From: Subject: =?Windows-1252?Q?Direct_isolation_of_human_central_nervous_system_stem_ce?= =?Windows-1252?Q?lls_=97_PNAS?= Date: Sun, 8 Nov 2009 10:08:57 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_003B_01CA605B.7C435B80" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5579 This is a multi-part message in MIME format. ------=_NextPart_000_003B_01CA605B.7C435B80 Content-Type: text/html; charset="utf-8" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.pnas.org/content/97/26/14720.long =EF=BB=BF Direct isolation of = human central nervous system stem cells =E2=80=94 PNAS

Direct isolation of human central nervous = system stem=20 cells

  1. Nobuko=20 Uchida*,=E2=80=A0,=
  2. David=20 W. Buck*, =
  3. Dongping=20 He*, =
  4. Michael=20 J. Reitsma*, =
  5. Marilyn=20 Masek*, =
  6. Thinh=20 V. Phan*, =
  7. Ann=20 S. Tsukamoto*, =
  8. Fred=20 H. Gage=E2=80=A1= , and
  9. Irving=20 L. Weissman=C2=A7
  1. *StemCells, Inc., 525 Del Rey Avenue, Suite C, = Sunnyvale,=20 CA 94085; =E2=80=A1Laboratory of Genetics, The Salk = Institute, 10010 North=20 Torrey Pines Road, La Jolla, CA 92037; and = =C2=A7Departments of=20 Pathology and Developmental Biology, Stanford University, Stanford, CA = 90305=20

  1. Contributed by Irving L. Weissman

Next=20 Section

Abstract

Stem cells, which are clonogenic cells with self-renewal and = multilineage differentiation properties, have the potential to replace = or repair=20 damaged tissue. We have directly isolated clonogenic human central = nervous=20 system stem cells (hCNS-SC) from fresh human fetal brain tissue, using=20 antibodies to cell surface markers and fluorescence-activated cell = sorting.=20 These hCNS-SC are phenotypically 5F3 (CD133)+, = 5E12+,=20 CD34=E2=88=92, CD45=E2=88=92, and = CD24=E2=88=92/lo. Single=20 CD133+ CD34=E2=88=92 CD45=E2=88=92 = sorted cells initiated=20 neurosphere cultures, and the progeny of clonogenic cells could = differentiate=20 into both neurons and glial cells. Single cells from neurosphere = cultures=20 initiated from CD133+ CD34=E2=88=92 = CD45=E2=88=92 cells were=20 again replated as single cells and were able to reestablish neurosphere=20 cultures, demonstrating the self-renewal potential of this highly = enriched=20 population. Upon transplantation into brains of immunodeficient neonatal = mice,=20 the sorted/expanded hCNS-SC showed potent engraftment, proliferation, = migration,=20 and neural differentiation.

Although much is known about the development of the brain = during=20 embryonic life, the pathways and lineages that neural precursors take in = neurogenesis during development and adult life are only partly = understood (1). = Until=20 recently, it was unclear whether neurogenesis proceeded like = hematopoiesis,=20 i.e., from pluripotent clonogenic stem cells that both self-renew to = produce=20 multipotent progenitors and give rise to progressively more committed = progeny=20 (2), or = if some=20 other developmental hierarchy, not including stem cells, was operative.=20 Recently, long-term cell culture systems have been developed for rodent = and=20 human central nervous system (CNS) cells that continuously propagate a=20 heterogeneous population of early neural stem and/or progenitor cells. = CNS=20 neural stem/progenitor cells can grow either in monolayers on = substrate-coated=20 tissue culture plates (3, 4) or as = self-adherent complexes of cells, forming clusters known as neurospheres = (5, 6). In = both=20 culture systems, these cells maintain the capacity to produce the three = main=20 mature cell classes of the CNS: neurons, astrocytes, and = oligodendrocytes (6, 7). = In=20 vivo analysis of transplanted cultured rodent cells shows = engraftment,=20 migration, and site-specific multilineage differentiation in mice and = rats (8, 9). = Human=20 long-term cultured neurosphere cells show similar engraftment, = migration, and=20 site-specific differentiation upon transplantation into immunosuppressed = rats=20 (10). = These rodent=20 and human studies suggest that the neural cultures support the growth of = stem=20 cells and/or progenitors capable of engraftment and differentiation = (3=E2=80=9314).=20 Recently, Roy et al. (13) = showed that=20 human primitive progenitor cells from the dentate gyrus of adult = hippocampus can=20 be selected by transfecting them with the reporter green fluorescent = protein=20 gene driven by the nestin enhancer. However, the direct isolation of = human=20 neural stem cells from fresh tissues through the identification of cell = surface=20 markers and fluorescence-activated cell sorting (FACS) of either the = central or=20 peripheral nervous system, to our knowledge, has not yet been reported. =

In the past decade, hematopoietic stem and progenitor cells = have been=20 identified by using mAbs against cell surface markers for enriching rare = subpopulations that are clonal self-renewing and multipotent stem cells = (15, 16) or=20 oligopotent progenitors (17, 18). = This=20 strategy was used successfully to isolate rat peripheral nervous system = stem=20 cells (14) = and, as we=20 report here, to identify and isolate a candidate human CNS stem cell = (hCNS-SC).=20 The mAb 5F3 recognizes the CD133 antigen.=C2=B6 = Antibodies to=20 CD133 define a five-transmembrane protein and have been used to enrich = for human=20 hematopoietic stem cells (19). = Here we show=20 that mAbs 5F3 and the novel mAb, 5E12, detect a distinct subset of human = fetal=20 brain (FBr) cells. FACS using these mAbs results in a subset of human=20 CD133+ FBr cells that are capable of neurosphere initiation,=20 self-renewal, and multilineage differentiation at the single-cell level. = Another=20 mAb, 8G1, which recognizes human CD24, can further enrich neurosphere = initiating=20 cell activity within the CD133+ CD24=E2=88=92/lo = fraction. Because=20 the CD133+ cells self-renew, significantly expand in = neurosphere=20 cultures, and differentiate in vitro to neurons and glial = cells, they=20 are candidate human CNS-SC. Moreover, transplantation of = CD133+=20 sorted/expanded neurosphere cells into the lateral ventricles of newborn = nonobese diabetic=E2=80=93severe combined immunodeficient (NOD-SCID) = mouse brains=20 resulted in specific engraftment in numerous sites of the brain, which = is=20 similar to results shown by Fricker et al. (10). = These cells=20 also demonstrated continued self-renewal, migration, and neural = differentiation=20 for at least 7 months.

Pr= evious=20 SectionNext=20 Section

Materials and Methods

Preparation of Human Neural Tissues.

Human fetal spinal cord (FSC) and brain = tissues (FBr)=20 were obtained from Advanced Bioscience Resources, in accordance with all = state=20 and federal guidelines. Minced FBr tissues were dissociated = enzymatically in=20 0.1% collagenase (Roche Molecular Biochemicals) and 0.1% hyaluronidase = (Sigma)=20 at 37=C2=B0C for 1 h. Dissociated cells were further treated with 0.05% = trypsin-0.53=20 mM EDTA (GIBCO) for 10=E2=80=9315 min to obtain a single cell suspension = for cell=20 sorting (=E2=89=881=E2=80=9310 =C3=97 108 cells per tissue). =

Source of mAbs and Staining FBr Cells for Cell Sorting.

mAb 5F3, also known as mAb CD133, identifies = the same=20 cell surface antigen recognized by two other mAbs, CD133/1 and CD133/2 = (Miltenyi=20 Biotec, Auburn, CA). The dissociated FBr cells were incubated with mAbs = against=20 CD45-FITC (Caltag, South San Francisco, CA), CD133/1 = CD133/2-PE, and=20 CD34-APC (Becton Dickinson Immunocytometry Systems) for 20=E2=80=9360 = min on ice. Washed=20 cells were resuspended in Hanks' balanced salt solution containing 0.5 = =CE=BCg/ml=20 propidium iodide, sorted with a FACS Vantage SE (Becton Dickinson=20 Immunocytometry Systems), and cultured for analysis.

Generation of Novel mAbs.

mAbs to human neural cells were produced by = using a=20 previously described decoy immunization strategy (19). = BALB/c mice=20 were immunized in the left footpad with decoy human peripheral blood = mononuclear=20 cells and in the contralateral footpad with enzyme-dissociated human = FBr. The=20 right popliteal lymph node cells were fused with mouse SP2/0 cells. This = fusion=20 resulted in 1,900 hybridoma-positive wells. Approximately 180 hybridomas = were=20 selected, expanded, and further tested for specific reactivity to = surface=20 epitopes on human FBr cells.

Assay for Neurosphere-Initiating Cells (NS-IC).

Sorted FBr cells were cultured either as = described (7) or in = neurosphere initiation medium consisting of Ex Vivo 15 (BioWhittaker) = medium=20 with N2 supplement (GIBCO), fibroblast growth factor-2 (20 ng/ml), = epidermal=20 growth factor (20 ng/ml), lymphocyte inhibitory factor (10 ng/ml), = neural=20 survival factor-1 (Clonetics, San Diego), and 60 =CE=BCg/ml = N-acetylcysteine=20 (Sigma). Cultures were fed weekly and passaged at =E2=89=882=E2=80=933 = weeks. In some cases,=20 sorted FBr cells were resorted, and different numbers of cells were = deposited by=20 the automated cell deposition unit into 96-well plates to evaluate = directly the=20 frequency of precursors that initiate neurosphere cultures. Plates were = fed=20 weekly, and wells were scored for neurosphere growth at 6=E2=80=938 = weeks. Linear=20 regression analysis of the proportion of negative wells at each cell=20 concentration was used to determine the frequency of NS-IC (20). =

Differentiation Conditions.

Neurosphere cells were harvested, = dissociated, and=20 cultured with differentiation medium [Ex=20 Vivo-15/N2/N-acetylcysteine/brain-derived neurotrophic factor = (10=20 ng/ml)/glial-derived neurotrophic factor (10 ng/ml)/laminin (1 = =CE=BCg/ml)] on=20 polyornithine-coated chamber slides. After 1=E2=80=932 weeks, chamber = slides were fixed=20 with 4% paraformaldehyde in PBS and stained to detect differentiation = into=20 neurons and astrocytes, with mAbs against =CE=B2-tubulin III (1:200; = Sigma)=20 conjugated with Alexa 488 (Molecular Probes) and glial fibrillary acidic = protein-Cy3=20 (GFAP-Cy3) (1:1,000; Sigma).

Transplantation of CD133+ Sorted/Expanded Neurosphere = Cells into=20 Neonatal NOD-SCID Mice.

Expanded CD133+ sorted = neurosphere cells at=20 passages 6=E2=80=9310 were harvested and gently dissociated with = collagenase. Neonatal=20 mice (<24 h after birth) were anesthetized by placing them in ice for = 5=E2=80=9310=20 min. Once cryoanesthetized, the pups were placed on a stereotaxic device = and=20 injected with 2 =CE=BCl of cells (ranging from 105 to = 106 cells=20 per injection) into the lateral ventricle. The injected mice were kept = for 7=20 months before they were tested for the engraftment of human cells. =

Immunocytochemical Analysis of Transplanted Mouse Brain.

Seven months after transplantation, the = injected mice=20 were perfused with 4% paraformaldehyde. The mouse brains were sectioned=20 sagitally at 5-=CE=BCm thickness for fluorescence microscopic analysis = or at 40-=CE=BCm=20 thickness for confocal microscopy (Bio-Rad MRC1024 UV confocal scanning=20 microscope). To detect human cells in the transplanted mouse brains, = sections=20 were stained with mouse mAbs against human nuclei (1:100; Chemicon) or = human=20 neural cell adhesion molecule (N-CAM) (1:20; D.W.B., unpublished work), = followed=20 by goat anti-mouse IgG conjugated with Alexa 488 (1:1000, Molecular Probes) or conjugated with Cy-3 (1:500; = Jackson=20 ImmunoResearch). To stain lineage-specific cell populations, sections = were=20 stained with guinea pig anti-GFAP (1:250; Advanced ImmunoChemical, Long = Beach,=20 CA), rabbit anti-Ki-67 (1:1,000, Novocastra), and rabbit antityrosine=20 hydroxylase (1:500; Pel-Freeze Biologicals), followed by donkey = anti-guinea pig=20 conjugated with Cy-5 (1:250; Jackson ImmunoResearch) or anti-rabbit = conjugated=20 with Cy-3 (1:250; Jackson ImmunoResearch).

Previou= s=20 SectionNext=20 Section

Results

Search Strategy for CNS-SC Markers.

To isolate the human CNS-SC prospectively, = we used the=20 approach that had proved successful for the isolation of several stem = and=20 progenitor cells, namely, mAb-based cell sorting of candidate = regenerative cells=20 followed by placement of the sorted cells into clonogenic assays in=20 vitro and in vivo (2, 15, 16, 21, 22). = Because=20 rodent and human neurospheres contain cells of both neuronal and glial = lineages=20 (6, 7), and = because=20 they contain cell populations capable of engraftment, migration, and=20 multilineage differentiation upon injection into rodent brains (10), we = tested=20 the possibility that each neurosphere derives from a clonogenic stem = cell. Thus,=20 a cell population that at the single-cell level initiates a neurosphere = could be=20 a candidate for hCNS-SC. Initially, we sought a mAb that cleanly = separated FBr=20 into two fractions: one fraction that established a neurosphere culture = and one=20 that did not. Such candidate hCNS-SC markers should be expressed on only = a minor=20 subset of FBr cells. Enzyme-dissociated FBr and long-term cultured = neurosphere=20 cells were stained with over 50 known mAbs. CD34 and CD45 were not = expressed on=20 the neurosphere cells, whereas they were expressed in FBr on endothelial = cells=20 and blood cells, respectively. Antibody screening revealed that another=20 hematopoietic stem cell marker, CD133 (19, 23, 24), = was=20 expressed on =E2=88=BC90=E2=80=9395% of long-term cultured neurosphere = cells but only on 1=E2=80=936% of=20 FBr cells derived from 16- to 20-gestational-week tissue (Fig. 1 = b).=20 Histochemical analysis of FSC tissue sections with the anti-CD133 mAb = 5F3 showed=20 staining of the ventricular zone surrounding the central canal, the roof = plate,=20 and the floor plate (Fig. 1 = a).

=20
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Figure 1=20

Isolation of human NS-IC from FBr based = on cell=20 surface markers. (a) Immunochemical staining of human FSC. = Cross=20 sections of human FSC (12 gestational weeks) were fixed with acetone and = stained=20 with mAb 5F3 (=C3=9710, Left; =C3=9740, Middle) and = 5E12 (=C3=9740,=20 Right), using peroxidase-labeled secondary antibody with=20 3-amino-9-ethylcarbazole substrate (red). (b) Flow cytometric = analysis=20 of fresh FBr. FBr cells were stained with mAbs as described. A rare = subset of=20 FBr cells coexpressed both CD133+ and 5E12+=20 (Left). The CD133+ FBr cells were further separated = based on=20 CD24 expression (Right). Sort regions were set based on isotype = controls of FBr. (c) Flow cytometric separation of FBr cells = based on=20 CD133 expression. Two cell populations,=20 CD133=E2=88=92CD34=E2=88=92CD45=E2=88=92= (CD133=E2=88=92) and=20 CD133+CD34=E2=88=92CD45=E2=88=92 = (CD133+), were=20 sorted. FACS preenrichment of CD133+ cells followed by a = second round=20 of cell sorting resulted in a distinct CD133+ population = (second=20 panel). In all subsequent experiments, both CD133=E2=88=92 = and=20 CD133+ subsets were sorted twice, resulting in = CD133=E2=88=92 and=20 CD133+ FBr cell fractions with high purity (right two = panels).

We also identified two mAbs, 5E12 and 8G1, by generating = hybridomas=20 to human FBr cells. The mAb 5E12 costained CD133+ FBr cells = (Fig. 1 = b) and=20 showed a staining pattern similar to that of 5F3 on FSC sections (Fig. = 1 = a). 5E12=20 recognized a protein of =E2=88=BC125 kDa, which, based on its tissue = distribution=20 pattern and molecular mass, appears to be different from CD133 (D.W.B.,=20 unpublished observation) and therefore appears to be novel. In contrast, = the mAb=20 8G1 stains the majority of total FBr cells (80=E2=80=9390%) at high = levels; among the=20 CD133+ cells, 8G1 expression is heterogeneous (Fig. 1 = b) and=20 defines two subsets of CD133+ cells: 8G1hi and=20 8G1=E2=88=92/lo (Fig. 1 = b).=20 Immunoprecipitation and blocking studies determined that 8G1 recognizes = the=20 human CD24 heat-stable antigen (D.W.B., unpublished observation).

mAbs Against CD133, 5E12, and CD24 Enriched for hCNS-SC.

To test whether hCNS-SC could be isolated = from FBr based=20 on the expression of CD133, 5E12, and/or CD24 alone or in combination, = two cell=20 populations, = CD133=E2=88=92CD34=E2=88=92CD45=E2=88=92= =20 (CD133=E2=88=92) and = CD133+CD34=E2=88=92CD45=E2=88=92=20 (CD133+), were sorted and cultured to evaluate their NS-IC = activity.=20 CD34 and CD45 were used to exclude contaminating vascular and blood = cells,=20 respectively, and bulk cultures of sorted CD45+ or = CD34+=20 cells from FBr failed to initiate long-term neurosphere cultures. Fig. = 1 = c shows=20 the analysis of FBr cells stained for CD133 and a reanalysis of the = sorted=20 CD133+ and CD133=E2=88=92 subsets. Single-cell = suspensions of=20 CD133+ cells proliferated and formed small neurospheres as = early as=20 7=E2=80=9310 days after culture initiation. In contrast, the sorted = CD133=E2=88=92=20 cells remained as a single-cell suspension, failed to initiate = neurospheres, and=20 eventually died. The CD133+-derived neurosphere cultures = expanded;=20 the number of CD133+ cells increased by at least 1,000-fold = by the=20 fifth passage (n =3D 5) and were capable of reinitiating = neurospheres=20 (see below).

To determine the frequency of NS-IC, unfractionated FBr = cells were=20 plated by limiting dilution and were cultured for 6=E2=80=938 weeks. = Fig. 2 = a shows a=20 representative limiting dilution analysis of control unseparated,=20 CD133+, and CD133=E2=88=92 subsets of FBr. Control = processed FBr=20 cells contained NS-IC activity at a frequency of 1/880 cells. In the=20 CD133=E2=88=92 subset, the NS-IC frequency dropped to 1/4,860 = cells. In contrast, NS-IC activity was highly enriched = in the=20 CD133+ subset, with a frequency of 1/32 cells (Fig. 2 = a). The=20 NS-IC frequencies from eight different FBr samples (16=E2=80=9320 = gestational weeks)=20 were evaluated and are summarized in Fig. 2 = b and=20 c and Table 1. On = average, about=20 1 in 730 unfractionated FBr cells contains NS-IC activity. The = CD133=E2=88=92=20 subset, representing >96% of FBr, contained <1 in 4,300 cells with = NS-IC.=20 In contrast, NS-IC activity was on average enriched 24-fold in the FBr=20 CD133+ subset with =E2=89=881/31 (range 1/6 to 1/72) cells = capable of=20 initiating a neurosphere. Because CD133+ cells represent = =E2=88=BC3.6% of FBr=20 (i.e., 1/28 cells), the enrichment of NS-IC activity virtually = paralleled the=20 frequency of CD133+ cells in FBr. Thus, CD133+ = cells=20 contained all detectable NS-IC activity in 16-to 20-gestational-week FBr = cell=20 suspensions. Similarly, 5E12+ FBr cells are enriched and=20 5E12=E2=88=92 cells are depleted in NS-IC activity (Table 1). As = shown in Table=20 1, the = frequency of=20 NS-IC is higher in the CD133+CD24=E2=88=92/lo = fraction compared=20 with the CD133+CD24hi subset. Thus, = CD133+=20 cells contained virtually all NS-IC activity but were still = heterogeneous; more=20 potent NS-IC activity was enriched in the subset of cells that expressed = negative to low levels of CD24. These data suggest that the=20 CD133+CD24hi cells could represent a subset of = multipotent=20 progenitor cells, such as the short-term or long-term stem cell subsets = defined=20 for the hematopoietic system (16, 25) =

=20
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Figure 2=20

Limiting dilution analyses of=20 CD133+-sorted hCNS-SC cells. (a) A representative = limiting=20 dilution analysis for NS-IC activity in FBr and sorted subsets. FBr = (=E2=97=8F),=20 CD133=E2=88=92 CD34=E2=88=92 = CD45=E2=88=92 (=E2=96=B4), and CD133+=20 CD34=E2=88=92 CD45=E2=88=92 (=E2=A7=AB, red line) = sorted cell populations were=20 plated in a series of limiting cell doses. FBr and = CD133=E2=88=92 cells=20 (100=E2=80=9310,000 cells per well) and CD133+ cells = (10=E2=80=93300 cells per well)=20 were plated into 96-well plates with a FACS=E2=80=93automated cell = deposition unit.=20 Cultures were carried for 6=E2=80=938 weeks, and wells that did not = contain neurospheres=20 were scored as negative. A linear regression analysis was used to = determine the=20 frequency of NS-IC (20). = (b)=20 Neurospheres generated from a single-cell event. CD133+=20 CD34=E2=88=92 CD45=E2=88=92 sorted cell = populations from eight different=20 FBr samples were plated in a series of limiting cell doses into 96-well = plates=20 with a automated cell deposition unit. Results from eight different = samples were=20 combined to generate a plot of the number of cells plated vs. log = percentage=20 negative wells with SEM (error bars). Linear correlation of log = percentage=20 negative wells and number of CD133+ cells plated indicate a=20 neurosphere initiated from a single-hit event. (c) NS-IC = frequencies=20 from unsorted, sorted, and cultured FBr cells. NS-IC frequency in FBr = control=20 (n =3D 8), CD133=E2=88=92 (n =3D 8), = CD133+=20 (n =3D 8) sorted, and CD133+ sorted/expanded = (passage 1,=20 n =3D 3) cell populations was determined. NS-IC frequencies of = individual=20 samples were calculated by linear regression analysis. The average NS-IC = frequency in a given population is shown alongside the bars.

View this table:=20
Table 1=20

NS-IC activity in FBr and sorted = populations

Clonal Neurosphere Expansion, Self-Renewal, and Differentiation from = Single=20 CD133+-Sorted Cells.

The limiting dilution analysis (Fig. 2=20 b) shows a linear correlation between the number of cells = plated and=20 the log percentage negative wells (20, 26), = indicating=20 that a neurosphere is generated from single CD133+-sorted = cells. To=20 confirm the clonality of neurospheres, single FBr-derived=20 CD133+-sorted cells were directly plated into wells of a = 96-well=20 plate with a FACS=E2=80=93automated cell deposition unit. After 8 weeks, = about 5=E2=80=9310% of=20 wells contained one neurosphere, confirming that neurospheres are = derived from=20 single CD133+-sorted cells (Fig. 3 = a). When=20 plated in medium with the growth factors, brain-derived neurotrophic = factor and=20 glial-derived neurotrophic factor, each of these clonally derived = neurospheres=20 differentiated into neurons and astrocytes (Fig. 3=20 b).

=20
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Figure 3=20

Clonal expansion from single = CD133+=20 cells. (a) Clonal expansion of neural stem/progenitor cells.=20 Neurospheres can be derived from a single sorted CD133+ cell = directly=20 isolated from fetal brain or from a single CD133+ cell from=20 sorted/expanded cultured neurospheres. (b) Differentiation = capacity of=20 clonally derived neurosphere cells. Progeny of single cell-derived = neurospheres=20 can be differentiated into neurons (=CE=B2-tubulin III, green) and = astrocytes (GFAP,=20 red). Nuclei were counterstained with Hoechst 33342 (blue).

We tested NS-IC self-renewal activity by examining the = replating=20 activity of single viable cells from the = CD133+-sorted/expanded=20 neurosphere cells. Nine of 96 wells had a single neurosphere in one = experiment.=20 Cells derived from neurosphere cultures initiated from = CD133+-sorted=20 cells could consistently reinitiate the formation of secondary = neurospheres, as=20 shown in Fig. 2 = c. Cells=20 from the primary neurosphere that became CD133=E2=88=92 = failed to reinitiate=20 secondary neurosphere cultures (data not shown). Together, the data = suggest that=20 the mAb against CD133 marks cells that can self-renew.

Potent Engraftment Ability of CD133+ Sorted/Expanded = Human=20 Neurosphere Cells.

To evaluate in vivo engraftment, migration = and the=20 differentiation capacity of sorted/expanded CD133+ hCNS-SC,=20 105 or 106 cells from CD133+-initiated=20 neurosphere cultures at passage 7=E2=80=9310 were injected into the = lateral ventricles=20 of neonatal NOD-SCID mice. Detailed analysis focused particularly on the = two=20 sites of the brain previously shown to be sites of active neurogenesis: = the=20 subventricular zone (SVZ) of the lateral ventricles and the dentate = gyrus of the=20 hippocampus (Figs. 4 and 5). Human = cells,=20 detected with an anti-human nuclear antibody, were found throughout the = brain of=20 mice transplanted with sorted/expanded human neural cells and were = abundant in=20 the SVZ 7 months after injection. Confocal microscopy indicated that = most of the=20 human cells were GFAP=E2=88=92, but occasional human = GFAP+ cells=20 also were detected (data not shown). Because stem/progenitor cells in = the SVZ=20 have been shown to proliferate continuously, it was important to test = whether=20 progeny of the transplanted human CD133+ sorted/expanded = neurosphere=20 cells were still proliferating in situ and coexpressed the=20 proliferation marker Ki-67, expressed on cells in late=20 G1/S/G2/M phases (27). As = shown in=20 Fig. 4 = a=E2=80=93c, a=20 cluster of human cells in the SVZ, nested in GFAP+ cells, = coexpressed=20 Ki-67; like their mouse counterparts, these cells continued to = proliferate 7=20 months after transplantation in the SVZ.

=20
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Figure 4=20

Detection of human cells in the = olfactory system.=20 CD133+-sorted/expanded human neurosphere cells were = transplanted into=20 the lateral ventricles of neonatal NOD-SCID mice. Engraftment of human = cells was=20 analyzed 7 months after transplantation. (a=E2=80=93c) SVZ: = detection of=20 proliferating human neural cells in the SVZ. Confocal images are of a = sagittal=20 section of the transplanted mouse brain stained with anti-human nuclear = antigen=20 (a, green) and Ki-67 (b, red) and GFAP (c, = blue).=20 Most of the human nuclear antigen positive cells in the SVZ also = coexpressed the=20 proliferation marker Ki-67 (c, merged). (d) Schematic = diagram=20 of adult mouse brain. (e) RMS: the array of human nuclear=20 antigen+ cells (red) in the RMS was colocalized with = =CE=B2-tubulin III=20 expression (green). Some of these cells were clearly double positive = (arrow).=20 (f and g) Confocal microscopic analysis of the = olfactory bulb.=20 Many human cells were distributed in the olfactory bulb, some of which = expressed=20 human N-CAM (green) on their neuronal processes (f). Very rare = human=20 dopaminergic neurons were detected [i.e., human nuclei (green), tyrosine = hydroxylase (red)] in the periglomerular layer (g, arrow).

=20
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Figure 5=20

Detection of proliferating and = differentiating=20 human cells in the dentate gyrus of hippocampus. (a=E2=80=93c) = Confocal=20 microscopic analysis of proliferating human neural cells in the = subgranular zone=20 of the dentate gyrus. Transplanted mouse brain was stained with = anti-human=20 nuclei (a, green), Ki-67 (b, red), and GFAP = (c,=20 blue). Some human nuclear antigen+ cells were costained with = Ki-67=20 (c, merged, arrow). (d) Detection of human neurons in = the=20 dentate gyrus. The sagittal section of the transplanted mouse brain was = stained=20 with anti-human nuclear antigen (red) and anti-=CE=B2-tubulin III = (green). One of two=20 human nuclear antigen-positive cells was also positive for = =CE=B2-tubulin III=20 (arrow).

In the olfactory system of rodents, the progeny of = stem/progenitor=20 cells that have proliferated in the SVZ enter the rostral migratory = stream (RMS)=20 and migrate to the olfactory bulb (8, 9). = These=20 endogenous rodent progenitors, the =E2=80=9Cchain of = neuroblasts=E2=80=9D in the RMS, express=20 both =CE=B2-tubulin III and N-CAM (10, 28). = Large=20 numbers of human cells were detected in mice transplanted 7 months = previously=20 with CD133+-sorted neurosphere cells, beginning in the SVZ = and=20 extending throughout the RMS (Fig. 4 = a=E2=80=93c, e).=20 Multiple cells were identified that were double positive for both = =CE=B2-tubulin III=20 and the human nuclear antigen (Fig. 4 = e, arrow=20 and Inset). In addition, many of these cells in the RMS = expressed the=20 human specific marker N-CAM (data not shown).

After migrating through the RMS, the progeny of = stem/progenitor cells=20 enter the olfactory bulb and extend toward the olfactory glomerulus to = the=20 periglomerular layers (8, 9). The=20 transplanted progeny of human cells distributed into the glomerular as = well as=20 the periglomerular layers. Some of these cells expressed human N-CAM, = indicating=20 that they were committed to neuronal lineages (Fig. 4=20 f). In a few instances, human dopaminergic neurons, defined by = tyrosine=20 hydroxylase expression, were observed (Fig. 4 = g).

Another critical site where neurogenesis takes place in = adult life is=20 the dentate gyrus of the hippocampus (29). We = found=20 numerous human cells in the dentate gyrus of the hippocampus. Some of = the human=20 cells in the subgranular cell zone coexpressed Ki-67, indicating that = they were=20 still able to proliferate 7 months after transplantation (Fig. 5 = a=E2=80=93c).=20 Distinct =CE=B2-tubulin III+ human cells also were detected = that had long=20 axonal processes extending toward the hilus of the hippocampus, as = expected for=20 developing granular neurons (Fig. 5 = d). These=20 results indicate that not only do these sorted/expanded hCNS-SC engraft, = migrate, continue to proliferate, and differentiate, but, furthermore, = their=20 behavior and cell fate are regulated by host cues in a site-specific = manner. In=20 no case did the injected cells form tumors even 1 year after transplant = into=20 immunodeficient SCID mice.

Previou= s=20 SectionNext=20 Section

Discussion

Previously, studies with rodent neural cultures showed that = retroviral-marked clones could self-renew and differentiate (30). = These cells=20 and rodent CNS-SC monolayer cells or v-myc immortalized rodent = neural=20 cells engraft, migrate, and differentiate appropriately upon = intracerebral=20 transplantation (9, 31). = Here we have=20 shown that human neurosphere cultures are initiated from a rare subset = of=20 clonogenic CD133+, 5E12+, = CD24=E2=88=92/lo,=20 CD34=E2=88=92 CD45=E2=88=92 cells that we define = as the hCNS-SC. In=20 vitro, these rare cells can initiate neurosphere cultures and = differentiate=20 into neurons and glia. It has been reported that the incidence of=20 differentiation to human oligodendrocytes is very rare (32). = The rarity=20 of detectable oligodendrocytes could be because stem cells do not = readily=20 differentiate toward that lineage, or it could be that the markers of = human=20 oligodendrocytes are inadequate both in vivo and in = vitro. The=20 latter seems possible, as in vitro cells with oligodendrocyte=20 morphology have been detected in cultures of our human stem cells. More=20 recently, our transplant data suggest that the progeny of human = neurosphere=20 cells were found in the fimbria of the hippocampus of engrafted mice, = with=20 oligodendrocyte-like morphology. Upon transplantation to the lateral = ventricles=20 of NOD-SCID newborn mice, this rare subset of sorted, then expanded,=20 CD133+ neurosphere cells engraft for at least 7 months and = give rise=20 to progeny that migrate and differentiate. Some grafted human cells = continue to=20 proliferate (i.e., express Ki-67+) in the SVZ and dentate = gyrus,=20 where counterpart mouse neural cells undergo the same events.

Rodent neurogeneic cells and ES cells grown in = vitro can=20 give rise to neurons and glia (33=E2=80= =9335). In = addition,=20 Frisen et al. provided evidence that mouse Notch+ = ependymal=20 cells also can initiate neurospheres (ref. 36; see = ref. 37 for = comments).=20 These neurospheres contain multipotent precursors for several nonneural = outcomes=20 when transplanted into blastocysts (38). In = apparent=20 contradiction, SVZ GFAP+ cells rather than ependymal cells = were=20 concluded to be rodent CNS-SC for the RMS-to-olfactory pathway (39, 40). We = could not=20 access fresh human adult brain tissue to determine the anatomical = location of=20 candidate adult CNS-SC.

Are there other CNS-SC candidates in the human FBr? The = data in Fig.=20 2 = b and=20 Table 1 = demonstrate that=20 there are no other detectable NS-IC cells outside of the = CD133+=20 subset in the human FBr, but this does not mean that there are no other = cells in=20 the human fetal or adult brain that have CNS-SC activity, or no other=20 progenitors that might engraft. It is not yet known which = cells=E2=80=94stem cells or=20 oligolineage-committed progenitors or even mature cells=E2=80=94engraft = best in the=20 brain. The transplant efficiencies of hCNS-SC versus highly enriched = progenitors=20 or mature cells in biological and preclinical models have yet to be = addressed.=20

It is striking that the progeny of transplanted = hCNS-SC-derived=20 neurosphere cells could be found adjacent to ventricular spaces, and in = the=20 hippocampus, cerebral cortex, and the corpus callosum, as well as in the = cerebellum, and in the SVZ, along the RMS, and within the olfactory bulb = in and=20 near the olfactory glomeruli. Previous experiments by Snyder et = al. (31) = using=20 v-myc immortalized mouse and human neural cell lines = demonstrated this=20 wide (=E2=80=9Cglobal=E2=80=9D) distribution of cells, and here we show = that this distribution=20 is a property not only of myc immortalized cells of murine = origin but=20 also of nongenetically modified hCNS-SC that have been considerably = expanded in=20 culture.

When analyzed 7=E2=80=9312 months after transplant, these = hCNS-SC appeared to=20 still respond to host microenvironmental cues and were not neoplastic.=20 Therefore, these cells are likely to reveal developmental and functional = pathways inherent in human neuronal systems not modified by the = functions of=20 oncogenes or already neoplastically transformed cell lines (11, 41, 42). = The=20 prospective isolation of human CNS-SC provides several opportunities to=20 delineate directly the lineages that derive from cells at a particular = stage of=20 differentiation, to obtain a gene expression profile of hCNS-SC, and = their=20 immediate downstream progeny in vitro or in vivo, and = to test=20 whether specific gene modification of the cells alters their = engraftment,=20 migration, differentiation, and/or functional integration. Finally, the=20 prospective isolation of hCNS-SC should allow the transplantation of = these cells=20 in mouse analogues of human disease (11, 31) as=20 preclinical tests for their transplantability, differentiation ability, = and lack=20 of tumorigenicity.

Previo= us=20 SectionNe= xt=20 Section

Acknowledgments

We are grateful to David Anderson, Ben Barres, and Eric = Lagasse for=20 stimulating discussions and advice; M. L. Gage for critical review of = the=20 manuscript; and Linda Kitabayashi for performing confocal microscopic = analysis.=20 We also thank the staff of StemCells, Inc., for their continuous = encouragement=20 and support.

Previou= s=20 SectionNe= xt=20 Section

Footnotes

  • =E2=86=B5= =E2=80=A0 To whom reprint requests should be = addressed. E-mail:=20 nuchida{at}stemcell.net.=20

  • =E2=86=B5= =C2=B6 7th Workshop and Conference on Human = Leucocyte=20 Differentiation Antigens, Harrogate, U.K., June 20=E2=80=9324, 2000. =

  • Abbreviations:=20
    CNS,
    central nervous system;
    hCNS-SC,
    human CNS stem cell;
    FACS,
    fluorescence-activated cell sorting;
    FBr,
    fetal brain;
    NOD-SCID,
    nonobese diabetic=E2=80=93severe combined immunodeficient;
    FSC,
    fetal spinal cord;
    NS-IC,
    neurosphere-initiating cells;
    GFAP,
    glial fibrillary acidic protein;
    N-CAM,
    neural cell adhesion molecule;
    SVZ,
    subventricular zone;
    RMS,
    rostral migratory stream
  • Copyright = =C2=A9 2000, The=20 National Academy of Sciences
Pr= evious=20 Section

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PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } CODE { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } DEL { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } DFN { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } EM { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } FONT { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } IMG { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } INS { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } KBD { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } Q { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } S { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } SAMP { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } SMALL { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } STRIKE { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } STRONG { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } SUB { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } SUP { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } TT { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } VAR { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } DL { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } DT { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } DD { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } OL { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } UL { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } LI { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } FIELDSET { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } FORM { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } LABEL { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } LEGEND { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } TABLE { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } CAPTION { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } TBODY { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } TFOOT { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } THEAD { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } TR { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } TH { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } TD { BORDER-BOTTOM: 0px; TEXT-ALIGN: inherit; BORDER-LEFT: 0px; = PADDING-BOTTOM: 0px; LINE-HEIGHT: inherit; FONT-STYLE: inherit; MARGIN: = 0px; OUTLINE-STYLE: none; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = FONT-FAMILY: inherit; FONT-SIZE: inherit; VERTICAL-ALIGN: baseline; = BORDER-TOP: 0px; FONT-WEIGHT: inherit; BORDER-RIGHT: 0px; PADDING-TOP: = 0px } A IMG { BORDER-BOTTOM: 0px; BORDER-LEFT: 0px; BORDER-TOP: 0px; BORDER-RIGHT: = 0px } :link IMG { BORDER-BOTTOM: 0px; BORDER-LEFT: 0px; BORDER-TOP: 0px; BORDER-RIGHT: = 0px } :visited IMG { BORDER-BOTTOM: 0px; BORDER-LEFT: 0px; BORDER-TOP: 0px; BORDER-RIGHT: = 0px } :focus { OUTLINE-STYLE: none; OUTLINE-COLOR: invert; OUTLINE-WIDTH: medium } BODY { LINE-HEIGHT: 1.25; BACKGROUND: white; COLOR: black } TABLE { BORDER-SPACING: 0px; BORDER-COLLAPSE: collapse } CAPTION { TEXT-ALIGN: left; FONT-WEIGHT: normal } TH { TEXT-ALIGN: left; FONT-WEIGHT: normal } TD { TEXT-ALIGN: left; FONT-WEIGHT: normal } BLOCKQUOTE:before { CONTENT: "" } BLOCKQUOTE:after { CONTENT: "" } Q:before { CONTENT: "" } Q:after { CONTENT: "" } BLOCKQUOTE { QUOTES: "" "" } Q { QUOTES: "" "" } BODY { LINE-HEIGHT: 1; FONT-FAMILY: "Lucida Grande", "Lucida Sans Unicode", = Tahoma, Verdana, Arial, Helvetica, sans-serif; BACKGROUND: white fixed = no-repeat; COLOR: black; FONT-SIZE: 100% } P { MARGIN: 1em 5% } PRE { MARGIN: 1em 5% } LABEL { MARGIN: 1em 5% } UL { MARGIN: 1em 5% } OL { MARGIN: 1em 5% } DL { MARGIN: 1em 5% } FIELDSET { MARGIN: 1em 5% } ADDRESS { MARGIN: 1em 5% } H1 { MARGIN: 0px 0px 0.4em } H2 { MARGIN: 0px 0px 0.4em } H3 { MARGIN: 0px 0px 0.4em } H4 { MARGIN: 0px 0px 0.4em } H5 { MARGIN: 0px 0px 0.4em } H6 { MARGIN: 0px 0px 0.4em } H1 { TEXT-ALIGN: left; FONT-WEIGHT: bold } H2 { TEXT-ALIGN: left; FONT-WEIGHT: bold } H3 { TEXT-ALIGN: left; FONT-WEIGHT: bold } H4 { TEXT-ALIGN: left; FONT-WEIGHT: bold } H1 { FONT-SIZE: 1.8em } H2 { FONT-SIZE: 1.5em } H3 { FONT-SIZE: 1.2em } BLOCKQUOTE { MARGIN: 1em 7% } LI { MARGIN-LEFT: 5% } DD { MARGIN-LEFT: 5% } FIELDSET { PADDING-BOTTOM: 0.5em; PADDING-LEFT: 0.5em; PADDING-RIGHT: 0.5em; = PADDING-TOP: 0.5em } EM { FONT-STYLE: italic } STRONG { FONT-WEIGHT: 700 } CODE { FONT-FAMILY: Courier New, Courier, monospace } VAR { FONT-FAMILY: Courier New, Courier, monospace } KBD { FONT-FAMILY: Courier New, Courier, monospace } SAMP { FONT-FAMILY: Courier New, Courier, monospace } PRE { FONT-FAMILY: Courier New, Courier, monospace } SUP { LINE-HEIGHT: 0; FONT-SIZE: 0.85em; VERTICAL-ALIGN: super } SUB { LINE-HEIGHT: 0; FONT-SIZE: 0.85em; VERTICAL-ALIGN: sub } DIV.hw-gen-page { PADDING-BOTTOM: 2em; PADDING-LEFT: 2em; PADDING-RIGHT: 4em; FONT-SIZE: = 0.8em; PADDING-TOP: 0.5em } .hw-gen-page #skip-link { POSITION: absolute; LEFT: -9999px } #content-block { BORDER-BOTTOM: 0px; BORDER-LEFT: 0px; PADDING-BOTTOM: 0px; LINE-HEIGHT: = 1.3; MARGIN: 0px; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; OVERFLOW: = hidden; BORDER-TOP: 0px; BORDER-RIGHT: 0px; PADDING-TOP: 0px } #content-block P { BORDER-BOTTOM: 0px; BORDER-LEFT: 0px; PADDING-BOTTOM: 0px; LINE-HEIGHT: = 1.5; MARGIN: 15px 0px; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = BORDER-TOP: 0px; BORDER-RIGHT: 0px; PADDING-TOP: 0px } #content-block { PADDING-BOTTOM: 0px; PADDING-LEFT: 5px; WIDTH: 518px; 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      • ' + $hdr + '
        • ';=0A= }=0A= }=0A= );=0A= }=0A= hdrList +=3D '
      ';=0A= if (hasItems) {=0A= var header =3D $this.find("div.most-header h3");=0A= if (header.length) {=0A= header.after(hdrList);=0A= }=0A= }=0A= mostHdrList =3D $this.find("div.most-header ul");=0A= }=0A= // 3) if none of the items are selected, make the first one be=0A= if (((mostHdrList !=3D null) && mostHdrList.length) && = (!hasSelected)) {=0A= modClass(mostHdrList.find("li:first"), "most-cur-sel", "");=0A= modClass($this.children("ul").children("li:first"), "most-cur-sel", = "");=0A= }=0A= // 4) bind a handler to each heading li a to toggle = class=3D"most-cur-sel" for =0A= // that item and its corresponding item in the list below=0A= if (((mostHdrList !=3D null) && mostHdrList.length)) {=0A= mostHdrList.find("li a").click(=0A= function(e) {=0A= var curA =3D $(this);=0A= toggleMostSelection(e, $this, curA.text());=0A= }=0A= );=0A= }=0A= }=0A= );=0A= if (hadMostBoxes) {=0A= fixColHeights(-1);=0A= }=0A= }=0A= =0A= function toggleMostSelection(e, mostItemDiv, groupName) {=0A= if (mostItemDiv.length) {=0A= var hdrItems =3D mostItemDiv.find("div.most-header ul li");=0A= if (hdrItems.length) {=0A= hdrItems.each(=0A= function() {=0A= var hdrName =3D $(this).text();=0A= var isChoice =3D (hdrName =3D=3D groupName);=0A= if (isChoice) {=0A= modClass($(this), 'most-cur-sel', '');=0A= }=0A= else {=0A= modClass($(this), '', 'most-cur-sel');=0A= }=0A= }=0A= );=0A= }=0A= var listItems =3D mostItemDiv.children("ul").find("li");=0A= if (listItems.length) {=0A= listItems.each(=0A= function() {=0A= var itemName =3D $(this).children("h4").text();=0A= var isChoice =3D (itemName =3D=3D groupName);=0A= if (isChoice) {=0A= modClass($(this), 'most-cur-sel', '');=0A= }=0A= else {=0A= modClass($(this), '', 'most-cur-sel');=0A= }=0A= }=0A= );=0A= }=0A= }=0A= fixColHeights(-1);=0A= e.preventDefault();=0A= }=0A= =0A= function toggleCollapse(collapsibleEl, hideTagExpr, modTextTagExpr, = expandStr, contractStr) {=0A= if (typeof(expandStr) =3D=3D "undefined") {=0A= expandStr =3D getSiteOption('expandString','+');=0A= }=0A= if (typeof(contractStr) =3D=3D "undefined") {=0A= contractStr =3D getSiteOption('contractString','-');=0A= }=0A= =0A= var $this =3D collapsibleEl;=0A= if ($this.hasClass('collapsed')) {=0A= modClass($this, 'collapsible', 'collapsed');=0A= }=0A= else if ($this.hasClass('collapsible')) {=0A= modClass($this, 'collapsed', 'collapsible');=0A= }=0A= var $toToggle =3D $this.find(hideTagExpr);=0A= if ($toToggle.length) {=0A= $toToggle.toggle();=0A= }=0A= var $rewrite =3D $this.find(modTextTagExpr);=0A= if ($rewrite.length) {=0A= $rewrite.each(=0A= function(i, e) {=0A= var txt =3D $(e).text();=0A= if (txt =3D=3D expandStr) {=0A= $(e).empty().append(contractStr);=0A= }=0A= else if (txt =3D=3D contractStr) {=0A= $(e).empty().append(expandStr);=0A= }=0A= }=0A= );=0A= }=0A= }=0A= =0A= function setupDockBlock(col, dockId, dockClass, contentRules) {=0A= if ((col =3D=3D 2) || (col =3D=3D 3)) {=0A= var navOrigHeight =3D ((col =3D=3D 2) ? gColOrigHeights.col2 : = gColOrigHeights.col3);=0A= var gColNum =3D col;=0A= var gNav =3D ((col =3D=3D 2) ? gCol2 : gCol3);=0A= var gNavHeight =3D ((gNav.length) ? navOrigHeight : 0);=0A= var gNavTopOffset =3D (gNavHeight > 0 ? gNav.offset().top : 0);=0A= var gScrollPos =3D 0;=0A= var gNavDocked =3D false;=0A= /* this won't work in IE 6, so skip it */=0A= /* article navigation docking block */=0A= $(window).scroll(function () {=0A= if (gNavHeight > 0 && !isIE6) {=0A= var offsets =3D getPageOffset();=0A= var origHeight =3D ((gColNum =3D=3D 2) ? gColOrigHeights.col2 : = gColOrigHeights.col3);=0A= var navBottom =3D (gNavTopOffset + origHeight) - offsets.y;=0A= if (gScrollPos !=3D offsets.y) {=0A= gScrollPos =3D offsets.y;=0A= if ((navBottom <=3D 0) && !gNavDocked) {=0A= /* dock the nav */=0A= gNavDocked =3D true;=0A= addDockBlock(col, dockId, dockClass, contentRules);=0A= }=0A= else if (gNavDocked && (navBottom > 0)) {=0A= /* undock the nav */=0A= gNavDocked =3D false;=0A= removeDockBlock(col, dockId);=0A= }=0A= }=0A= }=0A= });=0A= }=0A= }=0A= function addDockBlock(col, dockId, dockClass, contentRules) {=0A= // get direct children li elements only=0A= var colId =3D ("col-" + col);=0A= var newDiv =3D ' 1) {=0A= for (var i =3D 0; i < (contentRules.length - 1); i =3D i + 2) {=0A= var addTo =3D processDockingRule(contentRules[i]);=0A= var add =3D processDockingRule(contentRules[i + 1]);=0A= var addTarget, addVal;=0A= if (addTo.ruleType =3D=3D 'empty') {=0A= addTarget =3D newDivJQuery;=0A= }=0A= else if (addTo.ruleType =3D=3D 'jQuery') {=0A= addTarget =3D $(addTo.rule);=0A= }=0A= if (add.ruleType =3D 'jQuery') {=0A= var addJQEl =3D $(add.rule);=0A= if (addJQEl.length) {=0A= addVal =3D addJQEl.clone();=0A= }=0A= else {=0A= addVal =3D '';=0A= }=0A= }=0A= else if (add.ruleType =3D 'html') {=0A= addVal =3D add.rule;=0A= }=0A= if (addTarget.length && !(addVal =3D=3D '')) {=0A= addTarget.append(addVal);=0A= }=0A= else {=0A= }=0A= }=0A= }=0A= var newDockJQrule =3D "#" + colId + " #" + dockId;=0A= var newDock =3D $(newDockJQrule);=0A= if (newDock.length) {=0A= prepCollapsibles(newDockJQrule + " div.collapsible, " + newDockJQrule = + " div.collapsed", false);=0A= newDock.fadeIn(250);=0A= }=0A= }=0A= =0A= function processDockingRule(rule) {=0A= var retObj =3D new Object();=0A= retObj.empty =3D (((rule !=3D undefined) && (!(rule =3D=3D ''))) ? = false : true);=0A= if (rule =3D=3D '') {=0A= retObj.ruleType =3D 'empty';=0A= }=0A= else if (/^\$\(.+\)$/.test(rule)) {=0A= retObj.ruleType =3D 'jQuery';=0A= retObj.rule =3D rule.replace(/^\$\((.+)\)$/,"$1");=0A= }=0A= else {=0A= retObj.ruleType =3D 'html';=0A= retObj.rule =3D rule;=0A= }=0A= return retObj;=0A= }=0A= =0A= function removeDockBlock(col, dockId) {=0A= var dockedNav =3D $("div#" + dockId);=0A= if(dockedNav.length) {=0A= dockedNav.fadeOut(250, function() { dockedNav.remove(); });=0A= }=0A= }=0A= =0A= =0A= function updateFormInput(labelMatchString, inputMatchString, = defaultColorString, textColorString) {=0A= if ((defaultColorString =3D=3D null) || (defaultColorString =3D=3D '')) = {=0A= defaultColorString =3D "#A0A0A0";=0A= }=0A= if ((textColorString =3D=3D null) || (textColorString =3D=3D '')) {=0A= textColorString =3D "black";=0A= }=0A= var label =3D $(labelMatchString);=0A= var input =3D $(inputMatchString);=0A= if (input.length) {=0A= if ((label.length) && (input.val() =3D=3D '')) {=0A= input.val(label.text()).css("color",defaultColorString);=0A= }=0A= input.focus(=0A= function(e) {=0A= if ((label.length) && ($(this).val() =3D=3D label.text())) {=0A= $(this).val('').css("color",textColorString);=0A= }=0A= }=0A= );=0A= input.blur(=0A= function(e) {=0A= if ((label.length) && ($(this).val() =3D=3D '')) {=0A= $(this).val(label.text()).css("color",defaultColorString);=0A= }=0A= }=0A= );=0A= }=0A= var parentForm =3D label.parents("form").eq(0);=0A= if (parentForm.length) {=0A= parentForm.submit(=0A= function() {=0A= if ((label.length) && (input.length) && (input.val() =3D=3D = label.text())) {=0A= input.val('');=0A= }=0A= return true;=0A= }=0A= );=0A= }=0A= }=0A= =0A= function checkUnloadedImgs(lookupRule) {=0A= if (lookupRule =3D=3D null) {=0A= lookupRule =3D "img";=0A= }=0A= var imageNum =3D -1;=0A= var images =3D $(lookupRule);=0A= if (images.length) {=0A= imageNum =3D 0;=0A= images.each(=0A= function() {=0A= //if ($(this).height() =3D=3D 0) {=0A= if (this.offsetHeight =3D=3D 0) {=0A= imageNum++;=0A= }=0A= }=0A= );=0A= }=0A= return imageNum;=0A= }=0A= =0A= function fixHeightForImages(iter, numImagesToLoad, lookupRule) {=0A= if (lookupRule =3D=3D null) {=0A= lookupRule =3D "img";=0A= }=0A= if (numImagesToLoad > 0) {=0A= var lastNumImagesToLoad =3D numImagesToLoad;=0A= var newNumImagesToLoad =3D checkUnloadedImgs(lookupRule);=0A= if (newNumImagesToLoad < lastNumImagesToLoad) {=0A= fixColHeights(1);=0A= if ((newNumImagesToLoad > 0) && (iter < 10)) {=0A= setTimeout("fixHeightForImages(" + (iter+1) + "," + = newNumImagesToLoad + ",'" + lookupRule + "')", 1000);=0A= }=0A= }=0A= }=0A= }=0A= function getColInfo(colJQEl) {=0A= var colInfo =3D { valid: false, height: 0, bottom: 0, lastElBottom: 0, = extra: 0, fix: false };=0A= if (colJQEl.length) {=0A= var c =3D colJQEl.children();=0A= if (c.length) {=0A= colInfo.height =3D getObjHeight(colJQEl);=0A= colInfo.bottom =3D colJQEl.offset().top + colInfo.height;=0A= var last =3D c.eq(c.length - 1);=0A= colInfo.lastElBottom =3D last.offset().top + getObjHeight(last);=0A= colInfo.extra =3D colInfo.bottom - colInfo.lastElBottom;=0A= colInfo.valid =3D true;=0A= }=0A= }=0A= return colInfo;=0A= }=0A= function checkColHeights() {=0A= var colToFix =3D 0;=0A= if (gIsMultiCol && ((gColOrigHeights.main + gColOrigHeights.col2 + = gColOrigHeights.col3) > 0)) {=0A= var colMainInfo =3D getColInfo(gColMain);=0A= var col2Info =3D getColInfo(gCol2);=0A= var col3Info =3D getColInfo(gCol3);=0A= if (colMainInfo.valid && (gColLows.main =3D=3D 0)) { gColLows.main =3D = colMainInfo.lastElBottom; }=0A= if (col2Info.valid && (gColLows.col2 =3D=3D 0)) { gColLows.col2 =3D = col2Info.lastElBottom; }=0A= if (col3Info.valid && (gColLows.col3 =3D=3D 0)) { gColLows.col3 =3D = col3Info.lastElBottom; }=0A= // check if low point in any column has changed=0A= if ((colMainInfo.valid) && (colMainInfo.lastElBottom !=3D = gColLows.main)) {=0A= colMainInfo.fix =3D true;=0A= }=0A= if ((col2Info.valid) && (col2Info.lastElBottom !=3D gColLows.col2)) {=0A= col2Info.fix =3D true;=0A= }=0A= if ((col3Info.valid) && (col3Info.lastElBottom !=3D gColLows.col3)) {=0A= col3Info.fix =3D true;=0A= }=0A= // check for columns not having same bottom points=0A= if ((colMainInfo.valid) &&=0A= (((col2Info.valid) && (colMainInfo.height < col2Info.height)) ||=0A= ((col3Info.valid) && (colMainInfo.height < col3Info.height)))) {=0A= colMainInfo.fix =3D true;=0A= }=0A= if ((col2Info.valid) &&=0A= (((colMainInfo.valid) && (col2Info.height < colMainInfo.height)) ||=0A= ((col3Info.valid) && (col2Info.height < col3Info.height)))) {=0A= col2Info.fix =3D true;=0A= }=0A= if ((col3Info.valid) &&=0A= (((colMainInfo.valid) && (col3Info.height < colMainInfo.height)) ||=0A= ((col2Info.valid) && (col3Info.height < col2Info.height)))) {=0A= col3Info.fix =3D true;=0A= }=0A= =0A= if (colToFix > -1) {=0A= if (colMainInfo.valid && colMainInfo.fix) {=0A= if ((col2Info.valid && col2Info.fix) || (col3Info.valid && = col3Info.fix)) {=0A= colToFix =3D -1;=0A= }=0A= else { colToFix =3D 1; }=0A= }=0A= else if (col2Info.valid && col2Info.fix) {=0A= if (col3Info.valid && col3Info.fix) {=0A= colToFix =3D -1;=0A= }=0A= else { colToFix =3D 2; }=0A= }=0A= else if (col3Info.valid && col3Info.fix) {=0A= colToFix =3D 3;=0A= }=0A= }=0A= =0A= if (colToFix !=3D 0) {=0A= fixColHeights(colToFix);=0A= }=0A= if (colMainInfo.valid) { gColLows.main =3D colMainInfo.lastElBottom; }=0A= if (col2Info.valid) { gColLows.col2 =3D col2Info.lastElBottom; }=0A= if (col3Info.valid) { gColLows.col3 =3D col3Info.lastElBottom; }=0A= }=0A= setTimeout("checkColHeights()", 5000);=0A= }=0A= function fixColHeights(colChanged) {=0A= if (gIsMultiCol && getSiteOption('setColSizes', true)) {=0A= var setAll =3D false;=0A= if (colChanged =3D=3D 1) {=0A= if (gColMain.length) {=0A= setObjHeight(gColMain,'auto');=0A= gColOrigHeights.main =3D getObjHeight(gColMain);=0A= }=0A= }=0A= else if (colChanged =3D=3D 2) {=0A= if (gCol2.length) {=0A= setObjHeight(gCol2,'auto');=0A= gColOrigHeights.col2 =3D getObjHeight(gCol2);=0A= }=0A= }=0A= else if (colChanged =3D=3D 3) {=0A= if (gCol3.length) {=0A= setObjHeight(gCol3,'auto');=0A= gColOrigHeights.col3 =3D getObjHeight(gCol3);=0A= }=0A= }=0A= else {=0A= if (colChanged =3D=3D -1) {=0A= // force resize all 3=0A= if (gColMain.length) { setObjHeight(gColMain,'auto'); }=0A= if (gCol2.length) { setObjHeight(gCol2,'auto'); }=0A= if (gCol3.length) { setObjHeight(gCol3,'auto'); }=0A= }=0A= gColOrigHeights.main =3D ((gColMain.length) ? = (getObjHeight(gColMain)) : 0);=0A= gColOrigHeights.col2 =3D ((gCol2.length) ? (getObjHeight(gCol2)) : 0);=0A= gColOrigHeights.col3 =3D ((gCol3.length) ? (getObjHeight(gCol3)) : 0);=0A= setAll =3D true;=0A= }=0A= if ((gColOrigHeights.main + gColOrigHeights.col2 + = gColOrigHeights.col3) > 0) {=0A= var colInfo;=0A= var maxH =3D Math.max(Math.max(gColOrigHeights.main, = gColOrigHeights.col2), gColOrigHeights.col3);=0A= if ((gColMain.length) && ((gColOrigHeights.main < maxH) || = (getObjHeight(gColMain) > maxH) || setAll)) {=0A= setObjHeight(gColMain, maxH);=0A= colInfo =3D getColInfo(gColMain);=0A= if (colInfo.valid) { gColLows.main =3D colInfo.lastElBottom; }=0A= }=0A= if ((gCol2.length) && ((gColOrigHeights.col2 < maxH) || = (getObjHeight(gCol2) > maxH) || setAll)) {=0A= setObjHeight(gCol2, maxH);=0A= colInfo =3D getColInfo(gCol2);=0A= if (colInfo.valid) { gColLows.col2 =3D colInfo.lastElBottom; }=0A= }=0A= if ((gCol3.length) && ((gColOrigHeights.col3 < maxH) || = (getObjHeight(gCol3) > maxH) || setAll)) {=0A= setObjHeight(gCol3, maxH);=0A= colInfo =3D getColInfo(gCol3);=0A= if (colInfo.valid) { gColLows.col3 =3D colInfo.lastElBottom; }=0A= }=0A= }=0A= }=0A= }=0A= =0A= function getObjHeight(obj) {=0A= if (obj instanceof jQuery) {=0A= return obj.get(0).offsetHeight;=0A= }=0A= else if (obj instanceof Array) {=0A= return obj[0].offsetHeight;=0A= }=0A= else {=0A= return obj.offsetHeight;=0A= }=0A= }=0A= function getObjWidth(obj) {=0A= if (obj instanceof jQuery) {=0A= return obj.get(0).offsetWidth;=0A= }=0A= else if (obj instanceof Array) {=0A= return obj[0].offsetWidth;=0A= }=0A= else {=0A= return obj.offsetWidth;=0A= }=0A= }=0A= function setObjHeight(obj, h) {=0A= if (typeof(h) =3D=3D 'number') {=0A= h =3D "" + h + "px";=0A= }=0A= if (obj instanceof jQuery) {=0A= obj.get(0).style.height =3D h;=0A= }=0A= else if (obj instanceof Array) {=0A= obj[0].style.height =3D h;=0A= }=0A= else {=0A= obj.style.height =3D h;=0A= }=0A= }=0A= function setObjWidth(obj, w) {=0A= if (typeof(w) =3D=3D 'number') {=0A= w =3D "" + w + "px";=0A= }=0A= if (obj instanceof jQuery) {=0A= obj.get(0).style.width =3D w;=0A= }=0A= else if (obj instanceof Array) {=0A= obj[0].style.width =3D w;=0A= }=0A= else {=0A= obj.style.width =3D w;=0A= }=0A= }=0A= =0A= /* num pixels top of page has been scrolled offscreen */=0A= function getPageOffset() {=0A= var offset =3D new Object();=0A= if( typeof( window.pageXOffset ) =3D=3D 'number' ) {=0A= offset.x =3D window.pageXOffset;=0A= offset.y =3D window.pageYOffset;=0A= } else if( document.body && ( document.body.scrollTop ) ) {=0A= offset.x =3D document.body.scrollLeft;=0A= offset.y =3D document.body.scrollTop;=0A= } else if( document.documentElement && ( = document.documentElement.scrollTop ) ) {=0A= offset.x =3D document.documentElement.scrollLeft;=0A= offset.y =3D document.documentElement.scrollTop;=0A= }=0A= return offset;=0A= }=0A= function getViewportDim() {=0A= var dim =3D new Object();=0A= // non-IE=0A= if (typeof window.innerWidth !=3D 'undefined') {=0A= dim.x =3D window.innerWidth;=0A= dim.y =3D window.innerHeight;=0A= }=0A= // IE6 in standards compliant mode=0A= else if (typeof document.documentElement !=3D 'undefined'=0A= && typeof document.documentElement.clientWidth !=3D=0A= 'undefined' && document.documentElement.clientWidth !=3D 0) {=0A= dim.x =3D document.documentElement.clientWidth;=0A= dim.y =3D document.documentElement.clientHeight;=0A= }=0A= // older versions of IE=0A= else {=0A= dim.x =3D document.getElementsByTagName('body')[0].clientWidth;=0A= dim.y =3D document.getElementsByTagName('body')[0].clientHeight;=0A= }=0A= return dim;=0A= }=0A= =0A= function newWindowTargets() {=0A= var newWins =3D $("a.in-nw");=0A= if (newWins.length) {=0A= newWins.each(=0A= function() {=0A= var $this =3D $(this);=0A= $this.attr("target", "_blank");=0A= modClass($this, 'in-nw-vis', 'in-nw');=0A= }=0A= );=0A= }=0A= }=0A= =0A= function getSiteOption(optionName, defaultVal) {=0A= if (typeof(defaultVal) =3D=3D "undefined") {=0A= defaultVal =3D gSiteOptionUnknown;=0A= }=0A= if (gSiteOptions =3D=3D undefined) {=0A= return defaultVal;=0A= }=0A= else if (gSiteOptions[optionName] =3D=3D undefined) {=0A= return defaultVal;=0A= }=0A= else {=0A= return gSiteOptions[optionName];=0A= }=0A= }=0A= =0A= function addPartHeaders(req) {=0A= req.setRequestHeader('Accept', 'application/xhtml+xml');=0A= req.setRequestHeader('Range', 'part=3Dvariant-contents');=0A= }=0A= function addCommonHeaders(req) {=0A= if (typeof(callbackToken) !=3D 'undefined') {=0A= req.setRequestHeader('X-Token', callbackToken);=0A= }=0A= }=0A= function allowsCookies() {=0A= if (hasTestCookie()) {=0A= return true;=0A= }=0A= else {=0A= setTestCookie();=0A= return hasTestCookie();=0A= }=0A= }=0A= function setTestCookie() {=0A= setCookie('cks','allowed',7,'/',null,null);=0A= }=0A= function hasTestCookie() {=0A= var ckVal =3D getCookie('cks');=0A= if ((ckVal !=3D null) && (ckVal =3D=3D 'allowed')) {=0A= return true;=0A= }=0A= else {=0A= return false;=0A= }=0A= }=0A= =0A= var UIPrefsCk=3D{=0A= name: "UIPrefs",=0A= expDays: 3652,=0A= path: '/'=0A= };=0A= =0A= function prefDefined(name) {=0A= var val =3D getPrefValue(name);=0A= if (val =3D=3D null) {=0A= return false;=0A= }=0A= else {=0A= return true;=0A= }=0A= }=0A= function getPrefValue(name) {=0A= var prefArray =3D getPrefArray();=0A= if (prefArray) {=0A= name =3D convertPrefString(name);=0A= var match =3D name + ":";=0A= for (var i =3D 0; i < prefArray.length; i++) {=0A= if (prefArray[i].indexOf(match) =3D=3D 0) {=0A= return prefArray[i].substring(match.length);=0A= }=0A= else if (prefArray[i] =3D=3D name) {=0A= // name only, no value, return boolean true=0A= return true;=0A= }=0A= }=0A= }=0A= return null;=0A= }=0A= function convertPrefString(str) {=0A= if (str) {=0A= return str.replace(/[,:=3D]/g,"");=0A= }=0A= return str;=0A= }=0A= function removePref(name) {=0A= var prefArray =3D getPrefArray();=0A= name =3D convertPrefString(name);=0A= if (prefDefined(name)) {=0A= var newArray =3D new Array(prefArray.length - 1);=0A= var match =3D name + ":";=0A= var newArrIndex =3D 0;=0A= for (var i =3D 0; i 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------=_NextPart_000_003B_01CA605B.7C435B80 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.pnas.org/shared/js/pages/hw-content.js /* jQuery javascript functions for content page */=0A= var gIsFrameset =3D false;=0A= $(document).ready(function() {=0A= =0A= handleAuthAffil();=0A= =0A= fixWackyReflinksMarkup();=0A= =0A= var defaultDockedNavRules =3D [=0A= '', '

      This Article

        ',=0A= '$(#col-2 #docked-nav-views)', '$(#article-cb-main .cb-section = ol:has(li a[rel^=3D"view-"]) > li)',=0A= '$(#col-2 #docked-nav #docked-slug li)', '$(#slugline)',=0A= '', '$(#article-dyn-nav)'=0A= ];=0A= =0A= if (!(getSiteOption("suppressDockedNav") =3D=3D true)) {=0A= setupDockBlock(2, 'docked-nav', 'dockblock', defaultDockedNavRules);=0A= }=0A= =0A= var unloadedImgLookupRule =3D "div.article div.fig img";=0A= var numImagesToLoad =3D checkUnloadedImgs(unloadedImgLookupRule);=0A= =0A= /* ref rollover */=0A= if (!(getSiteOption("suppressRefPopups") =3D=3D true)) {=0A= setTimeout("addRefPops()", 25);=0A= }=0A= =0A= /* fig-expansion in page */=0A= if (!(getSiteOption("noInlineFigExpand") =3D=3D true)) {=0A= setTimeout("figExpandInline()", 10);=0A= }=0A= =0A= /* if configured, open ref links in new windows */=0A= if (getSiteOption("refLinksNewWindow") =3D=3D true) {=0A= setTimeout("refLinksNewWindowTarget()", 25);=0A= }=0A= =0A= /* 'new window' fig expansions */=0A= setTimeout("newWindowTargets()", 25);=0A= =0A= /* AJAX related article callbacks */=0A= setTimeout("getISIRelated()", 50);=0A= =0A= /* AJAX citing article callbacks */=0A= setTimeout("getHWCiting()", 50);=0A= setTimeout("getISICiting()", 50);=0A= =0A= /* AJAX Entrez Links callbacks */=0A= setTimeout("getEntrezLinks()", 50);=0A= =0A= /* Related content */=0A= setTimeout("getHWRelatedURLs()", 50);=0A= setTimeout("getPatientInformData()", 50);=0A= =0A= /* Fix col heights for images */=0A= setTimeout("fixHeightForImages(1" + "," + numImagesToLoad + ", '" + = unloadedImgLookupRule + "')",1000);=0A= =0A= /* Social Bookmarking enhancement */=0A= setTimeout("updateSBLinks()", 100);=0A= =0A= });=0A= =0A= function handleAuthAffil() {=0A= var authAffilMatch =3D getSiteOption('authAffilMatch','div.article = div.contributors ol.affiliation-list'); =0A= var authAffil =3D (authAffilMatch !=3D undefined) ? $(authAffilMatch) : = '';=0A= if (authAffil.length) {=0A= var expandStr =3D getSiteOption('authExpandString', null);=0A= if (expandStr =3D=3D null) {=0A= expandStr =3D getSiteOption('expandString', '+');=0A= }=0A= var contractStr =3D getSiteOption('authContractString', null);=0A= if (contractStr =3D=3D null) {=0A= contractStr =3D getSiteOption('contractString', '-');=0A= }=0A= var newP =3D '

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        No citing = articles
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n < length; n++) { var element =3D elements[n]; autoEllipseText(element, element.parentNode.parentNode); } } catch (ex) { } } function GetElementsByClassName(parentElement, className) { function _GetChildrenByClassName(elements, element, className) { for (var node =3D element.firstChild; node; node =3D node.nextSibling) { if (node.nodeType =3D=3D 1) { if (node.className =3D=3D className) elements[elements.length] =3D node; else _GetChildrenByClassName(elements, node, className); } } return elements; } return _GetChildrenByClassName(new Array(), parentElement, className); } function autoEllipseText(textElement, container) { if (textElement.offsetWidth > container.offsetWidth) { var i =3D 1; var text =3D textElement.innerHTML; var width =3D container.clientWidth - 10; textElement.innerHTML =3D ''; while (textElement.offsetWidth < width && i < text.length) { textElement.innerHTML =3D text.substr(0, i) + '...'; i++; } } } ------=_NextPart_000_003B_01CA605B.7C435B80 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.pnas.org/local/js/baynote-observer.js /* Last updated: 09/03/2009 */=0A= function bn_isNotEmpty(name) {=0A= return (name !=3D null) && (name !=3D "");=0A= }=0A= function bn_getOrderInfo() {=0A= if (typeof(bnOrderId) !=3D "undefined" && bn_isNotEmpty(bnOrderId))=0A= baynote_tag.attrs.purchaseId =3D bnOrderId;=0A= if (typeof(bnOrderTotal) !=3D "undefined" && = bn_isNotEmpty(bnOrderTotal))=0A= baynote_tag.attrs.totalPurchases =3D parseFloat(bnOrderTotal);=0A= if (typeof(bnOrderDetails) !=3D "undefined" && = bn_isNotEmpty(bnOrderDetails))=0A= baynote_tag.attrs.purchaseDetails =3D bnOrderDetails;=0A= }=0A= function bn_getMediaInfo() {=0A= if (typeof(bnMediaDuration) !=3D "undefined" && = bn_isNotEmpty(bnMediaDuration)) {=0A= baynote_tag.attrs.expectedDuration =3D bnMediaDuration;=0A= } =0A= }=0A= function baynote_setAttrs() {=0A= /*alert('setAttrs');*/=0A= var slugAhead =3D getElementsByClassName(document, = 'span','slug-ahead-of-print-date');=0A= if(slugAhead.length > 0){=0A= baynote_tag.docAttrs.slugAheadDate =3D slugAhead[0].innerHTML;=0A= }=0A= var slugDate =3D getElementsByClassName(document, = 'span','slug-pub-date');=0A= if(slugDate.length > 0){=0A= baynote_tag.docAttrs.slugPubDate =3D slugDate[0].innerHTML;=0A= }=0A= var onCover =3D getElementsByClassName(document, = 'div','from-the-cover-indicator');=0A= if(onCover.length > 0){=0A= baynote_tag.docAttrs.onCover =3D "true";=0A= }=0A= var metas =3D document.getElementsByTagName("meta");=0A= var DCPlace =3D "none";=0A= if (!metas) return;=0A= =0A= for (var i =3D 0; i < metas.length; i++) {=0A= if (!metas[i]) return;=0A= if (metas[i].name =3D=3D "DC.Date") {=0A= var tempMetaContent =3D metas[i].content;=0A= if(bn_isNotEmpty(tempMetaContent)) {=0A= baynote_tag.docAttrs.date =3D tempMetaContent;=0A= }=0A= } else if (metas[i].name =3D=3D "citation_volume") {=0A= var tempMetaContent =3D metas[i].content;=0A= if(bn_isNotEmpty(tempMetaContent)) {=0A= baynote_tag.docAttrs.citeVolume =3D tempMetaContent;=0A= }=0A= } else if (metas[i].name =3D=3D "citation_doi") {=0A= var tempMetaContent =3D metas[i].content;=0A= if(bn_isNotEmpty(tempMetaContent)) {=0A= baynote_tag.docAttrs.citationDoi =3D tempMetaContent;=0A= }=0A= } else if (metas[i].name =3D=3D "citation_issue") {=0A= var tempMetaContent =3D metas[i].content;=0A= if(bn_isNotEmpty(tempMetaContent)) {=0A= baynote_tag.docAttrs.citeIssue =3D tempMetaContent;=0A= }=0A= } else if (metas[i].name =3D=3D "citation_authors") {=0A= var tempMetaContent =3D metas[i].content;=0A= if(bn_isNotEmpty(tempMetaContent) ) {=0A= var commaIndex =3D tempMetaContent.indexOf(",");=0A= var semiIndex =3D tempMetaContent.indexOf(";");=0A= var str =3D tempMetaContent.substring(0,commaIndex);=0A= if(semiIndex >1){=0A= str =3D str + " et al."=0A= }=0A= baynote_tag.docAttrs.author =3D str;=0A= =0A= }=0A= }else if (metas[i].name =3D=3D "citation_abstract_html_url") {=0A= var tempMetaContent =3D metas[i].content;=0A= if(bn_isNotEmpty(tempMetaContent)) {=0A= baynote_tag.url =3D tempMetaContent;=0A= }=0A= }else if (metas[i].name =3D=3D "DC.Title") {=0A= var tempMetaContent =3D metas[i].content;=0A= if(bn_isNotEmpty(tempMetaContent)) {=0A= baynote_tag.docAttrs.docTitle =3D tempMetaContent;=0A= }=0A= }=0A= =0A= }=0A= }=0A= =0A= function getElementsByClassName(oElm, strTagName, strClassName) {=0A= var arrElements =3D (strTagName =3D=3D "*" && oElm.all) ? oElm.all : = oElm=0A= .getElementsByTagName(strTagName);=0A= var arrReturnElements =3D new Array();=0A= strClassName =3D strClassName.replace(/\-/g, "\\-");=0A= var oRegExp =3D new RegExp("(^|\\s)" + strClassName + "(\\s|$)");=0A= var oElement;=0A= for ( var i =3D 0; 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