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An Intermediate Level of BMP Signaling Directly
Specifies Cranial Neural Crest Progenitor Cells in
Jennifer A. Schumacher.¤, Megumi Hashiguchi., Vu H. Nguyen., Mary C. Mullins*
Department of Cell and Developmental Biology, University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania, United States of America
The specification of the neural crest progenitor cell (NCPC) population in the early vertebrate embryo requires an elaborate
network of signaling pathways, one of which is the Bone Morphogenetic Protein (BMP) pathway. Based on alterations in
neural crest gene expression in zebrafish BMP pathway component mutants, we previously proposed a model in which the
gastrula BMP morphogen gradient establishes an intermediate level of BMP activity establishing the future NCPC domain.
Here, we tested this model and show that an intermediate level of BMP signaling acts directly to specify the NCPC. We
quantified the effects of reducing BMP signaling on the number of neural crest cells and show that neural crest cells are
significantly increased when BMP signaling is reduced and that this increase is not due to an increase in cell proliferation. In
contrast, when BMP signaling is eliminated, NCPC fail to be specified. We modulated BMP signaling levels in BMP pathway
mutants with expanded or no NCPCs to demonstrate that an intermediate level of BMP signaling specifies the NCPC. We
further investigated the ability of Smad5 to act in a graded fashion by injecting smad5 antisense morpholinos and show that
increasing doses first expand the NCPCs and then cause a loss of NCPCs, consistent with Smad5 acting directly in neural
crest progenitor specification. Using Western blot analysis, we show that P-Smad5 levels are dose-dependently reduced in
smad5 morphants, consistent with an intermediate level of BMP signaling acting through Smad5 to specify the neural crest
progenitors. Finally, we performed chimeric analysis to demonstrate for the first time that BMP signal reception is required
directly by NCPCs for their specification. Together these results add substantial evidence to a model in which graded BMP
signaling acts as a morphogen to pattern the ectoderm, with an intermediate level acting in neural crest specification.
Citation: Schumacher JA, Hashiguchi M, Nguyen VH, Mullins MC (2011) An Intermediate Level of BMP Signaling Directly Specifies Cranial Neural Crest Progenitor
Cells in Zebrafish. PLoS ONE 6(11): e27403. doi:10.1371/journal.pone.0027403
Editor: Bruce Riley, Texas A&M University, United States of America
Received September 16, 2011; Accepted October 16, 2011; Published November 15, 2011
Copyright: ? 2011 Schumacher et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: This work was supported by an National Institutes of Health (NIH) grant to MCM (GM56326), NIH training grants (T32 HD0075165 and T32 GM07229-28)
to JAS, and an NIH predoctoral fellowship to VHN. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the
Competing Interests: The authors have declared that no competing interests exist.
* E-mail: email@example.com
. These authors contributed equally to this work.
¤ Current address: Division of Developmental Biology, Cincinnati Children's Hospital Medical Center, Cincinnati, Ohio, United States of America
Neural crest cells are a multipotent population derived from
embryonic ectoderm. During neurulation neural crest cells
undergo an epithelial-to-mesenchymal transition, delaminate from
the dorsal neural tube, and migrate throughout the embryo,
contributing to a variety of tissues including craniofacial skeleton,
pigment cells, and the peripheral nervous system. In both frog and
chick, juxtaposition of explanted neural and non-neural ectoderm
gives rise to neural crest cells [1–3]. Consistent with this neural-
non-neural tissue interaction generating the neural crest, the
zebrafish fate map reveals that neural crest cells are derived from
lateral regions of the gastrula, where prospective neural tissue
meets prospective epidermis [4]. Similarly in Xenopus, fate map
analysis reveals that the prospective neural crest population lies
adjacent to the dorsolateral marginal zone at an early gastrula
stage [5]. Consistent with these studies, the earliest genes
specifically expressed within the neural crest progenitor cells
(NCPC), e.g. snail, AP2, and foxd3, are localized to lateral regions of
the neural plate adjacent to the non-neural ectoderm [6–12].
Gain-of-function studies in chick and Xenopus have addressed
the molecular nature of the signals that are involved in the
induction of the neural crest. These studies have implicated Bone
Morphogenetic Protein (BMP) signaling, among other signals such
as Wnt and FGF, as necessary in this inductive process [9,13,14].
BMPs are postulated to pattern the ectoderm of zebrafish and
Xenopus in a gradient fashion, such that high levels of activity
induce epidermis, intermediate levels induce neural crest, and the
absence of BMP activity is required for neurectoderm formation.
In support of this idea, when zebrafish embryos are treated with a
high concentration of dorsomorphin, a small molecule that inhibits
type I BMP receptor activity, neural crest cells are absent, whereas
a low concentration of dorsomorphin causes expansion of neural
crest cells [15]. When Xenopus animal caps are excised and treated
with intermediate levels of Noggin, they express the early neural
crest marker slug, although this also requires the presence of FGF
[9]. These results indicate that modest attenuation of endogenous
BMP signaling can lead to neural crest induction. Other evidence
for a BMP signaling gradient in the ectoderm, and evidence for an
intermediate level of BMP signaling patterning lateral regions of
PLoS ONE | www.plosone.org1 November 2011 | Volume 6 | Issue 11 | e27403
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the embryo, particularly neural crest, comes from genetic analysis
in zebrafish [16–18]. In the strongly dorsalized swirl/bmp2b
mutant, foxd3, AP2, and snail expression in neural crest during
somitogenesis is absent, consistent with a requirement for BMP
signaling in neural crest specification. In more weakly dorsalized
somitabun (sbn)/smad5 and snailhousety68a(snh)/bmp7a mutants, neural
crest is greatly and moderately expanded, respectively, suggesting
that these mutants retain an intermediate level of BMP signaling in
an expanded region sufficient to specify neural crest [17].
However, the extent of the expansions has not been characterized,
nor has the residual signaling in these mutants been demonstrated.
Furthermore, the gradient model predicts that neural crest
progenitors directly respond to the intermediate level of BMP
signaling, however, this has not been addressed experimentally.
Here, we quantified the effects of reduction in BMP signaling on
the number of neural crest cells by counting the number of Foxd3-
positive cells in wild-type, swirl, sbn, and snhty68aembryos to show
that the expansion of the neural crest domain is not due to
impaired morphogenesis but rather an increase in neural crest cell
number. We modulate BMP signaling levels by over-expression of
BMP antagonists in wild type and various mutant conditions to
demonstrate that different levels of BMP signaling remain in sbn
and snhty68amutants. We further investigate the ability of Smad5 to
act in a graded fashion by injecting smad5 antisense morpholinos
and show that its dose-dependent loss recapitulates the BMP
mutant phenotypes, consistent with Smad5 acting directly in
neural crest progenitor specification. Using Western blot analysis,
we show that P-Smad5 levels are reduced in smad5 morphants in a
dose-dependent manner, consistent with an intermediate level of
BMP signaling acting through Smad5 to specify the neural crest
progenitors. Finally, we perform chimeric analysis to show that
BMP signaling is directly required within neural crest progenitor
cells for their specification. Together these results add substantial
evidence to a model in which graded BMP signaling acts as a
morphogen to pattern the ectoderm, with an intermediate level
responsible for neural crest specification.
Reduction of BMP signaling in the swr, sbn, and snh
mutants affects NCPC specification
We reported previously that the NCPC domain is decreased in
swr/bmp2b, and increased in sbndtc24/smad5 and snhty68a/bmp7a
mutant embryos [17]. This analysis was done during early
somitogenesis stages, several hours after foxd3, the earliest neural
crest marker, is expressed. To investigate if these defects are due to
defects in NCPC specification, we examined foxd3 expression at the
end of gastrulation (bud stage) in these BMP pathway mutants. We
found that NCPCs were greatly reduced to absent in swr/bmp2b
mutants (Fig. 1B); greatly expanded in sbn/smad5 (Fig. 1C); and
moderately expanded in snhty68a/bmp7a mutant embryos (Fig. 1D)
compared to wild-type (Fig. 1A). These results are consistent with
the hypothesis that mutations in BMP pathway components affect
specification, rather than maintenance of neural crest.
BMP signaling has been implicated in both promoting and
inhibiting cell proliferation (reviewed in [19–22]). To determine if
the expanded number of NCPC observed in sbn mutants reflects an
increase in their proliferation, we examined the expression of
Phospho-Histone H3, a marker of proliferating cells, in wild-type
and sbn/smad5 mutant embryos. By gross inspection, we did not
detect a difference between the mutants and wild type at any point
throughout gastrulation (50%, 70% and 90% epiboly stages, and
bud stage, data not shown). We counted the number of Phospho-
Histone H3 positive cells at mid-gastrulation (70-80% epiboly) in
wild-type (n=3, Fig. 1E) and sbn/smad5 mutants (n=3, Fig. 1F) and
found no correlation between expanded NCPCs and an increase in
proliferation (Fig. 1G). Hence, an increase solely in cell proliferation
cannot account for the large increase in NCPCs in sbn/smad5
mutants. Rather these results are consistent with an enlarged
domain of cells that is specified as NCPC in sbn/smad5 mutants.
The number of NCPC is increased in sbn/smad5 and snh/
bmp7a mutants
In 5-somite stage wild-type embryos, the anterior neural crest
population is 2–3 cell layers thick, and thins to a single cell layer in
the posterior (data not shown). It has previously been shown that
dorsal convergence is impaired in several BMP pathway mutants
[23,24], thus the apparent increase in the neural crest population
in these mutants could be due in part or entirely to a failure of the
NCPCs to converge into a multilayer tissue rather than an actual
increase in cell number. To determine the number of NCPC in
sbn/smad5 and snhty68a/bmp7a mutants compared to wild-type, we
counted the number of Foxd3-positive NCPCs (Fig. 1H–J). Foxd3
protein localizes to the nucleus, allowing one to easily count
individual cells, particularly in regions with multiple cell layers
[25]. Foxd3 protein detection is delayed compared to foxd3 RNA.
Thus, we counted Foxd3-positive nuclei in 2-somite stage
embryos, the earliest time point exhibiting strong Foxd3
expression (Fig. 1H). The average number of neural crest cells in
wild-type was 403 (n=4), and in snhty68awas 1160 (n=4), or a 2.8-
fold increase over wild-type. sbn/smad5 embryos averaged 1965
neural crest cells (n=4), or a 4.8-fold increase over wild-type.
Thus, the larger apparent neural crest domain in snhty68a/bmp7a
and sbn/smad5 mutant embryos reflects a larger number of
NCPCs. The NCPCs in these mutants also occupy a larger area
than expected based on their increased number due to reduced
dorsal convergence of the NCPC in these mutants, i.e. most of the
NCPC were found in a single rather than a multi-cell layer.
Together, these results support a model in which an intermediate
level of BMP signaling is present in a larger domain in these
mutants than in wild-type embryos.
Reducing BMP signaling in wild-type embryos dose-
dependently affects NCPC phenotypes
Based on our previously proposed model, we predict that
moderate or strong inhibition of BMP signaling in wild-type
embryos will lead to an expansion or loss of NCPC, respectively.
To test this prediction, we decreased BMP signaling in wild type
embryos by injecting mRNA encoding either a truncated Xenopus
BMP receptor (tBR, [26]) or zebrafish chordin (Fig. 2A, [27]), an
extracellular BMP antagonist [28]. Both of these methods yielded
similar results. As expected, over-expression of either inhibitor
produced a range of dorsalized phenotypes. We classified weakly
dorsalized embryos that displayed a roughly normal NCPC
phenotype as ''wild-type''. Upon injection of 50 pg of chordin
mRNA into wild-type embryos (n=89), we observed that 67% of
the embryos exhibited a wild-type (WT) NCPC phenotype, 21%
the ''snh'' and 12% the ''sbn'' phenotype (Fig. 2A, see Fig 1A–D for
the classification of ''wild-type'', ''snh'', ''sbn'', and ''swr''
phenotypes). When we increased the amount of chordin mRNA
injected to 200 pg (n=87), 40% of the embryos displayed the
''swr'' phenotype; 32% the ''sbn''; 18% the ''snh'' phenotype; and
only 10% exhibited a wild-type NCPC phenotype.
The sbndtc24mutation used in these neural crest studies is an
antimorphic allele of smad5, raising the possibility that the large
expansion of neural crest in these embryos is due to dominant-
negative effects interfering with Smad proteins that are used by
Intermediate BMP Directly Specifies Neural Crest
PLoS ONE | www.plosone.org2November 2011 | Volume 6 | Issue 11 | e27403
Page 3
other signaling pathways acting in NCPC specification. To
confirm that the sbndtc24phenotype is due to a reduction in
BMP-responsive Smad5 activity only, and to determine if Smad5
itself can act in a dose-dependent fashion, we injected various
amounts of a previously described translation-blocking smad5
antisense morpholino oligonucleotide into wild-type embryos [29].
Injection of 2.5 ng of smad5 MO1 led to embryos with a range of
moderately to largely expanded neural crest populations, whereas
4 ng of smad5 MO1 led to a large expansion or loss of neural crest
cells (Fig. 2B). Taken together, these results indicate that reducing
BMP signaling dose-dependently recapitulates the NCPC pheno-
types observed in the BMP mutants. Furthermore, the results
indicate that the gradient of BMP signaling that specifies NCPCs
acts primarily through Smad5, and that Smad5 itself can act in a
graded fashion to specify NCPCs.
The NCPC phenotypes of swr/bmp2b, sbn/smad5, and
snh/bmp7a mutants correlate with distinct residual BMP
signaling levels
We previously proposed a model in which an intermediate level
of BMP signaling specifies NCPCs, and that BMP signaling is
lowered below this level in swr/bmp2b mutants, resulting in a loss of
NCPCs. Furthermore, we predict that different intermediate
amounts of residual BMP signaling are present in sbn/smad5 and
snh/bmp7a mutants, leading to the great and moderate expansion
of NCPCs, respectively, in these mutants [17]. To test our
hypothesis that the NCPC phenotypes of swr/bmp2b, sbn/smad5,
and snh/bmp7a reflect the relative amounts of BMP signaling in
these mutants, we decreased or increased the amount of BMP
signaling in these mutants and examined the effects on NCPCs.
Having established that tBR and chordin over-expression can
recapitulate the BMP mutant phenotypes in a dose-sensitive
fashion, we used this technique to further decrease BMP signaling
in the BMP pathway component mutants. We predicted that
decreasing BMP signaling in sbn/smad5 or snhty68a/bmp7a mutants
would phenocopy the NCPC phenotype of swr mutants or sbn and
swr mutants, respectively, in a dose-dependent manner. Fig. 3A
shows a representative experiment in which we injected tBR into
embryos from a cross between two sbn/smad5 heterozygous fish.
Since sbn/smad5 is a fully penetrant dominant maternal-effect
mutation [30], all offspring from this cross exhibit the mutant
phenotype. In the uninjected group (n=48), 90% of the embryos
exhibit a ''sbn'' NCPC phenotype in which there is a great
Figure 1. Reduction of BMP signaling in BMP pathway mutants affects the number of NCPC specified. foxd3 expression at bud stage in
wild-type (A), swr (B), sbn (C), and snhty68a(D). P-histone H3 expression at 80% epiboly stage in wild-type (E) and sbn (F). (G) Quantification of the
number of P-histone H3 positive cells in wild-type (n=3) and sbn (n=3). Foxd3 protein expression at the 2-somite stage in wild-type (H), sbn (I), and
Intermediate BMP Directly Specifies Neural Crest
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expansion of NCPC and 10% display a ''snh'' NCPC phenotype, a
moderate expansion of NCPC. However, injection of 150 pg of
tBR into these mutants (n=86) resulted in a reduction in the
number of NCPCs. Only 27% of embryos exhibited the ''sbn''
NCPC phenotype, whereas the majority (73%) showed a great
reduction of NCPCs, resembling swr mutants.
It is possible that the loss of foxd3 expression reflects a general
deleterious effect on development and not a stronger dorsalization
of sbn/smad5 mutant embryos. To address whether over-expression
of tBR caused a general reduction in gene expression, we examined
the expression of krox20, a marker of prospective hindbrain
rhombomeres 3 and 5 [31]. We found that sbn/smad5 mutant
embryos injected with the tBR mRNA expressed krox20 and, in
fact, both rhombomeres were greatly expanded in these embryos
in a manner similar to that of swr/bmp2b mutants (data not shown,
[17]). From these results, we conclude that there is residual BMP
signaling in sbn/smad5 embryos, thus reducing BMP signaling in
these embryos phenocopies the ''swr'' NCPC phenotype.
We next reduced the level of BMP signaling in snh/bmp7a
mutant embryos to ask whether this would result in the NCPC
phenotype observed in sbn/smad5 or swr/bmp2b mutants. We
injected embryos from a cross between two rescued homozygous
Figure 2. Reduction of BMP signaling in wild-type embryos
causes expansion or loss of NCPC in dosage-sensitive manner.
foxd3 expression in chd mRNA injected embryos (A) and smad5 MO
injected embryos (B) at the end of gastrulation. (A) Injection of a low
dose of chordin mRNA (50 pg) generates weaker NCPC phenotypes
(WT=normal or very mild expansion, ''snh'' = moderate expansion),
whereas a high dose (200 pg) leads to strong phenotypes (''sbn''=large
expansion, or ''swr'' =loss). (B) Injection of a low 2.5 ng dose of smad5
MO leads to ''snh'' and ''sbn'' phenotypes. Injection of a high 4 ng dose
of smad5 MO leads to ''sbn'' and ''swr'' phenotypes exclusively.
Figure 3. NCPC domains when BMP signaling is reduced in
somitabun and snailhouse and increased in swirl embryos. foxd3
expression at the end of gastrulation in tBR mRNA injected embryos of
sbn(A)andof snh (B), andsmad5 mRNA injectedintoswr embryos (C). (A)
Injection of tBR mRNA into sbn mutants leads to the majority of embryos
displaying a ''swr'' phenotype. (B) Injection of a low 15 pg dose of tBR
mRNA into snhmutants leads tonearlyequal numbers of''snh'' and''sbn''
phenotypes. Injection of a higher 100 pg dose leads to the majority of
embryos displaying the ''sbn'' phenotype and also a percentage
displaying a stronger ''swr'' phenotype. (C) Injection of a low 30 pg dose
of murine smad5 mRNA results in nearly half of embryos displaying a
''sbn'' phenotype. Injection of a higher 150 pg dose results in a small
percentage of embryos displaying the ''swr'' phenotype, and the rest of
the embryos divided between ''sbn'', ''snh'', and WT phenotypes.
Intermediate BMP Directly Specifies Neural Crest
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snh/bmp7a adult fish; thus all progeny display a mutant phenotype.
As expected, the uninjected embryos exhibited a moderate
expansion of NCPC typical of the ''snh'' phenotype (Fig. 3B).
When we injected a low amount (15 pg) of tBR mRNA (n=61),
52% of the embryos showed the ''sbn/smad5'' NCPC phenotype;
whereas only 48% exhibited the ''snh'' NCPC phenotype (Fig. 2B).
When we injected a high amount (100 pg) of tBR mRNA, the
strength of the NCPC phenotype increased: 17% of the snh
homozygous embryos exhibited the ''swr'' NCPC phenotype; 71%
displayed the ''sbn'' phenotype; and only 12% showed the ''snh''
phenotype seen in the uninjected siblings. Thus, we find that
lowering BMP signaling in snhty68amutant embryos can phenocopy
the great expansion of the NCPCs observed in sbn/smad5 mutants,
and that a further reduction results in loss of NCPCs in a small
percentage of embryos. We note here that with all of these
experiments, we consistently found embryos that appeared
intermediate between either the "swr" and ''sbn" or the "sbn" and
"snh " NCPC phenotypes. In these cases, we classified the embryos
into the group they most closely resembled.
We also increased BMP signaling in a dose-dependent manner
in the swr/bmp2b mutant by over-expressing smad5 mRNA to
determine if this could produce the NCPC phenotype of sbn/smad5
and snhty68a/bmp7a mutants. Following injection of 30 pg of murine
smad5 mRNA (n=82), we found that 44% of the embryos
displayed the expanded NCPC phenotype of ''sbn'' mutants, and
the remaining mutants appeared unchanged. When we increased
the amount to 150 pg (n=93), we observed a range of phenotypes
from the "swr" to wild-type NCPC phenotypes (Fig. 3C).
Therefore, we conclude that increasing BMP signaling in swr/
bmp2b mutants leads to an expansion of NCPC in a dosage-
sensitive manner.
Phosphorylated Smad1/5 levels correlate with the
strength of NCPC phenotype in smad5 morphant
To further examine the extent of reduction in BMP signaling in
smad5 morphant embryos, we examined phosphorylated Smad1/5
(P-Smad1/5) levels. We injected smad5 translation-blocking
morpholinos (MOs) into wild-type embryos at different concen-
trations and examined P-Smad1/5 levels by Western blot analysis
at the 60% epiboly stage, just after cranial NCPC are specified
[32]. For each injection dose, a fraction of the embryos were used
for Western blot analysis and the remaining embryos were
examined for the NCPC phenotype.
Compared with wild-type embryos, all smad5 morphant
embryos displayed lower P-Smad5 levels that correlated with the
smad5 MO dose injected. Embryos injected with 6ng of smad5
MOs had no detectable P-Smad1/5 compared to embryos
injected with 5 ng, 4 ng and 3 ng of smad5 MOs, which had
progressively increasing levels of P-Smad1/5 (Fig. 4A).
We examined expression of foxd3, one of the earliest markers of
NCPC, at the end of gastrulation in the remaining embryos of
each group to determine the extent of expansion of, or loss of
NCPC at each smad5 MO dose. This allowed us to correlate the
NCPC phenotype with P-Smad1/5 levels during gastrulation.
Injection of a low dose of smad5 MOs (3 ng) caused primarily WT
and ''snh'' phenotypes, whereas higher doses of smad5 MOs (5 ng,
6 ng) resulted in ''sbn'' and ''swr'' phenotypes almost exclusively
(Fig. 4B, 4C). We find that increasing amounts of smad5 MOs leads
to decreasing P-Smad1/5 levels, which correlates with expansion
of NCPCs at intermediate P-Smad1/5 levels and then a loss of
NCPCs at undetectable P-Smad1/5 levels, consistent with the
BMP mutant results.
BMP signaling is autonomously required for neural crest
progenitor cell specification
Models of BMP gradient activity predict that each cell type
within the gradient field responds directly to the level of BMP
signaling, thus neural crest progenitor cells are expected to
respond directly to an intermediate level of BMP signaling. The
direct response of NCPCs to BMP signaling has not been
addressed in any organism. We addressed the cell autonomy of
BMP signaling in NCPCs by placing donor cells that cannot
respond to a BMP signal into a wild-type host environment where
BMP signaling is intact and determined if the donor cells can
express Foxd3. If the donor cells can express Foxd3, this indicates
that BMP signaling is not acting directly to specify NCPCs.
Alternatively, if donor cells do not express Foxd3 this would
indicate that BMP signaling directly induces NCPCs.
To generate donor cells that cannot respond to BMP signaling,
we used a combination of smad5 morpholinos to inhibit Smad5
translation. We then transplanted 5–15 cells from blastula-stage
donors to a region above the margin of a blastula-stage wild type
host embryo, in a region predicted to become neural crest based
on fate-mapping studies [4]. Donor embryos were analyzed
individually for foxd3 mRNA expression, and only chimeras
derived from donors that completely lacked foxd3 expression were
further analyzed. As a control, we transplanted wild-type cells into
the same region of blastula-stage wild-type host embryos, and
examined Foxd3 expression at the 3-somite stage. In 7 of 7
chimeras analyzed, these wild-type donor cells readily expressed
Foxd3 (Fig. 5A–D). In 8 of 9 embryos with smad5 morphant clones
in the neural crest area, donor cells did not express Foxd3. Clones
in two separate embryos from two different donors are shown in
Figure 5E–N. In the single embryo in which donor cells expressed
Foxd3, donor cells became neural crest cells only ectopically in a
more ventral region to the normal domain (Fig. 5O–Q). Ectopic
neural crest induction was never observed with wild-type donor
cells. It is possible that the morphant-derived cells retained a slight
amount of Smad5, insufficient in the donor to specify neural crest
but sufficient to weakly respond to a surrounding high level of
BMP signaling ventrally in the chimera; responding at the lower
level appropriate for neural crest rather than epidermis, which
would normally occupy the domain. Taken together, these results
indicate that NCPCs directly respond to BMP signaling.
A low level of BMP signaling is required for specification
of neural crest progenitors
Temporal inhibition of BMP signaling using Tg(hsp70:chd) [32]
and treatment of embryos with dorsomorphin at different time
points [15] show that BMP signaling acts in cranial neural crest
specification at an early gastrula stage. Cells become committed in
the zebrafish embryo between the shield stage and 80% epiboly
[33,34]. NCPC phenotypes are evident in BMP pathway mutants
at bud stage, shortly after cell commitment, suggesting that BMP
signaling is involved in the primary specification of these cells
rather than maintenance of this cell type.
We previously reported that there are no differences in cell
death between wild-type and BMP pathway mutants during
blastula and gastrula stages [17]. Thus, cell death likely does not
contribute to reduction of NCPCs in swr/bmp2b mutants. Our
phospho-histone H3 data revealed no correlation between the
increase in number of NCPCs in sbn/smad5 mutants and cell
proliferation. Taken together, these results indicate that cell
proliferation and cell death do not contribute to the expansion or
Intermediate BMP Directly Specifies Neural Crest
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text:
Professor Simon Brown
Professor of Physics
Department's "Innovation Champion"
On sabbatical 2nd Semester 2015
Ph.D. (Cambridge)
608, Physics and Astronomy
Contact Details
Phone: +64 3 364-2507
Internal Phone: 6507
Postal address:
Department of Physics and Astronomy
University of Canterbury,
Private Bag 4800,
New Zealand
Simon was an undergraduate at Victoria University of Wellington and has a Ph.D. from Cambridge in the U.K. He spent 6 years as a postdoctoral fellow at the University of NSW in Sydney, and a year at the University of British Columbia in Vancouver before moving to Canterbury in 1998. His research interests have evolved from magnetism to semiconductor devices to atomic clusters.
Simon is also the Deputy Director for Commercialisation and Industry Engagement at The MacDiarmid Institute.
Research Interests
UC Research Profile
Physics PhD scholarship available-deadline 7 June 2015
Scanning Tunneling Microscopy studies of rare earth nitrides and related materials
This is an opportunity to explore how new rare earth nitride materials can be made and how they can be probed using scanning tunneling microscopy.
The successful candidate will have enthusiasm, a good honours or masters degree in physics (or related subject), and a desire to work in a multi-institutional, multi-disciplinary, collaborative environment. The successful candidate will be a member of the MacDiarmid Institute for Advanced Materials and Nanotechnology, a national Center of Research Excellence the provides may collaborative opportunities and a thriving environment to work in. The scholarship is worth $27,000 per annum and includes all student fees. The research will take place at the University of Canterbury in Christchurch , which has recently been rated as one the world's top 10 cities to visit by Lonely Planet. http://www.lonelyplanet.com/new-zealand/christchurch-and-canterbury/christchurch/travel-tips-and-articles/77531.
Please note that all applications must include:
- A full Curriculum Vitae, INCLUDING your University transcript (i.e. list of grades awarded).
- The names of at least two people who are prepared to act as referees.
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Only applications received before the deadline of 7 June 2015 will be considered. All applications should be emailed to email@example.com.
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- Dr Nadia Grillet (December 2012)
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Newly Appointed Cardinals Pay Courtesy Visit at Regia and Ducale Halls
Archbishop of Sao Paulo Odilo Scherer poses with diocesans of the... Nieuwsfoto'sBezoek,Bijwonen,Geloof,Goede manieren,Internationaal monument,Kardinaal,Nieuw,Odilo Scherer,Paus,Paus Benedictus XVI,Portret,Spiritualiteit,São Paulo,Ten voeten uit,Vaticaanstad,Vaticaanstad - Staat,VerticaalPhotographer Franco OrigliaCollection: Getty Images News 2012 Franco OrigliaVATICAN CITY, VATICAN - NOVEMBER 24: Archbishop of Sao Paulo Odilo Scherer poses with diocesans of the newly appointed cardinal John Olorunfemi Onaiyekan, archbishop of Abuja Nigeria, attend the courtesy visits at the Sala Regia Hall at the end of the concistory held by Pope Benedict XVI on November 24, 2012 in Vatican City, Vatican. The Pontiff installed 6 new cardinals during the ceremony, who will be responsible for choosing his successor. (Photo by Franco Origlia/Getty Images)


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South Park
Helen Keller! The Musical
- Episode Description
The 4th grade is doing their own version of Helen Keller's story, but when they're told the kindergarteners are putting on a "Thanksgiving extravaganza," they decide to turn their production into a musical. Timmy is set to star as Helen in the play, but when Timmy's disabled turkey is passed over for a part in the play, Timmy has second thoughts about participating.
- Cast & Crew
Trey Parker
Matt Stone
Isaac Hayes
Eliza Schneider
Mona Marshall
- Fan Reviews (4)
4th graders battle against the Kindergartners for Thanksgiving supremacy.
No, no, no. I did not like this at all.
Hellen Keller! The Musical!
Gobbles! Timm-ah!
- Trivia & Quotes
Quotes (7)
Mr. Mackey: Every year the fourth graders do "The Miracle Worker", and every year we have to sit and watch it. Principal Victoria: Yeah, I swore that if I had to see it one more time I'd put a bullet in my head. But luckily, I got really stoned before I came.
(When Cartman is told to kill Gobbles) Cartman: I can't dude, Timmy has a boner for it.
Cartman: Kyle, why do you do these things to me? Kyle: I didn't do anything to you; Timmy saw the turkey and wanted to get it, what did you want me to say to him? Cartman: You say: "NO TIMMY, YOU CAN'T HAVE THAT TURKEY! BAD TIMMY!"
Actor: (singing) Why not use a trick we actors use, cover your eyes and ears to see what it would be like to be blind and deaf. See what plays inside your mind. (Cartman does so and it is followed by a montage of horrific scenes, the holocaust, someone on fire, grizzly operations, a dead rat being eaten by another rat, etc.) Cartman: (takes off blindfold and ear muffs) WHOA! Actor: Anything useful? Cartman: No, just the stuff i usually see when I close my eyes.
Stan: Where's Gobbles? Cartman: I don't know. (Stan and Kyle look at him doubtfully) Cartman: I DON'T KNOW! I might have tried to have Gobbles crushed by a stage light but that's it, I'm not an asshole.
Notes (2)
Originally instead of a disabled turkey it was going to be a disabled puppy, but Trey and Matt found it too sad.
This is the first Thankgiving themed episode since "Starvin' Marvin."
Trivia (12)
When Kyle says: 'Timmy we already got a turkey' it was in Stan's voice, not Kyle's
Timmy's mother's name is Helen, and Timmy plays Helen in the play.
When Kyle said "Timmy, we already got a turkey" Look at Clyde in the next scene, his clothes have been changed.
Timmy's house was small in Timmy 2000, now he lives in a big house.
When Timmy brings out the lighter to set fire to the hoop, if you look carefully, you can see that the lighter was already lit before he flips it open and strikes it.
Allusions (8)
The Miracle Worker: The play the kids are performing, albeit in a very free adaptation, is "The Miracle Worker" by William Gibson. This tells the story of an episode in the real life of Helen Keller, a woman who became blind an deaf when she was infant. "The Miracle Worker" specifically describes the relationship between Keller -- deaf, blind, and initially almost feral -- and Annie Sullivan, the teacher who aided her into education, activism, and intellectual celebrity.
The Phantom of the Opera: Cartman makes a reference to Tim Rice writing the lyrics to "The Phantom of the Opera." However, these lyrics were written by Charles Hart and Richard Stilgoe.
Simpsons Did It!: There was an episode of The Simpsons, I Love Lisa, in which Milhouse complains to Miss Hoover that his Abraham Lincoln beard itching him. Just like Stan did in this South Park episode.
Les Miserables: Most of the songs Jeffrey Maynard sung are similar to ones from the musical "Les Miserables". When Kenny dies, Maynard sings a parody of "Bring Him Home."
Air Bud: The way Timmy attempts to abandon his beloved pet turkey is very similar to a scene in the 1997 movie Air Bud. In the movie, the main character, a young boy, takes his golden retriever out away from his house and walks away from the dog with tears in his eyes.
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Making plans to spend Christmas in Stockholm? Here's a selection of Christmas markets to make a visit to.
Christmas market, Old town in Stockholm
Henrik Trygg/Stockholm Visitors Board
Welcome to The Royal Mews and a very special Christmas Fair in a unique environment. Besides the usual selling of quality handicrafts and delicacies, there will be stroll-rounds in the Royal Mews buildings. See the stables, harnesses, and carriages!Dates: 27 - 29 November
Sigtuna's Christmas market is an open-air market with Christmas items, handicraft, traditional Swedish food items and much more. Four Sundays before Christmas Eve, Sigtuna lights up with a traditional Christmas market.Dates: November 30, December 7, 14 and 21Time: 11-16
Welcome to visit this famous, oldest and most traditional Christmas Market in Stockholm with a special cheery festive atmosphere!Dates: The Old Town Christmast Market is openfrom Saturday 21 November until 23 December 2015.Open daily between 11am and 6pm
Traditional Christmas market at SkansenAt the Bollnäs Square and in the Town Quarter. Enjoy our traditional Christmas market with stands filled with Swedish handicraft, sweets and lovely food.Dates: 28- 29 Nov; 5-6 Dec; 12 - 13 Dec; 19 - 20 Dec.10am - 4pm
Pink Christmas Week . More info to come for 2015. Pink Christmas Week is organised by Stockholm Gay Life. Over the course of the week, there will be Christmas markets, concerts and pink offerings at gay bars and clubs, as well as lectures and the Rainbow Fund's prize presentation.
More about Christmas in Stockholm
Order brochures over different activities and destinations in Sweden.
Partner with us. Contact us and we'll tell you more. In Swedish only!
Go here for press releases, images and press contacts.
Find your contacts for Meetings & Incenitives in Sweden
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$4.99Two almost-exes battle sharks, saboteurs, and treacherous desire when they become caught up in a dangerous search for a legendary shipwreck Salvage diver Roc Trellyn has just hauled up the biggest catch of his career. But the creature in his net isn't a prize fish—or a mermaid. It's Melinda Davenport, his ex-wife. Except they're still married—just a little glitch they both seem to have overlooked.
$9.99Collects RAT QUEENS #1-5Who are the Rat Queens? A pack of booze-guzzling, death-dealing battle maidens-for-hire, and theyÍre in the business of killing all godÍs creatures for profit. ItÍs also a darkly comedic sass-and-sorcery series starring Hannah the Rockabilly Elven Mage, Violet the Hipster Dwarven Fighter, Dee the Atheist Human Cleric and Betty the Hippy Smidgen Thief. This modern spin on an
Linear CMOS RF Power AmplifiersThe work establishes the design flow for the optimization of linear CMOS power amplifiers from the first steps of the design to the final IC implementation and tests. The authors also focuses on design guidelines of the inductor's geometrical characteristics for power applications and covers their measurement and characterization. Additionally, a model is proposed which would facilitate designs in
Brotherhood of Misfits (Book #1)What would you do with a second chance at life?Would you go back to the same things that you were doing before, or would you embrace that chance and take a new path?Being the byproduct of a single parent home growing up, I learned quickly that life wasn't easy and it damn sure wasn't fair.I was faced with a choice back then; continue to stand back and watch my mother struggle with working three
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$4.99Roc Isle: Tempest is a heroic fantasy novel based on an epic sword-and-sorcery battle of good versus evil.Lord Azure commands the Northern Army in a war against the Trade Lords, who are a class of evil conspirators. He expects victory for he is aided by veterans of war and Ankah, a battle strategist and master swordsman. However, his leadership is failing because of his son's insolence and
Roc and Roe's Twelve Days of ChristmasNick Cannon and Mariah Carey's twins, Roc and Roe, decorate their Christmas tree with their "pip" version of "The Twelve Days of Christmas." From an angel with sparkly, shiny wings to four skiing snowmen to twelve chugging choo-choos, Roc and Roe have a frolicking time getting ready for the holidays.
The Story of Roc-A-Fella RecordsWhen Shawn Carter, an aspiring rapper with the stage name Jay-Z, couldn't get his career off the ground at a major record label, he didn't get discouraged. He and his partners Damon Dash and Kareem "Biggs" Burke created a hip-hop label of their own. They named the label Roc-A-Fella Records. Originally intended as a one-time project to release Jay-Z's full-length debut album, Reasonable Doubt, Roc
$4.99Roc Isle: The Descent is a heroic fantasy novel based on a tale of revenge and destiny.It is now Year One, foretold to be the final year before The End of Roc Isle.In the North, Lord Azure of the Azure-Cloud Clan vows revenge on those responsible for ordering the assassination of his parents. Lord Azure will invade the South in reprisal, and to rid the land of the infidel. But first he needs to
$8.89Do you long to see the area where you live transformed, but haven't a clue how to make it happen? Would you like to see relationships in your community change for the better? Do you want to be a part of God's miracle in your neighbourhood and beyond? In this book, author Debra Green shares insights gained from 20 years' experience of community transformation and ...
WHAT HAPPENS WHEN THE KINGDOM OF GOD INVADES THE HOODRemember the parable of the Good Samaritan that Jesus told over two thousand years ago? Well, live from Bedford-Stuyvesant is the Hood Samaritan, Donteer Brown… Fresh out the pens, Donteer reunites with his family with new and exciting plans for the future. He's a new man in Christ, just trying to do the right thing and can't afford to waste anymore time. However, before he can even begin to put
Signal Detection Theory and ROC Analysis in Psychology and DiagnosticsSignal detection theory--as developed in electrical engineering and based on statistical decision theory--was first applied to human sensory discrimination 40 years ago. The theoretical intent was to provide a valid model of the discrimination process; the methodological intent was to provide reliable measures of discrimination acuity in specific sensory tasks. An analytic method of detection


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You should follow Drew on Twitter and Facebook, you should read Toothpaste For Dinner and Married To The Sea, and you should absolutely listen to CRUDBUMP like right now.
Vince Staples's Summertime in the City
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We bring Miguel together with artist Connor Olthuis to examine how globalization has changed the way we think about resources, and how music has been affected by similar issues.


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10 reasons why Halloween is the absolute best holiday
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Missed connections
In Sixteen Candles, Sam
Baker is totally crushing on cutie Jake Ryan, but he barely knows she exists
(or so she thinks). In reality, Jake is chasing after Sam throughout the dance
and into the rest of the night, but they completely miss each other and Sam
spends her night being unsuccessfully wooed by a pushy geek. Yikes! We're
hoping your Prince Charming has no trouble finding you come dance night.
Food fights
This movie packs a
double-whammy! In Never Been Kissed, Josie is asked to prom by total hottie Billy
Prince. But when he comes to pick her up, he reveals he asked her as a joke,
then pelts Josie with eggs alongside his real date. Josie gets a chance at
school-dance redemption when working undercover for the newspaper, but it still
doesn't go so great. Mean girls almost pour dog food all over Josie's friend as
an act of revenge, but she steps in and pours it all over the mean girls
instead. We're hoping you have an easier time avoiding food meltdowns during
your big event.
Lousy bets…with lame dudes
In She's All That, Dean bets
Zack that Zack cannot get a girl to be prom queen within six weeks. Dean says
Zack must go with Laney, an independent and feisty art student. While trying to
win the bet, both Zack and Dean end up developing feelings for Laney, but Dean
is a pig and tries to make advances on her. Hopefully your story has a lucky
ending like this one. (Spoiler alert!) Laney makes it out of Dean's hotel room
and ends up with Zack in the end.
A paid gig
In the classic 10
Things I Hate About You, bad boy Patrick is getting paid to win
over—and date—bitter older sister Kat so that shy-guy Cameron can date younger
sister Bianca. It's complicated, we know, but not as messy as when Kat finds
out the guy she's slowly been falling for was only with her for some extra
dough! In the end, everything works out because Patrick and Kat realize they
have genuine feelings for each other. But talk about a total night-ruiner—finding out at the dance that your date was paid to be there? Ugh.
An MIA date
You've worked for months to
grab the attention of the cutie from math class, and you were pretty sure you
succeeded. He agreed to be your date, so you're in for a night of fun, right?
Well, maybe not so much. In Napoleon Dynamite, Napoleon crafts a
drawing of the girl he's crushing on. She agrees to go to the dance with him
only because her mom makes her…then promptly ditches Napoleon to (ouch!) dance
with her buds. Word to the wise, ladies: If you say yes to going with a guy, at
least have some respect and spend some of your night with him. And if a dude
ditches you on the dance floor? His loss!
An unexpected crush
If you've got a BGF, you're
probably thinking the friendship is totally platonic. But careful, because he
may just be crushing on ya. In Harry Potter and the Goblet of Fire, Hermione is thrilled to have been
asked to the Yule Ball by one of the most in-demand guys visiting Hogwarts. But
her night quickly becomes a bust when Ron, realizing he has feelings for
Hermione, mocks her and her date. Poor Hermione leaves the dance shouting,
"You've ruined everything!" We're hoping your BGF has more common sense than
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WIN BIG! Snag a $450 sandals 'n'
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INSTA-AWESOME! GL is now on Instagram.
POSTED IN entertainment, movies, movies we love, dance
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Articles published on November 3, 2011
Alabama Supreme Court ruling puts Gadsden in playoffs
Alabama Supreme Court ruling puts Gadsden in playoffs
Staff Report
Alabama women's tennis splits for competition
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All in the family
By Alex Scarborough Sports Writer
Breaking down fats
By Monica Eng McClatchy Newspapers
Charley Pell Classic is Friday
From staff reports
County workers to get 2 percent pay increase
By Robert DeWitt Senior Writer
City Council hires contractor to clear damaged homes
By Jason Morton Staff Writer
COMMENTARY: Occupy Wall Street standing against GOP obstructionism
By Oscar Tucker
Crimson Tide, Tigers excel at stopping the run
By Aaron Suttles Sports Writer
Deputies evict man, find marijuana plants
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EDITORIAL: Eyes of nation return to town on Saturday
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Guardrail repairs scheduled for I-20/59
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Holt students use art to heal storm's wounds
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LETTER: Thanks to Tide fans for civility
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Retired Supreme Court justice to speak at UA
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School wants out of civil suit involving coach
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Stillman trying to right the ship
By Andrew Carroll Sports Writer
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McClatchy Newspapers
Tigers quarterbacks take final shot at Alabama
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Teen hits hole-in-one at Ol' Colony
From staff reports
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From combined reports
Two retailers raising money for charities
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Tuscaloosa man whose case changed mental health care in U.S. has died
By Dana Beyerle Montgomery Bureau
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University officials: UA-LSU game calls for 'more of everything'
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Worker verification likely to stand
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Images of the DayAlabama defensive lineman Josh Chapman (99) sacks LSU quarterback Jarrett Lee (12) at Tiger Stadium in Baton Rouge, La on Nov. 6, 2010.
More than 100 University of Alabama at Birmingham students, supporters and fans attended the UA Board of Trustees meeting Thursday to protest their refusal to consider a new stadium for UAB.
From left, Ja'Darius Woods, 9, Quentin Williams, 10, and Jonaad Jenkins, 10, work on their paintings during art class at Holt Elementary School, which is now being housed at the former Lloyd Wood Middle School, in Northport on Wednesday.
From left, Dominic Branch, Montell Dent, D'Leisha Dent, Daniel Dent, Damion Dent and DeMonte Dent form an athletic group of cousins making their marks on Tuscaloosa County sports. Daniel is a track star at Stillman, while the other five are in high school: Branch at Paul W. Bryant, Montell at Northridge, D'Leisha at Central and Damion and DeMonte at Brookwood.
Alabama linebacker Dont'a Hightower (30) closes in on Ole Miss running back Jeff Scott in Oxford, Miss., Oct. 15, 2011. The University of Alabama and LSU football teams are ranked first and third nationally in run defense, respectively.
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Is this attraction free to the public?
Is gratuity suggested at this attraction?
Would this be a good rainy day activity?
Is there food available at this attraction?
Is a car required to access this attraction?
Is this attraction accessible using public transportation?
Is this attraction closed during certain parts of the year?
Does this attraction require payment to access?
Do you recommend that visitors take a guided tour at this attraction?
Is this attraction wheelchair accessible?
Thanks for helping!
Good place for rest after hiking or skiing.
Nice view for mountains from the pool. Little crowdy in winter season.
Review collected in partnership with Deutsche Telekom
I really recommend this place after a day of hiking. On the first floor there is a large pool which is inside as well as outside. The view outside is amazing. We visited the place twice, and the first time is was very noisy on this first floor due to all the kids in the area. On the second floor,...
The spa and therme around Vienna are famous and some of them offer remarkable landscape like St. Martin Lodge in Neusiedl am see or in therme Laa and even in Vienna itself you can find therme Wien and other that are sooo much better then Romerbad Thermal Spa. There is a sort of tram that bring you to Baden in...
We went here to spent our free skiing day. We expected a little more than just definetely nice pool with outside swimming... But it was nice and we and our kids enjoyed it. By the way, in case you are hungry go to the pools cafeteria, they cook great and tasty!
Very attractive design, facilities excellent in particular wellness area i.e. sauna, steamroom etc. Very clean in all areas, not cheap but worth the money
definitely quality good facility but be aware those who have never experienced mixed male and female spa.
부산 센텀 스파 정도로 꽤 규모있고 괜찮은 시설의 스파예요.
하지만 수영장을 제외하고 사우나는 모두 남녀공용이고 전라로 입장해야합니다. 사우나 너무 그리웠는데 저는 도저히 용기가 안나서 바로 나와버렸네요ㅠㅠ. 컬쳐쇼크 이런거 없이 자연스럽게 전라로 이용할 의향이 있으신 용자님들에겐 강추합니다!
I'm used to visit several thermal centers around Europe and I never felt so cold in such an un-relaxed place with over-payed bill.
It's definitely too small and too less for the price they ask.
We spent 3 hours at Romerbad Spa and it was well worth the time. The spa is beautifully designed and a great place to relax. Their swimming pool is huge and has a great outdoor section, plus a (small) water slide. Their sauna area is also great and their scheduled services are incredible. We did a mint-chocolate steam room scrub...
Very good thermal bad for adults and for very small kids. But no fun for older kids (3 nad more years). Expensive!
I find this complex to be the right choice to relax if you are in the area of Bad Kleinkirchheim. I'm not saying it's the best, but definitely I recommend a visit, especially in the winter time.
The pool area is quite limited, so don't expect to swim there like a pro. Even so, the outside pools, thermal water combined...
Own or manage this property? Claim your listing for free to respond to reviews, update your profile and much more.


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The summer program/camp they run at the Fishers YMCA is what I deem as an accident waiting to happen. For the entire summer they will allow your child run wild with minimual supervison (handful of teenagers for 50 or so kids) through the woods located behind (north) of the building. I have seen children in the woods in rain, storms, 90+ degree weather for almost the entire day. Oh sure they'll tell you we bring them ... view moreLike this review?Yes|Flag AbuseProvided by Citysearch
Today:5:00 am - 10:00 pm
Tomorrow:7:00 am - 7:00 pm
9012 E 126th St,
Fishers, IN 46038
(317) 595-9622
Business Details
YP Qualified
- Hours:
Regular Hours
- General Info:
- With more than 2,600 YMCA is one of the largest nonprofit community service organizations in America that works to meet the health and human service needs of more than 20 million men, women and children in nearly 10,000 communities. YMCAs are at the heart of community life since 42 million families and 72 million households are located within three miles of a YMCA. YMCA stands for Young Men's Christian Association. From its start more than 150 years ago, when George Williams founded the YMCA as a substitute Bible study and prayer for life on the streets, the YMCA was unusual because it crossed the lines that separated all the different social classes in England. YMCAs offer its members a variety of programs that include adventure guides, aquatics, arts and humanities, camping, child care, community development, family, health and fitness, older adults, SCUBA, sports and teen leadership. YMCA has a location in Fishers, Ind.
- Location:
- Fishers Branch YMCA
- Other Link:
- Categories:
- Social Service Organizations, Health Clubs


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The Coast Guard medevaced a 50-year-old woman from a cruise ship 225 miles southeast of Southwest Pass, Saturday.
Watchstanders at Coast Guard District 8 received a call at approximately 9 a.
Plans to terminate commercial tours to the North Pole on the nuclear-powered icebreakers of the Atomflot company in 2016 have surged the demand for these tours.
The Palaszczuk Government has appointed one of Queensland's most respected businesswomen, Renita Garard as the new Chair of the Port of Townsville.
In a statement released earlier today,
South Korean shipbuilders have ranked first in the world outpacing Chinese and Japanese rivals in terms of new orders in the third quarter of the year, reports
The General Ship Repair Corporation, a fixture on the Baltimore, Maryland waterfront for nearly a century since its founding by Charles "Buck" Lynch in 1924, has
MARAD provides funding to further study LNG as marine fuel for containerships
The U.S. Maritime Administration (MARAD) announced it has entered a $900,000 cooperative


text:
Royalty Free Music Unique To Your Project - Every Time
Royalty Free Music — Unique To Your Project – Every Time
Try SmartSound's Award Winning and Patented Technology Yourself
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Or Browse By (Click to add multiple browse filters)
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The following tracks match: New Age, Energetic, Action
New RitualLively rhythms surge through the night air on this compelling drum piece. Underscore your next chase scene or set an exotic tone in your island footage.https://www.smartsound.com/producer/images/ps11-210px.jpgSingle31109-ST$ 39.9531109-EX$ 69.9531109-W$ 149.95Keywords: Driving, Energetic, Incessant, Lively, Urgent, Action Genre: World, New Age Production Types: Action-Drama Instrument: Ethnic Ensemble, Percussion, Instrumental
New Ritual from the album Mystical / Percussive / Soundscapes (PS11)
Zimba DanceFestive drums, children's voices and rhythmic clapping paint an authentic picture of African celebration under starry skies.https://www.smartsound.com/producer/images/ps11-210px.jpgSingle31114-ST$ 39.9531114-EX$ 69.9531114-W$ 149.95Keywords: Driving, Energetic, Lively, Urgent, Action Genre: World, New Age Production Types: Action-Drama Instrument: Ethnic Ensemble, Percussion, Instrumental
Zimba Dance from the album Mystical / Percussive / Soundscapes (PS11)
The following tracks match: Energetic, Action
One Shall RiseA meticulous opening with dark electronic sounds gives way to a high-intensity action sequence ideal for scenes with frenetic motion held together with an undeniable drive.https://www.smartsound.com/artist/images/ASMM01_200.jpgSingle70707-ST$ 39.9570707-EX$ 69.9570707-W$ 149.95Keywords: Driving, Energetic, Exciting, Atmospheric, Bold, Competitive, Fierce, Heroic, Intense, Thrilling, Action Genre: Futuristic, Orchestral, Electronica, Dubstep Production Types: Cinematic, Film Trailer, Video Games, Epic, Action-Drama Instrument: Electronic, Orchestra, Studio Orchestra, Strings, Synthesizer, Instrumental
One Shall Rise from the album Massive Edge Orchestral
ResurrectAn ominous string ostinato sets up a driving drumbeat with a soaring, heroic melody. A peer into the abyss is provided through a breakdown section with growling, angry synthesizers.https://www.smartsound.com/artist/images/ASMM01_200.jpgSingle70709-ST$ 39.9570709-EX$ 69.9570709-W$ 149.95Keywords: Energetic, Exciting, Bold, Competitive, Fierce, Heroic, Intense, Powerful, Thrilling, Action Genre: Futuristic, Orchestral, Electronica, Dubstep Production Types: Sports, Cinematic, Film Trailer, Video Games, Epic, Action-Drama Instrument: Electronic, Orchestra, Studio Orchestra, Strings, Synthesizer, Instrumental
Resurrect from the album Massive Edge Orchestral
StrongAs the title suggest, everything is heavy in this track. Crushed and Heavy Drums, Rockin Guitars, WARNING! this track will increase the Rock & Roll Factor of your production.https://www.smartsound.com/strata/images/STRATCOE_200.jpgSingle69208-ST$ 39.9569208-EX$ 69.9569208-W$ 149.95Keywords: Driving, Energetic, Exciting, Edgy, Bold, Competitive, Extreme, Fierce, Intense, Powerful, Punchy, Striving, Action, Angry, Heavy, Slammin Genre: Rock, Sports FX, Heavy Metal, Rockin Production Types: Sports, Cinematic, Commercials, Reality TV, Video Games, Action-Drama Instrument: Rock/Pop Band, Instrumental
Strong from the album Core Evolution
Escape CarTurbulent drums, nonstop energy and a churning pulse make this track perfect for the final scene, where it's all on the line.https://www.smartsound.com/strata/images/STRATCRS_200.jpgSingle69404-ST$ 39.9569404-EX$ 69.9569404-W$ 149.95Keywords: Driving, Energetic, Competitive, Extreme, Incessant, Intense, Powerful, Somber, Suspenseful, Tense, Action, Drama, Chase Genre: Electronica, Electronic Production Types: TV/ Film Drama, Reality TV, Suspense, Action-Drama Instrument: Electronic, Synthesizer, Instrumental
Escape Car from the album Crime Scene
Driving My BassGritty bass-lines and furious drums help your character prepare for the big day or for the race itself.https://www.smartsound.com/filmscore/images/FSCSG01_200.jpgSingle68506-ST$ 39.9568506-EX$ 69.9568506-W$ 149.95Keywords: Driving, Energetic, Exciting, Edgy, Bold, Competitive, Extreme, Fierce, Intense, Pulsating, Punchy, Tense, Action, Drama, Chase Genre: Rock Production Types: Sports, Commercials, Suspense, Action-Drama Instrument: Electronic, Rock/Pop Band, Bass (Electric), Synthesizer, Instrumental
Driving My Bass from the album Craig Stuart Garfinkle Vol 1 - Emotive & Dramatic
Bring It OnYou've just won the big game or received the gold medal, and are being hoisted up above your team and fans. Featuring brass, strings and guitar melody.https://www.smartsound.com/strata/images/STRATCSP_200.jpgSingle66902-ST$ 39.9566902-EX$ 69.9566902-W$ 149.95Keywords: Energetic, Competitive, Fierce, Heroic, Intense, Powerful, Pulsating, Striving, Thrilling, Action Genre: Olympic, Specialty Production Types: Sports, Epic Instrument: Orchestra, Brass Section, Drums, French Horn, Guitar (Electric), Instrumental, Guitar
Bring It On from the album Classic Sports
Breaking BondsThis uptempo groove is ideal for movie trailers and TV spots, as well as a getaway or chase theme.https://www.smartsound.com/strata/images/STRATUD_200.jpgSingle66802-ST$ 39.9566802-EX$ 69.9566802-W$ 149.95Keywords: Driving, Energetic, Bold, Action, Upbeat Genre: Rap/Hip-Hop, Urban Production Types: Action-Drama Instrument: R&B Band, Synthesizer, Instrumental
Breaking Bonds from the album Urban Drama
Connection FailureThis hard-hitting percussive score lends itself to video games, trailers, storming the castle, or when good finally faces evil.https://www.smartsound.com/filmscore/images/FSAPF05_200.jpgSingle67901-ST$ 39.9567901-EX$ 69.9567901-W$ 149.95Keywords: Driving, Energetic, Exciting, Ominous, Bold, Competitive, Dramatic, Fierce, Intense, Powerful, Pulsating, Tense, Thrilling, Action, Drama Genre: Orchestral Production Types: TV/ Film Drama, Cinematic, Film Trailer, Video Games, Epic, Action-Drama Instrument: Percussion, Vocal
Connection Failure from the album Amotz Plessner & Friends Vol 5 - Rise To Power
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text:
We have a genuine respect for the art and science of massage therapy, and are committed to providing a supportive environment where you can further develop your...
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Autocad Operator
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Data Analyst/Excel Operations
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Industrial Hygienist
Our client, located in Norcross, Georgia, has an immediate need to fill the position of Industrial Hygienist. Job Description This Consulting Engineering firm looking for an Industrial Hygienist. Certified Industrial Hygienist (CIH) or Industrial Hygienist (IH) who meets the certification requirements to get certified. A Bachelor of Science in engineering or the sciences (environmental, ...
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Electronic Technician
Hello, My name is Mark and I represent The Fountain Group. We are a national staffing firm and are currently seeking an Electronic Technician for a prominent client of ours. This position is located in Norcross, GA.. Details for the positions are as follows:Job Description· Provide test and fabrication support for electronic circuitry on development programs. · Lay out ...
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Laboratory Assistant
URGENT NEED!!!! Laboratory Technician; Atlanta, GA Kelly Services is currently seeking a Laboratory Technician for one of our top clients in Atlanta, GA. As an Laboratory Technician placed with Kelly Services, you would be responsible for executing the assigned food safety, quality, regulatory compliance, and laboratory program tasks within the location to ensure compliance with company, ...
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Sr. Food Scientist 479.15KO
:: If you meet the qualifications listed below and would like more information contact Kristine at 815-756-1221. Attach resume when responding to this posting ::\
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-BS in food science
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Candidates with a Master's Degree in Biological or Chemical Science with 4+ years of clinical laboratory experience are also eligible....
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Product Development Technologist - 1st shift 478.15KO
:: If you meet the qualifications listed below and would like more information contact Kristine at 815-756-1221. Attach resume when responding to this posting ::
The Product Development Technologist will be responsible for supporting Product Development Scientists.
-BS in food science
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Bachelor of Science in Nursing (BSN). Imagine a place where you can improve lives every day -- this place is Northside Hospital....
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Environmental Due Diligence Consultant
Nova is seeking experienced environmental professionals to conduct Environmental Site Assessments (ESAs) and architectural and engineering professionals to conduct Property Condition Assessments (PCAs). These positions entail:Environmental Applicants:Primary responsibilities will be to prepare Phase I Environmental Site Assessments (ESAs) in accordance with ASTM Phase I standards as well as ...
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Laboratory Assistant in Cornelia, GA
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Medical Affairs - Hepatology - Global Medical Director - Atlanta, GA
Lead Medical Affairs efforts surrounding the approval of novel therapeutics treating chronic liver diseases, develop long-term product and lifecycle management plans aimed at ensuring product has adequate support, and manage interactions with key healthcare professionals in multiple global markets.
*** Develop medical communication and dissemination strategies aimed at ...
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Endocrine Product Specialist (Sales Rep) - GA/AL/MS
We are seeking a dynamic and motivated ENDOCRINE PRODUCT SPECIALIST/SALES REP to join our Sales and Marketing team - geography will include Georgia/Alabama/Mississippi.The PositionYou are a highly-motivated individual with the ability to build/grow our business and manage accounts, requiring effective development and maintenance of a customer base, selling Veracyte's solutions and aligning ...
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Hello, my name is Tony Zarlingo. Kelly Scientific® has been providing outstanding employment opportunities to the most talented individuals in the marketplace. We are proud to offer a long-term temporary opportunity to work as a Lab Supervisor at a prestigious Fortune 500® Pharmaceutical company working near Atlanta, GA.This is a 1 year temporary position at 40 hours per week.Important ...
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text:
Enter your Email Address and click 'Continue' Example: firstname.lastname@example.com Submit Cancel Forgot Password Close /GarnetHill/US/Global/Popups/forgot_password Enter your email address:
Women's Fashion
false pdp {"scheme":"product1_cr"}
false pdp {"scheme":"product2_cr"}
Rated 4.5 out of 5 by 93 reviewers.
Rated 5 out of 5 by ExtraOlives Soft Basic Nice soft merino wool - I've got pretty sensitive skin and some merino wools still make me itchy, but this one isn't itchy for me. Plus, I am so OVER 3/4 sleeves! I've got long arms, and these sleeves actually fit. I am 5'5", 114lb, and bought the XS, which has a narrow tailored fit, but is not form-fitting. October 5, 2015
Rated 2 out of 5 by musekings VERY thin, translucent sweater The hand on the merino is very nice but be warned that the cloth is so thin that it required wearing a shirt underneath-- not by itself as pictured on the models. September 27, 2015
Rated 5 out of 5 by domestique Bigger than the cardigan This is a great sweater. Be aware though, if you love the cardigans, they run small and this does not. I think it might even run a little big. Otherwise this is a similar weight and has the same broad range of uses. September 26, 2015
Rated 5 out of 5 by CAMTED Love the color This is a wonderful sweater. A great color/pattern and the fit and material are terrific. September 26, 2015
Rated 3 out of 5 by sunchild disappointed I ordered the Signature Merino Crewneck Sweater on sale because I was drawn to the Aqua Ikat, and owning several of the GH Merino cardigans I knew the quality would be great. It is still a good quality product. Not itchy, well made, sizing is right, BUT! The aqua is almost a fluorescent blue- green and I am one who likes more muted colors. The brightness makes it look cheap. The other thing that turned me off is that the sleeves are very long (down to the middle of my fingers) and they are made for someone with sizable biceps. I would have put up with the sleeves as they are if the color had been more like on my screen and in the catalog. I am sorry to return this. The sale price was right. September 26, 2015
Rated 5 out of 5 by Ampersand5 High style So soft in texture with lovely soft coloring. Classic fit. I am 5'2 about 120 so normally I have to order petite however this sweater had a universal cut to it. Works very well for me. September 25, 2015
(4 Questions : 4 Answers)
Product Q&A
Ask your questions. Share your answers.
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1 answer
Does the new long sleeved merino crew sweater have the same boxy fit and short length as the original signature cardigan?
Age: 45-54
Gender: Female
1 year, 8 months ago
sw wa
Answer:This sweater has a close to the body fit and is longer than the signature merino cardigan.
1 year, 8 months ago
1 answer
Does this crew neck have the same slim fit as the Signature Merino cardigan? Is the length the same or longer than the cardigan style?
Do you suggest ordering up a size?
Age: 55-64
Gender: Female
1 year, 8 months ago
Answer:This sweater has a close to the body fit and is longer than the signature merino cardigan.
1 year, 8 months ago
4 months ago
Answer:Per the manufacturer, we suggest dry cleaning this sweater.
4 months ago
Has anyone had success with hand washing this? I don't understand why the cardigan version is hand washable but the crew neck is dry-clean only…?
Age: Under 25
Gender: Female
4 weeks ago
West Coast
Answer:They should both be Dry Clean (except for the Decorative Ivory Floral cardigan which is the only one that is hand wash).
3 weeks, 5 days ago
1 of 1
1 of 1
Product:29947-Signature Merino Crewneck Sweater


text:
Dr. Jean Y. Park graduated from the Case Western Reserve University School of Medicine in 2002. She works in Carmel, NY and 4 other locations and specializes in Rheumatology. Dr. Park is affiliated with Northern Dutchess Hospital, Northern Westchester Hospital and Putnam Hospital Center.
Medical School: Case Western Reserve University School of Medicine; Graduated 2002
*This physician has multiple office locations. See Office Information for details.
664 Stoneleigh Ave Ste 100
Carmel, NY 10512
400 E Main St
Mount Kisco, NY 10549
500 E Main St
Jefferson Valley, NY 10535
26 S Greeley Ave Ste 2
Chappaqua, NY 10514
15 Fulton Ave
Poughkeepsie, NY 12603


text:
El cónsul de Sodoma
Genre: Biography Year: 2009 DOWNLOAD • WATCH ONLINE
Actors: Jordi Mollà / Bimba Bosé / Àlex Brendemühl / Josep Linuesa / Isaac de los Reyes…
Fascinating journey through the life and work of the prestigious Catalan poet Jaime Gil de Biedma, both marked by sexuality and eroticism. Charismatic and somewhat eccentric, brilliant intellectual with extraordinary sensitivity and member of Barcelona's 'gauche divine' in the 60s, Gil de Biedma liked to describe himself as a 'poet of experience' while he suffered dreadfully from the dichotomy strangling him: bourgeois and executive for a multinational by day, communist and homosexual poet by night.
Watch El cónsul de Sodoma online. El cónsul de Sodoma download for free with trailer and subtitles. El cónsul de Sodoma putlocker, novamov, videobb.
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El cónsul de Sodoma is a really emotional and dramatical movie. If you want to watch an interesting movie online and you like to cry at the end, I think El cónsul de Sodoma online will not disappoint you. I watched this movie and I was just like WOW, it was incredible story and the soundtrack... Great!


text:
Creating a 2-D rendition on canvas requires either the ability to masterfully wield a paintbrush or the strength to flatten a fruit bowl. Discover your artful brawn with this Groupon.
Choose from Three Options
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Armed with any drinks or snacks they'd like to bring, artists aged 12 and older render their own versions of paintings via easy-to-follow instruction. See the class calendar.
Quench Your Palette
The walls of their studio brim with paintings from their classes, ranging from sunlit beach scenes to bolder, more abstract compositions. Specialty classes reach beyond the canvas to adorn household ornaments such as wine glasses, candleholders, and boring old family tintypes.
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837 NE Jensen Beach Blvd
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+15619513589Get Directions


text:
1900 10C PR67 PCGS Secure. CAC....Click the image to load the highest resolution version.
Superb Gem Proof 1900 Barber Dime
1900 10C PR67 PCGS Secure. CAC. Darkly patinated in shades
of green and cobalt-blue, this splendid Superb Gem displays
razor-sharp details on the ribbon bow and wreath. The fields are
mirrored and the devices show pronounced frosty texture. This piece
might qualify as a Cameo if the toning did not dampen the contrast.
Population: 9 in 67, 1 finer (3/11).(Registry
values: N2998) (NGC ID# 23GD, PCGS# 4884)
Vivid Patina, Outstanding Eye Appeal
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text:
Credits & Info
- Date
- 09/10/2006
- File Info
- Song
- 5 min 19 sec
- Score
- 4.39 / 5.00
Related Content
Heard in 10 Entries!
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Licensing Terms
You may not use this work for commercial purposes without making specific arrangements with the artist UNLESS your work is a web-based game or animation, in which case you may use this freely.
- Score:
- Rated 4.39 / 5 stars
- Plays & Downloads:
- 182,140 Plays | 138,315 Downloads
- Share Links:
- Genres:
- Electronic - Techno
- None
Author Comments
Rated 5 / 5 stars
I love it~~ Good work!
Rated 5 / 5 stars
this song alone is what made me start listening to techno and i come back to it after 8 years to thank you for getting me started
Rated 5 / 5 stars
nice :D reminds me of super Mario galaxy super slimy galaxy
Rated 5 / 5 stars
I gotta say I love this from the beginning of it I felt like I was place in a empty sense of isolation in a urban scene and boom the pickup just surges through me like a rushing tidal wave. And I love also how it transcends from calm to active. Nice work.
Rated 5 / 5 stars
This is epic. This would be a great soundtrack for a madness combat animation! :)


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Views Archive
Get Your Quad Charts Ready!
Get Your Quad Charts Ready!The US DoD has put out this Broad Agency Announcement (PDF) asking for bids for new technology that will help combat a variety of forms of terrorist attacks. Specific technology that the DoD is looking to…
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MIT's Music Service Shut Down
Last week I relayed that two MIT students had set up a new music service that sent music over MIT's analog cable TV network and so was legal under existing laws as MIT was paying normal music distribution fees for…
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Kinsley: Bush Was Wrong
I normally don't blog politics, but Michael Kinsley's column on stem cell policy is just too good to pass up. In this column, Kinsley argues that Bush's famous August 2001 compromise allowing stem cell research on existing cell lines, but…
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Are National Television Systems Obsolete?
Emerging digital and satellite technologies are transforming the world's experience of television. News and information channels such as CNN and al-Jazeera reach audiences across national and regional boundaries. Entertainment formats, programming, films, and popular music are being produced with an…
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More Voting Software Online
Wired News reports that the source code to the Sequoia Voting Systems electronic voting machine was found on an FTP server operated by Jaguar Computer Systems. This happened to Diebold Election Systems' AccuVote-TS software earlier this year. But whereas the…
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Blogger gets fired for posting photo
This came out on October 23rd, but I missed it. A blogger who worked at the MSCopy print shop (on Microsoft campus)eclecticism: snapped this photograph showing a delivery of Apple G5 computers to "deep in the bowels of Redmond."According to…
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There's More Stuff Out There!
Peter Lyman and Hal Varian at Berkeley's School of Information Management and Systems have surveyed the amount of "new" information in the world and concluded that it has doubled in the past three years — that is, the rate that…
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Media Use Enters the Playpen
A new study released this week from the Kaiser Family Foundation, based on telephone interviews with more than 1000 parents of 2-6 year olds, raises new concerns about the amount of time even the youngest American children spend consuming media….
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Reruns on Demand?
Sony's Soap City, a popular website for fans of daytime dramas, will now offer a package of "the most scandalous, memorable and joyous episodes" from the archives of The Young and the Restless, Days of our Lives and As the…
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Napster 2.0
The new "legal" version of Napster launches today. The question is: will anyone care? The latest incarnation of the service aims to capitalize on the file-sharing mania that the original version so controversially unleashed. Napster boasts a database of 500,000…
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text: Alana and Abigail Martinez were two average sisters who were sent from there comfy home in Manhattan, New York to London, England; for the whole summer. The girls thought it was going to be another average summer with their cousins Bethany, Caroline, and Catherine but there is a surprise in store for this family. International Pop Sensations One Direction just so happens to be on break that exact same summer and when a small accident in the London Airport occurs there life is turned upside down as they take a interest in certain girls, Can you guess? Yes, it's Alana, Abigail, Bethany, Caroline, and Catherine who just so happen to be big time One Direction fans but the guy who falls for them are not who they suspect it will be. This will make each girl blinded by their emotions and one action can probably be the wrong one. It is just Five Girls, Five Boys, One place, and One Story.


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3 posts tagged with comedy by amberglow.
Displaying 1 through 3 of 3.
"But there was no cheese. Jeeves died a little inside"
Monkey Fluids --20th century book and magazine illustrations with new text. ; >
Mr. Humphries has measured his last inside leg.
John Inman, RIP --better known as the campy stereotype Mr. Humphries on Are You Being Served? (a gay icon?)
well, they were a big hit at Plato's Laugh Shack
A man, just back from a trip abroad, went to an incompetent fortune-teller. He asked about his family, and the fortune-teller replied: "Everyone is fine, especially your father." When the man objected that his father had been dead for ten years, the reply came: "You have no clue who your real father is."--that's one of the jokes from The Laughter Lover (Philogelos), an ancient Greek joke book published in the 4th or 5th century AD. The New Yorker commented on it, and other old jokes here, stating about one of the possible authors: ... there is some scholarly speculation that the Hierocles in question was a fifth-century Alexandrian philosopher of that name who was once publicly flogged in Constantinople for paganism, which, as one classicist has observed, "might have given him a taste for mordant wit."
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