From: Subject: Migration and differentiation of neural precursor cells can be directed by microglia -- Aarum et al. 100 (26): 15983 -- Proceedings of the National Academy of Sciences Date: Thu, 17 Jan 2008 18:29:54 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C85936.F43A32E0" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.3198 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C85936.F43A32E0 Content-Type: text/html; charset="Windows-1252" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.pnas.org/cgi/content/full/100/26/15983 Migration and differentiation of neural precursor = cells can be directed by microglia -- Aarum et al. 100 (26): 15983 -- = Proceedings of the National Academy of Sciences
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Published online before print December = 10, 2003,=20 10.1073/pnas.2237050100
PNAS | = December 23,=20 2003 | vol. 100 | no. 26 | 15983-15988


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Neuroscience
Migration and=20 differentiation of neural precursor cells can be directed by microglia=20

Johan Aarum *, = Kristian=20 Sandberg 3D{dagger}, Samantha L. Budd = Haeberlein ,=20 and Mats A. A. Persson * 3D{ddagger}=20=20

*Karolinska Institutet, Department of Medicine and Center = for=20 Molecular Medicine, Karolinska Hospital, 171 76 Stockholm, Sweden; and = Astrazeneca R&D, Novum, S-141 86 Huddinge, Sweden=20

Communicated by Tomas H=F6kfelt, Karolinska Institutet, Stockholm, = Sweden,=20 October 30, 2003 (received for review October 10, 2003)


3D"   =20 Abstract
3D"Top
3D"Abstract
Materials and Methods
Results
Discussion
References
 
R= ecent=20 reports have supported the existence of neural stem cells in = the=20 adult mammalian CNS. Important features of such cells are=20 self-renewal and multipotency, i.e., they can give rise to = neurons,=20 astrocytes, and oligodendrocytes and thus in principle = replace lost=20 cells in the CNS. Observations in several animal models of = CNS=20 diseases have shown that by unknown mechanisms endogenous as = well as=20 exogenous precursor cells preferentially migrate to damaged = areas.=20 Microglia are immunoreactive cells of nonneural lineage = resident in=20 the CNS. After injury to the CNS, microglia are rapidly = activated and=20 found concentrated at the sites of injury. In the present = article we=20 show, in two different assays, that soluble factors released = from=20 mouse microglial cells direct the migration of neural CNS = precursor=20 cells. We also provide evidence that microglia have the = capacity to=20 influence the differentiation of both adult and embryonic = neural=20 precursor cells toward a neuronal phenotype. Given that an = invariant=20 feature of pathological processes in CNS is the activation of = microglia, these results indicate an important and unique = role for=20 microglia in directing the replacement of damaged or lost = cells in=20 the CNS.

The discovery of neural stem cells and their capacity to regenerate = neural cells in the adult CNS have altered old dogmas in = neurobiology=20 and raised hopes for novel treatment in medicine. = Nonetheless,=20 it is clear that damage to the CNS in, e.g., = neurodegenerative=20 diseases is not matched by the ability of endogenous = precursor=20 cells to replace lost cells. To reach this therapeutic ideal, = much additional knowledge has to be gained regarding the = processes=20 regulating replacement of damaged cells.

In the adult CNS, there are two principle locations of neural = stem=20 cells: in the subventricular zone and in the subgranular zone = of the=20 dentate gyrus in the hippocampus (reviewed in ref. 1). = In=20 addition, precursor cells with varying capacity for self-renewal = and=20 differentiation have also been isolated from other regions of = the=20 CNS, including from human white matter (2=96<= A=20 href=3D"http://www.pnas.org/cgi/content/full/100/26/15983#REF4">4). = In=20 avian and rodent CNS, the new cells formed in the = subventricular=20 zone migrate in the rostral migratory stream from the = ventricular=20 walls to the olfactory bulb where they replace granule cells=20 and periglomerular cells, whereas precursor cells in the = subgranular=20 zone mainly differentiate to granular cells (5, = 6). = In=20 response to a number of experimental lesions in rodent's = brain,=20 including stroke, ischemia, and seizure, precursor cells in = the=20 neurogenic centers start to proliferate and migrate (7=96<= A=20 href=3D"http://www.pnas.org/cgi/content/full/100/26/15983#REF9">9).=20 Interestingly, by unknown mechanisms, new cells = preferentially end up=20 at the sites of pathology, indicating that agents present in = the=20 damaged tissue guide the migration of precursor cells (7, = 10).= =20 Besides the proliferation and migration of endogenous = precursor=20 cells, exogenously propagated cells are similarly capable of=20 migrating and repopulating damaged areas of the adult CNS (11=96= 14).= =20

The signals that direct the migration of precursor cells are=20 largely unknown. In the rostral migratory stream, the = secreted=20 protein Slit has been suggested to function as a repellant = for=20 young migrating GABAergic neurons (15).= Other=20 molecules shown to guide neuronal precursor cells include = SDF-1,=20 which guides embryonic cerebellar neurons (16, = 17),= and=20 recently unidentified signals from the prenatal cortex have = been=20 shown to guide interneurons to the cerebral cortex (18).= The=20 effect of these and other signals have mainly been studied on = committed, immature neurons, whereas the signals that direct = the=20 migration of multipotent neural precursor cells remain = elusive.=20

Another important issue is the correct differentiation of = precursor=20 cells to the cell type that should be replaced to regain = function.=20 Recent studies show a remarkable flexibility of stem cells = regarding=20 the cell type they will differentiate to. For example, = exogenously=20 propagated neural precursor cells administrated to an = experimental=20 model of multiple sclerosis were observed as oligodendrocytes = in the damaged white matter (19, = 20),= whereas=20 other studies have shown that precursor cells injected in = neurogenic=20 areas of the adult brain develops into neurons, gaining=20 attributes appropriate to their location (21, = 22).= It is=20 reasonable to believe that interactions between precursor = cells and=20 the microenvironment faced by the new cells influence their=20 differentiation. In accordance with this, astrocytes were = recently=20 shown to induce a neuronal phenotype on adult stem cells from = mice,=20 both as a result of cell=96cell contact and by secreted = factors (23).= =20

However, little is known about how a pathological environment=20 affects the differentiation of precursor cells. An invariant=20 feature of damage to the CNS is the migration of microglia = cells=20 to the site of injury and their subsequent activation; also=20 astrocytes may be activated. The activation of microglia can=20 result in either neuroprotective or neurotoxic effects, or = both.=20 Among such protective features, microglia can express and = secrete=20 neurotrophins such as brain-derived neurotrophic factor, as=20 well as several cytokines and chemokines (24, = 25).= Because=20 of the relocation of microglia to the site of injury and = their=20 rapid activation as response to CNS abnormalities, we = reasoned=20 that these cells may be involved in the attraction of = precursor=20 cells as well as in effecting their differentiation; we here=20 present data that support this hypothesis.


3D"   =20 Materials and Methods
3D"Top
3D"Abstract
3D"Materials and Methods
Results
Discussion
References
 
<= B>Cell=20 Culture and Conditioned Media. All cell culture reagents = were=20 from Invitrogen unless otherwise stated in the text. Cell = cultures=20 were kept at 37=B0C in 10% CO2 and 90% relative = humidity.=20 Primary microglia (PM) was prepared from embryonic day (E) 16 = to E17 C57/BL mice, with some modifications, as described (26).= =20 In summary, isolated tissue was mechanically dissociated = between=20 the frosted ends of two glass slides, filtered trough a = 70-=B5m=20 nylon mesh (Falcon), and washed in culture medium (DMEM 10%=20 FCS/0.45% glucose) before seeded in poly(L)lysine-coated = flasks.=20 When confluence was reached, microglial cells were harvested = by=20 gently shaking the flasks and transferring the resulting cell = suspension, after washing, to poly(L)-lysine coated petridishes,=20 changing one-half the medium to serum-free Neurobasal medium=20 supplemented with B27 (NB27). After seeding (1 day), the = enriched=20 microglial {90=9695% microglia [Iba-1+ cells]/<5% = astrocytes=20 [glia fibrillary acidic protein (GFAP)-positive cells]} = cultures=20 were washed with fresh NB27 and then cultivated in NB27 with = or=20 without lipopolysaccharide (LPS, Sigma) for 24 h. Conditioned = media=20 was harvested, cleared from free-floating cells by centrifugation = for=20 5 min at 1,000 x g, and = sterile=20 filtered. Conditioned media was kept at 4=B0C until use. =

The mouse microglia cell lines BV-2 and N13 and the astrosarcoma = cell line C6 were cultured in NB27. When reaching confluence, = fresh media were added with or without LPS (10 ng/ml), = harvested=20 after 24 h, centrifuged as described above, and passed = through=20 a 0.2-=B5m filter.

Conditioned medium (CM) taken from the cell lines (BV-2, N13, = and=20 C6) will be termed BV-2/N13-CM and C6-CM, respectively, = whereas=20 medium taken from PM cultures will be referred to as PM-CM.=20 Conditioned media from BV-2 and N13, irrespective of LPS = treatment,=20 gave very similar results and will, therefore, for sake of = clarity,=20 be treated as one.

Aggregate Cultures and Isolation of Embryonic Precursor = Cells.=20 Cortical aggregate of total CNS cells and isolation of = precursor=20 cells were performed as described (C. M. D. Berglund, = S.L.B.H.,=20 J. R. Nyengaard, T. H=F6kfelt, K.S., J. N=E4slund, and = M.A.A.P.,=20 unpublished data). In summary, cortical tissue from E16=96E17 = C57/BL=20 mice were isolated and mechanically dissociated into = single-cell=20 suspensions, seeded in Erlenmyer flasks, and incubated on a = gyratory=20 shaker. After seeding (7 days), spherical aggregates formed, = which=20 contained the principal brain cells; astrocytes, neurons, = microglia,=20 and oligodendrocytes. After seeding (11=9625 days), the = culture medium=20 was supplemented with 20 ng of epidermal growth factor = (EGF)/ml=20 (Becton Dickinson) and one-half the medium was then changed = to fresh=20 EGF-containing medium every third day.

Precursor cells were isolated from EGF-treated aggregates by=20 incubating the aggregates in 0.25% trypsin for 5 min at = 37=B0C=20 followed by gently pipetting to release cells from the = aggregate.=20 The cell suspension was seeded in bacterial petridishes = (Sarstedt)=20 in NB27 supplemented with 20 ng of EGF/ml. Formed = neurospheres=20 were divided as described below for adult precursor cells. =

Adult Precursor Cell Culture. Adult precursor cultures = were=20 prepared essentially as described by Johansson et al. (27).= =20 In brief, the lateral ventricles were dissected from adult = female=20 C57/BL mice (>20 weeks), and the tissue was dissociated in = trypsin solution [0.25% Trypsin/EDTA/15 mM Hepes (Sigma)/0.5% = glucose in 1x Hanks' balanced = salt=20 solution (Sigma)] at 37=B0C in 10% CO2 for 25 min, = gently=20 triturated, and incubated for 5 more min. The trypsin was = quenched by=20 the addition 0.5 vol of 2% BSA and 20 mM Hepes in 1x Hanks' balanced salt solution, = and the=20 cell solution was filtered through a 70-=B5m nylon mesh. = Cells were=20 washed in NB27 before seeding in bacterial petridishes in = NB27=20 containing 20 ng of EGF/ml. One week after seeding, the = cultures (now=20 neurospheres) were divided by trypsination for 5 min and = thereafter=20 once a week.

Precursor Cell Migration. Migration from aggregates.=20 Conditioned media were concentrated 30 times by centricon = filtration=20 (Millipore; cut-off 50 kDa). Concentrated medium was then = mixed 1:1=20 with 4% Baculovirus agarose (Invitrogen) in Hanks' balanced = salt=20 solution at 48=B0C and 5-=B5l aliquoted to 96-well plates = (Costar).=20 Plates were then placed on ice and allowed to set before the = addition=20 of 48 =B5l of Matrigel (Becton Dickinson)/NB27 at a ratio of = 1:3.=20

EGF-treated aggregates were washed extensively with fresh NB27 = and=20 cultivated for 24 h without EGF. Single aggregates were then = picked=20 and placed in the Matrigel-containing wells (on ice) close to = the=20 agarose-mixed conditioned media (Fig.=20 2H). Twenty-four hours later, 152 =B5l of NB27 was = added,=20 and after 48 h, the cultures were fixed for 15 min in 4%=20 paraformaldehyde, washed with PBS, and kept at 4=B0C in 0.5% = BSA/0.02%=20 NaN3/PBS until determination.



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  		Fig. 2. = Migration=20 of precursor cells from EGF-treated aggregates in Matrigel.=20 (A) Precursor cells migrate radially from EGF-treated = aggregate when embedded in Matrigel. (B) Cells = migrated both=20 as single cells frequently attached to long processes = (arrows) and=20 tightly packed together. (C) No cells migrated from=20 aggregates not pretreated with EGF. (D=96G)=20 Pseudocolored immunofluorescence micrographs of the same = EGF-treated=20 aggregate 6 days after embedding in Matrigel. Aggregates = were=20 immunostained for 3D{beta}=20III = (neurons, red)=20 (D) and GFAP (astrocytes, green) (E) and=20 counterstained with DAPI (nuclei, blue) (F). = (G)=20 Overlay picture of D=96F. (Scale bars, 50 = =B5m.)=20 (H) Schematic drawing showing the experimental = principle and=20 the overlaid quadrants used to quantify precursor migration = from the=20 embedded aggregate in the centre. The proximal (p) and the = distal=20 (d) quadrants relative to the conditioned media. =

=

Fixed cells in Matrigel were either stained with Abs, Tuj-1=20 against 3D{beta}=20 class III tubulin (3D{beta}III) (rabbit, Convance) at a dilution of 1:3,000 = and=20 anti-GFAP (mouse, Sigma) at a dilution of 1:500 for 48 h at = 4=B0C,=20 washed for 4 h at room temperature, and stained with = secondary Abs=20 for 24 h or only stained with 4', 6-diamidino-2-phenylindole = (DAPI,=20 Calbiochem). Migrating cells were quantified by dividing the = area=20 around the embedded aggregate into four quadrants, as = described by Wu=20 et al. (15) = (Fig.=20 2H).

Transwell migration. Transwell membranes (Costar, = 8-=B5m=20 pore size) were coated 4 h on both sides with Matrigel in = NB27=20 (100 =B5g/ml), washed with NB27, and placed in 24-well = plates.=20 Below the membrane, 320 =B5l of cell-free conditioned medium = was=20 added and diluted in NB27 to represent a concentration of = conditioned=20 media of 0.75 x 106 cells = per ml.=20 To the upper chamber, 100 =B5l of precursor cells, incubated = for 2 h at=20 37=B0C and 10% CO2 in NB27 (0.65 x 106 cells per ml) were added, = and=20 the plate was incubated over night at 37=B0C and 10% = CO2.=20 The inserts were removed and the upper surface was carefully=20 cleansed with cotton pads. Cells on the lower surface were = air=20 dried, counterstained with Harris' hematoxylin for 20 min, = and=20 washed. Stained inserts were placed on object slides and the=20 number of cells on the lower surface assessed at x200 in an inverted = bright-field=20 microscope. Eight x200 fields per = insert=20 were counted. Migration is expressed as a percentage (of = cells=20 per field) of the spontaneous migration toward NB27.

Precursor Cell Differentiation. For differentiation = studies,=20 neurospheres (passage 2=9610) were dissociated to single = cells in=20 trypsin, quenched with 4% BSA, washed with NB27, and seeded = as single=20 cells on 9-mm poly(L)-ornithine (0.01%)-coated glass cover = slips at a=20 density of 35 to 50 x 103 = cells per=20 cm2. Alternatively, cells were seeded, at the same = concentration, in poly(L)-lysine-coated 96-well plates = (Biocat,=20 Becton Dickinson). Cells were grown in conditioned media = [diluted 1:1=20 in NB27 containing 4 ng of EGF/ml (only adult precursor = cells)] or=20 only NB27 with or without LPS (10 ng/ml). All experiments = were=20 performed two to four times with two to four wells per=20 treatment.

Immunocytochemistry. Cover slips or 96 wells were fixed in = 4%=20 paraformaldehyde and washed in PBS. For identification of = neurons,=20 we used Tuj-1 (3D{beta}III) at 1:6,000 and for astrocytes anti-GFAP at = 1:5,000.=20 Microglia was identified with rabbit anti-Iba-1 at 1:500. All = primary=20 Abs were diluted in 0.3% Triton X-100, 5% normal goat serum = (Vector=20 Laboratories) in PBS. Primary Abs were present at 4=B0C = overnight=20 followed by wash in PBS. Secondary reagents were goat Alexa = Fluor 488=20 and 594 (Molecular Probes) both at 1:750, diluted in 0.3% = Triton=20 X-100/PBS and incubated for 2=963 h at room temperature. = Cells were=20 counterstained with DAPI and, in the case of cover slips, = mounted=20 with Moviol (Calbiochem).

Immunoreactivity was examined at x200=20 magnification in a Zeiss microscope, and images from 6=9610 = random=20 fields per coverslip were captured. The numbers of DAPI+ = cells were=20 counted by the CARNOY program (http://www.carnoy.org/), and the = number of=20 neurons and astrocytes were counted manually. All images were = counted=20 blind. Images of immunostained cells in 96-well format were=20 automatically captured in an Image-Xpress (Axon Instruments). = Cells=20 were automatically identified by an algorithm designed to = detect=20 DAPI-stained nuclei and the presence of 3D{beta}=20III or GFAP = immunoreactivity in=20 the cytoplasm immediately surrounding the DAPI nuclei. Five = fields=20 per well from two different wells were captured. The number = of=20 different cells is expressed as percentage of DAPI+ = cells.=20

Statistical Analysis. Data were analyzed and tested for=20 significance between groups by using the Student's t = test=20 assuming equal variance.


3D"   =20 Results
3D"Top
3D"Abstract
Materials and Methods
3D"Results
Discussion
References
 
<= B>Precursor=20 Cell Migration in Matrigel. Cortical aggregate cultures = were=20 established by cultivating mixed mouse CNS cells on a gyratory=20 shaker. When the resulting aggregates (3D{approx}=20500 =B5m in = diameter)=20 were treated with EGF, an outer layer of proliferating cells=20 were formed. They expressed the neuroepithelial marker nestin = while lacking expression of more mature markers for = astrocytes=20 or neurons (Fig. 1=20 A=96C; C. M. D. Berglund, J.A., S.L.B.H., = J. R.=20 Nyengaard, T. H=F6kfelt, K.S., J. N=E4slund, and M.A.A.P., = unpublished=20 data). The identity of these cells as precursor cells was = previously=20 shown by clonal analysis where clones derived from single = cells gave=20 rise to astrocytes, neurons, and oligodendrocytes (C. M. D. = Berglund,=20 J.A., S.L.B.H., J. R. Nyengaard, T. H=F6kfelt, K.S., J. = N=E4slund, and=20 M.A.A.P., unpublished data). Migration of these precursor = cells was=20 readily studied by placing the aggregates in Matrigel. =



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  		Fig. 1.=20 EGF-treated mixed CNS cell aggregates contain neural = precursor=20 cells. (A=96C) Bright-field micrographs = displaying=20 EGF-treated aggregates, with an outer ring area of = nestin-positive=20 precursor cells (A). The outer ring of cells lack the = expression of the neuronal marker 3D{beta}=20III = (B) and=20 the astrocytic marker GFAP (C). Consecutive sections = from the=20 same aggregate are shown. (Scale bar, 50 =B5m.) =

=

After being placed in Matrigel (48 h), cells were migrating = out=20 from the EGF-treated aggregates, both as single cells with = one or two=20 thin processes or as chains of cells (Fig. 2 = A=20 and B). Close to the aggregate surface, the single = cells=20 were frequently observed associated to long, thin processes = extending=20 radially from the aggregate (Fig.=20 2B). Cells with more extended processes and = morphology=20 were observed further away from the aggregates (not shown). = In=20 contrast, aggregates not treated with EGF, and hence lacking = the=20 outer ring of precursor cells, failed to display any = migrating cells=20 when placed in Matrigel (Fig.=20 2C).

When aggregates embedded in Matrigel were fixed after 6 days = and=20 immuno-stained for markers of neurons and astrocytes, most of = the=20 stained migrating cells expressed markers of early neurons = (III), = whereas very=20 few were positive for the astrocytic marker GFAP (Fig. 2 = D=20 and E). However, a large proportion of the migrating = cells=20 were not stained by any of the used markers (Fig. 2=20 F and G).

In initial studies, we observed that if EGF-treated aggregates=20 were placed in Matrigel together with microglial-conditioned=20 media, more cells migrated out from the aggregates than when=20 using NB27 or conditioned media from the astrosarcoma cell = line=20 C6 (C6-CM) (not shown). This suggested that = microglial-conditioned=20 media contained factors that promoted the migration of = precursor=20 cells, either by increasing their mobility (motogen) or by = attracting=20 them (attractant).

To test whether microglia cells express molecules that could=20 direct the migration of precursor cells, EGF-treated = aggregates=20 were embedded in Matrigel close to a defined source of = concentrated=20 microglial-conditioned media. To quantify cell migration, the = area surrounding the aggregates was divided into quadrants, = and=20 the number of cells in the proximal (p, closest to the concentrated = media) and distal (d, opposite side from the concentrated = media)=20 quadrants were determined (see Fig.=20 2H).

Dividing the number of cells in the proximal zone with the = number=20 of cells in the distal zone revealed that there were up to = two=20 times as many cells moving toward conditioned media (CM) from = the mouse microglia cell lines BV-2 or N13 (BV-2/N13-CM), = whereas=20 no such bias was found in the control wells (NB27) (Fig. = 3).=20 A similar relationship as for BV-2/N13-CM and NB27 was found=20 when comparing the migration of cells toward conditioned = media=20 from LPS-stimulated PM (LPS PM-CM) vs. nonstimulated PM = (PM-CM)=20 (Fig.=20 3). After embedding (48 h), migrating cells were observed = up to=20 800 =B5m from the aggregate surface, frequently, in the case = of=20 microglial-conditioned media, migrating on top of the=20 agarose-embedded conditioned media (data not shown).



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  		Fig. 3. = Directed=20 migration of precursor cells in Matrigel toward=20 microglial-conditioned media. Ratio between number of cells = in the=20 proximal quadrant (p) and distal quadrant (d) as described = in Fig.=20 2H. Data are the mean from 12 aggregates per = treatment =B1=20 SEM. *, P < 0.05 by two-tailed t test. =

=

Transwell Migration of Precursor Cells. To confirm that = microglia=20 cells were capable of directing the migration of precursor = cells,=20 we also used the modified Boyden chamber where precursor = cells=20 and conditioned media are separated by a membrane containing=20 pores large enough to allow cell passage. More than three = times=20 as many precursor cells migrated toward BV-2/N13-CM as = compared=20 to controls [NB27 and conditioned media from a astrosarcoma=20 cell line, C6 (C6-CM)] (Fig. = 4).=20 Placing BV-2/N13-CM in both the upper and lower chamber, so = as to=20 remove any potential gradient, decreased the migration of = precursor=20 cells consistent with a chemotactic model of migration (Fig. = 4).=20 Conditioned media from LPS PM-CM or PM-CM were both able to = attract=20 more precursor cells than controls (Fig. = 4).=20



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  		Fig. 4. = Transwell=20 migration of precursor cells in modified Boyden chambers. = Percentage=20 of cells per microscopic field migrating toward conditioned = media,=20 relative to the spontaneous migration toward NB27 (100%).=20 BV-2/N13-CM U/L represent the case where conditioned media = were=20 placed in both the upper (U) and lower (L) chamber. Values = are given=20 as mean =B1 SEM of two to four independent experiments. *, = P=20 < 0.05 by two-tailed t test. =

=

Differentiation of Embryonic Precursor Cells. To test = whether=20 microglia also affected the differentiation of embryonic = precursor=20 cells, BV-2/N13-CM and conditioned media from PM cells were=20 prepared and added to precursor cells. In cultures treated = with=20 BV-2/N13-CM, almost six times more neurons (3D{beta}=20III-positive = cells,=20 3D{beta}III+)=20 were found as compared to control cultures (C6-CM or NB27) = (Fig.=20 5A). Similarly, a higher proportion of neurons were = found=20 in cultures cultivated in LPS PM-CM (Fig.=20 5A). Also, the proportion of astrocytes (GFAP+ = cells) in=20 BV-2/N13-CM and LPS PM-CM were reduced compared to control = cultures=20 (C6-CM and NB27) or precursor cells in nonstimulated PM-CM = (Fig.=20 5B).



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  		Fig. 5. = Increased=20 proportion of neurons from embryonic neural precursor cells = in=20 microglial-conditioned media. (A and B) = Percentages of=20 neurons (3D{beta}III+ cells) (A) and astrocytes (GFAP+ = cells)=20 (B) of DAPI+ cells, after 7 days in conditioned media = from=20 microglial cells or controls. Data are expressed as mean =B1 = SEM of=20 positively stained cells per microscopic field and were = counted=20 blind from two to four independent experiments. ***, = P <=20 0.001; *, P < 0.05 (by two-tailed t test).=20 (C and D). Fluorescence overlay micrographs = showing=20 the morphology of neurons (red) and astrocytes (green) in = BV-2/N13=20 (C) or C6 (D) conditioned media. Cells were=20 counterstained with DAPI (blue). (Scale bar, 50 =B5m.) =

=

We saw a morphological difference between 3D{beta}=20III+ cells in = BV-2/N13-CM=20 or LPS PM-CM and control medium, with most of the 3D{beta}=20III+ cells = in=20 microglial-conditioned media displaying long, frequently = branched=20 processes, whereas the 3D{beta}III+ cells in control (NB27 and C6-CM) had smaller = processes=20 (Fig. = 5=20 C and D). Furthermore, when precursor cells = were=20 treated with BV-2/N13-CM or LPS PM-CM, cells positive for = both GFAP=20 and 3D{beta}III could=20 be found (2=968% of total), whereas such double-positive = cells were=20 never observed in control cultures or cultures in = nonstimulated=20 PM-CM.

Differentiation of Adult Precursor Cells. We also asked = how=20 microglia might affect the differentiation of precursor cells = isolated from adult mice, defined here as older than 20 = weeks.=20 Morphologically, day 7 BV-2/N13-CM and LPS PM-CM-treated = adult=20 precursor cells resembled embryonic precursor cells (compare = Fig. 5 = C=20 and D to Fig. 6 = A=20 and B), although the astrocytes in adult cultures = treated=20 with BV-2/N13-CM or LPS PM-CM frequently projected longer = processes=20 and were not as compact as the ones in nonstimulated PM-CM or = control=20 (Fig. = 6=20 A and B).



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  		Fig. 6.=20 Differentiation of adult neural precursor cells in=20 microglial-conditioned media. (A and B) = Fluorescence=20 overlay micrographs of adult neural precursor cells cultured = for 7=20 days in BV-2/N13-CM (A) or NB27 (B). Neurons = were=20 identified with anti-3D{beta}=20III = (red) and=20 astrocytes with anti-GFAP (green). Cells were counterstained = with=20 DAPI (blue). (Scale bar, 50 =B5m.) (C and D) = Proportions=20 of neurons (C) and astrocytes (D) of DAPI+ = cells in=20 cultures of adult neural precursor cells treated with = conditioned=20 media from microglia or control cells. Bars represent mean = =B1 SEM=20 from two to four independent experiments. **, P < = 0.01; *,=20 P < 0.05 (by two-tailed t test). =

=

The survival rate of adult cells was low, both in control = cultures=20 and in nonstimulated PM-CM-treated cultures (not shown). To=20 increase the survival of adult precursor cells, = differentiating=20 cultures were supplemented with low amounts of EGF (2 ng/ml). = When such cultures were treated with BV-2/N13-CM, >30% of=20 the cells were positive for the neuronal marker 3D{beta}=20III after 7 = days,=20 whereas significantly less neurons were found in control = cultures (Fig.=20 6C). There was no significant difference in the = proportion=20 of astrocytes among the treatments, although somewhat less = astrocytes=20 were observed in LPS PM-CM (Fig.=20 6D). As observed in cultures of embryonic = precursor cells,=20 adult cultures treated with BV-2/N13-CM or LPS PM-CM had a = small=20 fraction of double-positive cells (1=963% of total, not = shown).=20


3D"   =20 Discussion
3D"Top
3D"Abstract
Materials and Methods
Results
3D"Discussion
References
 
I= n the=20 present paper, we have examined how mouse microglia cells = affect the=20 migration and differentiation of neural precursor cells. We = report=20 that precursor cells isolated from the embryonic brain = migrated=20 toward a gradient of microglial-conditioned media. In = addition,=20 soluble factors from microglia influenced the differentiation = of=20 precursor cells so that the proportion of neurons was increased = in=20 cultures treated with microglial-conditioned media.

Directed migration of precursor cells was observed both with=20 precursor cells emanating from aggregate cultures embedded in = a=20 protein matrix, as well as with purified precursor cells in a = modified Boyden chamber. In both experimental systems precursor = cells=20 preferentially migrated toward microglial-conditioned media, = clearly=20 showing that microglia have the capacity to direct the = migration of=20 precursor cells. Confirming this is our finding that, when = purified=20 precursor cells were placed in the modified Boyden chamber = with=20 microglial-conditioned media in both the upper and lower = chamber, the=20 migration was decreased, indicating that the observed = directed=20 migration was not merely a result of increased motility of = the=20 precursor cells (Fig. = 4).=20

The cellular identity of migrating cells remains to be firmly=20 established, although the presence of nestin immune = reactivity=20 and the lack of detectable expression of neuronal or = astrocytic=20 markers in the proliferating cells constituting the outer rim = of the EGF-treated aggregates, makes it highly probable that=20 these cells are true, uncommitted neural precursor cells. = Such=20 an outer rim of cells was not seen in non-EGF-treated = aggregate=20 cultures, nor did we see any migration from such aggregates = (Fig. = 2).=20 Moreover, the cells migrating out from EGF-treated aggregates = did=20 acquire neuronal, and to lesser extent, astrocytic markers = only when=20 the EGF had been removed. Taken together, we reason that the=20 migration seen from the aggregates was mainly derived from=20 uncommitted neural precursor cells. In addition, in the = experiments=20 performed in the modified Boyden chamber, purified precursor = cells=20 were used, and the results corroborated the ones from the = Matrigel=20 assay. However, we cannot at present rule out the possibility = that=20 microglial factors attract a specific population of precursor = cells.

Studying microglial effects on differentiation, we found to = our=20 surprise that precursor cell cultures, grown in conditioned = media=20 from microglia cells, contained a higher proportion of = neurons than=20 would be expected from their spontaneous differentiation = alone. This=20 effect was observed on precursor cells isolated from both the = embryonic as well as the adult brain, with the strongest = microglial=20 effect on embryonic precursor cells. For adult precursor = cells, such=20 differentiation was readily induced by conditioned media from = the=20 microglia cell lines, whereas we did not observe any = increased=20 proportion of neurons in cultures treated with conditioned = media from=20 PM cells, even after LPS activation. Possibly, LPS activation = may be=20 a rather artificial way of activating microglia compared to = the=20 conditions in CNS in vivo. Accordingly, more = physiological=20 activation strategies may provide other results. In this = regard, we=20 note the consistent and highly significant effect of the = microglia=20 cells lines BV-2 and N13.

There are at least three different mechanisms that could be = the=20 basis for the increased proportion of neurons in microglial-treated = cultures. First, a purely instructive role, inducing the = precursor=20 cells to be committed for a neuronal fate. Secondly, = microglia=20 may selectively promote the survival of neuronal cells e.g., = by=20 providing neurotrophins. Finally, microglia could produce = factors=20 that are toxic to astrocytes. In preliminary experiments on = adult=20 precursor cells, we have not seen any significant difference = in the=20 proportion of terminal deoxynucleotidyltransferase-mediated = dUTP nick=20 end labeling-positive cells (apoptosis and necrosis) between = the=20 treatments (not shown), arguing against but not disproving = that=20 microglial factors increase the survival of neuronal cells. = If=20 microglia released factors were toxic to astrocytes, we would = expect=20 a more pronounced difference in the proportion of astrocytes = between=20 microglial- and control-treated cultures, a difference we do = not=20 observe (see Figs. 5 = and 6).=20 Accordingly, we currently favor an interpretation with an = instructive=20 role of the factors. Still, we would like to emphasize that = the=20 mechanism(s) responsible for the increased proportion of = neurons in=20 microglial-treated cultures remains to be formally = proven.=20

The observation of cells stained positive for both astrocytic=20 (GFAP) and neuronal (3D{beta}=20III) markers may be = explained=20 by recent findings showing that many precursor cells, = although=20 displaying all of the characteristics of neural stem cells, = express=20 GFAP (28, = 29).= =20

It is tempting to speculate that different pathological = processes=20 in the CNS may induce distinct modes of microglial activation = which may result in different migration and differentiation=20 patterns of precursor cells. For instance, several recent = studies=20 have shown that precursor cells preferentially migrate to = sites=20 of inflammation in animal models of multiple sclerosis (MS),=20 and that these new cells preferentially differentiate into=20 oligodendrocytes (19, = 20, = 30).= =20 Contrary, in experimental models of more acute damages with = neuronal=20 loss, precursor cells, both extrinsically provided as well as = endogenous precursor cells, migrated to the damaged area and=20 differentiated to neurons (7, = 10=96= 14).= =20 Microglia activation at the site of injury is firmly = established=20 in all of these experimental lesions (31, = 32).= =20

During embryonic development fetal macrophages/microglia are=20 detected in the CNS around E8 in mice and E12 in rat (33, = 34),= =20 a time-point well before the end of neurogenesis but before = the=20 onset of gliogenesis. The role of microglia cells during CNS=20 development is generally believed to be to remove dead cells = by=20 phagocytosis. However, the strong effects of microglia cells = on=20 embryonic precursor cells, as shown here, may point to other=20 developmental roles of microglia then just to remove dead = cells=20 and debris.

In conclusion, despite the apparent imbalance between = degeneration=20 and renewal in the CNS, several recent reports have shown = migration=20 and regeneration of neural cells to sites of injury or other=20 pathological processes in the CNS (7, = 10, = 13, = 14, = 19, = 20, = 30).= =20 Microglia cells are invariably activated at such locations. = The=20 present report may add to these observations because it = indicates=20 that microglia have the capacity to direct the migration of = neural=20 precursor cells as well as effect their differentiation. A = role for=20 activated microglia in the attraction as well as = differentiation of=20 precursor cells may well represent a more general route for = cell=20 replacement in the CNS, both during development and in = disease=20 states.


3D"   =20 Acknowledgements
 
We are = grateful to=20 Niklas Holmstr=F6m for technical advice and Jonas Fris=E9n, = Jan N=E4slund,=20 and Darius Matusevicius for fruitful discussions. The cell = line N13=20 was kindly donated by Paola Ricciardi-Castagnoli, and Abs to = Iba1=20 were provided by Yoshinori Imai. This work was supported by = the=20 Swedish Cancer Society, the Foundation for Strategic Research = (NNN-program), and AstraZeneca R&D.


3D"   =20 Footnotes
 
Abbreviations: 3D{beta}=20III, 3D{beta}=20 class III tubulin; = CM,=20 conditioned medium or media; DAPI, 4', = 6-diamidino-2-phenylindole;=20 EGF, epidermal growth factor; En, embryonic day = n;=20 GFAP, glia fibrillary acidic protein; LPS, = lipopolysaccharide; NB27,=20 Neurobasal medium supplemented with B27; PM, primary = microglia.=20

3D{ddagger}=20 To whom = correspondence=20 should be addressed. E-mail: mats.persson{at}cmm.ki.se .=20


3D"   =20 References
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Results
Discussion
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Opera = requires=0A= the Sun Java Runtime Environment .=0A= =0A= by Andrew Gregory=0A= http://www.scss.com.au/family/andrew/webdesign/xmlhttprequest/=0A= =0A= This work is licensed under the Creative Commons Attribution License. To = view a=0A= copy of this license, visit = http://creativecommons.org/licenses/by-sa/2.5/ or=0A= send a letter to Creative Commons, 559 Nathan Abbott Way, Stanford, = California=0A= 94305, USA.=0A= =0A= Attribution: Leave my name and web address in this script intact.=0A= =0A= Not Supported in Opera=0A= ----------------------=0A= * user/password authentication=0A= * responseXML data member=0A= =0A= Not Fully Supported in Opera=0A= ----------------------------=0A= * async requests=0A= * abort()=0A= * getAllResponseHeaders(), getAllResponseHeader(header)=0A= =0A= */=0A= // IE support=0A= if (window.ActiveXObject && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= var msxmls =3D new Array(=0A= 'Msxml2.XMLHTTP.5.0',=0A= 'Msxml2.XMLHTTP.4.0',=0A= 'Msxml2.XMLHTTP.3.0',=0A= 'Msxml2.XMLHTTP',=0A= 'Microsoft.XMLHTTP');=0A= for (var i =3D 0; i < msxmls.length; i++) {=0A= try {=0A= return new ActiveXObject(msxmls[i]);=0A= } catch (e) {=0A= }=0A= }=0A= return null;=0A= };=0A= }=0A= // Gecko support=0A= /* ;-) */=0A= // Opera support=0A= if (window.opera && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= this.readyState =3D 0; // = 0=3Duninitialized,1=3Dloading,2=3Dloaded,3=3Dinteractive,4=3Dcomplete=0A= this.status =3D 0; // HTTP status codes=0A= this.statusText =3D '';=0A= this._headers =3D [];=0A= this._aborted =3D false;=0A= this._async =3D true;=0A= this._defaultCharset =3D 'ISO-8859-1';=0A= this._getCharset =3D function() {=0A= var charset =3D _defaultCharset;=0A= var contentType =3D = this.getResponseHeader('Content-type').toUpperCase();=0A= val =3D contentType.indexOf('CHARSET=3D');=0A= if (val !=3D -1) {=0A= charset =3D contentType.substring(val);=0A= }=0A= val =3D charset.indexOf(';');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= val =3D charset.indexOf(',');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= return charset;=0A= };=0A= this.abort =3D function() {=0A= this._aborted =3D true;=0A= };=0A= this.getAllResponseHeaders =3D function() {=0A= return this.getAllResponseHeader('*');=0A= };=0A= this.getAllResponseHeader =3D function(header) {=0A= var ret =3D '';=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= if (header =3D=3D '*' || this._headers[i].h =3D=3D header) {=0A= ret +=3D this._headers[i].h + ': ' + this._headers[i].v + '\n';=0A= }=0A= }=0A= return ret;=0A= };=0A= this.getResponseHeader =3D function(header) {=0A= var ret =3D getAllResponseHeader(header);=0A= var i =3D ret.indexOf('\n');=0A= if (i !=3D -1) {=0A= ret =3D ret.substring(0, i);=0A= }=0A= return ret;=0A= };=0A= this.setRequestHeader =3D function(header, value) {=0A= this._headers[this._headers.length] =3D {h:header, v:value};=0A= };=0A= this.open =3D function(method, url, async, user, password) {=0A= this.method =3D method;=0A= this.url =3D url;=0A= this._async =3D true;=0A= this._aborted =3D false;=0A= this._headers =3D [];=0A= if (arguments.length >=3D 3) {=0A= this._async =3D async;=0A= }=0A= if (arguments.length > 3) {=0A= opera.postError('XMLHttpRequest.open() - user/password not = supported');=0A= }=0A= this.readyState =3D 1;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= };=0A= this.send =3D function(data) {=0A= if (!navigator.javaEnabled()) {=0A= alert("XMLHttpRequest.send() - Java must be installed and = enabled.");=0A= return;=0A= }=0A= if (this._async) {=0A= setTimeout(this._sendasync, 0, this, data);=0A= // this is not really asynchronous and won't execute until the = current=0A= // execution context ends=0A= } else {=0A= this._sendsync(data);=0A= }=0A= }=0A= this._sendasync =3D function(req, data) {=0A= if (!req._aborted) {=0A= req._sendsync(data);=0A= }=0A= };=0A= this._sendsync =3D function(data) {=0A= this.readyState =3D 2;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= // open connection=0A= var url =3D new java.net.URL(new = java.net.URL(window.location.href), this.url);=0A= var conn =3D url.openConnection();=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= conn.setRequestProperty(this._headers[i].h, this._headers[i].v);=0A= }=0A= this._headers =3D [];=0A= if (this.method =3D=3D 'POST') {=0A= // POST data=0A= conn.setDoOutput(true);=0A= var wr =3D new = java.io.OutputStreamWriter(conn.getOutputStream(), this._getCharset());=0A= wr.write(data);=0A= wr.flush();=0A= wr.close();=0A= }=0A= // read response headers=0A= // NOTE: the getHeaderField() methods always return nulls for me :(=0A= var gotContentEncoding =3D false;=0A= var gotContentLength =3D false;=0A= var gotContentType =3D false;=0A= var gotDate =3D false;=0A= var gotExpiration =3D false;=0A= var gotLastModified =3D false;=0A= for (var i =3D 0; ; i++) {=0A= var hdrName =3D conn.getHeaderFieldKey(i);=0A= var hdrValue =3D conn.getHeaderField(i);=0A= if (hdrName =3D=3D null && hdrValue =3D=3D null) {=0A= break;=0A= }=0A= if (hdrName !=3D null) {=0A= this._headers[this._headers.length] =3D {h:hdrName, = v:hdrValue};=0A= switch (hdrName.toLowerCase()) {=0A= case 'content-encoding': gotContentEncoding =3D true; break;=0A= case 'content-length' : gotContentLength =3D true; break;=0A= case 'content-type' : gotContentType =3D true; break;=0A= case 'date' : gotDate =3D true; break;=0A= case 'expires' : gotExpiration =3D true; break;=0A= case 'last-modified' : gotLastModified =3D true; break;=0A= }=0A= }=0A= }=0A= // try to fill in any missing header information=0A= var val;=0A= val =3D conn.getContentEncoding();=0A= if (val !=3D null && !gotContentEncoding) = this._headers[this._headers.length] =3D {h:'Content-encoding', v:val};=0A= val =3D conn.getContentLength();=0A= if (val !=3D -1 && !gotContentLength) = this._headers[this._headers.length] =3D {h:'Content-length', v:val};=0A= val =3D conn.getContentType();=0A= if (val !=3D null && !gotContentType) = this._headers[this._headers.length] =3D {h:'Content-type', v:val};=0A= val =3D conn.getDate();=0A= if (val !=3D 0 && !gotDate) this._headers[this._headers.length] = =3D {h:'Date', v:(new Date(val)).toUTCString()};=0A= val =3D conn.getExpiration();=0A= if (val !=3D 0 && !gotExpiration) = this._headers[this._headers.length] =3D {h:'Expires', v:(new = Date(val)).toUTCString()};=0A= val =3D conn.getLastModified();=0A= if (val !=3D 0 && !gotLastModified) = this._headers[this._headers.length] =3D {h:'Last-modified', v:(new = Date(val)).toUTCString()};=0A= // read response data=0A= var reqdata =3D '';=0A= var stream =3D conn.getInputStream();=0A= if (stream) {=0A= var reader =3D new java.io.BufferedReader(new = java.io.InputStreamReader(stream, this._getCharset()));=0A= var line;=0A= while ((line =3D reader.readLine()) !=3D null) {=0A= if (this.readyState =3D=3D 2) {=0A= this.readyState =3D 3;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= }=0A= reqdata +=3D line + '\n';=0A= }=0A= reader.close();=0A= this.status =3D 200;=0A= this.statusText =3D 'OK';=0A= this.responseText =3D reqdata;=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onload) {=0A= this.onload();=0A= }=0A= } else {=0A= // error=0A= this.status =3D 404;=0A= this.statusText =3D 'Not Found';=0A= this.responseText =3D '';=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onerror) {=0A= this.onerror();=0A= }=0A= }=0A= };=0A= };=0A= }=0A= // ActiveXObject emulation=0A= if (!window.ActiveXObject && window.XMLHttpRequest) {=0A= window.ActiveXObject =3D function(type) {=0A= switch (type.toLowerCase()) {=0A= case 'microsoft.xmlhttp':=0A= case 'msxml2.xmlhttp':=0A= case 'msxml2.xmlhttp.3.0':=0A= case 'msxml2.xmlhttp.4.0':=0A= case 'msxml2.xmlhttp.5.0':=0A= return new XMLHttpRequest();=0A= }=0A= return null;=0A= };=0A= }=0A= ------=_NextPart_000_0000_01C85936.F43A32E0 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.pnas.org/javascript/ajax/utility.js /************************************************************************= *****=0A= * javascript/ajax/utility.js=0A= *=0A= * Utility functions for working with XMLHttpRequest data.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Copy XML nodes into an HTMLElement. This effectively=0A= * clones XML markup which uses XHTML naming conventions=0A= * into an HTML DOM.=0A= */=0A= function copy_xml_to_html(src, dst) {=0A= if (src.nodeType =3D=3D 1) { /* Node.ELEMENT_NODE */=0A= var e =3D document.createElement(src.nodeName);=0A= for (var i =3D 0; i < src.childNodes.length; i++) {=0A= copy_xml_to_html(src.childNodes[i], e);=0A= }=0A= for (var i =3D 0; i < src.attributes.length; i++) {=0A= var n =3D src.attributes[i].name;=0A= var v =3D unescape_xml_string(src.attributes[i].value); =0A= e.setAttribute(n, v);=0A= if (n =3D=3D "class") {=0A= e.className =3D v;=0A= }=0A= else if (n =3D=3D "style") {=0A= set_css_style(v, e, "");=0A= }=0A= }=0A= dst.appendChild(e);=0A= }=0A= else if (src.nodeType =3D=3D 3) { /* Node.TEXT_NODE */=0A= dst.appendChild(document.createTextNode(src.nodeValue));=0A= }=0A= }=0A= =0A= /* =0A= * It is unclear that this is the right thing to be calling=0A= * from copy_xml_to_html, but it appears that Safari decides=0A= * to convert & to the NCR #, and then encodes that=0A= * NCR to &%26%2338;. So, I'm going to treat the DOM Attr=0A= * value as a plain string, and run our XML string input=0A= * through the decoding routine below.=0A= */=0A= function unescape_xml_string(s) {=0A= return s.replace(/'/g, "'")=0A= .replace(/'/g, "'")=0A= .replace(/"/g, "\"")=0A= .replace(/"/g, "\"")=0A= .replace(/>/g, ">")=0A= .replace(/>/g, ">")=0A= .replace(/</g, "<")=0A= .replace(/</g, "<")=0A= .replace(/&/g, "&")=0A= .replace(/&/g, "&");=0A= }=0A= =0A= /*=0A= * Parse set of CSS rules and apply them to an element.=0A= * This is quite horrifying, but I'm unable to determine=0A= * how else to handle this with IE 6. FireFox and other=0A= * sane browsers let you simply set the style attribute=0A= * or use e.style.setProperty(rule, value, priority),=0A= * IE 6 appears to have neither of these capabilities..=0A= */=0A= function set_css_style(css, e, priority) {=0A= var rules =3D css.split(";");=0A= for (var i =3D 0; i < rules.length; i++) {=0A= var nvpair =3D rules[i].split(":");=0A= if (nvpair.length =3D=3D 2) {=0A= try {=0A= var name =3D nvpair[0]; /* style attribute */=0A= var value =3D nvpair[1]; /* attribute value */=0A= =0A= /*=0A= * For each possible style attribute, set the=0A= * appropriate style property in the element.=0A= */=0A= if (name =3D=3D "background") {=0A= e.style.background =3D value;=0A= }=0A= else if (name =3D=3D "background-attachment") {=0A= e.style.backgroundAttachment =3D value;=0A= }=0A= else if (name =3D=3D "background-color") {=0A= e.style.backgroundColor =3D value;=0A= }=0A= else if (name =3D=3D "background-image") {=0A= e.style.backgroundImage =3D value;=0A= }=0A= else if (name =3D=3D "background-position") {=0A= e.style.backgroundPosition =3D value;=0A= }=0A= else if (name =3D=3D "background-position-x") {=0A= e.style.backgroundPositionX =3D value;=0A= }=0A= else if (name =3D=3D "background-position-y") {=0A= e.style.backgroundPositionY =3D value;=0A= }=0A= else if (name =3D=3D "background-repeat") {=0A= e.style.backgroundRepeat =3D value;=0A= }=0A= else if (name =3D=3D "behavior") {=0A= e.style.behavior =3D value;=0A= }=0A= else if (name =3D=3D "border") {=0A= e.style.border =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom") {=0A= e.style.borderBottom =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-color") {=0A= e.style.borderBottomColor =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-style") {=0A= e.style.borderBottomStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-width") {=0A= e.style.borderBottomWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-collapse") {=0A= e.style.borderCollapse =3D value;=0A= }=0A= else if (name =3D=3D "border-color") {=0A= e.style.borderColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left") {=0A= e.style.borderLeft =3D value;=0A= }=0A= else if (name =3D=3D "border-left-color") {=0A= e.style.borderLeftColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left-style") {=0A= e.style.borderLeftStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-left-width") {=0A= e.style.borderLeftWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-right") {=0A= e.style.borderRight =3D value;=0A= }=0A= else if (name =3D=3D "border-right-color") {=0A= e.style.borderRightColor =3D value;=0A= }=0A= else if (name =3D=3D "border-right-style") {=0A= e.style.borderRightStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-right-width") {=0A= e.style.borderRightWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-style") {=0A= e.style.borderStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top") {=0A= e.style.borderTop =3D value;=0A= }=0A= else if (name =3D=3D "border-top-color") {=0A= e.style.borderTopColor =3D value;=0A= }=0A= else if (name =3D=3D "border-top-style") {=0A= e.style.borderTopStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top-width") {=0A= e.style.borderTopWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-width") {=0A= e.style.borderWidth =3D value;=0A= }=0A= else if (name =3D=3D "bottom") {=0A= e.style.bottom =3D value;=0A= }=0A= else if (name =3D=3D "clear") {=0A= e.style.clear =3D value;=0A= }=0A= else if (name =3D=3D "clip") {=0A= e.style.clip =3D value;=0A= }=0A= else if (name =3D=3D "color") {=0A= e.style.color =3D value;=0A= }=0A= else if (name =3D=3D "cssText") {=0A= e.style.Sets =3D value;=0A= }=0A= else if (name =3D=3D "cursor") {=0A= e.style.cursor =3D value;=0A= }=0A= else if (name =3D=3D "direction") {=0A= e.style.direction =3D value;=0A= }=0A= else if (name =3D=3D "display") {=0A= e.style.display =3D value;=0A= }=0A= else if (name =3D=3D "font") {=0A= e.style.font =3D value;=0A= }=0A= else if (name =3D=3D "font-family") {=0A= e.style.fontFamily =3D value;=0A= }=0A= else if (name =3D=3D "font-size") {=0A= e.style.fontSize =3D value;=0A= }=0A= else if (name =3D=3D "font-style") {=0A= e.style.fontStyle =3D value;=0A= }=0A= else if (name =3D=3D "font-variant") {=0A= e.style.fontVariant =3D value;=0A= }=0A= else if (name =3D=3D "font-weight") {=0A= e.style.fontWeight =3D value;=0A= }=0A= else if (name =3D=3D "height") {=0A= e.style.height =3D value;=0A= }=0A= else if (name =3D=3D "ime-mode") {=0A= e.style.imeMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-flow") {=0A= e.style.layoutFlow =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid") {=0A= e.style.layoutGrid =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-char") {=0A= e.style.layoutGridChar =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-line") {=0A= e.style.layoutGridLine =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-mode") {=0A= e.style.layoutGridMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-type") {=0A= e.style.layoutGridType =3D value;=0A= }=0A= else if (name =3D=3D "left") {=0A= e.style.left =3D value;=0A= }=0A= else if (name =3D=3D "letter-spacing") {=0A= e.style.letterSpacing =3D value;=0A= }=0A= else if (name =3D=3D "line-break") {=0A= e.style.lineBreak =3D value;=0A= }=0A= else if (name =3D=3D "line-height") {=0A= e.style.lineHeight =3D value;=0A= }=0A= else if (name =3D=3D "list-style") {=0A= e.style.listStyle =3D value;=0A= }=0A= else if (name =3D=3D "list-style-image") {=0A= e.style.listStyleImage =3D value;=0A= }=0A= else if (name =3D=3D "list-style-position") {=0A= e.style.listStylePosition =3D value;=0A= }=0A= else if (name =3D=3D "list-style-type") {=0A= e.style.listStyleType =3D value;=0A= }=0A= else if (name =3D=3D "margin") {=0A= e.style.margin =3D value;=0A= }=0A= else if (name =3D=3D "margin-bottom") {=0A= e.style.marginBottom =3D value;=0A= }=0A= else if (name =3D=3D "margin-left") {=0A= e.style.marginLeft =3D value;=0A= }=0A= else if (name =3D=3D "margin-right") {=0A= e.style.marginRight =3D value;=0A= }=0A= else if (name =3D=3D "margin-top") {=0A= e.style.marginTop =3D value;=0A= }=0A= else if (name =3D=3D "min-height") {=0A= e.style.minHeight =3D value;=0A= }=0A= else if (name =3D=3D "overflow") {=0A= e.style.overflow =3D value;=0A= }=0A= else if (name =3D=3D "overflow-x") {=0A= e.style.overflowX =3D value;=0A= }=0A= else if (name =3D=3D "overflow-y") {=0A= e.style.overflowY =3D value;=0A= }=0A= else if (name =3D=3D "padding") {=0A= e.style.padding =3D value;=0A= }=0A= else if (name =3D=3D "padding-bottom") {=0A= e.style.paddingBottom =3D value;=0A= }=0A= else if (name =3D=3D "padding-left") {=0A= e.style.paddingLeft =3D value;=0A= }=0A= else if (name =3D=3D "padding-right") {=0A= e.style.paddingRight =3D value;=0A= }=0A= else if (name =3D=3D "padding-top") {=0A= e.style.paddingTop =3D value;=0A= }=0A= else if (name =3D=3D "page-break-after") {=0A= e.style.pageBreakAfter =3D value;=0A= }=0A= else if (name =3D=3D "page-break-before") {=0A= e.style.pageBreakBefore =3D value;=0A= }=0A= else if (name =3D=3D "pixelBottom") {=0A= e.style.pixelBottom =3D value;=0A= }=0A= else if (name =3D=3D "pixelHeight") {=0A= e.style.pixelHeight =3D value;=0A= }=0A= else if (name =3D=3D "pixelLeft") {=0A= e.style.pixelLeft =3D value;=0A= }=0A= else if (name =3D=3D "pixelRight") {=0A= e.style.pixelRight =3D value;=0A= }=0A= else if (name =3D=3D "pixelTop") {=0A= e.style.pixelTop =3D value;=0A= }=0A= else if (name =3D=3D "pixelWidth") {=0A= e.style.pixelWidth =3D value;=0A= }=0A= else if (name =3D=3D "posBottom") {=0A= e.style.posBottom =3D value;=0A= }=0A= else if (name =3D=3D "posHeight") {=0A= e.style.posHeight =3D value;=0A= }=0A= else if (name =3D=3D "position") {=0A= e.style.position =3D value;=0A= }=0A= else if (name =3D=3D "posLeft") {=0A= e.style.posLeft =3D value;=0A= }=0A= else if (name =3D=3D "posRight") {=0A= e.style.posRight =3D value;=0A= }=0A= else if (name =3D=3D "posTop") {=0A= e.style.posTop =3D value;=0A= }=0A= else if (name =3D=3D "posWidth") {=0A= e.style.posWidth =3D value;=0A= }=0A= else if (name =3D=3D "right") {=0A= e.style.right =3D value;=0A= }=0A= else if (name =3D=3D "ruby-align") {=0A= e.style.rubyAlign =3D value;=0A= }=0A= else if (name =3D=3D "ruby-overhang") {=0A= e.style.rubyOverhang =3D value;=0A= }=0A= else if (name =3D=3D "ruby-position") {=0A= e.style.rubyPosition =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-3dlight-color") {=0A= e.style.scrollbar3dLightColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-arrow-color") {=0A= e.style.scrollbarArrowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-base-color") {=0A= e.style.scrollbarBaseColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-darkshadow-color") {=0A= e.style.scrollbarDarkShadowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-face-color") {=0A= e.style.scrollbarFaceColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-highlight-color") {=0A= e.style.scrollbarHighlightColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-shadow-color") {=0A= e.style.scrollbarShadowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-track-color") {=0A= e.style.scrollbarTrackColor =3D value;=0A= }=0A= else if (name =3D=3D "float") {=0A= e.style.styleFloat =3D value;=0A= }=0A= else if (name =3D=3D "table-layout") {=0A= e.style.tableLayout =3D value;=0A= }=0A= else if (name =3D=3D "text-align") {=0A= e.style.textAlign =3D value;=0A= }=0A= else if (name =3D=3D "text-align-last") {=0A= e.style.textAlignLast =3D value;=0A= }=0A= else if (name =3D=3D "text-autospace") {=0A= e.style.textAutospace =3D value;=0A= }=0A= else if (name =3D=3D "text-decoration") {=0A= e.style.textDecoration =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationBlink") {=0A= e.style.textDecorationBlink =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationLineThrough") {=0A= e.style.textDecorationLineThrough =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationNone") {=0A= e.style.textDecorationNone =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationOverline") {=0A= e.style.textDecorationOverline =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationUnderline") {=0A= e.style.textDecorationUnderline =3D value;=0A= }=0A= else if (name =3D=3D "text-indent") {=0A= e.style.textIndent =3D value;=0A= }=0A= else if (name =3D=3D "text-justify") {=0A= e.style.textJustify =3D value;=0A= }=0A= else if (name =3D=3D "text-kashida-space") {=0A= e.style.textKashidaSpace =3D value;=0A= }=0A= else if (name =3D=3D "text-overflow") {=0A= e.style.textOverflow =3D value;=0A= }=0A= else if (name =3D=3D "text-transform") {=0A= e.style.textTransform =3D value;=0A= }=0A= else if (name =3D=3D "text-underline-position") {=0A= e.style.textUnderlinePosition =3D value;=0A= }=0A= else if (name =3D=3D "top") {=0A= e.style.top =3D value;=0A= }=0A= else if (name =3D=3D "unicode-bidi") {=0A= e.style.unicodeBidi =3D value;=0A= }=0A= else if (name =3D=3D "vertical-align") {=0A= e.style.verticalAlign =3D value;=0A= }=0A= else if (name =3D=3D "visibility") {=0A= e.style.visibility =3D value;=0A= }=0A= else if (name =3D=3D "white-space") {=0A= e.style.whiteSpace =3D value;=0A= }=0A= else if (name =3D=3D "width") {=0A= e.style.width =3D value;=0A= }=0A= else if (name =3D=3D "word-break") {=0A= e.style.wordBreak =3D value;=0A= }=0A= else if (name =3D=3D "word-spacing") {=0A= e.style.wordSpacing =3D value;=0A= }=0A= else if (name =3D=3D "word-wrap") {=0A= e.style.wordWrap =3D value;=0A= }=0A= else if (name =3D=3D "writing-mode") {=0A= e.style.writingMode =3D value;=0A= }=0A= else if (name =3D=3D "z-index") {=0A= e.style.zIndex =3D value;=0A= }=0A= else if (name =3D=3D "zoom") {=0A= e.style.zoom =3D value;=0A= }=0A= }=0A= catch (e) {=0A= /* ignore error on attempt to set e.style.[property] */=0A= }=0A= }=0A= }=0A= }=0A= ------=_NextPart_000_0000_01C85936.F43A32E0 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.pnas.org/javascript/entrez/callback.js /************************************************************************= *****=0A= * javascript/entrez/callback.js=0A= *=0A= * Entrez Linking callback to populate content box.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Execute callback to fill content box with Entrez Linking information.=0A= */=0A= function entrez_callback(pmid, callback_url) {=0A= /*=0A= * MSIE 5.5 and below have issues with the JavaScript=0A= * used for Entrez Linking. 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i < src.childNodes.length; i++) {=0A= copy_xml_to_html(src.childNodes[i], dst);=0A= }=0A= var tbl =3D tr.parentNode;=0A= tbl.replaceChild(dst, tr);=0A= }=0A= }=0A= req.open('GET', callback_url, true);=0A= req.send(null);=0A= }=0A= ------=_NextPart_000_0000_01C85936.F43A32E0 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www.google-analytics.com/urchin.js //-- Google Analytics Urchin Module=0A= //-- Copyright 2007 Google, All Rights Reserved.=0A= =0A= //-- Urchin On Demand Settings ONLY=0A= var _uacct=3D""; // set up the Urchin Account=0A= var _userv=3D1; // service mode (0=3Dlocal,1=3Dremote,2=3Dboth)=0A= =0A= //-- UTM User Settings=0A= var _ufsc=3D1; // set client info flag (1=3Don|0=3Doff)=0A= var _udn=3D"auto"; // (auto|none|domain) set the domain name for cookies=0A= var _uhash=3D"on"; // (on|off) unique domain hash for cookies=0A= var _utimeout=3D"1800"; // set the inactive session timeout in seconds=0A= var _ugifpath=3D"/__utm.gif"; 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