From: Subject: =?Windows-1252?Q?The_nuclear_receptor_tailless_is_required_for_neurogenes?= =?Windows-1252?Q?is_in_the_adult_subventricular_zone_=97_Genes_&_Developm?= =?Windows-1252?Q?ent?= Date: Wed, 5 May 2010 19:53:45 +0200 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01CAEC8C.AC04E920" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5579 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01CAEC8C.AC04E920 Content-Type: text/html; charset="utf-8" Content-Transfer-Encoding: quoted-printable Content-Location: http://genesdev.cshlp.org/content/22/18/2473.full =EF=BB=BF The nuclear receptor = tailless is required for neurogenesis in the adult subventricular zone = =E2=80=94 Genes & Development
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The nuclear receptor tailless is required for=20 neurogenesis in the adult subventricular zone

  1. Hai-Kun=20 Liu1,=20
  2. Thorsten=20 Belz1,=20
  3. Dagmar=20 Bock1,=20
  4. Andrea=20 Takacs1,=20
  5. Hui=20 Wu2,=20
  6. Peter=20 Lichter2,=20
  7. Minqiang=20 Chai1, = and=20
  8. G=C3=BCnther=20 Sch=C3=BCtz1,3 =
  1. 1 Division of Molecular Biology of the Cell I, = German=20 Cancer Research Center (DKFZ), Heidelberg 69120, Germany;
  2. 2 Division of Molecular Genetics, German Cancer = Research=20 Center (DKFZ), Heidelberg 69120, Germany
Ne= xt=20 Section

Abstract

The tailless (Tlx) gene encodes an orphan = nuclear=20 receptor that is expressed by neural stem/progenitor cells in the adult = brain of=20 the subventricular zone (SVZ) and the dentate gyrus (DG). The function = of Tlx in=20 neural stem cells of the adult SVZ remains largely unknown. We show here = that in=20 the SVZ of the adult brain Tlx is exclusively expressed in = astrocyte-like B=20 cells. An inducible mutation of the Tlx gene in the adult brain = leads=20 to complete loss of SVZ neurogenesis. Furthermore, analysis indicates = that Tlx=20 is required for the transition from radial glial cells to astrocyte-like = neural=20 stem cells. These findings demonstrate the crucial role of Tlx in the = generation=20 and maintenance of NSCs in the adult SVZ in vivo.

Keywords

The subventricular zone (SVZ) of the lateral ventricle and = the=20 subgranular zone (SGZ) of the dentate gyrus (DG) are the largest = germinal zones=20 of sustained neurogenesis during adulthood in the mammalian central = nervous=20 system (Gage = 2000;=20 Alvarez-= Buylla=20 and Garcia-Verdugo 2002). Astrocyte-like type B cells in the adult = SVZ are=20 thought to be multipotent neural stem cells (NSCs) (Doetsch = et al.=20 1999). These cells give rise to transient amplifying type C cells, = which in=20 turn differentiate into type A cells (neuroblasts) that migrate to the = olfactory=20 bulb (OB) through the rostral migratory stream (RMS). The SVZ-OB system = (Alvarez-= Buylla=20 and Garcia-Verdugo 2002) has recently been reported to also exist in = the=20 adult human brain (Curtis = et al.=20 2007).

Tailless (Tlx) is an orphan nuclear receptor, which is = expressed in=20 the periventricular neurogenic zone during mouse embryonic development = (Monagha= n et al.=20 1995). In the absence of this transcription factor, mutant animals = survive,=20 but suffer specific anatomical deficits in the cortex and the limbic = system (Monagha= n et al.=20 1997). Late-developing structures such as the upper cortical layers = and the=20 DG, are reduced in size. These phenotypic changes indicate that Tlx has = an=20 important role for brain development in the young postnatal stage. The = loss of=20 adult neurogenesis in Tlx-null mice does not exclude the possibility = that this=20 phenotype, lack of neurogenesis, is caused by the absence of Tlx during=20 development (Shi et = al.=20 2004). Recently, a study demonstrated that inactivation of Tlx leads = to a=20 decrease but not complete loss of neural stem cell proliferation in the = DG (Zhang = et al.=20 2008). The role of Tlx in the adult neurogenic region of the SVZ, = however,=20 remains largely unknown.

Previous=20 SectionNe= xt=20 Section

Results and Discussion

Generation of Tlx=E2=80=93CreERT2 mice leads to inducible = Cre activity in=20 the SVZ and RMS

To investigate the expression of Tlx in the adult brain, we = took=20 advantage of the BAC technology to obtain faithful expression of the=20 tamoxifen-inducible Cre recombinase (CreERT2 fusion protein) = under=20 the control of the Tlx gene regulatory sequences. Using = homologous=20 recombination in Escherichai coli, part of exon1 of the = Tlx=20 gene was replaced by the CreERT2 cassette (Supplemental Fig. = S1A).=20 The modified BAC will express CreERT2 under the control of=20 Tlx gene regulatory sequences contained in 104 kb of 5=E2=80=B2 = flanking=20 sequences and 22 kb of 3=E2=80=B2 flanking sequences. We assayed for = inducible Cre=20 recombinase translocation by tamoxifen in the F1 progeny from six = founders. Cre=20 recombinase is consistently found in the cytoplasm before injection and=20 translocated into the nucleus after injection of tamoxifen (data not = shown). We=20 analyzed three lines with one, two, or three copies of the transgene = (data not=20 shown). They showed efficient Cre translocation to the nucleus after = tamoxifen=20 treatment and an identical Cre expression pattern that follows that of = the=20 endogenous Tlx gene (see below).

Tlx is expressed in the developing forebrain (Monagha= n et al.=20 1995; Roy et = al.=20 2004). Interestingly, it was found that the SVZ where adult = neurogenesis=20 occurs is strikingly decreased in Tlx-deficient animals (Roy et = al.=20 2004). Since an antibody for Tlx is not available for adult brain = tissue, we=20 used an antibody against the recombinase as a reporter to follow the=20 Tlx gene expression pattern in the adult brain via = immunohistochemistry=20 (IHC). We find activity of this reporter exclusively in sites where the=20 Tlx gene has been found to be expressed (Monagha= n et al.=20 1995; Shi et = al.=20 2004). Cre is highly expressed in the SVZ (Supplemental Fig. S1B,C), = as well=20 as in the RMS (Supplemental Fig. S1B,D). We describe these cells as=20 Tlx+.

Tlx is specifically expressed by B cells in the SVZ-OB system

NSCs and progenitors continuously produce neuroblasts, which = migrate=20 to the OB via the RMS. The cellular composition of the SVZ and RMS has = been=20 analyzed previously (Doetsch = et al.=20 1997). The SVZ contains type B, C, and A cells and the main cell = populations=20 in the RMS are type B and A cells, although few C cells may also be = found in the=20 RMS (Doetsch = et al.=20 2002). According to the Cre expression pattern, we found that only a = subpopulation of the SVZ cells is Tlx-Cre-positive cells. To identify = the=20 identity of these cells on the basis of Cre expression, three=20 Tlx-CreERT2 mice were used for the following experiments; at = least 10=20 sections were analyzed for each experiment per animal. We first analyzed = sections coimmunostained for nuclear-localized Cre recombinase and SVZ = cell=20 type-specific markers. GFAP+ cells in the adult SVZ have been = shown=20 to be type B cells (Doetsch = et al.=20 1999), which has led to the concept of glial identity of NSCs. Here = we show=20 that these GFAP+ cells are also Cre+, which = indicates that=20 Tlx is expressed by astrocyte-like B cells (Fig. = 1A). Most=20 EGFR+ (epidermal growth factor receptor) cells in the SVZ are = type C=20 cells (Doetsch = et al.=20 2002). Cre staining was neither detected in the EGFR+ = cells (Fig. = 1B), nor=20 did the staining colocalize with that of doublecortin (DCX) (Fig.=20 1C) or PSA-NCAM (data not shown), both of which mark A cells (Doetsch = et al.=20 1999). It has been reported that the ependymal cells that surround = the=20 lateral ventricle may also be NSCs (Johanss= on et al.=20 1999). To determine if ependymal cells do express the Cre = recombinase, we=20 analyzed the sections with the confocal microscope. The ependymal cells = that=20 surround the lateral ventricle very tightly can be stained by DAPI (Fig. = 1D), but=20 these cells did not stain for Cre. Cre+ cells are always = located=20 below the ependymal layer (Fig. = 1D),=20 establishing Cre expression in the SVZ.

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Figure 1.=20

Tlx-CreERT2 is expressed by B = cells in=20 the adult SVZ and RMS. Tlx-CreERT2 mice were treated with = tamoxifen.=20 (A) Confocal optical sections of coronal sections of the mouse = SVZ=20 coimmunostained for Cre recombinase (green) and GFAP (red). = Cre+=20 cells also express GFAP. Bar, 20 =CE=BCm. (B) Cre recombinase = (green) and=20 EGFR (red). Cre is not costained with EGFR indicating that=20 Tlx-CreERT2 is not expressed by C cells in the mouse SVZ. = Bar, 10 =CE=BCm.=20 (C) Cre (green) and DCX (red). No colabled cells were found in = A cells.=20 Bar, 10 =CE=BCm. (D) Cre (green) and DAPI (blue). The ependymal = cells (E,=20 arrow) do not express Cre and Cre+ cells are found to reside = only in=20 the subependymal region. Bar, 10 =CE=BCm. (E) Confocal optical = sections of=20 coronal sections of the mouse RMS coimmunostained for Cre (green) and = GFAP=20 (red). Cre+ cells are costained with GFAP as in the SVZ. Bar, = 10 =CE=BCm.=20 (F) Cre (green) and DCX (red). Note the absence of = double-positive=20 cells. Bar, 10 =CE=BCm.

We next analyzed sections coimmunostained for Cre and RMS = cell=20 type-specific markers. In the RMS, GFAP+ cells (type B cells) = are=20 Cre+ (Fig. = 1E). No=20 cells costained for DCX and Cre (Fig. = 1F) were=20 detected. Since as described above the main cell populations of the RMS = are type=20 B and A cells, this means that Cre is only expressed by type B cells in = the RMS.=20

To determine whether Tlx is also expressed in mature = astrocytes in=20 the cortex and OB, we also analyzed these regions. Cells colabeled for = Cre and=20 GFAP were not found, either in the cortex (data not shown), or in the OB = outside=20 the RMS region. Since Cre is only expressed in the RMS region in the OB=20 (Supplemental Fig. S1D), we strongly conclude that Tlx is not a general = marker=20 for astrocytes.

Tlx+ cells in the SVZ are multipotent

Stem cells have the ability to self-renew and they are = multipotent.=20 To demonstrate that Tlx+ cells are NSCs we marked these cells = in vivo=20 and determined whether they continuously give rise to different neural = cell=20 type. Mice carrying a floxed stop cassette before the knockin = =CE=B2-galactosidase=20 (=CE=B2-gal+) gene at the ROSA locus (Soriano= =20 1999), were crossed with Tlx-CreERT2 mice to allow = genetic fate=20 mapping. After tamoxifen treatment of these Tlx-CreERT2; R26R = mice,=20 all the Tlx+ cells and their progeny will express = =CE=B2-galctosidase in=20 the cytoplasm.

Tlx-CreERT2; R26R mice were injected daily with = tamoxifen=20 and 5 d after the last injection (in total, 10 d from the first = injection),=20 X-gal staining indicated efficient recombination in the SVZ and the OB = where Cre=20 is expressed (Fig. = 2A). To=20 follow the long-term fate of the marked cells, we injected tamoxifen and = analyzed the animals 1 mo later. X-gal staining in the OB showed that = the=20 =CE=B2-gal+ cells were radially distributed in the whole OB = (Fig. = 2A). To=20 identify the =CE=B2-gal+ cell types outside of the Cre = expression region,=20 we analyzed sections coimmunostained for =CE=B2-gal and other neural = cell markers. In=20 the OB, the vast majority of =CE=B2-gal+ cells are = NeuN+ (Fig. = 2B). A few=20 =CE=B2-gal+ cells were found in the corpus callosum (CC) = costained with=20 O4, an oligodendrocyte marker (Fig. = 2C). Some=20 =CE=B2-gal+ cells outside of the RMS in the OB are costained = with GFAP (Fig. = 2D). These=20 GFAP+ cells showed more processes compared with the = GFAP+=20 in the cells of the RMS or SVZ, which are bipolar or unipolar (Garcia = et al.=20 2004).

<= IMG=20 alt=3D"Figure 2."=20 src=3D"http://genesdev.cshlp.org/content/22/18/2473/F2.small.gif">
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Figure 2.=20

Adult Tlx=E2=80=93Cre+ cells = are multipotent and=20 self-renewing NSCs in vivo. (A) Two-month-old mice carrying a = floxed=20 ROSA26 locus and expressing Tlx-CreERT2 were injected with = tamoxifen=20 for 5 d, and X-gal staining of brain sections was done. Note that 5 d = after the=20 last tamoxifen injection, X-gal staining shows efficient recombination = in the=20 SVZ and RMS. One month after the last tamoxifen injection, X-gal = staining in OB=20 shows radial migration of cells with the recombined locus. Bar:=20 B,C, 500 =CE=BCm. (B) Two-month-old=20 Tlx-CreERT2;R26R mice were injected with tamoxifen for 5 d = and 2 mo=20 after the last tamoxifen injection, and confocal analysis of brain = coronal=20 sections were performed after coimmunostaining for = =CE=B2-gal+ (green) and=20 NeuN (red). In OB, most =CE=B2-gal+ cells are costained with = NeuN outside=20 of the RMS. Bar, 20 =CE=BCm. (C) In the corpus callosum (CC) = some=20 =CE=B2-gal+ cells (green) are colabeled with O4 (red), an = oligodentrocyte=20 marker. Bar, 10 =CE=BCm. (D) In OB, some =CE=B2-gal+ = cells (green)=20 outside of the RMS can be found costained with GFAP (red). Note that = these cells=20 have multipolar morphology. Bar, 10 =CE=BCm. Confocal optical sections = of 2-mo-old=20 tamoxifen-treated Tlx-CreERT2 mice, treated with BrdU 2 h = before=20 sacrifice, were coimmunostained for Cre (green) and BrdU (red). = (E) A=20 low percentage (18.5%) of BrdU+ cells is costained with Cre = (arrows)=20 in SVZ. Note that the population is simlar in size to the B-cell = population.=20 Bar, 20 =CE=BCm. (F) Few cells double-labeled for BrdU and Cre = can also be=20 found in RMS. Bar, 10 =CE=BCm. (G) Some BrdU LRCs express Cre = and GFAP.=20 Tlx-CreERT2 mice received BrdU continuously in the drinking = water for=20 1 wk followed by a 3-wk survival. Triple staining for Cre (green), GFAP = (red)=20 and BrdU (blue). Bar, 20 =CE=BCm.

Recently, a report showed mosaic organization of NSCs in = the SVZ of=20 the adult brain (Merkle = et al.=20 2007), suggesting that the NSC population in the adult SVZ is = heterogeneous.=20 To investigate if Tlx+ cells include the entire B cell = population or=20 only a subpopulation, the genetic cell fate mapping approach was carried = out=20 using the Tlx-CreERT2; Z/EG mice (Novak = et al.=20 2000), 2 mo after the last induction. We found that Tlx+ = cells=20 can give rise to different kinds of neuronal cell types using neuronal = markers=20 (TH, calretinin, and calbindin) costained with EGFP (Supplemental Fig. = S2A=E2=80=93C).=20 To further investigate whether all the SVZ NSCs express Tlx, we = administered=20 BrdU to Tlx-CreERT2;R26R mice in the drinking water for 1 wk = 2 mo=20 after tamoxifen treatment. Two weeks after the BrdU treatment the mice = were=20 analyzed. If Tlx+ cells will indicate all B cells within the=20 population, all the BrdU+ cells should be colabeled with = =CE=B2-gal.=20 Coimmunostaing for BrdU and =CE=B2-gal proved this hypothesis, since all = the=20 BrdU+ cells are also =CE=B2-gal+ (Supplemental = Fig. S2D). These=20 results indicate that cells in the heterogeneous NSC population all = express Tlx.=20

Tlx+ cells are self-renewing stem cells in vivo

Since Tlx is only expressed by B cells; cells costained = with Cre and=20 BrdU should be found after BrdU treatment if Tlx+ cells can = indeed=20 self-renew. Tamoxifen-treated mice were injected with bromodeoxyuridine = (BrdU),=20 and sections were coimmunostained for Cre and BrdU 2 h after the = injection. A=20 small percentage (18.5% when determined in five slides) of = BrdU+=20 cells are costained with Cre in SVZ (Fig. = 2E), which=20 corresponds to the B cell population as previously reported (Doetsch = et al.=20 1997). To determine whether the B cells in the RMS are also = self-renewing in=20 vivo, we analyzed the RMS regions by coimmunostaining for Cre and BrdU. = Some=20 Tlx-Cre+ B cells (3.6% when determined in 15 slides) do = incorporate=20 BrdU (Fig. = 2F), which=20 indicates that the B cells in the RMS are active NSCs in vivo, although = they are=20 less active compared with the SVZ B cells. This is consistent with = previous=20 reports demonstrating that B cells are also active in the adult RMS (Gritti = et al.=20 2002; Hack = et al.=20 2005; Merkle = et al.=20 2007).

Several studies reported that the slowly dividing, = label-retaining=20 cells (LRCs) correspond to the resident stem cell population (Bickenba= ch=20 1981; Morris = and=20 Potten 1994; Jackson= et al.=20 2006). BrdU was administered in the drinking water for 1 wk and then = removed=20 for 3 wk. This is sufficient time for the labeled C and A cells to = migrate to=20 the OB (Jackson= et al.=20 2006). We found that the BrdU+ cells in the SVZ were also = positive for GFAP and Cre (Fig. = 2G); the=20 percentage of label-retained cells among the Cre-expressing cells is = =E2=88=BC6.3% (when=20 determined in 15 slides), which confirm that they are long-term = self-renewing=20 stem cells.

Induction of Tlx ablation leads to complete loss of the self-renewal = ability=20 of NSCs in the adult SVZ

The specificity of Tlx expression suggests that Tlx might = be a=20 crucial regulator of neurogenesis in the adult SVZ. We therefore = generated mice=20 with a conditional Tlx allele (Belz et = al.=20 2007) that, after crossing to Tlx-CreERT2 mice, ablated = the=20 Tlx gene in the adult.

We first analyzed the targeting efficiency after tamoxifen=20 administration. PCR analysis for the DNA isolated from the = microdissected SVZ=20 cells indicates that the recombination is efficient after induction=20 (Supplemental Fig. S3A). The time measured after the last tamoxifen = injection is=20 expressed as weeks post-induction (WPI). To investigate the effect on = cell=20 proliferation in SVZ after induction, BrdU was injected to the mice at 4 = WPI,=20 and the proliferation frequency analyzed 2 h after the injection. = Strikingly,=20 cell proliferation in the SVZ is almost completely depleted in the Tlx = mutant as=20 indicated by the number of BrdU+ cells (Fig.=20 3A,B). Further analysis of DCX-stained brain sections at 4 WPI shows = a=20 strong decrease in the number of migrating type A cells in the SVZ and = in the OB=20 of the Tlx mutant (Fig. = 3C) as=20 expected from the BrdU incorporation assay. To confirm the effect of = loss of Tlx=20 in SVZ by another approach independent of BrdU incorporation, we = introduced the=20 R26R reporter line in the Tlxfl/fl; = Tlx-CreERT2=20 background. As described in the genetic cell fate mapping experiment, = the=20 Tlx+ cells and their progeny will carry the activated = =CE=B2-galactosidase=20 gene for the entire life after tamoxifen treatment (Fig.=20 2A). If the loss of Tlx in the adult SVZ leads to loss of = neurogenesis, we=20 should not see any more =CE=B2-galactosidase-expressing cells = continuously migrating=20 to the OB after tamoxifen induction of the Tlx=20 fl/fl;Tlx-CreERT2;R26R mice. X-gal = staining at 4=20 WPI confirmed our hypothesis (Fig. = 3D). This=20 proves that loss of Tlx indeed causes the depletion of neurogenesis in = the adult=20 mouse SVZ. Since Tlx is exclusively expressed by type B cells, these = results=20 indicate that Tlx is crucial to maintain the self-renewal ability of = adult SVZ=20 NSCs. To investigate whether Tlx has a cell-autonomous function to NSCs, = Tlxfl/fl; Tlx-CreERT2;R26R mice=20 (Tlx-CreERT2; R26R mice as control), 5 d after the last = tamoxifen=20 injection, the cells that lose Tlx expression will carry =CE=B2-gal = expression, and=20 BrdU was injected to the mice to label the SVZ proliferating cells that = will=20 include the C and A cells generated before the recombination which are=20 =CE=B2-gal-negative. No significant difference was found about the = number of=20 BrdU+;=CE=B2-gal=E2=88=92 cells between the = control and mutant mice 2 h=20 after BrdU injection (Supplemental Fig. S3B) exclude the cell-autonomous = function of Tlx to the adult NSCs.

<= IMG=20 alt=3D"Figure 3."=20 src=3D"http://genesdev.cshlp.org/content/22/18/2473/F3.small.gif">
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Figure 3.=20

Tlx is essential for maintaining the = self-renewal=20 of adult NSCs in the SVZ. (A) Control and Tlx mutant mice at 4 = WPI were=20 administrated with BrdU and sacrificed 2 h later; coronal brain sections = were=20 stained for BrdU. Note that only a few positive cells can be found in = the=20 mutant. Bar, 50 =CE=BCm. (B) Number of BrdU+ cells in the SVZ = per brain=20 section of control and mutant at 4 WPI. (C) Coronal brain = sections=20 containing the SVZ or OB were stained for DCX. Note that a dramatic = decrease of=20 DCX+ cells is found in the SVZ or the OB of the Tlx mutant. Bar, 10 = =CE=BCm.=20 (D) Two-month-old Tlx-CreERT2;R26R and Tlx-CreERT2;R26R and Tlx = fl/fl=20 mice were injected with tamoxifen for 5 d, and after 4 wk, X-gal = staining of=20 brain sections containing the OB was done. Note that blue cells outside = of the=20 RMS in the Tlx fl/fl;Tlx-CreERT2;R26R mice are rarely found. Bar, 200 = =CE=BCm.=20 (E) Coronal brain sections of control and Tlx mutant mice at 4 = WPI were=20 stained for GFAP. Note that GFAP-expressing cells are seen in the SVZ of = the=20 mutant. Bar, 20 =CE=BCm.

To investigate the fate of NSCs after loss of Tlx = expression, we=20 analyzed GFAP-stained brain sections at 4 WPI. There is no significant=20 difference between the control and Tlx mutant (Fig. = 3E),=20 indicating that GFAP+ cells are still found in the SVZ. We = also did=20 not observe increased cell apoptosis in the mutant, which suggests that = the=20 astrocyte-like B cells are not lost.

Loss of Tlx causes absence of NSCs in the adult brain: Tlx is = required by=20 radial glial cells to give rise to astrocyte-like B cells

Inactivation in the adult shows that Tlx is crucial for = maintaining=20 neurogenesis in the SVZ. We could confirm the recently published = findings that=20 loss of Tlx in the DG leads to a decrease but not loss of neurogenesis = (Zhang = et al.=20 2008). Tlx-null mice, however, lack neurogenesis in both regions. = These=20 findings suggest that the phenotype in the null mutant results from the = absence=20 of Tlx during development. Adult NSCs are derived from radial glial = cells at an=20 early postnatal stage (Anthony = et al.=20 2004; Merkle = et al.=20 2004; Ventura= and=20 Goldman 2007), which are the progenitors of the early developing = brain.

To investigate whether the function of Tlx in radial glial = cells is=20 necessary to give rise to astrocyte-like B cells, we first analyzed = brain=20 sections from adult Tlx=E2=88=92/=E2=88=92 mice stained for = GFAP and LeX/ssea1. No=20 GFAP-expressing cells or LeX/ssea1-positive cells (data not shown) are = found in=20 the SVZ=E2=80=94not even in the dorsal SVZ where most GFAP+ = are located (Fig. = 4A)=E2=80=94but=20 GFAP+ cells can still be found in other regions (Fig.=20 4A, arrows). To further investigate if the SVZ cells can still give = rise to=20 new neurons in Tlx=E2=88=92/=E2=88=92 mice, BrdU was = administrated to=20 Tlx=E2=88=92/=E2=88=92 mice in the drinking water for 1 wk. = No BrdU+ cells=20 were found in the brain sections of Tlx=E2=88=92/=E2=88=92 = mice (Fig.=20 4B), suggesting an absence of neurogenesis. Interestingly, = Tlx+/=E2=88=92=20 mice have 38.2% less BrdU+ cells than wild-type mice in the = BrdU=20 incorporation assay, which indicates a threshold requirement for Tlx in = adult=20 neurogenesis (Fig. = 4C).=20 Phosphatase and tensin homolog (PTEN), a tumor suppressor that = negatively=20 regulates neural stem/progenitor cell proliferation by modulating = G0=E2=80=93G1 cell=20 cycle entry (Groszer= et al.=20 2006), has been shown to be a target gene of Tlx (Zhang = et al.=20 2006). We therefore analyzed the level of PTEN mRNA in wild-type and = Tlx+/=E2=88=92 mice by real-time PCR of laser-captured = microdissected cells=20 from the SVZ. Loss of one Tlx allele leads to a proportionate increase = in the=20 level of PTEN mRNA in microdissected cells of the SVZ (Fig.=20 4D,E). These findings, together with chromatin immunoprecipitation=20 (ChIP)-based analyses (data not shown), confirm that PTEN is a direct = target=20 gene of Tlx and explains the decrease in adult neurogenesis in the=20 Tlx+/=E2=88=92 mice.

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Figure 4.=20

Tlx is required for the generation of = adult NSCs.=20 (A) Coronal brain sections of 2-mo-old Tlx+/+ and=20 Tlx=E2=88=92/=E2=88=92 mice were stained for GFAP (red). Note = that no=20 GFAP+ cells were found in SVZ of = Tlx=E2=88=92/=E2=88=92 mice, but normal=20 astrocytes can still be found outside the SVZ (arrows). Bar, 30 =CE=BCm. = (B)=20 BrdU. BrdU was administrated to mice for 2 h. Note that no = BrdU+=20 cells in SVZ of Tlx=E2=88=92/=E2=88=92 mice are detected. = Bar, 30 =CE=BCm. (C)=20 Number of BrdU+ cells in SVZ per brain section of 2-mo-old=20 Tlx+/+ and Tlx+/=E2=88=92 mice. Note that = Tlx+/=E2=88=92 mice=20 have 38.2 =C2=B1 6.7% less labeled cells than Tlx+/+ mice. = Mean =C2=B1 standard=20 deviation (SD). (D) Quantitative PCR validates decreased = expression of=20 Tlx mRNA in Tlx+/=E2=88=92 (n =3D 3) versus = Tlx+/+=20 (n =3D 3) laser-microdissected SVZ cells. Note that expression = of Tlx=20 mRNA in Tlx+/=E2=88=92 is 48 =C2=B1 11% of the wild type. = P < 0.01,=20 mean =C2=B1 standard deviation (SD). (E) Quantitative PCR = validates=20 increased expression of PTEN mRNA in Tlx+/=E2=88=92 = (n =3D 3) versus=20 Tlx+/+ (n =3D 3) laser microdissected SVZ cells. = Note that=20 expression of PTEN mRNA in Tlx+/=E2=88=92 is 78 =C2=B1 34% = higher than that in=20 Tlx+/+. P < 0.01, mean =C2=B1 standard deviation = (SD).=20 (F, G) Coronal brain sections of P9 Tlx+/+ = and=20 Tlx=E2=88=92/=E2=88=92 mice were stained for GFAP (red in = F). Note that=20 almost no GFAP+ cells were found in the SVZ of = Tlx=E2=88=92/=E2=88=92=20 mice. Bar, 10 =CE=BCm. (G) DCX (red). Note that almost no DCX = +=20 cells were found in the SVZ and RMS of Tlx=E2=88=92/=E2=88=92 = mice. Bar, 30 =CE=BCm.=20 (H=E2=80=93J) Tamoxifen were injected to Tlx = fl/fl;=20 Tlx-CreERT2 and Tlxfl/fl control mice = from P5 to=20 P8, the P10 brain sections were stained for GFAP (red in H), = Ki67=20 (green in I), and BLBP (green in J). Note that less = GFAP-,=20 Ki67-, and BLBP-expressing cells in the mutant SVZ than the control = mice. Bar,=20 10 =CE=BCm.

We show here that there are neither GFAP+ B = cells nor=20 neurogenesis in SVZ of Tlx=E2=88=92/=E2=88=92 mice. Adult = NSCs are derived from=20 radial glial cells around postnatal day 9 (P9) during postnatal = development (Merkle = et al.=20 2004). RC2-reactive cells and fibers appear morphologically normal = in=20 Tlx=E2=88=92/=E2=88=92 mice (Roy et = al.=20 2004), suggesting that radial glial cells are functioning normally = in the=20 absence of Tlx. This leads us to hypotheze that Tlx is involved in the=20 transition from radial glia to astrocyte-like stem cells. To investigate = if this=20 is indeed the case, we decided to analyze = Tlx=E2=88=92/=E2=88=92 mice at postnatal=20 stages. Brain sections from P9 Tlx=E2=88=92/=E2=88=92 mice = were stained for GFAP and=20 DCX. GFAP-expressing cells were rarely found in the SVZ of = Tlx=E2=88=92/=E2=88=92=20 mice. In contrast, GFAP+ cells already appeared at this time = in the=20 SVZ of wild-type mice (Fig. = 4F). The=20 same result was obtained for DCX staining, which indicates that P9=20 Tlx=E2=88=92/=E2=88=92 mice have a strong decrease in = neurogenesis (Fig.=20 4G). To further confirm Tlx is required for the transition from = radial glia=20 to astrocyte-like B cells during the early postnatal development, the=20 Tlxfl/fl; Tlx=E2=80=93CreERT2 mice were = used to=20 inactivate Tlx specifically in postnatal brain. Tamoxifen was injected = from P5=20 to P8 to inactivate Tlx in radial glial cells, P10 injected mice brain = were=20 analyzed by IHC. GFAP-expressing cells are strongly decreased in the = mutant (Fig. = 4H), which=20 is consistent with the results from the = Tlx=E2=88=92/=E2=88=92 mice. Ki67 staining=20 indicates that loss of Tlx in radial glial cells leads to a decrease in = cell=20 proliferation (Fig. = 4I). Brain=20 lipid binding protein (BLBP) is expressed by radial glial cells in = development=20 and by astrocytes in the adult brain (Mori = et al.=20 2005). To analyze whether the radial glial cells still exist in the = mutant=20 if they cannot transit into astrocyte-like B cells, BLBP antibody was = used to=20 stain the brain sections from the tamoxifen-injected mice. The SVZ cells = in the=20 mutant do not express BLBP like the cells from the control mice (Fig. = 4J), which=20 suggest the absence of radia glia and astrocyte-like stem cells in the = SVZ of=20 the Tlx mutant. We also tried to inactivate Tlx from P10 to P14, which = means=20 loss of Tlx after the transition from radial glia to astrocyte-like B = cell, we=20 found a similar phenotype to the adult inactivation of Tlx. The SVZ NSCs = did not=20 lose the expression of GFAP but lost the proliferation ability (data not = shown).=20 Taken all the above results together, we strongly suggest that Tlx is = also=20 required for the generation of astrocyte-like B cells in the adult = brain. These=20 results can explain why late-developing structures are also reduced in = size in=20 the Tlx-null mutant (Monagha= n et al.=20 1997).

That GFAP+ cells in the adult SVZ are = multipotent NSCs is=20 by now well accepted. However, we do not know the fundamental = characteristics=20 that distinguish neurogenic astrocytes from the vast population of = nonneurogenic=20 astrocytes elsewhere in the brain. Obviously, there is an urgent need = for=20 markers to distinguish neurogenic astrocytes from nonneurogenic = astrocytes (Doetsch = 2003;=20 Merkle = and=20 Alvarez-Buylla 2006). It is also very important to conceptually = distinguish=20 stem and progenitor cells (Seaberg= and van=20 der Kooy 2003). Here, using Cre as a reporter for Tlx expression, we = demonstrate that Tlx is specifically expressed by type B cells in the = SVZ-OB=20 neurogenic system. C and A cells do not express Tlx. Whether all = astrocyte-like=20 B cells in the SVZ are NSCs is still not clear, because it is difficult = to study=20 individual stem cells in vivo. Current methods to define NSCs are = strongly=20 dependent on in vitro experiments, and it has been reported that the = majority of=20 neurospheres correspond to rapidly cycling transit-amplifying cells = rather than=20 to primary and less proliferative in vivo stem cells (Doetsch = et al.=20 2002). Therefore, the number of quiescent primary stem cells in the = SVZ most=20 likely cannot be determined by in vitro studies. The expression of Tlx = in the=20 developing and adult NSCs strongly suggests that the Tlx+ = cells in=20 SVZ are indeed primary stem cells. In the SVZ of the adult brain, using = an=20 inducible knockout system, we show that Tlx is essential for = self-renewal of the=20 SVZ NSCs. This underscores the importance of this nuclear receptor in = the adult=20 NSCs of the SVZ (see the Supplemental Material).

In summary we show in this study that the nuclear receptor = Tlx is=20 expressed with an unprecedented exclusivity in astrocyte-like B cells of = the=20 SVZ-OB system, and that it is a crucial determinant of the generation of = adult=20 NSCs in the SVZ and the SGZ. Tlx is essential for the maintenance of=20 self-renewal of adult NSCs in the SVZ. This study also provides insights = of the=20 different identities of NSCs in the postnatal and adult brain, as well = as in=20 different neurogenic regions of the adult brain.

Pr= evious=20 SectionNe= xt=20 Section

Materials and methods

Animals

Mice were housed according to international standard = conditions, and=20 all animal experiments conformed to local and international guidelines = for the=20 use of experimental animals.

Tamoxifen, BrdU administration

Tamoxifen (Sigma) was dissolved in sunflower seed oil = (Sigma) with=20 10% EtOHabs to prepare the 10 mg/mL solution. To follow Cre=20 translocation by staining, 1-mg of tamoxifen was injected = intraperitoneally and=20 the animals were sacrificed 12 h later. For the analysis by X-gal = staining and=20 induction of loss of the Tlx conditional allele, tamoxifen was injected = for 5 d,=20 twice per day. BrdU (Sigma) was dissolved in sterile NaCl to prepare the = 15=20 mg/mL solution and was injected once 2 h before sacrifice. For provision = of BrdU=20 by feeding, BrdU was dissolved in the drinking water to prepare a 1 = mg/mL=20 solution.

The full description of materials and methods is available = in the=20 Supplemental Material.

Pr= evious=20 SectionNext=20 Section

Acknowledgments

We are grateful to Wolfgang Schmid for comments on the = manuscript. We=20 thank Monika Bock for excellent secretarial assistance. This work was = supported=20 by the =E2=80=9CDeutsche Forschungsgemeinschaft=E2=80=9D through = CollaborativeResearch Centres=20 SFB 488 and SFB 636, FOR Ot 165/2-2, GRK 791/1.02, and Sachbeihilfe Schu = 51/7-2;=20 by the European Union through grant LSHM-CT-2005-018652 (CRESCENDO); by = the=20 =E2=80=9CFonds der Chemischen Industrie=E2=80=9D; the Bundesministerium = f=C3=BCr Bildung und=20 Forschung (BMBF) through NGFN grants FZK 01GS01117, 01GS0477, and=20 KGCV1/01GS0416; and through project number 0313074C (Systems Biology).=20

Pr= evious=20 SectionNext=20 Section

Footnotes

  • =E2= =86=B53 Corresponding author.

    =E2= =86=B53 E-MAIL g.schuetz{at}dkfz.de; FAX = 49-6221-42-34-70.=20

  • Supplemental material is available at http://www.genesdev.org/. =

  • Article is online at http://www.ge= nesdev.org/cgi/doi/10.1101/gad.479308.=20

    • Received March 12, = 2008.=20
    • Accepted July 9, = 2008.=20
  • Copyright = =C2=A9 2008, Cold=20 Spring Harbor Laboratory Press
Previous=20 Section

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0px } #col-2 .cover IMG { BORDER-RIGHT: #333300 1px solid; BORDER-TOP: #333300 1px solid; = MARGIN-BOTTOM: 10px; BORDER-LEFT: #333300 1px solid; WIDTH: 134px; = BORDER-BOTTOM: #333300 1px solid; HEIGHT: 178px } #col-2 .toc-navigation CITE { FONT-SIZE: 0.85em; FONT-STYLE: normal } #col-2 .this-issue CITE { FONT-SIZE: 0.85em; FONT-STYLE: normal } #col-2 .toc-navigation .toc-links LI { PADDING-RIGHT: 0px; BACKGROUND-POSITION: 0pt center; DISPLAY: block; = PADDING-LEFT: 9px; FONT-SIZE: 0.85em; BACKGROUND-IMAGE: = url(../../shared/img/misc/rightchevron.gif); PADDING-BOTTOM: 0px; = MARGIN-LEFT: 0px; PADDING-TOP: 2px; BACKGROUND-REPEAT: no-repeat; = WHITE-SPACE: normal } #col-2 .toc-navigation .data-supp-links LI { PADDING-RIGHT: 0px; BACKGROUND-POSITION: 0pt center; DISPLAY: block; = PADDING-LEFT: 9px; FONT-SIZE: 0.85em; BACKGROUND-IMAGE: = url(../../shared/img/misc/rightchevron.gif); PADDING-BOTTOM: 0px; = MARGIN-LEFT: 0px; PADDING-TOP: 2px; BACKGROUND-REPEAT: no-repeat; = WHITE-SPACE: normal } #col-2 .this-issue .toc-links LI { PADDING-RIGHT: 0px; BACKGROUND-POSITION: 0pt center; DISPLAY: block; = PADDING-LEFT: 9px; FONT-SIZE: 0.85em; BACKGROUND-IMAGE: = url(../../shared/img/misc/rightchevron.gif); PADDING-BOTTOM: 0px; = MARGIN-LEFT: 0px; PADDING-TOP: 2px; BACKGROUND-REPEAT: no-repeat; = WHITE-SPACE: normal } #col-2 .issue-sections { BORDER-TOP: #b9c28d 1px solid } ------=_NextPart_000_0000_01CAEC8C.AC04E920 Content-Type: text/css; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://genesdev.cshlp.org/content/22/18/hw-global-elements.css HTML { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } BODY { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } DIV { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } SPAN { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } APPLET { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } OBJECT { BORDER-TOP-WIDTH: 0px; 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inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } H3 { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } H4 { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } H5 { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } H6 { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } P { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } BLOCKQUOTE { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } PRE { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } A { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } ABBR { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } ACRONYM { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } ADDRESS { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } BIG { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } CITE { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } CODE { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } DEL { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } DFN { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } EM { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } FONT { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; PADDING-LEFT: 0px; = BORDER-LEFT-WIDTH: 0px; BORDER-BOTTOM-WIDTH: 0px; PADDING-BOTTOM: 0px; = MARGIN: 0px; VERTICAL-ALIGN: baseline; PADDING-TOP: 0px; FONT-FAMILY: = inherit; BORDER-RIGHT-WIDTH: 0px; outline-style: none } IMG { BORDER-TOP-WIDTH: 0px; PADDING-RIGHT: 0px; 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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 || gColTempResize) && ((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 || gColTempResize) && 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= if (typeof(offset.x)=3D=3D"undefined") { offset.x =3D 0; }=0A= if (typeof(offset.y)=3D=3D"undefined") { offset.y =3D 0; }=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; 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';expires=3D'+expires_date.toGMTString() : '' ) +=0A= ( ( path ) ? ';path=3D' + path : '' ) +=0A= ( ( domain ) ? ';domain=3D' + domain : '' ) +=0A= ( ( secure ) ? ';secure' : '' );=0A= }=0A= =0A= function deleteCookie( name, path, domain ) {=0A= if ( getCookie( name ) ) document.cookie =3D name + '=3D' +=0A= ( ( path ) ? ';path=3D' + path : '') +=0A= ( ( domain ) ? 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        ' + expandStr + ' Author Affiliations

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'&' : '?') = + 'baseURI=3D' + ((baseAjaxUrl.indexOf('?') > 0) ? = baseAjaxUrl.substring(0, baseAjaxUrl.indexOf('?')): baseAjaxUrl);=0A= //var ajaxUrl =3D baseAjaxUrl + ((href.indexOf('?') >=3D 0) ? '&' : = '?') + 'baseURI=3D' + baseAjaxUrl;=0A= $.ajax({=0A= url: ajaxUrl,=0A= dataType: "html",=0A= type: "GET",=0A= error: ajaxErr,=0A= beforeSend: addFigHeaders,=0A= success: function(xhtml) {=0A= addFig(xhtml, figWrapperEl);=0A= },=0A= complete: ajaxComplete=0A= });=0A= e.preventDefault();=0A= }=0A= function addFigHeaders(req) {=0A= addCommonHeaders(req);=0A= addPartHeaders(req);=0A= }=0A= =0A= function addFig(xhtmlData, figWrapperEl) {=0A= if (xhtmlData && !(xhtmlData.indexOf('=3D 0)) {=0A= figWrapperEl.replaceWith(xhtmlData);=0A= newWindowTargets();=0A= var lookupRule =3D "div.article div.fig img";=0A= var numImagesToLoad =3D checkUnloadedImgs(lookupRule);=0A= setTimeout("fixHeightForImages(1" + "," + numImagesToLoad + ",'" + = lookupRule + "')", 1000);=0A= fixColHeights(1);=0A= }=0A= }=0A= =0A= function refLinksNewWindowTarget() {=0A= $("div.ref-list div.cit-extra a").each(=0A= function(i) {=0A= var origTitle =3D $(this).attr("title");=0A= var newTitle =3D '';=0A= if ((origTitle =3D=3D undefined) || (!origTitle)) {=0A= origTitle =3D '';=0A= }=0A= else {=0A= newTitle =3D origTitle + ' ';=0A= }=0A= newTitle +=3D '[opens in a new window]';=0A= $(this).attr("target", "_blank").attr("title", newTitle);=0A= }=0A= );=0A= }=0A= =0A= function addRefPops() {=0A= var numMissed =3D 0;=0A= var maxToSkip =3D getSiteOption('refPopsMaxToSkip', 10);=0A= var i =3D 1;=0A= var idroot =3D "#xref-ref-";=0A= var el =3D $(idroot + i + '-1');=0A= /* not all refs appear in text; compensate */=0A= while (numMissed < maxToSkip) {=0A= if (el.length) {=0A= numMissed =3D 0;=0A= el.hover(dispRef, hideRef);=0A= var j =3D 2;=0A= var el2 =3D $(idroot + i + "-" + j);=0A= while (el2.length) {=0A= el2.hover(dispRef, hideRef);=0A= j++;=0A= el2 =3D $(idroot + i + "-" + j);=0A= }=0A= }=0A= else {=0A= numMissed++;=0A= }=0A= i++;=0A= el =3D $(idroot + i + "-1");=0A= }=0A= }=0A= =0A= function dispRef(e) {=0A= var link =3D $(this).attr("href");=0A= if($("div#hovering-ref").length) {=0A= $("div#hovering-ref").remove();=0A= //alert("hovering-ref div removed on new hover!");=0A= }=0A= var linkEl =3D $(link);=0A= if (linkEl.length) {=0A= var citHtml =3D linkEl.next("div").children("div.cit-metadata");=0A= if (!(citHtml.length)) {=0A= citHtml =3D linkEl.parent().next("div").children("div.cit-metadata");=0A= }=0A= if (citHtml.length) {=0A= var newDiv =3D '
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        ';=0A= $("body").append(newDiv);=0A= var elH =3D getObjHeight($("div#hovering-ref"));=0A= $("div#hovering-ref").css("left", e.pageX+10).css("top", = e.pageY-elH).css("position", "absolute");=0A= }=0A= }=0A= }=0A= =0A= function hideRef(e) {=0A= if($("div#hovering-ref").length) {=0A= $("div#hovering-ref").remove();=0A= }=0A= }=0A= =0A= function getHWCiting() {=0A= var citingA =3D $("#cb-hw-citing-articles");=0A= if (citingA.length) {=0A= var newA =3D 'Loading citing = article data...';=0A= citingA.replaceWith(newA);=0A= var href =3D citingA.attr("href");=0A= var id =3D '';=0A= if (href && (href.indexOf('?') > 0)) {=0A= var args =3D href.substring(href.indexOf('?') + 1).split('&');=0A= for (var i =3D 0; i < args.length; i++) {=0A= if (args[i].toLowerCase().indexOf('legid=3D') =3D=3D 0) {=0A= id =3D args[i].substring(args[i].indexOf('=3D') + 1);=0A= if (id.indexOf('#') > 0) {=0A= id =3D id.substring(0, id.indexOf('#'));=0A= }=0A= }=0A= }=0A= if (!(id =3D=3D '')) {=0A= var host =3D document.location.protocol + "//" + = document.location.host;=0A= var ajaxUrl =3D host + '/cited-by/' + id.replace(/;/,'/');=0A= $.ajax({=0A= url: ajaxUrl,=0A= dataType: "html",=0A= type: "GET",=0A= error: ajaxErr,=0A= success: addHWCiting,=0A= complete: ajaxComplete=0A= });=0A= =0A= }=0A= }=0A= }=0A= }=0A= function getHWRelatedURLs() {=0A= var relatedURLsA =3D $("#cb-related-urls");=0A= if (relatedURLsA.length) {=0A= var newA =3D 'Loading = related web page data...';=0A= relatedURLsA.replaceWith(newA);=0A= var href =3D relatedURLsA.attr("href");=0A= var id =3D '';=0A= if (href && (href.indexOf('?') > 0)) {=0A= var args =3D href.substring(href.indexOf('?') + 1).split('&');=0A= for (var i =3D 0; i < args.length; i++) {=0A= if (args[i].toLowerCase().indexOf('legid=3D') =3D=3D 0) {=0A= id =3D args[i].substring(args[i].indexOf('=3D') + 1);=0A= if (id.indexOf('#') > 0) {=0A= id =3D id.substring(0, id.indexOf('#'));=0A= }=0A= }=0A= }=0A= if (!(id =3D=3D '')) {=0A= var host =3D document.location.protocol + "//" + = document.location.host;=0A= var ajaxUrl =3D host + '/related-web-pages/' + id.replace(/;/,'/');=0A= $.ajax({=0A= url: ajaxUrl,=0A= dataType: "html",=0A= type: "GET",=0A= error: ajaxErr,=0A= success: addRelatedURLs,=0A= complete: ajaxComplete=0A= });=0A= =0A= }=0A= }=0A= }=0A= }=0A= function getPatientInformData() {=0A= var pInformA =3D $("#cb-patientinform");=0A= if (pInformA.length) {=0A= var newA =3D 'Loading = patientINFORMation...';=0A= pInformA.replaceWith(newA);=0A= var href =3D pInformA.attr("href");=0A= var id =3D '';=0A= if (href && (href.indexOf('?') > 0)) {=0A= var args =3D href.substring(href.indexOf('?') + 1).split('&');=0A= for (var i =3D 0; i < args.length; i++) {=0A= if (args[i].toLowerCase().indexOf('legid=3D') =3D=3D 0) {=0A= id =3D args[i].substring(args[i].indexOf('=3D') + 1);=0A= if (id.indexOf('#') > 0) {=0A= id =3D id.substring(0, id.indexOf('#'));=0A= }=0A= }=0A= }=0A= if (!(id =3D=3D '')) {=0A= var host =3D document.location.protocol + "//" + = document.location.host;=0A= var ajaxUrl =3D host + '/related-web-pages/patientinform/' + = id.replace(/;/,'/');=0A= $.ajax({=0A= url: ajaxUrl,=0A= dataType: "html",=0A= type: "GET",=0A= error: ajaxErr,=0A= success: addPatientInform,=0A= complete: ajaxComplete=0A= });=0A= =0A= }=0A= }=0A= }=0A= }=0A= function getISIRelated() {=0A= var relatedA =3D $("#cb-isi-similar-articles");=0A= if (relatedA.length) {=0A= var newA =3D 'Loading Web = of Science article data...';=0A= relatedA.replaceWith(newA);=0A= var href =3D relatedA.attr("href");=0A= var id =3D '';=0A= if (href) {=0A= var hrefDec =3D decodeURI(href);=0A= if ((hrefDec.indexOf('?') > 0)) {=0A= var args =3D hrefDec.substring(hrefDec.indexOf('?') + 1).split('&');=0A= for (var i =3D 0; i < args.length; i++) {=0A= var argDec =3D decodeURIComponent(args[i]);=0A= if (argDec.toLowerCase().indexOf('access_num=3D') =3D=3D 0) {=0A= id =3D argDec.substring(argDec.indexOf('=3D') + 1);=0A= if (id.indexOf('#') > 0) {=0A= id =3D id.substring(0, id.indexOf('#'));=0A= }=0A= }=0A= }=0A= if (!(id =3D=3D '')) {=0A= var host =3D document.location.protocol + "//" + = document.location.host;=0A= var ajaxUrl =3D host + '/isi-links/has-related/' + = id.replace(/;/,'/');=0A= $.ajax({=0A= url: ajaxUrl,=0A= dataType: "html",=0A= type: "GET",=0A= error: ajaxErr,=0A= success: addISIRelated,=0A= complete: ajaxComplete=0A= });=0A= }=0A= }=0A= }=0A= }=0A= }=0A= function getCiting(service, msg, pathseg, successFn) {=0A= var citingA =3D $("#cb-" + service + "-citing-articles");=0A= if (citingA.length) {=0A= var newA =3D '' = + msg + '';=0A= citingA.replaceWith(newA);=0A= var href =3D citingA.attr("href");=0A= var id =3D '';=0A= if (href) {=0A= var hrefDec =3D decodeURI(href);=0A= if ((hrefDec.indexOf('?') > 0)) {=0A= var args =3D hrefDec.substring(hrefDec.indexOf('?') + 1).split('&');=0A= for (var i =3D 0; i < args.length; i++) {=0A= var argDec =3D decodeURIComponent(args[i]);=0A= if (argDec.toLowerCase().indexOf('access_num=3D') =3D=3D 0) {=0A= id =3D argDec.substring(argDec.indexOf('=3D') + 1);=0A= if (id.indexOf('#') > 0) {=0A= id =3D id.substring(0, id.indexOf('#'));=0A= }=0A= }=0A= }=0A= if (!(id =3D=3D '')) {=0A= var host =3D document.location.protocol + "//" + = document.location.host;=0A= var ajaxUrl =3D host + '/' + service + '-links/' + pathseg + = id.replace(/;/,'/');=0A= $.ajax({=0A= url: ajaxUrl,=0A= dataType: "html",=0A= type: "GET",=0A= error: ajaxErr,=0A= success: successFn,=0A= complete: ajaxComplete=0A= });=0A= }=0A= }=0A= }=0A= }=0A= }=0A= =0A= function getISICiting() {=0A= var citingA =3D $("#cb-isi-citing-articles");=0A= if (citingA.length) {=0A= var newA =3D 'Loading Web of = Science citing article data...';=0A= citingA.replaceWith(newA);=0A= var href =3D citingA.attr("href");=0A= var id =3D '';=0A= if (href) {=0A= var hrefDec =3D decodeURI(href);=0A= if ((hrefDec.indexOf('?') > 0)) {=0A= var args =3D hrefDec.substring(hrefDec.indexOf('?') + 1).split('&');=0A= for (var i =3D 0; i < args.length; i++) {=0A= var argDec =3D decodeURIComponent(args[i]);=0A= if (argDec.toLowerCase().indexOf('access_num=3D') =3D=3D 0) {=0A= id =3D argDec.substring(argDec.indexOf('=3D') + 1);=0A= if (id.indexOf('#') > 0) {=0A= id =3D id.substring(0, id.indexOf('#'));=0A= }=0A= }=0A= }=0A= if (!(id =3D=3D '')) {=0A= var host =3D document.location.protocol + "//" + = document.location.host;=0A= var ajaxUrl =3D host + '/isi-links/' + id.replace(/;/,'/');=0A= $.ajax({=0A= url: ajaxUrl,=0A= dataType: "html",=0A= type: "GET",=0A= error: ajaxErr,=0A= success: addISICiting,=0A= complete: ajaxComplete=0A= });=0A= }=0A= }=0A= }=0A= }=0A= }=0A= function getEntrezLinks() {=0A= var entrezDiv =3D $("#cb-entrez-links-placeholder");=0A= if (entrezDiv.length) {=0A= var entrezA =3D entrezDiv.children("a");=0A= if (entrezA) {=0A= var host =3D document.location.protocol + "//" + = document.location.host;=0A= var ajaxUrl =3D host + entrezA.attr("href");=0A= $.ajax({=0A= url: ajaxUrl,=0A= dataType: "html",=0A= type: "GET",=0A= error: ajaxErr,=0A= success: addEntrezLinks,=0A= complete: ajaxComplete=0A= });=0A= }=0A= }=0A= }=0A= =0A= function ajaxErr(req, msg, e) {=0A= }=0A= function ajaxComplete(req, msg) {=0A= }=0A= function updateCBItem(cbItem, newHTML, hasData) {=0A= var parentItem =3D cbItem.parents("li").eq(0);=0A= cbItem.replaceWith(newHTML);=0A= if (!hasData) {=0A= // hide the parent li=0A= if (parentItem.length) {=0A= modClass(parentItem,"nodata","");=0A= // check if there are any siblings still being displayed=0A= var otherItems =3D parentItem.siblings();=0A= var allItemsEmpty;=0A= if (otherItems.length) {=0A= if (otherItems.length =3D=3D otherItems.filter(".nodata").length) {=0A= allItemsEmpty =3D true=0A= }=0A= else {=0A= allItemsEmpty =3D false;=0A= }=0A= }=0A= else {=0A= allItemsEmpty =3D true;=0A= }=0A= if (allItemsEmpty) {=0A= var cbsection =3D parentItem.parents("div.cb-section").eq(0);=0A= if (cbsection.length) {=0A= modClass(cbsection,"nodata","");=0A= }=0A= }=0A= }=0A= }=0A= // in frameset fix targets on child links, forms=0A= fixFrameLinks(parentItem.find("a,form"));=0A= fixColHeights(2);=0A= }=0A= function fixFrameLinks(jqItems) {=0A= if ((gIsFrameset !=3D null) && gIsFrameset) {=0A= jqItems.each(=0A= function(i) {=0A= var href =3D $(this).attr("href");=0A= var action =3D $(this).attr("action"); // if form=0A= if ((href !=3D null) || (action !=3D null)) {=0A= var inFrameAnchor =3D ((href !=3D null) && = (((/frameset=3D/.test(href)) && (/#/.test(href))) || = (href.substring(0,1) =3D=3D '#')));=0A= if ((!inFrameAnchor) || (action !=3D null)) {=0A= $(this).attr("target", "_top");=0A= }=0A= }=0A= }=0A= );=0A= }=0A= }=0A= function addRelatedURLs(xhtmlData) {=0A= var cbA =3D $("#cb-loading-related-urls");=0A= if (gIsFrameset) {=0A= if (cbA.length) {=0A= updateCBItem(cbA, '', false);=0A= }=0A= }=0A= else if (xhtmlData && !(xhtmlData.indexOf('=3D 0)) {=0A= $("#related-urls").replaceWith(xhtmlData);=0A= var relatedWebPagesLabel =3D getSiteOption('relatedWebPagesLabel', = 'Related Web Pages');=0A= fixColHeights(1);=0A= if (cbA.length) {=0A= updateCBItem(cbA, '' + relatedWebPagesLabel = + '', true);=0A= }=0A= }=0A= else {=0A= if (cbA.length) {=0A= updateCBItem(cbA, '', false);=0A= }=0A= }=0A= }=0A= function addPatientInform(xhtmlData) {=0A= var cbA =3D $("#cb-loading-patientinform");=0A= if (gIsFrameset) {=0A= if (cbA.length) {=0A= updateCBItem(cbA, '
        Not = available in this view
        ', false);=0A= }=0A= }=0A= else if (xhtmlData && !(xhtmlData.indexOf('=3D 0)) {=0A= $("#patientinform-links").replaceWith(xhtmlData);=0A= fixColHeights(1);=0A= if (cbA.length) {=0A= updateCBItem(cbA, 'patientINFORMation', true);=0A= }=0A= }=0A= else {=0A= if (cbA.length) {=0A= updateCBItem(cbA, '
        No = patientINFORMation available for this article
        ', false);=0A= }=0A= }=0A= }=0A= function addHWCiting(xhtmlData) {=0A= var cbA =3D $("#cb-loading-hw-cited");=0A= if (gIsFrameset) {=0A= if (cbA.length) {=0A= updateCBItem(cbA, '
        Not available = in this view
        ', false);=0A= }=0A= }=0A= else if (xhtmlData) {=0A= $("#content-block").append(xhtmlData);=0A= var hwCitingLabel =3D getSiteOption('hwCitingLabel', 'View citing = article information');=0A= fixColHeights(1);=0A= if (cbA.length) {=0A= updateCBItem(cbA, '' + hwCitingLabel + '', = true);=0A= }=0A= }=0A= else {=0A= if (cbA.length) {=0A= updateCBItem(cbA, '
        No citing = articles
        ', false);=0A= }=0A= }=0A= }=0A= function addISIRelated(xhtmlData) {=0A= var cbA =3D $("#cb-isi-similar-articles");=0A= if (xhtmlData && !(xhtmlData.indexOf('=3D 0)) {=0A= if (cbA.length) {=0A= updateCBItem(cbA, xhtmlData, true);=0A= }=0A= }=0A= else {=0A= if (cbA.length) {=0A= updateCBItem(cbA, '', false);=0A= }=0A= }=0A= }=0A= function addISICiting(xhtmlData) {=0A= var cbA =3D $("#cb-loading-isi-cited");=0A= if (xhtmlData && !(xhtmlData.indexOf('=3D 0)) {=0A= if (cbA.length) {=0A= updateCBItem(cbA, xhtmlData, true);=0A= }=0A= }=0A= else {=0A= if (cbA.length) {=0A= updateCBItem(cbA, '
        No Web of = Science citing articles
        ', false);=0A= }=0A= }=0A= }=0A= function addScopusCiting(xhtmlData) {=0A= var cbA =3D $("#cb-loading-scopus-cited");=0A= if (xhtmlData && (xhtmlData.indexOf('=3D 0)) {=0A= if (cbA.length) {=0A= updateCBItem(cbA, xhtmlData, true);=0A= }=0A= }=0A= else {=0A= if (cbA.length) {=0A= updateCBItem(cbA, '
        No Scopus = citing articles
        ', false);=0A= }=0A= }=0A= }=0A= function addEntrezLinks(xhtmlData) {=0A= var entrezDiv =3D $("#cb-entrez-links-placeholder");=0A= =0A= if (xhtmlData && (xhtmlData.indexOf('        =3D 0)) {=0A= if (entrezDiv.length) {=0A= updateCBItem(entrezDiv, xhtmlData, true);=0A= }=0A= }=0A= else {=0A= if (entrezDiv.length) {=0A= updateCBItem(entrezDiv, '', false);=0A= }=0A= }=0A= }=0A= =0A= function updateSBLinks() {=0A= var fbLink =3D $("a.sb-facebook");=0A= if (fbLink.length) {=0A= fbLink.click(=0A= function(e) {=0A= window.open(this.href, 'sharer', = 'toolbar=3D0,status=3D0,width=3D626,height=3D436');=0A= e.preventDefault();=0A= }=0A= );=0A= }=0A= }=0A= =0A= function addDockedNav() {=0A= var slugEl =3D $("#col-2 #slugline");=0A= // get direct children li elements only=0A= var artViews =3D $("#article-cb-main .cb-section ol:has(li = a[rel^=3D'view-']) > li").clone();=0A= var newDiv =3D '
        ';=0A= $("#col-2").append(newDiv);=0A= $("#col-2 #docked-nav").hide();=0A= var newDivJQuery =3D $("#docked-nav");=0A= newDivJQuery.append('

        This Article

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