and R.=20 Weissleder
Imaging of Stem Cell Recruitment to Ischemic = Infarcts in a Murine Model
Stroke, = April 1, 2004;=20 35(4): 952 - 957.
[Abs= tract]=20 [Full=20 Text] [PDF]=20
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J.=20 Aarum, K. Sandberg, S. L. B. Haeberlein, and M. A. A.=20 Persson
Migration and differentiation of neural = precursor cells=20 can be directed by microglia
PNAS,=20 December 23, 2003; 100(26): 15983 - 15988.
[Abstract]= =20 [Full=20 Text] [PDF]=20
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 3D""=20 3D"Cancer3D"Home
J.=20 Lee, A. G. Elkahloun, S. A. Messina, N. Ferrari, D. Xi, C. L. = Smith, R.=20 Cooper Jr., P. S. Albert, and H. A. Fine
Cellular and = Genetic=20 Characterization of Human Adult Bone Marrow-Derived Neural = Stem-Like=20 Cells: A Potential Antiglioma Cellular Vector
Cancer = Res.,=20 December 15, 2003; 63(24): 8877 - 8889.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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 3D""=20 3D"Physiol.3D"Home
J.=20 Imitola, E. Y. Snyder, and S. J. Khoury
Genetic = programs and=20 responses of neural stem/progenitor cells during demyelination: = potential=20 insights into repair mechanisms in multiple = sclerosis
Physiol=20 Genomics, August 15, 2003; 14(3): 171 - 197.
[Abstract]=20 = [Full=20 Text] [PDF]= =20
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D.=20 Kaushal, J. J. A. Contos, K. Treuner, A. H. Yang, M. A. Kingsbury, = S. K.=20 Rehen, M. J. McConnell, M. Okabe, C. Barlow, and J.=20 Chun
Alteration of Gene Expression by Chromosome Loss = in the=20 Postnatal Mouse Brain
J. Neurosci., = July 2, 2003;=20 23(13): 5599 - 5606.
[Abstra= ct]=20 [Full=20 Text] [PDF] =
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R A=20 Barker, M Jain, R J E Armstrong, and M A Caldwell
Stem = cells=20 and neurological disease
J. Neurol. Neurosurg. = Psychiatry,=20 May 1, 2003; 74(5): 553 - 557.
[Full=20 Text] [PDF] =
=20 		3D""=20
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 3D""=20 3D"Stem3D"Home
K. E.=20 Mitchell, M. L. Weiss, B. M. Mitchell, P. Martin, D. Davis, L. = Morales, B.=20 Helwig, M. Beerenstrauch, K. Abou-Easa, T. Hildreth, et=20 al.
Matrix Cells from Wharton's Jelly Form Neurons = and=20 Glia
Stem Cells, January 1, 2003; 21(1): 50 = - 60.=20
= [Abstract]=20 [Ful= l=20 Text] [PDF]= =20
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 3D""=20 3D"Cancer3D"Home
M.=20 Ehtesham, P. Kabos, M. A. R. Gutierrez, N. H. C. Chung, T. S. = Griffith, K.=20 L. Black, and J. S. Yu
Induction of Glioblastoma = Apoptosis=20 Using Neural Stem Cell-mediated Delivery of Tumor Necrosis = Factor-related=20 Apoptosis-inducing Ligand
Cancer Res.,=20 December 15, 2002; 62(24): 7170 - 7174.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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 3D""=20 3D"Cancer3D"Home
M.=20 Ehtesham, P. Kabos, A. Kabosova, T. Neuman, K. L. Black, and J. S. = Yu
The Use of Interleukin 12-secreting Neural Stem = Cells for=20 the Treatment of Intracranial Glioma
Cancer Res.,=20 October 15, 2002; 62(20): 5657 - 5663.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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 3D""=20 3D"Cancer3D"Home
C. R.=20 Miller, C. R. Williams, D. J. Buchsbaum, and G. Y.=20 Gillespie
Intratumoral 5-Fluorouracil Produced by = Cytosine=20 Deaminase/5-Fluorocytosine Gene Therapy Is Effective for = Experimental=20 Human Glioblastomas
Cancer Res., = February 1, 2002;=20 62(3): 773 - 780.
= [Abstract]=20 [Ful= l=20 Text] [PDF]= =20
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A.=20 Gritti, L. Bonfanti, F. Doetsch, I. Caille, A. Alvarez-Buylla, D. = A. Lim,=20 R. Galli, J. M. G. Verdugo, D. G. Herrera, and A. L.=20 Vescovi
Multipotent Neural Stem Cells Reside into the = Rostral=20 Extension and Olfactory Bulb of Adult Rodents
J. = Neurosci.,=20 January 15, 2002; 22(2): 437 - 445.
[Abstract= ]=20 [Full=20 Text] [PDF] =
=20 		3D""=20
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 3D""=20 3D"Cancer3D"Home
T.=20 Ichikawa and E. A. Chiocca
Comparative Analyses of = Transgene=20 Delivery and Expression in Tumors Inoculated with a=20 Replication-conditional or -defective Viral = Vector
Cancer=20 Res., July 1, 2001; 61(14): 5336 - 5339.
[Abstract]=20 [F= ull=20 Text] [PDF]=20
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E. A.=20 Maher, F. B. Furnari, R. M. Bachoo, D. H. Rowitch, D. N. Louis, W. = K.=20 Cavenee, and R. A. DePinho
Malignant glioma: genetics = and=20 biology of a grave matter
Genes & Dev.,=20 June 1, 2001; 15(11): 1311 - 1333.
[Full = Text]=20 [PDF]=20
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K.=20 Jin, M. Minami, J. Q. Lan, X. O. Mao, S. Batteur, R. P. Simon, and = D. A.=20 Greenberg
Neurogenesis in dentate subgranular zone and = rostral=20 subventricular zone after focal cerebral ischemia in the=20 rat
PNAS, April 10, 2001; 98(8): 4710 - = 4715.
[Abstract]=20 [Full = Text]=20 [PDF] =
=20 		3D""=20
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 3D""=20 3D"Hum3D"Home
P.=20 Y.P. Lam and X. O. Breakefield
Potential of gene = therapy for=20 brain tumors
Hum. Mol. Genet., = April 1, 2001; 10(7):=20 777 - 787.
[Abs= tract]=20 [Full=20 Text] [PDF]=20
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M.=20 Noble
Can neural stem cells be used to track down and = destroy=20 migratory brain tumor cells while also providing a means of = repairing=20 tumor-associated damage?
PNAS, = November 7, 2000;=20 97(23): 12393 - 12395.
[Full = Text] [PDF] =
=20 		3D""=20

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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_01C85862.60200690 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 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quoted-printable Content-Location: http://www.pnas.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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i < src.childNodes.length; i++) {=0A= copy_xml_to_html(src.childNodes[i], dst);=0A= }=0A= var tbl =3D tr.parentNode;=0A= tbl.replaceChild(dst, tr);=0A= }=0A= }=0A= req.open('GET', callback_url, true);=0A= req.send(null);=0A= }=0A= ------=_NextPart_000_0000_01C85862.60200690 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.google-analytics.com/urchin.js //-- Google Analytics Urchin Module=0A= //-- Copyright 2007 Google, All Rights Reserved.=0A= =0A= //-- Urchin On Demand Settings ONLY=0A= var _uacct=3D""; // set up the Urchin Account=0A= var _userv=3D1; // service mode (0=3Dlocal,1=3Dremote,2=3Dboth)=0A= =0A= //-- UTM User Settings=0A= var _ufsc=3D1; // set client info flag (1=3Don|0=3Doff)=0A= var _udn=3D"auto"; // (auto|none|domain) set the domain name for cookies=0A= var _uhash=3D"on"; // (on|off) unique domain hash for cookies=0A= var _utimeout=3D"1800"; // set the inactive session timeout in seconds=0A= var _ugifpath=3D"/__utm.gif"; // set the web path to the __utm.gif file=0A= var _utsp=3D"|"; // transaction field separator=0A= var _uflash=3D1; // set flash version detect option (1=3Don|0=3Doff)=0A= var _utitle=3D1; // set the document title detect option = (1=3Don|0=3Doff)=0A= var _ulink=3D0; // enable linker functionality (1=3Don|0=3Doff)=0A= var _uanchor=3D0; // enable use of anchors for campaign = (1=3Don|0=3Doff)=0A= var _utcp=3D"/"; // the cookie path for tracking=0A= var _usample=3D100; // The sampling % of visitors to track (1-100).=0A= =0A= //-- UTM Campaign Tracking Settings=0A= var _uctm=3D1; // set campaign tracking module (1=3Don|0=3Doff)=0A= var _ucto=3D"15768000"; // set timeout in seconds (6 month default)=0A= var _uccn=3D"utm_campaign"; // name=0A= var _ucmd=3D"utm_medium"; // medium (cpc|cpm|link|email|organic)=0A= var _ucsr=3D"utm_source"; // source=0A= var _uctr=3D"utm_term"; // term/keyword=0A= var _ucct=3D"utm_content"; // content=0A= var _ucid=3D"utm_id"; // id number=0A= var _ucno=3D"utm_nooverride"; 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path=3D"+_utcp+";"+nx+_udo;=0A= _ubd.cookie=3D"__utmb=3D"+_udh+"; path=3D"+_utcp+";"+x+_udo;=0A= _ubd.cookie=3D"__utmc=3D"+_udh+"; path=3D"+_utcp+";"+_udo;=0A= _ufns=3D1;=0A= } else if (a>=3D0 && b>=3D0 && c>=3D0) {=0A= _ubd.cookie=3D"__utmb=3D"+_udh+"; path=3D"+_utcp+";"+x+_udo;=0A= } else {=0A= if (a>=3D0) a=3D_uFixA(_ubd.cookie,";",_ust);=0A= else a=3D_udh+"."+_uu+"."+_ust+"."+_ust+"."+_ust+".1";=0A= _ubd.cookie=3D"__utma=3D"+a+"; path=3D"+_utcp+";"+nx+_udo;=0A= _ubd.cookie=3D"__utmb=3D"+_udh+"; path=3D"+_utcp+";"+x+_udo;=0A= _ubd.cookie=3D"__utmc=3D"+_udh+"; path=3D"+_utcp+";"+_udo;=0A= _ufns=3D1;=0A= }=0A= if (_ulink && xx && xx!=3D"" && xx!=3D"-") {=0A= xx=3D_uUES(xx);=0A= if (xx.indexOf(";")=3D=3D-1) _ubd.cookie=3D"__utmx=3D"+xx+"; = path=3D"+_utcp+";"+nx+_udo;=0A= }=0A= if (_ulink && v && v!=3D"" && v!=3D"-") {=0A= v=3D_uUES(v);=0A= if (v.indexOf(";")=3D=3D-1) _ubd.cookie=3D"__utmv=3D"+v+"; = path=3D"+_utcp+";"+nx+_udo;=0A= }=0A= _uInfo(page);=0A= _ufns=3D0;=0A= _ufno=3D0;=0A= if (!page || page=3D=3D"") _uff=3D1;=0A= }=0A= function _uInfo(page) {=0A= var p,s=3D"",dm=3D"",pg=3D_udl.pathname+_udl.search;=0A= if (page && page!=3D"") pg=3D_uES(page,1);=0A= _ur=3D_ubd.referrer;=0A= if (!_ur || _ur=3D=3D"") { _ur=3D"-"; 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