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Endothelial Notch4 signaling induces hallmarks
of brain arteriovenous malformations in mice
Patrick A. Murphy*, Michael T. Y. Lam*†, Xiaoqing Wu*‡, Tyson N. Kim*, Shant M. Vartanian*, Andrew W. Bollen§,
Timothy R. Carlson*¶, and Rong A. Wang*?
*Pacific Vascular Research Laboratory, Division of Vascular Surgery, Department of Surgery, and Department of Anatomy and§Department of Pathology,
University of California, San Francisco, CA 94143
Edited by Michael A. Gimbrone, Jr., Harvard Medical School, Boston, MA, and approved May 29, 2008 (received for review March 24, 2008)
Brain arteriovenous malformations (BAVMs) can cause devastating
stroke in young people and contribute to half of all hemorrhagic
stroke in children. Unfortunately, the pathogenesis of BAVMs is
in the endothelium during brain development causes BAVM in
mice. We turned on constitutively active Notch4 (int3) expression
in endothelial cells from birth by using the tetracycline-regulatable
system. All mutants developed hallmarks of BAVMs, including
cerebral arteriovenous shunting and vessel enlargement, by 3
weeks of age and died by 5 weeks of age. Twenty-five percent of
the mutants showed signs of neurological dysfunction, including
ataxia and seizure. Affected mice exhibited hemorrhage and neu-
ronal cell death within the cerebral cortex and cerebellum. Strik-
ingly, int3 repression resolved ataxia and reversed the disease
progression, demonstrating that int3 is not only sufficient to
induce, but also required to sustain the disease. We show that int3
expression results in widespread enlargement of the microvascu-
lature, which coincided with a reduction in capillary density,
linking vessel enlargement to Notch's known function of inhibiting
vessel sprouting. Our data suggest that the Notch pathway is a
molecular regulator of BAVM pathogenesis in mice, and offer hope
that their regression might be possible by targeting the causal
molecular lesion.
angiogenesis ? cell signaling ? endothelial cell ? stroke ? cerebrovascular
rupture, leading to stroke and death. Historically, 50% of
BAVMs first present with hemorrhage (1). Development of
noninvasive imaging methods has increased BAVM detection
before hemorrhage, but the efficacy of current treatments,
primarily surgical resection, is questionable. Accompanying risk
of mortality (3%) and morbidity (9%) may outweigh the risk of
rupture itself (2). An ongoing clinical trial (ARUBA) is testing
the hypothesis that invasive treatment may not provide a signif-
icant improvement over the natural course of the disease (2).
BAVMs disrupt normal vessel hierarchy and are caused by the
replacement of capillary beds, which separate arteries from
veins, with enlarged and tangled vessels. The retention of
to the speculation that they form during early brain development
(3). Studies of a more accessible skin arteriovenous malforma-
tion (AVM) in human beings suggests that enlargement of
postcapillary venules precedes AVM formation (4), but exper-
imental evidence is required to validate such mechanisms. An
animal model that mimics the characteristics of human BAVMs,
principally arteriovenous shunting, would open avenues to un-
derstanding their cellular and molecular pathogenesis.
The primary factors regulating vessel hierarchy were once
thought to be hemodynamic forces (5). It is now known that
genetic programs contribute to the development of vascular
emerged as a critical genetic mediator in the differentiation of
arteries and veins. Originally discovered in Drosophila, the
pproximately 600,000 people worldwide harbor brain arte-
riovenous malformations (BAVMs) that can potentially
transmembrane Notch receptor is conserved throughout the
animal kingdom and is best known for its function in dichoto-
mous cell fate decisions through cell–cell communication (7).
Ligand binding to its extracellular domain results in sequential
cleavage events and release of an active intracellular domain
(ICD), which then translocates to the nucleus and initiates
of Notch-ICD is a hallmark of Notch activation.
In the vascular system, Notch receptors and ligands are
expressed in arteries but not veins (8) and are necessary and
sufficient to induce ephrin-B2, a faithful marker of arterial
identity (9–11). Notch loss-of-function mutations impair vascu-
lar development, resulting in arteriovenous shunting in both
zebrafish and mouse embryos (10, 12). Notch gain-of-function
also results in abnormal vascular remodeling in embryos, dem-
onstrating that proper spatial and temporal patterns of Notch
activity are critical for this process (13). Because BAVMs are
thought to occur during brain development (3), we hypothesized
that expression of constitutively active Notch4 would induce
BAVMs during the neonatal period, when areas of the brain, such
as the neocortex and cerebellum, expand by nearly 10-fold (14).
Expression of Constitutively Active Notch4 (int3) in the Endothelium
Results in Hemorrhage, Neurological Damage, and Death in Neonates.
To test our hypothesis that Notch4 activation in the endothelium
of the developing brain would cause BAVM, we expressed int3
specifically in the endothelium from birth in tetracycline-
regulated mice (Tie2-tTA;TRE-int3) (11). Endothelial-specific
expression was demonstrated by a TRE-LacZ reporter (11, 15)
[supporting information (SI) Fig. S1]. In addition, Notch4-ICD
was detected in the nuclei of a subset of mutant but not control
endothelial cells (ECs) (Fig. 1A and B). Because int3 is a portion
of Notch4, the anti-Notch4-ICD staining could not distinguish
int3 from endogenous Notch4. However, the staining did not
detect endogenous levels of Notch4 in the control, suggesting
that staining in the mutant represents int3 expression. The
activity specifically in ECs of mutant mice.
M.T.Y.L., X.W., T.N.K., and T.R.C. performed research; S.M.V. contributed new reagents/
analytic tools; P.A.M., M.T.Y.L., X.W., T.N.K., A.W.B., T.R.C., and R.A.W. analyzed data; and
P.A.M. and R.A.W. wrote the paper.
The authors declare no conflict of interest.
This article is a PNAS Direct Submission.
†Present address: Medical Scientist Training Program, University of California, San Diego,
‡Present address: Vaccine Basic Research, Merck and Company Inc., West Point, PA 19486.
¶Present address: Abbott Vascular, Abbott Laboratories, Abbott Park, IL 60064.
?To whom correspondence should be addressed at: HSW 1618, Box 0507, 513 Parnassus
Avenue, San Francisco, CA 94143-0507. E-mail: firstname.lastname@example.com.
This article contains supporting information online at www.pnas.org/cgi/content/full/
© 2008 by The National Academy of Sciences of the USA
www.pnas.org?cgi?doi?10.1073?pnas.0802743105PNAS ?
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Mutant mice died between 2 and 5 weeks of age (Fig. 1C).
Signs of neurological dysfunction, including ataxia and seizure,
were evident in 27 of 107 mutant mice. Ataxia ranged in severity,
from mice unable to right themselves to those with splayed rear
legs. We occasionally noticed seizure episodes in which the mice
ran wildly before collapsing. All but one of the cases of ataxia
(23/24) and seizure (5/6) occurred before P22, suggesting that
younger mice are more susceptible to these phenotypes.
To investigate potential causes for the neurological dysfunc-
tion, we examined mutant brains and found prominent hemor-
rhage by gross inspection (Fig. 2B) and by histology (Fig. 2 C–F).
Hemorrhage occurred most often in the cerebellum, followed by
the neocortex, but never in brainstem. We examined six of the
ataxic mice and found hemorrhage in the cerebellum (see Fig. 2
C and D). Hemorrhage appeared more widespread than the
neurological phenotype, because we also observed hemorrhage
in all ill mutant mice without detectable ataxia (13 of 13; see Fig.
2E) and none in their littermate controls. Thrombosis, often a
consequence of hemorrhage (16), was seen in hemorrhagic areas
(see Fig. 2 D and F). As with the neurological phenotype,
hemorrhage was most severe in mice ill before P22. Blood was
found in the ventricle of 4 of 11 ill mice before P21 (see Fig. 2F).
Neuronal cell death consistently occurred adjacent to hemor-
rhage (see Fig. 2E), suggesting that hemorrhage led to neuronal
damage. Foci of pyknotic and karyorectic nuclei, often associ-
ated with macrophage infiltrations, were detected in the Pur-
kinje and granular layers of the cerebellum, as well as areas of
the neocortex and midbrain (see Fig. 2E). Signs of calcification,
an indication of earlier cerebral damage, occurred in multiple
cases (data not shown). Therefore, increased endothelial Notch4
activation results in hemorrhage, which likely causes neuronal
damage and ataxia in the mutant mice.
Vessel Enlargement and Hallmarks of BAVMs in Mutants Before
Hemorrhage. Because increased Notch4 activation was targeted
to ECs specifically, we investigated lesions underlying hemor-
rhage and found vascular abnormalities resembling BAVMs.
Vascular perfusion with casting agents (Fig. 3B) or fluorescent
lectins (Fig. 3D and Fig. S2) revealed enlarged and tangled
vessels, hallmarks of BAVMs (1, 17). We verified that arteries,
arterial and venous branches of the middle-cerebral vessels (see
Fig. S2). To determine whether the enlarged vessels had smooth
muscle coverage, we performed immunofluorescence with anti-
smooth muscle ?-actin (SMA) and found strong staining in these
vessels (Fig. S3). Vessel enlargement occurred before hemor-
rhage, neuronal cell death, or signs of neurological dysfunction
(Fig. S4), suggesting that it is a primary defect.
Because hemorrhage occurred primarily in the cerebellum,
followed by the neocortex, and never in brainstem, we tested
whether vessel enlargement correlated with hemorrhage. A
decrease in the proportion of small vessels and an increase in the
proportion of larger vessels were most significant in the cere-
bellum, followed by the neocortex, and least in brainstem (Fig.
4). Therefore, regional enlargement correlated with hemor-
Fig. 1.
mice. (A and B) Increased Notch4 activation specifically in the ECs lining
lectin-perfused vessels of P29 mutant brain was revealed by nuclear staining
of Notch4 intracellular domain (N4-ICD). DAPI-labeled EC nuclei (white ar-
rowheads). (C) Kaplan–Meier survival curve shows that all mutants died by
P36. (Scale bar, 50 ?m.)
Endothelial expression of int3 causes ataxia and death in neonatal
Fig. 2.
hemorrhage (arrowheads) revealed in mutants after vascular perfusion. (C
and D) H&E stained sagittal sections of perfusion fixed cerebellum from a
mutant with ataxia show hemorrhage (green asterisk) and thrombosis (green
arrowhead). (E) H&E stained sagittal section of cerebellar folia from a mutant
without ataxia shows hemorrhage (green asterisk) and adjacent dropout of
granular and Purkinje neurons (yellow arrowhead). Normal neuronal archi-
tecture (yellow arrow) is away from the hemorrhage. (F) H&E stained axial
section of a severely affected mutant shows intraventricular hemorrhage
(white arrow) immediately adjacent to a large, thrombosed vessel (green
arrowhead) and parenchymal hemorrhage (green asterisk) in the cerebrum.
(Scale bars: C and F, 400 ?m; D, 100 ?m; E, 200 ?m.)
Brain hemorrhage occurred in all mutant mice. (A and B) Multifocal
Fig. 3.
and midbrain by vascular casting at P27 (B; white arrowheads) and in the
(A and C) Abnormal vessels were not observed in littermate controls. (Scale
bar, 200 ?m.)
Enlarged and tangled blood vessels developed in all mutant mice.
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www.pnas.org?cgi?doi?10.1073?pnas.0802743105Murphy et al.
Page 3
rhage, supporting the hypothesis that vessel enlargement con-
tributes to hemorrhage.
Endothelial Notch activity decreases vessel density by inhib-
iting sprouting angiogenesis (18). To investigate vessel density,
we quantified CD31 positive vessels. Vessel density was de-
creased significantly in the cerebellum, and progressively less so
in the neocortex and brainstem (see Fig. 4). Vessel density was
inversely correlated with the proportion of large vessels (?50
?m) across all of these regions (Fig. S5), suggesting that vessel
enlargement is linked to the reduction in vessel density.
Arteriovenous Shunting Occurs in all Mutants. The fundamental
defect in human BAVMs is high-flow arteriovenous shunting.
Blood velocity in carotid arteries can double in BAVM patients,
a clinical indication of arteriovenous shunting (19). We mea-
sured carotid blood velocity with high-resolution ultrasound and
found increased carotid blood velocity in all mutants by 3 weeks
of age (Fig. 5A). This noninvasive approach allowed for pro-
gressive measurements in the same animal, demonstrating that
carotid blood velocity increased in a short time window between
P17 and P21 (see Fig. 5A). Furthermore, at P21, velocity in
mutants with neurological dysfunction was significantly higher
than those without neurological dysfunction (2.6 ? 0.5 m/sec;
n ? 6 versus 1.9 ? 0.5 n ? 20, respectively; P ? 0.007). Because
of limitations of this assay, we were unable to examine sick
mutants before this age, and they often exhibited severe neu-
rological defects. As a result, the cohort of mice examined did
not display severe neurological defects, which may contribute to
the minimal increase at P15.
To more directly test for arteriovenous shunts, we performed
a microsphere passage assay (11). Microspheres lodged within
the control brain (Fig. 5B) but bypassed the mutant brain and
lodged in the downstream lung (Tie2-tTA n ? 6; Tie2-tTA;TRE-
int3 n ? 5). This result demonstrates the existence of arterio-
venous shunts in the brain, mimicking the principal defect of
human BAVMs.
To directly visualize arteriovenous shunts, we distinguished
arterial from venous branches with the arterial marker ephrin-
B2. Expression of the reporter in ephrin-B2?/tLacZmice (6) was
strong in the middle cerebral arteries and weak in the middle
cerebral veins (Fig. 6A). In mutants, direct connections between
the arteries and veins were prominent at P21 (Fig. 6B). Similar
arteriovenous connections within the cerebellum were detected
by using a nuclear GFP reporter in ephrin-B2?/GFPmice (20).
Cerebellar interfolial arteries (GFP?) in the control follow a
typical pattern of branching and ramifying into a fine capillary
bed within the granular layer before coalescing into the draining
veins (GFP?) (Fig. 6C) (21). However, in mutants there were
enlarged vascular connections, replacing normal capillaries,
between the interfolial arteries and veins (Fig. 6D). Therefore,
direct arteriovenous shunts were visible in the surface and deep
cerebellar vasculature of mutant mice.
Repression of int3 Rescues Moribund Mice. Taking advantage of the
tetracycline-regulated expression system, we tested whether
frequency of hemorrhage. Shown is immunofluorescence in sagittal sections
CD31 immunostained (black) vessels in each region (per squared millimeter).
? Proportion of vessel with given diameter represents the percent change in
the proportion of small (?7.5 ?m), medium (7.5–20 ?m), and large (?20 ?m)
diameter vessels in each region, relative to average controls. (A and B) Tie2-
tTA (n ? 6) and Tie2-tTA;TRE-int3 (n ? 3); (C, D, E, and F), Tie2-tTA (n ? 6) and
Fig. 5.
(A) Increased carotid blood flow was detected by P21. Maximal (systolic)
carotid blood velocity was measured by pulsed-wave Doppler ultrasound.
Tie2-tTA and Tie2-tTA;TRE-int3 (n ? 9 and n ? 6) at P15; (n ? 12 and n ? 14)
at P17; (n ? 16 and n ? 26) at P21; (n ? 18 and n ? 12) at P23; (n ? 14 and n ?
P21 (***, P ? 0.00005). Typical Doppler traces from mice at P21 (A Right). (B)
Brain arteriovenous shunts developed by P19 as shown by fluorescent micro-
sphere passage. The microspheres bypassed the brain and lodged in the lung
in the mutants but not controls. BF, Bright-field; FITC, green fluorescent
All mutants developed shunting and arteriovenous malformations.
Murphy et al.
August 5, 2008 ?
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Page 4
repression of the causal genetic lesion could reverse disease
progression in mutant mice. Vascular enlargement and shunting
occurred in all mutant animals by P21, so this time-point was
chosen for a regression study. Littermates were allowed to
express the int3 transgene from birth until P20 or P21, at which
time sick mutants were fed doxycycline (Dox), a more stable
derivative of tetracycline (Tet), to repress int3 expression. No
mutants survived to P32 without Dox treatment (n ? 19), but 8
of 9 mutants fed Dox appeared outwardly healthy and active at
P35, and were healthy until killed by P64 (Fig. 7A). The one
exception was a severely affected mutant that died in the first day
of treatment.
To investigate the reversal of brain abnormalities specifically,
examined was resolved within just days of transgene repression
(Fig. 7B). These findings suggest that repression of endothelial
int3 expression, the causal stimulus, recovers brain function in
mutant mice.
We demonstrate that increased Notch4 activity causes vascular
abnormalities with the characteristics of BAVM, implying that
increased Notch4 activation is a potential molecular cause of
human BAVM. Our findings also suggest that regression of this
devastating disease may be possible by targeting the causal
molecular lesion. The activated Notch4 model provides a pow-
erful platform for the study of the pathogenesis and regression
of BAVM-like lesions. In addition, our studies of pathogenesis
in this model provide experimental support for the hypothesis
that BAVMs occur in the developing brain and that they can be
induced by vessel enlargement.
Notch4 Activation Is a Candidate in the Molecular Regulation of BAVM
Pathogenesis. Molecular etiology of BAVM is unknown. Muta-
tions in the TGF-? signaling pathway, primarily loss-of-function
ENG (endoglin) and ACVRL1 (ALK-1) alleles, have been asso-
ciated with BAVMs (22). However, these mutations are impli-
cated in only ?2% of all BAVMs (23). We found that consti-
tutively active Notch4 causes BAVM-like lesions promptly in
neonates, identifying activation of the Notch4 pathway as a
potent molecular candidate in the development of human
The primary characteristic of human BAVMs is the develop-
ment of arteriovenous shunting (1). In patients, this is detected
by arteriography, the gold standard in the diagnosis of this
vascular lesion (24), and also by ultrasound measurement of
increased carotid blood flow (25). We demonstrated the pres-
ence of arteriovenous shunting by similar methods in mice,
including anatomical connections between arteries and veins,
functional passage of beads too large for capillaries, and in-
creased carotid blood velocity.
Enlarged and tangled vessels are another key characteristic of
BAVMs (1). We detected massively enlarged and tangled vessels
by perfusion labeling with a casting agent and fluorescently
labeled lectins. In contrast to human BAVMs, which usually
develop as a single nidus (23), the vascular abnormalities in
because of the expression of the int3 transgene throughout the
Hemorrhage, often resulting in neuronal damage and ataxia,
is the most feared consequence of human BAVM (26). All of the
sick mutants examined developed brain hemorrhage, often with
adjacent neuronal cell death. Approximately 25% of the mice
developed neurological deficits, including ataxia and seizure.
Repression of int3 Expression Leads to Recovery of Sick Mice. It is
commonly thought that natural regression of BAVMs does not
occur (27); instead they increase in size and eventually rupture
(26). Regression of BAVMs has been reported in only a handful
of cases, which are thought to be caused by thrombotic occlusion,
a naturally occurring event similar to the embolizing treatments
used clinically (27). The recovery of brain function in our mutant
mice that resulted from molecular intervention is provocative.
One potential caveat is that Dox, which is used to repress
transgene expression, can also inhibit both vessel growth and
hemorrhage (28–30), and may therefore contribute to recovery.
Because the dose of Dox for transgene repression is similar to
Fig. 6.
surface and in the parenchyma of mutant brains. (A and B) X-gal staining of
ephrin-B2?/tLacZbrains revealed the middle cerebral artery (MCA, red arrow-
arterial branches of the MCA (A) and venous vessels (V) in Tie2-tTA;TRE-
int3;ephrin-B2?/tLacZmice (B?). (C and D) Fluorescent images of sagittal cere-
bellar 100-?m thick sections show enlarged connections (white arrowheads)
between GFP-labeled interfolial arteries of Tie2-tTA;TRE-int3;ephrin-B2?/tLacZ
50 ?m.)
Enlarged arteriovenous connections developed on the meningial
Fig. 7.
sion of int3 with Dox at P20 or P21 allowed the survival of mutant mice, as
shown by the Kaplan–Meier curve. Tie2-tTA;TRE-int3 Dox-off (n ? 23) and
Tie2-tTA;TRE-int3 Dox-on (n ? 15) at 20 days; Tie2-tTA;TRE-int3 Dox-off (n ?
0) and Tie2-tTA;TRE-int3 Dox-on (n ? 5) at 40 days. (B) Ataxia was resolved by
int3 repression. Still frames were taken from a movie of a severely ataxic
mutant at P19, and upon its recovery at P22, after 3 days of Dox treatment.
Repression of int3 resolved ataxia and prevented death. (A) Repres-
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Page 5
that used for inhibition of vessel growth, we are currently unable
to delineate these two effects. However, similar Tet-regulated
mouse genetic approaches were used to show that tumors regress
after repression of transgenic oncogene expression, inspiring the
oncogene addiction theory (31–33). It is possible that Notch4-
induced BAVM-like lesions require continuous Notch4 activa-
tion for their maintenance.
repression of transgene expression or because of its direct effect
on vessel growth and hemorrhage, the recovery of the ill mice is
evocative. Although further studies are required to determine
whether this is a result of regression of the BAVM-like lesions,
lesion may lead to regression of the disease.
High Penetrance and Rapid Onset of BAVM-Like Lesions in the Acti-
vated Notch4 Mutants Provide a Robust Platform to Model Human
BAVM Pathogenesis. Our transgenic mice provide a much needed
mouse model for the study of BAVM pathogenesis. The study of
the cellular and molecular etiology of BAVMs is limited by the
lack of a good animal model for this disease (34). Mice with
mutations in endoglin or ALK-1 have advanced the understand-
ing of molecular contributions to AVMs. Autosomal dominant
mutations in these genes are responsible for the human hered-
itary hemorrhagic telengectasia (HHT) (22), which involves
AVMs primarily in the skin, liver and lung, along with a 4%
incidence in the brain (35). Of 22 endoglin?/?mice that devel-
oped an HHT phenotype, 8 showed signs of brain hemorrhage
by ?40 weeks of age (36). Vascular brain casts showed small
clusters of capillaries or aneurismal dilations in 3 of 10 endog-
lin?/?mice (37). Cerebrovascular abnormalities are also ob-
served in ?2% of Alk-1?/?mice (38). However, it is unknown
whether either the endoglin?/?or Alk-1?/?mice develop arte-
riovenous shunting. Therefore, our activated Notch4 mice pro-
vide a model for molecular understanding of both the develop-
ment and potentially the regression of BAVMs.
Notch4 Activity May Selectively Induce BAVM-Like Lesions in Growing
Brains. BAVMs typically affect children and younger people.
Based primarily on the belief that BAVMs retain embryonic
vascular structures, it has been speculated that BAVMs occur
during brain growth (26). Our data supports the theory that the
growing brain is more vulnerable to BAVM formation. First, we
found that hallmarks of BAVM are prominent when the trans-
gene is expressed in the neonatal period, when the brain grows
rapidly, but not when it is expressed in the postweaning mouse
(11), when brain growth is reduced. Second, we found that vessel
enlargement was greatest in the cerebellum and cortex and least
in the brainstem, which correlates with the greater growth in the
cortex and cerebellum than in the brainstem at this stage (39).
Third, when AVMs occur in older mice, they appear to affect the
liver, skin, and uterus, three actively remodeling tissues in adult
mice (11). These data support the hypothesis that Notch func-
tions in a temporal and spatial specific manner, and that the
developing brain is particularly susceptible to Notch4-mediated
formation of BAVM-like lesions.
cellular mechanism of BAVM formation is currently unknown.
A few models, however, are proposed. The most favored model
is that BAVMs represent a failed transition from embryonic
brain vasculature to adult brain vasculature (3). Supporting
evidence for this theory has rested heavily on the interpretation
of the morphological structure of adult BAVMs. Another theory
is that BAVMs are not necessarily congenital abnormalities, but
can form de novo through vessel enlargement. This theory is
based on the observation that enlargement of postcapillary
venules is the first observable phenotype in the development of
AVMs in human skin (4).
Our data support the theory of de novo formation of BAVMs
through vessel enlargement in the context of growing brain. We
are able to measure carotid blood flow, a sensitive indicator of
brain arteriovenous shunting, longitudinally in our mutant mice.
By this measurement, we were able to show that in individual
mutant mice, there was no shunting at P15, but that shunting
increased dramatically over the subsequent 6-day period. In
addition, the diameters of all vessels, including those at the
capillary level, were enlarged. The enlargement of vessels before
brain hemorrhage or neurological phenotypes suggests that it
plays a causal role in the development of BAVM-like lesions.
Thus, our data support the hypothesis that vessel enlargement
promotes the development of BAVMs.
An alternative possibility is that Notch activation disrupts the
arteriovenous interface by increasing arterial specification. We
and others have shown that Notch activity promotes arterial
specification in ECs in vivo in embryonic and adult mice, as
determined by the expression of arterial markers, such as
ephrin-B2 (9–11). In this study we found that in neonatal brain
endothelium, Notch4 activity increased ephrin-B2 expression in
capillaries and veins. Interestingly, expression was most mark-
edly increased in direct arteriovenous connections, suggesting
that induction of arterial specification in these ECs may disrupt
the proper formation of the capillary bed connecting the artery
and vein.
Notch Activity May Promote Vessel Enlargement by Inhibiting Sprout-
ing. We previously reported that Notch activity promotes vessel
enlargement (11), and others subsequently demonstrated a
reduction in vessel density (18), but the relationship of these
functions is unclear. We observed a correlation between vessel
enlargement and vessel density, suggesting that Notch's role in
enlargement and BAVM development. Supporting our finding,
it was recently reported that retroviral expression of the Notch
ligand Dll4 increases lumen diameters and reduces vessel density
in grafted tumors in mice (40). The observation that impaired
endothelial sprouting can result in vessel enlargement was first
made in mice exclusively expressing the VEGF120isoform (41).
In both the VEGF mutant and our previous studies (11),
proliferation was not significantly altered, suggesting that in-
creased vessel size may simply reflect the retention of cells that
would have otherwise contributed to sprouts. Therefore, Notch
activation may promote vessel enlargement by inhibiting vessel
In summary, we believe that this work opens an area of
research to advance the molecular understanding of BAVM
pathogenesis and provides hope for the treatment of this dev-
astating disease.
Mice. Tet sucrose solution (0.5-mg/ml Tet, 50-mg/ml sucrose, Sigma) was
administered to pregnant mothers from plugging, and withdrawn from pups
at birth, and Dox (200-mg/kg diet, Bio-Serv) diet was administered to mutant
mice at P20–21 as we described (11, 42). All animals were treated in accor-
dance with the guidelines of the University of California San Francisco Insti-
tutional Animal Care and Use Committee.
as we described (11). Casting and FITC-lectin staining were performed as we
described (11). Cy3-streptavidin (Jackson Immuno) was used to detect biotin-
?g of each in 200-?l PBS). Immunostaining was performed with anti-CD31,
anti-?-SMA, and anti-Notch4 (Upstate) according to our published protocols
(43, 44).
Murphy et al.
August 5, 2008 ?
vol. 105 ?
no. 31 ?


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*PLEASE NOTE WHILE THIS POSITION IS VIRTUAL IT DOES REQUIRE THE CANDIDATE LIVE IN THE SAN FRANCISCO BAY AREA.* Do you love developing relationships, partnerships and exceeding sales targets? Do you ha…
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Graduate With No Debt — Virtual
Graduate With No Debt is a 501 (c)3 non-profit organization offering educational entrepreneurial training through paid internships giving students of all ages real world experience. We are seeking mot…
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Accounting Manager
cox-enterprises — Virtual
Must have a high school diploma or equivalent Must be at least 18 years old Must have excellent written and verbal communication skills with an attention to detail and ability to adapt communication …
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Belkin Smooth FolioSmooth Folio for iPad 2
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Top Network Posts
- 25Proof of triangle inequality
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- 13Is every element in a set also an element of the powerset?
- 13Embeddings $A → B → A$, but $A \not\cong B$?
- 12Taylor expansion of $(1+x)^α$ to binomial series – why does the remainder term converge?
- 11Prove that G\S generates G
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Actually, I have already moved on, to B&W, C-41, MF (and even digital for some things.... ).
I've had a great year with K-14, it's demise made me make the effort to go to shoot many places, events and subjects, and to enjoy two extra holidays, which I might not have done otherwise. How about you?


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$15.99Close-up and macro photography is one of the most popular types of photography among digital photographers today. After the big-range zoom, a macro lens is typically one of the next purchases considered by serious outdoor photographers and amateur photographers looking for close-up capability in all zoom lenses. Yet hobbyist photographers struggle. Many macro shots just don't make it–not sharp,
Choosing the Right Lens for Nature and Landscape PhotographyIn Choosing the Right Lens for Nature and Landscape Photography, author and photographer Rob Sheppard walks you through selecting the lens you need for capturing beautiful shots, whether in your own garden or farther afield in the great outdoors. He defines the basic focal lengths you have to choose from, including wide-angle and telephoto lenses, and he demonstrates how zoom features offer
The Power of Color in Nature and Landscape PhotographyIn The Power of Color in Nature Photography, author and photographer Rob Sheppard teaches you how to master color and create compelling and effective photos every time. He shows how you can use colors to change emphasis in a photo to affect where the viewer looks, and he walks you through the effects of light on color, from front to back light, and gentle to dramatic, so you can get just the
The Power of Black and White in Nature and Landscape PhotographyIn The Power of Black and White in Nature and Landscape Photography, author and photographer Rob Sheppard shows you that creating beautiful black-and-white images involves so much more than just removing color. He begins by discussing how the history and tradition of black-and-white photography affect you as a photographer. He then shares the key elements to making dramatic images in shades of
Photoshop Elements 11 Top 100 Simplified Tips and Tricks
Top 100 Simplified Tips & TricksGet the most out of Photoshop Elements 11 with this simplified guide If you already understand the basics of Photoshop Elements and are eager try out some unique tips and techniques, then you'll love this collection of 100 must-know tips and tricks. Two-page tutorials, full-color screen shots, and step-by-step instructions make it easy to see and follow the directions, helping you to get the very
$15.99Now that you've bought that amazing new DSLR, you need a book that goes beyond the camera manual to capture stunning images. For photographers interested in landscape photography, this guide will help you conquer the fundamentals and take amazing outdoor shots. Pro photographer Rob Sheppard starts with the basics–composition, light, and perspective–and describes how to connect with your scene. He
A Totally Biased Close-Up and Macro Photography Book
Secrets of a working pro that anyone can use, part 1Rob Sheppard put this book together to give you solid information about shooting close-ups, yet not require you to spend days reading in order to use it. In this e-book, Sheppard helps you become a better close-up photographer and not simply throw a lot of information at you that is hard to take in and use. Sheppard knows what that is like. He says he spent years trying to learn as much as he
Adobe Camera Raw for Digital Photographers OnlyExpert photographer Rob Sheppard explains the details of Camera Raw, the steps for using it, the workflow process, and certain best practices that demonstrates how Camera Raw can empower the digital photographer. Encouraging you to use it as you see fit, he explores the enhancements in the newest generation and helps you deal with RAW's limitations, manage white balance and exposure, reduce noise
Digital Photography SimplifiedGo from digital photography set up to print out with this easy-to-understand visual resource Digital photography makes taking pictures easier, but learning the tools to develop and manipulate those pictures can be a bit overwhelming. This full-color guide demystifies the latest digital camera models and their features and inspires you to be more creative with your camera and in your image editing.
Top 100 Simplified Tips and Tricks
Top 100 Simplified Tips & Tricks (Book #22)You already know Photoshop Elements 7 basics. Now you'd like to go beyond with shortcuts, tricks, and tips that let you work smarter and faster. And because you learn more easily when someone shows you how, this is the book for you. Inside, you'll find clear, illustrated instructions for 100 tasks that reveal cool secrets, teach timesaving tricks, and explain great tips guaranteed to make you more
Top 100 Simplified Tips and Tricks
Top 100 Simplified Tips & Tricks (Book #31)A visual guide to innovative techniques with Photoshop Elements 9 Photographers who are familiar with the basics of using Photoshop Elements 9 will love this collection of 100 tips and tricks to getting more from the software. Highly visual two-page tutorials and step-by-step screen shots make it easy to see and follow the directions, enabling you to coax even more functionality from this top
Secrets of selling photos to publications in today's competitive marketSelling Photos is a guide to selling photos to publications and is written by an industry insider, Rob Sheppard. Sheppard has been on both sides of sales, both selling images to publications as well as buying them as an editor. He has seen how the industry has changed and lays it out straight for any photographer who wants to sell photos. This is information that you can't find elsewhere and it


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This report, published by the UK Gay Men's Health Network, highlights the diverse range of health issues affecting lesbians, gay men and bisexual people living in the UK.
In 2000 sexuality orientation was recognised for the first time as a social determinant of health in the U.S. 10-year public health plan 'Healthy People 2010'. This report is the first-ever comprehensive document on the state of LGBT health in the US.
Produced by the Gay and Lesbian Medical Association (GLMA)it lists specific recommendations in the following areas: quality health services, mental health, public health infrastructure, HIV, immunization and infectious diseases, tobacco, injury and violence prevention, and substance abuse.
This article was written by Lee Vickers on Family violence law and legislation; Sexual orientation Volume 3, Number 4 (December 1996)
Australian website focusing on domestic violence with a section specific to gay men and lesbians.
This PDF file is a diagrammatic representation of the effects of social justice issues on GLBTT-SQ health and wellbeing.
by Michael Flood and Clive Hamilton
This study identifies the extent of homophobia in Australia, using data from 24,718 self-completion interviews with respondents aged 14 and over.
Edited by Bec Hanley for the Toronto Seminar Group "(www.jrf.org.uk<)":http://www.jrf.org.uk.
This review of the involvement of service users in the development of an 'evidence base' for health and social care, including guidance on good practice for researchers and research funders.
This publication summarises discussions that took place at a series of seminars held around the UK to discuss a number of key issues:
* involving service users in traditional or mainstream research;
* involving service users in peer review;
There is a dearth of research into the mentalhealth of gay men, lesbians and bisexualmen and women in the UK.
The aims of this paper are to assess rates and possible predictors of mental illness in these groups.
A comprehensive assessment was made of the psychological and social well-being of a sample of gay men, lesbians and bisexual men and women, identified
The hypothesis of this study is that lesbians, gay men and bisexuals (LGB) in England and Wales have a different experience of mental health, quality of life and mental health services than heterosexual people. This study examines these three areas of experience.
This lecture given by The Hon Justice Michael Kirby AC CMG Justice of the High Court of Australia (pictured) at the R G Myers Memorial Lecture 2000.
Looking for a specific report, training resource or poster? Use the GLHV Resource Navigator below to find what you want in our Online Library.
A poster created by GLHV for use in health service provider settings.
More GLHV resources<
Our eNewsletter is sent out every two months with news and information about GLHV's work and other relevant issues.
A woman whose primary emotional and sexual attraction is for other women. The term is often used to designate women who identify as same-sex attracted as opposed to women who have sex with other women but who do not self-identify as lesbian.


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Barbra Streisand Duets With Jimmy Fallon [VIDEO]
On her new album, 'Partners,' Barbra Streisand teams up with some of the greatest voices in music. Her latest duet "partner"? Jimmy Fallon!
On her new album, 'Partners,' Barbra Streisand teams up with some of the greatest voices in music. Her latest duet "partner"? Jimmy Fallon!
Last night (Sept. 9) on 'The Tonight Show,' Britney Spears stopped by to promote her new lingerie line ... and to let Jimmy Fallon sign her up for Tinder.
Most of the time, Jimmy Fallon confines the 'Tonight Show' games to just one or two participants, enough to keep things clean and easy to follow, with plenty of attention for his various guests. On last night's edition, however, the host decided to pump things up just a bit with an all-star take on 'Family Feud' that crowded the show's stage with a metric ton of Hollywood talent.
Just when we thought we couldn't be any more obsessed with Meghan Trainor's 'All About That Bass,' Jimmy Fallon had to come along and make it positively epic.
Everyone knows that heartthrob Adam Levine has the voice of an angel, but did you know that the Maroon 5 frontman also did crazy good impressions?
By now, it seems that every celebrity under the sun has taken the ALS Ice Bucket Challenge -- even your favorite Marvel stars -- but there is still a handful of big names who have yet to take the plunge. One obvious choice? Lindsay Lohan, who seems to have so far slipped through the cracks. Well, at least until one random fan challenged her to the task and then employed Jimmy Fallon to make it happen on 'The Tonight Show.' Oh, you're done for now, Lohan.
Since Jimmy Fallon had former 'Bond' star Pierce Brosnan in the studio, he figured the two go head-to-head in a round of GoldeneEye -- in front of the studio audience!
Jimmy Fallon has never been a teenage girl. Taylor Swift, however, has. Still, the veracity that Fallon brings to his performance as a teen in his fake-talk-show-within-a-talk-show, 'Ew!,' is pretty striking -- he somehow seems like more of a teenage girl than his guest, who was a teenage girl mere years ago. It's madness!
A choked-up Jimmy Fallon took to 'The Tonight Show' last night to pay tribute to the recently passed Robin Williams, a highly emotional affair that should help squeeze one last round of tears out of Williams' many shocked and saddened fans.
Poor Nick Cannon. Perhaps the 'Tonight Show' guest realized he'd want to hide after last night's show, thus deciding to kit himself out in a camouflage suit before he even hit the stage. Cannon was paired up with Megan Fox during the latest round of Jimmy Fallon-approved Pictionary, and the pair turned in a ... well, it was a pretty dismal performance.


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Cameron 3 Drawer Nightstand - Traditional CherryItem # HN-ELET024
- Large moldings, felt-lined hidden top drawer
- Select hardwoods and birch veneers
- Traditional cherry finish
- 3 drawers with antiqued brass knobs
- Dimensions: 30W x 18D x 30H inches
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The Elements Fine Home Furnishings Cameron Collection
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Comments about Cameron 3 Drawer Nightstand - Traditional Cherry:
We received two nightstands by FEDEX promptly after ordering online, but one had some flaws in the finish so we notified Hayneedle. They immediately sent a replacement and provided the shipping label to return the flawed item. Unfortunately, the replacement also had flaws in the finish. Hayneedle did not hesitate and sent a third one that was found to be perfect for our bedroom suite. We're very satisfied with our purchase. Just be careful and inspect your product.
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- Beautiful finish
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- Full Slide-out Drawers
- Secret Top Drawer
- Slides Need Lubricating
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Comments about Cameron 3 Drawer Nightstand - Traditional Cherry:
We received the 2 nightstands we ordered and when assembling it found that the bottom drawer of one was not operating properly. And any attempt to get the slides working was futile. Hayneedle was able to contact the manufacturer and get a replacement nightstand to us within a week. We had to return the flawed one before we got that order refunded. But we're now very satisfied with our order and the furniture is absolutely beautiful.
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Visit our other Wisconsin Historical Society websites!
Refine or narrow your results by clicking any term in the gray boxes below.
Expand your results by removing any breadcrumb in a gray outlined box above. Click on the blue "X" to remove the term.
Narrow your results by entering a keyword into the search box above. Then click the "Search Within" checkbox, then click on the orange "Search" button.
If you did not find what you are looking for or you did not get any results, try changing your search term(s). See our Search Help page for more tips.
Date: 1901
Description: Interior of the Wisconsin Historical Society's library with a view of the card catalogs.
Date: 1969
Description: Father Groppi speaking at demonstration in the Wisconsin State Capitol Assembly Chamber.
Date: 1936-02-26
Description: Three men in CUNA (Credit Union National Association) mailing room, Raiffeisen House, located at 142 East Gilman Street.
Date: 1952-07-30
Description: A new, two-color, lithograph press, capable of running off over 7,000 impressions an hour with color perfectly registered, is examined by Walter Larson (le...
Date: 1972-01
Description: Exhibition poster for Maryam Gossling's M.F.A. show in the 7th Floor Gallery in the University of Wisconsin-Madison Humanities building. Held January 1 - 8...
Date: 1899 ca.
Description: East entrance of State Historical Society of Wisconsin showing the decorative details of the building. This detail shows the four lion's heads and columns.
Date: 1955-02-02
Description: Estah Cummings and Evelyn Milliff, members of the Ways and Means Committee of the Madison Business and Professional Women's Club, in the kitchen preparing...
Date: 1950-08-08
Description: Portrait of Professor Joseph Hirschfelder, University of Wisconsin chemist. He also is project director for the U. W. Naval Research Laboratory, member of...
Date: 1952-04-16
Description: Portrait of two cousins, grandchildren of Dr. and Mrs. Frank L. Weston. On the left is Peter Johnson, age 5-months, son of Mr. and Mrs. William E. Johnson....
Date: 1958-12-29
Description: A check for $400 is being handed over to the State Department of Public Welfare by the Associated Optimists Clubs to help in a study of "hard-to-reach yout...
Date: 1955 ca.
Description: A nighttime view of the Wisconsin State Capitol as seen from Monona Avenue (South Wisconsin Avenue until December 1, 1877; Martin Luther King, Jr. Boulevar...
Date: 1958-08
Description: Group portrait of Badeau Plumbing's Midget baseball team, winners of the East Midget Central No. 3 championship. Batboy seated in front: Steve Badeau. Knee...
Date: 1935-10-26
Description: The intersection of State, Johnson and Henry streets, 245 State Street.
Date: 1960-07-17
Description: The Sun Prairie Home Talent baseball team has a record of 10 Eastern Sectional victories this season in its bid for a third straight Home Talent League cha...
Date: 1954-02-27
Description: Group portrait of the Wisconsin State Journal staff with correspondents who were recognized for at least 20 years of service at the Wisconsin State Journal...
Date: 1918 ca.
Description: Wisconsin State Capitol from Monona Avenue, now Martin Luther King, Jr. Boulevard. Automobiles are parked in the center of the avenue.
Date: 1948-05-24
Description: County traffic officers Ray Case (left) and Henry Gerry are placing Elmer C. Evans, with head and face lacerations, in an ambulance to take him to the Jack...
Date: 1990
Description: Norwegian-born Colonel Hans Christian Heg of the 15th Wisconsin Infantry was mortally wounded at Chickamauga on 19 September 1863 and died the next day. Th...
Date: 1950-11-17
Description: Susie Burns and Judy Berger dance the roles of the Russian dancers in 'Trepak' in 'The Nutcracker' on stage at the Masonic auditorium.
Date: 1921 ca.
Description: This map, oriented with north to the upper right, shows roads, railroads, and points of interest. Lake Mendota, Wingra, Monona, Waubesa, Mud, Hook, Island,...
If you didn't find the material you searched for, our Library Reference Staff can help.
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© 1996-2015 Wisconsin Historical Society, 816 State Street, Madison, WI 53706


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Hotel Merida Santiago
No prices found through our partners. Please contact the business directly or check some of our recommended alternatives.
How has this b&b rated in the past?
- Continental Breakfast
- Minbar in room
- Airport Transportation
- Suites
- Wireless internet connection in public areas
- Refrigerator in room
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Travel Blogs from Merida
Bus Day
... portable. The best taquerias are those with long lineups, which means high turnover so the meat won't have been sitting out in the sun all day. Definitely avoid those by the Oaxaca bus station. (See Friday's entry for why.)
Speed Bumps
Mexicans sure love their speed bumps. They lay down thousands of them, on side ...
UXMAL Mayan SIte
... recorded. It was popularised after one of these was recounted by John Lloyd Stephens in his influential 1841 book, Incidents of Travel in Central America, Chiapas, and Yucatan. According to Stephens' version, the pyramid was magically built overnight during a series of challenges issued to a dwarf by the gobernador (ruler or king) of Uxmal. The dwarf's mother (a bruja, or witch) arranged the trial of strength and magic to compete against the king.1
The Nunnery Quadrangle ...
I am finally here
... from food to small wooden sculptures.
We arrived at the Casa Azul Hotel Munumento Historico and went straight to bed. As soon as i woke up the next morning and looked outside this was the view from just outside my door. The hotel is a lovely little place were everyone seems very friendly and kind. If all the rooms were booked up by the a school group this hotel would work well because then ...
... surreal!! Our guide instructed us to get out and do a pin drop into the 10 metre deep crystal clear water below. I was like "are you mad", but then I was like "ok lets do this". With only a small torch lighting the water and the cave wall so close I could almost touch it, I jumped in with a yelp that echoed through the cave. Everyone cheered and I looked up and saw Stuie's face, he looked so proud, it was really cute. As we left the cave I though to myself "ok ...
Last day at the worksite
... scintillate) words, and tried to explain southern accents while Eduardo laughed at us. Then sure enough, Eduardo's sister and brother-in-law go zooming by on their motorcycle, and his teenage nephew ends up waiting for the bus with us and we say goodbye to him as he gets off near our houses to go to school (school hours are very short and go in chunks throughout the day with rotations of kids to accommodate all of the students) closer to Merida.


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Woodridge, NY – BD"E: Orthodox Jewish Child Dies After Being Struck By Vehicle
A 6-year-old girl, a resident from Lakewood, passed away Sunday afternoon after being struck by a vehicle on Tabaczynski Road, in front of the Village Park Colony in Woodridge, NY.
Hatzolah responded to a call of a child struck by a vehicle shortly after 3:10 pm and found the child unconscious.
Paramedics first requested a chopper to transport the child to Westchester hospital, but canceled it moments later. The child was transported to Harris Hospital where she was pronounced dead.
The Fallsburg Police Department and State Police are on the scene investigating.
Update, 7:30 pm, the Lavaya for Rivka Neiman will be 11:30PM in Lakewood at 7th St Chapel.
More from Catskills News
Lamm Scores Victory in Annexation of Mamakating Land Onto Bloomingburg
After years of local infighting, lawsuits and accusations of discrimination, developer Shalom Lamm has scored a final victory in his bid to expand the village of…
Fed Judge Sides with Hasidic Community In Discriminatory Suit Against Mamakating, Bloomingburg
(Reuters). A federal judge ordered two New York municipalities in the Catskills foothills to face a lawsuit accusing them of discriminating against Hasidic Jews by trying to stop…


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Finish Ordering to Complete Your Exchange Request
Your prepaid return-shipping label will be sent only after you complete your replacement order.
1. Request an Exchange
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Brand: Softspots
Style: Softspots Marathon
Color: White
To email a friend about the Marathon, click here
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Designed in 1746 by Francesco Maria Preti, using the harmonic mean proportion
Foursquare © 2015 Lovingly made in NYC & SF
Cinema Hesperia
Movie Theater
Via San Pio X, 5
"Happy Halloween"
an Architectural Trail in Treviso province
Created by Visit Treviso18 items • 5 followers
Veneto best places
Created by Ms. Ele Elena200 items • 4 followers
Make sure your information is up to date. Plus use our free tools to find new customers.
Download Foursquare for your smart phone and start exploring the world around you!


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For the second home game in a row Bills fans left the Ralph with hopes of a win falling short. It was an in state rival game for New Yorkers at Ralph Wilson Stadium. The Bills took on the Giants but came up short, losing 24 to 10.
Buffalo city officals are applauding the implosion of the former Millard Fillmore Gates Circle Hospital because is clears the way for new residential and retail space. But not all residents are celebrating the future plan.Time Warner Cable News Reporter Kevin Jolly explains why some residents are worried the move could open the door for over-development.
Updated 10/04/2015 07:15 PM


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Toll Free: 1-877-489-7333
(Click here for Popcorn and Ornamental corn)
This sugar-enhanced variety has an exceptional sweet flavor. Because Bodacious slowly converts sugar to starch, it will maintain its outstanding quality up to 10 days longer than most other corns. Stalks grow to 7 feet and produce 8-inch well-filled ears. Packet is 1 ounce, about 150 seeds.
A remarkably sweet, sugar-enhanced hybrid with 8-9 inch ears covered with tender deep yellow, or white kernels. The rugged, easy to grow plants will reach 6-7 feet tall. Will maintain its quality much longer than regular corns.
Organically grown. Originating from an early 1900's heirloom, this tasty bicolor corn matures early and germinates well in cooler soils, making it a great choice for gardeners with short growing seasons. The first open-pollenated bicolor corn, it produces tasty 7 inch ears with yellow and white kernels on sturdy, 5 foot plants.
Direct sow 5-6 seeds per foot, about 1-2 inches deep in rows (a minimum of 4 rows will ensure pollination) 3 feet apart. Thin to 12 inches apart. Make 3 or 4 additional plantings at two week intervals to extend the season. Do not plant near popcorn. When the kernels ooze a milky juice when pierced, they are ready to harvest. If birds are a problem, sow in cell packs and transplant into the garden 1 foot apart. Planting is done from spring through early summer, beginning after all danger of frost, and when the ground has warmed.


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Ibis Mons Centre Gare
No prices found through our partners. Please contact the business directly or check some of our recommended alternatives.
How has this hotel rated in the past?
- Free High-Speed Internet
- Restaurant
- Bar/lounge
- Wheelchair accessibility
- Pets allowed
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TripAdvisor Reviews Ibis Mons Centre Gare
Travel Blogs from Mons
Tracing our family history
... now and if it was not for the small memorial you would drive through the area admiring the scenery and not realising the sacrifice made by our forefathers to keep this country free.
After an emotional visit we made our way to the Thiepval Memorial Visitor Centre where there is an excellent display outlining the history of WW1 and in particular the western front. A short distance from the Memorial Centre is the Thiepval Memorial which is ...
Day 8 - Mons
... depicting the battle of St George and the dragon, and then headed up to the belfry. Sadly the belfry wasn't open so we couldn't climb the 360-odd steps to the top to see the views over Mons, but we were atop the hill that gives Mons its name (Mons = mountain) so the view wasn't all bad! It reminded me of Mary Poppins, being at roof height and seeing the chimneys at eye level. Further out, there are a couple of small hills which are forested ...
"Don't chase me I'm full of chocolate!"
By Paul I think our Brussels, Belgium experience can be summed up a bit after one of our hosts this morning, (on our third day here) said to me "I think we need to make breakfast". I replied "We have some muesli we can eat". Both our hosts then laughed. I replied "WE CAN'T JUST EAT WAFFLES, CHOCOLATE, AND DRINK BEER ALL THE TIME. I NEED SOMETHING ...
... gable that looked like the back end of a sailing ship belonged to the shipowners' guild. Even though the Grand Place is a large open space, it feels closed in and a bit claustrophobic (though that could be because of all the people milling around).
I ate lunch at Exki with Christian, Rolinka, Heidi and four or five others. I had a nice pasta salad and lemon water, but I was eyeing Heidi's yummy-looking quiche. Ventured out after that to find the HEMA ...
A Baby Visit To Brussels
... Caroline was very displeased with this brute.
We spent the morning wondering around Brussels and picking up some last minute souvenirs before heading to the train to catch our Eurostar. You might be thinking that the only thing we did in Brussels this time around was have our stuff get puked on. If so, you are correct. Needless to say, we are looking forward to staying at Caroline's grandparents' flat in London...hostels are getting a bit old!


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There's more power in the Photos for iOS app than you might realize.
If you delete a photo from iCloud Photo Library, is it gone forever?
A reader wonders if accidental deletions have huge repercussions.
Photo thumbnails on your Mac turned to black? Here's a fix
A reader finds her photo thumbnails are black rectangles instead of little images. How can this be fixed?
How to merge large iPhoto and Photos libraries
How can you take two photo libraries, one created for iPhoto and one for Photos, with duplicates removed and merge them?
How to create a twirl effect from a photo in Photoshop and Photoshop Elements
Quickly convert an ordinary image into a spectacular series of colorful swirls and twirls
How to transfer a photo album from your iPhone to your Mac
A reader wants to import a single photo album, not everything, to her Mac. But it can't be easily done.
Stop Photos from launching when you plug your iPhone into your Mac
You plug your iPhone into your Mac, and Photos launches—but you don't want that! There's a way to stop this behavior.
How to remove objects in your photos with Photoshop's content-aware tools
It doesn't take expert Photoshop skills to make unwanted objects vanish from your photos.
How to find Photos' image files in the Finder
Where does Photos for OS X stash pictures?
How to AirDrop photos and videos between Macs and iOS devices
The Photos apps for OS X and iOS make it super simple to share photos and videos, even without an Internet connection.
How to delete images from Photos
Readers ask about how to clear out space used by multimedia in the Photos app through deletion and optimization.
More Photos Q&A: Dealing with limitations in multiple libraries, smart albums, and more
Readers (still!) have (many, many!) questions about several features in Photos for OS X.
The hidden editing power of Photos for OS X
Beneath that user-friendly surface, you'll find a set of image-editing tools to help your photos look their best.
How to use the Retouch tool in Photos for Mac
OS X's Photos app has a more powerful Retouch tool than the one in iPhoto.
How to be sure Photos for Mac stores full-resolution images
Photos for OS X when coupled with iCloud Photo Library means you're not sure where high-resolution images live for purposes of local backups.
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Right about now, N.W.A. court is in full effect
Judge Dre presiding
In the case of N.W.A. vs. the Police Department
Prosecuting attorneys are MC Ren, Ice Cube
And Eazy-motherfucking-E
Lyric Meaning
N.W.A is about to have a different style of law and order: One where the Judge is Dr. Dre and the prosecuting attorneys are MC Ren, Ice Cube, and Eazy E. And the Police Department is on trial.
Improve the quality of Fuck tha Police Lyrics by leaving a suggestion at the bottom of the page


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nureign Photographer
Angel LashayModel Female Las Vegas, Nevada, US My MM URL: http://www.modelmayhem.com/angellashay
Mayhem # 522054
About Me
What story will our picture tell? Modeling is not about putting on a baithing suit or some lingerie and taking a sexy picture. Modeling is an art. Every picture does not have to be whats considered pretty or sexy. There are so many things a picture could say and so many stories a great picture can tell. Im interested in working with people that know the true meaning of being a great photographer or a great model.Im looking to expand my port with all sorts of styles.Glamour,Fashion,Urban,Casual,Swimwear etc.Please Email me for faster responses if interested in working with me:email@example.com please no scams , no asking for nude photos etc
Thannk you
nureign Photographer
C E Wiley Studios Photographer
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About StumbleDNA StumbleDNA
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Being a new Vice-Chancellor of this University, it is a matter of great pride and consolation that the University has successfully entered into its 8th year of its age. I would fully utilize all my energies, experience and potentials to make this University a great seat of learning. Most of the infrastructure of the University has been completed and it is my pleasure to express the achievements made -- More...
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From a simple cup of coffee in a Calistoga bakery has sprung an idea that could change the lives of 16 children, and could go on to change the way all 6 million of California's public school students are taught.
In January, Superintendent Esmeralda Mondragon met with Chuck McMinn, president of the nonprofit NapaLearns, to figure out a way to help a small group of pre-kindergartners whose families couldn't afford private preschools yet who were not poor enough to qualify for state-funded classes.
Out popped a simple idea: why not try giving the kids educational technology to help get them ready for school?
The fruit of that conversation wrapped up late last week — an experimental class in which 16 4- and 5-year-olds spent two weeks learning their letters and numbers, and important social skills, armed with brand new iPad portable tablets.
"It's magical," said Tom Torlakson, California's State Superintendent of Public Instruction, after a tour of the pilot class on Thursday. "It's a great way to have students begin to learn languages in a rich way, but it also teaches them 21st-century skills, to use technology to improve themselves."
The idea will expand to a county-wide program next summer and Torlakson said he intends to incorporate lessons from the experiment in a new state-wide educational plan.
For such a grand-sounding venture, the class itself was surprisingly unassuming. It looked pretty much like any preschool class, with blocks, toys, and books scattered around.
But there is a difference.
After reading a book to the class, teacher Kim Floyd asked students to walk up to one of the slim iPads and use their fingers to draw parts of an imaginary monster.
Later, she uses the same machine hooked to a projector to show a short animated version of another story, with words and cartoons flashing by on the screen as the children giggle at the silly pictures and noises.
In between, the kids sat at their desks, eagerly sketching their own imaginary creatures using a simple drawing program. At other times, the kids used their iPads to read books, or play educational games, part of an online package called "Fooststeps 2 Brilliance," designed to teach small children language and number skills while entertaining them as well.
The technology doesn't really change the basics of teaching preschool in terms of the kinds of things kids do or the sorts of lessons they learn, said Floyd, a veteran teacher from Yountville who came to Calistoga Elementary to lead the experimental class. Instead, it allows her to give more individual attention to students, since the software tracks the abilities of students, adjusts the games accordingly, and reports to the teacher how they are doing.
"In the olden days, I would have to teach to the middle of the class, pulling out kids who needed higher or lower attention," she said.
Now she can progress more quickly and confidently through the lessons, knowing which kids are responding well and which need some extra help.
And the children can take control of their own education, she said. When they are done with a particular lesson, they are free to explore the learning software, read books, play games and fill their time. The kids have even discovered things about the computers she didn't know.
One child was typing his spelling words in a colorful font she had never seen. When she asked him where it came from, he was delighted to teach his teacher something new.
"He found a drop-down menu I didn't know about," Floyd said. "He found this whole package of tools."
The computers do not, however, replace traditional pre-school lessons. They still plant seeds to learn about nature, build with blocks to learn about shapes and structure, and read paper books.
"Because at this age, they're not abstract learners," Floyd said. "They're concrete learners and they need concrete things."
Parents seem pleased with the experiment. The children were allowed to take home the machines over one weekend, exposing the families to technology many could not afford. The children also seem to be more excited about learning.
"They seem to be catching onto things faster," said parent Domi Caldrea, whose son Edgar is in the class.
Such technology doesn't come cheap. The iPads for teacher and students cost about $25,000, money provided by NapaLearns. The expanded program next summer will cost about $175,000 for the technology alone.
The remainder of the cost was relatively modest, Mondragon said — about $3,000 to hire Floyd, a few thousand more for supplies and equipment. That cost was defrayed by a private fundraiser held in Calistoga in May that brought in about $8,000.
But whatever the cost, it was worth it, Mondragon said. Many of these children come to school with little background in English, and often the families cannot afford books and technology to give the children a head start on learning.
Without some preschool education, these children would be at a significant disadvantage relative to other students coming into Kindergarten, a gap that can be difficult to make up as children progress to higher grades.
"They would not be integrating quickly into the classroom quickly" in Kindergarten, she said. "To me that's what it's all about — giving them the confidence" to learn.


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, Volume 34, Issue 2, pp 155-165
First online:
Influence of Protozoa and Nutrient Availability on Nitrification Rates in Subsurface Sediments
- E.A. StraussAffiliated withDivision of Biology, Kansas State University, Manhattan, KS 66506, USA
- , W.K. DoddsAffiliated withDivision of Biology, Kansas State University, Manhattan, KS 66506, USA
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Protozoan abundance, nitrification potential, and related factors in saturated subsurface sediments and the overlying soil were compared at a nonfertilized grassland and an agricultural cropland site. In a 6-week laboratory experiment, DOC, ammonium, and protozoan abundance were manipulated in flasks containing groundwater-sediment slurries. Microbial abundance (protozoa, actively respiring bacteria, and total bacteria) and nutrient concentrations (extractable ammonium and nitrate) were measured.
Results from the soil profile analysis showed that protozoan abundance declined with depth at both sites, but significant numbers (392 cells g−1dw) were found in groundwater sediments at the cropland site. Nitrification potential declined with depth at the grassland site and increased with depth at the cropland site. In the laboratory experiment, treatment responses generally were observed within 3 weeks, but had diminished by 6 weeks. Protozoa reduced bacterial populations through the first 3 weeks, but this effect was not significant by week 6. In the cropland sediments, increased net nitrate production occurred in the two reduced protozoa treatments that received ammonium, suggesting that nitrification was occurring and was limited by ammonium. High protozoan abundance in the cropland sediments increased the nitrate flux response, unless DOC was added; in this case, no response occurred. No such responses were recorded in the grassland sediments.
Apparently, appreciable nitrification can occur in some groundwater sediments, if sufficient ammonium is present and DOC availability is low. Furthermore, nitrification can be enhanced when protozoan abundance is elevated. Finally, our results suggest that surface land use practices can alter subsurface nitrification rates and microbial community structure.
- Title
- Influence of Protozoa and Nutrient Availability on Nitrification Rates in Subsurface Sediments
- Journal
Microbial Ecology
Volume 34, Issue 2 , pp 155-165
- Cover Date
- 1997-09
- 10.1007/s002489900045
- Print ISSN
- 0095-3628
- Online ISSN
- Publisher
- Springer-Verlag
- Additional Links
- Industry Sectors
- Authors
- E.A. Strauss (A1)
- W.K. Dodds (A1)
- Author Affiliations
- A1. Division of Biology, Kansas State University, Manhattan, KS 66506, USA, US


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< Return to Religious Affiliation of U.S. Presidents
< Return to Famous Unitarians
The Religious Affiliation of U.S. President
Millard Fillmore previously served as U.S. Vice-President from 1849-50 under Pres. Taylor.
Millard Fillmore was Unitarian. He joined the Unitarian Church in 1831, nearly two decades before he became President upon the death of Pres. Taylor.
From: Peter Roberts, "Millard Fillmore" page in "God and Country" section of "Science Resources on the Net" website (http://www.geocities.com/peterroberts.geo/Relig-Politics/MFillmore.html; viewed 23 November 2005):
Religious Affiliation: Unitarian
Summary of Religious Views:
Fillmore [originally] had no formal religious ties, and indeed had shown relatively little interest in religion, until he joined the Unitarian church in about 1831.
"In my opinion, Church and State should be separate, not only in form, but fact -- religion and politics should not be mingled." -- from a series of campaign speeches, Summer of 1856
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Preparing to spend one more night in Turin.
Push back operation, ready for start up engines in a beautiful day
Flight BMS132 to Bucharest (OTP/LROP) departs runway 26.
A new company for us, connecting Athens (ATH) to Larnaca (LCA) taxies to the active 03L for departure during heavy snowfall! [1200pix]
For the first 10 years of operation, this B737 was applied the sticker of the anniversary.
YR-BAF "Blue Air 10 Years" [1600px]