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I have always approached life and business with passion. I believe that in order to live a fulfilled life one needs to be passionate about all aspects of life; family, spiritualty, hobbies, community, fun time, and of course one's work.&nb...Learn more about me.
I have always approached life and business with passion. I believe that in order to live a fulfilled life one needs to be passionate about all aspects of life; family, spiritualty, hobbies, community, fun time, and of course one's work. We spend a good portion of our waking lives at work, so it made sense to me that one had better love what they do and endeavor to make a difference in this world.
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Life was trying to tell me something. " Judy you love travel, you like to talk about travel, you are detail oriented, it's time to starting helping folks with another type of investment; dream vacations!" I listened. Retiring from financial services in 2005 I embarked on a new journey. The mild mannered funds specialist from Monument Colorado had arrived at her true calling: travel professional for Cruise Holidays. It's been 10 wonderful years, 25 plus cruises, and countless land vacations and resorts since that fateful day.
Someone once asked me "What's Your Dream Vacation?" For me it's about the memories! Having experienced the world of travel both as a customer and a travel professional my first love in travel has to be a river cruise: the five star boutique ships of Uniworld are breathtaking; sipping a glass of fine wine while relaxing on the sundeck of AmaWaterways' AmaCello as she gently traverses the Rhine Gorge; and of course waking to the stunning French balcony views on Avalon Waterways. My love of river cruising has prompted me to specialize in all of the river cruise lines and destinations available to explore.
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As a travel professional I have been able to bring the personal touch to all my clients, from the time they begin thinking about that next luxury river cruise or resort vacation, through the research, booking process, bon voyage, and welcome home, I am with you every step of the way. Perhaps my greatest satisfaction is having repeat clients call me and simply say "book my next vacation" safe in the knowledge that I will customize a wonderful itinerary that will exceed their expectations, be it a river cruise, ocean cruise or premium luxury resort.
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Defined by its expansive urban skyline and deep natural harbor, Hong Kong is one of the most diverse and exciting destinations in the world. Verdant mountains provide a majestic backdrop to the over 1,300 skyscrapers in the world's most vertical city – more people live or work above the 14th floor than anywhere else on earth.
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Hop on a ferry to Lamma Island and spend the day enjoying quaint fishing villages, hiking trails and open-air seafood restaurants.
Visit the inspiring Po Lin Buddhist Monastery on Lantau Island to pay your respects to the tallest seated bronze Buddha statue in the world.
Enjoy a guided Foodie Walking Tour in Central Hong Kong, where you will taste delicious classic Hong Kong cuisine at six local restaurants through SITA World Tours.
Look upon the Terracotta Warriors in Xi'an, roughly 8,000 soldiers, chariots, and horses started in 210 BC—all to help China's first emperor in the afterlife.
Walk along the expansive Great Wall of China and imagine what it was like to construct the wall—at a time without modern machinery to help.
Lan Kwai Fong is one of Hong Kong's most popular nightlife hot spots and home to over 90 restaurants and bars with food diverse as the clientele.
Contact one of our travel experts to start planning your own Hong Kong getaway.
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Skeletal Muscle From Myogenesis to Clinical Relations .pdf
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Chapter 1
Molecular and Cellular Mechanism
of Muscle Regeneration
Kunihiro Sakuma and Akihiko Yamaguchi
Additional information is available at the end of the chapter
1. Introduction
Skeletal muscle contractions power human body movements and are essential for
maintaining stability. Skeletal muscle tissue accounts for almost half of the human body
mass and, in addition to its power-generating role, is a crucial factor in maintaining
homeostasis. Given its central role in human mobility and metabolic function, any
deterioration in the contractile, material, and metabolic properties of skeletal muscle has an
extremely important effect on human health.
Several possible mechanisms for age-related muscle atrophy have been described; however
the precise contribution of each is unknown. Age-related muscle loss is a result of
reductions in the size and number of muscle fibers [1] possibly due to a multi-factoral
process that involves physical activity, nutritional intake, oxidative stress, and hormonal
changes [2-4]. The specific contribution of each of these factors is unknown but there is
emerging evidence that the disruption of several positive regulators [Akt and serum
response factor (SRF)] of muscle hypertrophy with age is an important feature in the
progression of sarcopenia [5-7]. In addition, sarcopenia seems to include the defect of
muscle regeneration probably due to the repetitive muscular damage. Indeed, the group of
Conboy [8-10] indicates that Notch-dependent signaling is impaired in sarcopenic muscle.
Upon tissue injury, the cues released by the inflammatory component of the regenerative
environment instruct somatic stem cells to repair the damaged area [11]. The elucidation of
the molecular events underpinning the interplay between the inflammatory infiltrate and
tissue progenitors is crucial to devise new strategies toward implementing regeneration of
diseased or injured tissues. Regeneration of diseased muscles relies on muscle stem cells
(satllite cells) located under the basal lamina of muscle fibers [12], which are activated in
response to cytokines and growth factors [13]. The current lack of knowledge of how
© 2012 Sakuma and Yamaguchi licensee InTech. This is an open access chapter distributed under the terms
of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits
unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
4 Skeletal Muscle – From Myogenesis to Clinical Relations
external cues coordinate gene expression in these cells precludes their selective
manipulation through pharmacological interventions.
The inflammatory infiltrate is a transient, yet essential, component of the satellite cell niche
and provides the source of locally released cytokines, such as interleukin (IL)-1, IL-6, and
tumor necrosis factor-α (TNF-α), which regulate muscle regeneration [14]. As an inducible
element of the satellite cell niche, the inflammatory infiltrate provides an ideal target for
selective interventions aimed at manipulating muscle regeneration [15]. However, because
local inflammation regulates multiple events within the regeneration process, global antiinflammatory interventions have both positive and negative effects on satellite cells [16].
Thus, it is important to elucidate the intracellular signaling by which inflammatory
cytokines deliver information to individual genes in satellite cells.
Similarly to the embryonic stem cells that build organs, adult stem cells that regenerate
organs are capable of symmetric and asymmetric division, self-renewal, and differentiation.
This precise coordination of complex stem cell responses throughout adult life is regulated
by evolutionally conserved signaling networks that cooperatively direct and control (1) the
breakage of stem cell quiescence, (2) cell proliferation and self-renewal, (3) cell expansion
and prevention of premature differentiation and finally, (4) the acquisition of terminal cell
fate. This highly regulated process of tissue regeneration recapitulates embryogenic
organogenesis with respect to the involvement of interactive signal transduction networks
such as hepatocyte growth factor (HGF), Notch, MyoD, calcineurin, and SRF [17, 18]. This
review aims to outline the molecular and cellular mechanisms of muscle regeneration.
2. Early immune response
Two distinct macrophage populations exist. Classically activated (or type I) macrophages
are induced by interferon (IFN)-γ, alone or in concert with microbial stimuli (e.g.
lipopolysaccharide) or selected cytokines (e.g. TNF-α and granulocyte macrophage colonystimulating factor). They have pro-inflammatory functions: classically activated
macrophages produce effector molecules (reactive oxygen and nitrogen intermediates) and
inflammatory cytokines (IL-1β, TNF-α, IL-6), participate as inducer and effector cells in
polarized Th1 responses, and mediate resistance against intracellular parasites and tumors.
Type I macrophages characteristically and selectively express pro-inflammatory
chemokines, in particular CCL [chemokine (C-C motif) ligand] 3. Alternatively activated (or
type II) macrophages comprise cells exposed to IL-4 or IL-13, immune complexes, IL-10, and
glucocorticoid; they participate in polarized Th2 reactions, promote killing and
encapsulation of parasites, and are present in established tumors, where they promote
progression. Moreover, alternatively activated macrophages are involved in wound healing
and have immunoregulatory functions [18]. The expression of membrane receptors, like the
hemoglobin scavenger receptor CD163, unambiguously identifies type II macrophages [19].
Studies in the rat have shown that type I macrophages are associated with muscle necrosis,
whereas type II macrophages are associated with regenerative myofibers [20]. Of striking
Molecular and Cellular Mechanism of Muscle Regeneration 5
interest, these cells, once within the muscle, apparently acquire a type II phenotype,
revealing a previously ignored plasticity. What are the signals that trigger the shift?
Recognition and phagocytosis of muscle cell debris is probably a critical event. Indeed while
type I macrophages enhance the proliferation of local myogenic precursor cells, type II
macrophages stimulate their fusion and differentiation [21]. Some molecular interactions are
required for macrophage recruitment and function in damaged muscles. The muscle tissue
of mice with a null mutation of CCR2, the CCL2 receptor, undergoes regenerating defects
including fibrosis and calcification after muscle damage. In addition, uPA (urokinase-type
plasminogen activator)-/- macrophages fail to infiltrate damaged muscle [22]. This failure is
associated with defective muscle regeneration, demonstrating that uPA is required for the
homeostatic response to injury. Mice lacking an inhibitor of uPA, PAI-1 (plasminogen
activator inhibitor 1), exhibit increased uPA activity: injured muscle of PAI-1-/- mice shows
evidence of increased macrophage accumulation, and of accelerated muscle repair [23].
Expression of uPA is apparently required for the expression of insulin-like growth factor-I
(IGF-I), a central regulator of muscle regeneration [24]. IGF-I suppresses the expression and
activity of macrophage migration inhibitory factor and the transcription factor NF-κB,
possibly directly regulating the persistence of inflammatory responses [25, 26].
3. Hepatocyte growth factor and neuronal nitric oxide synthase
By 24 hours after muscle injury, satellite cells enter the G1/S phase of the cell cycle [27].
Two factors have been demonstrated to activate quiescent satellite cells. The first is HGF.
Early experiments using single muscle fibers with associated quiescent satellite cells have
shown that growth factors, such as IGF-I and fibroblast growth factor (FGFs), do not
activate satellite cells in fibers [28, 29]. Although IGF-I and FGFs are reported to activate
satellite cells, the studies involved typically used cultures of muscle cells that were not
quiescent; IGF-I and FGFs increase the proliferative activity of satellite cells once they are
activated, even when that activation results during the cell isolation process, i.e. prior to
the plating of cells or fibers for culture. Moreover, platelet-derived growth factor BB,
transforming growth factor-β (TGF-β), and epidermal growth factor do not stimulate
quiescent cells to enter the cell cycle in vitro [30, 31]. Therefore, HGF is the only growth
factor that has been established to have the ability to stimulate quiescent satellite cells to
enter the cell cycle early in a culture assay and in vivo [32, 33]. HGF is localized to the
extracellular domain of un-injured skeletal muscle fibers through a possible association
with glycosaminoglycan chains of proteoglycans that are essential components of the
extracellular matrix, and following injury, quickly associates with satellite cells [34] by
binding to its receptor, c-Met [33].
The second component shown to be involved in satellite cell activation is nitric oxide (NO),
possibly through activation of matrix metalloproteinases (MMP), which induce the release
of HGF, from the extracellular matrix [34, 35]. Studies in vitro and in vivo using rodent
muscle have shown HGF and NO to regulate the activity of many satellite cells [33, 34, 36,
37]. Intriguingly, inhibition of NO production inhibits HGF release, c-Met/HGF co-
6 Skeletal Muscle – From Myogenesis to Clinical Relations
localization, and satellite cell activation [34]. NO is a short-lived free radical that is well
known as a freely diffusible and ubiquitous molecule produced by nitric oxide synthase
(NOSs) from the L-arginine of substrates. In skeletal muscle, neuronal NOS (nNOS, also
called NOS-1) is localized to the sarcolemma of muscle fibers by association at its amino
terminus with alpha1-syntrophin linked to the dystrophin cytoskeleton [38]. The NO radical
is normally produced in very low level pulses by muscles under conditions where satellite
cells are quiescent [39], and the expression and activity of constitutive NOS (nNOS and
eNOS) are up-regulated by exercise, loading injury, shear force, and mechanical stretch. NO
also induces expression of follistatin [40], a fusigenic secreted molecule, known to
antagonize myostatin, thus possibly contributing to the exit of satellite cells from quiescence.
More recently, Tatsumi and Allen [37] proposed the intriguing hypothesis that HGF has
another role in satellite cells. Although, in culture, a low level of HGF (2.5 ng/ml) optimally
stimulates the activation of satellite cells, high levels of HGF (10-500 ng/ml) promote the reentering of quiescence through a concentration-dependent negative feedback mechanism.
Such a role seems to be regulated by the induction of the cyclin-dependent kinase (CDK)
inhibitor p21 in a myostatin-dependent manner. Further descriptive analysis is needed to
elucidate whether HGF and myostatin really do interact in skeletal muscle in vivo. Tatsumi
and Allen [37] suggested the importance and difficulty of monitoring whether or not
extracellular HGF concentrations reach a threshold (over 10 ng/ml) in muscle of living
4. The proliferating process of satellite cells
4.1. Leukemia inhibitory factor
Leukemia inhibitory factor (LIF) is a newly discovered myokine [41], originally identified by
its ability to induce the terminal differentiation of myeloid leukemic cells. Today, LIF is
known to have a wide array of functions, including acting as a stimulus for platelet
formation, the proliferation of hematopoietic cells, bone formation, neural survival and
formation, muscle satellite cell proliferation and acute phase production by hepatocytes [42].
LIF is a long chain four α-helix bundle cytokine, which is highly glycosylated and may be
present with a weight of 38-67 kDa, which can be deglycosylated to ~20 kDa [43, 44]. Several
tissues, including skeletal muscle, express LIF. LIF is constitutively expressed at a low level
in type I muscle fibers [45, 46] and is implicated in conditions affecting skeletal muscle
growth and regeneration [45-47]. Indeed, LIF knockout mice showed a decrease in the area
occupied by regenerating myofibers after crush injury compared to wild-type mice, which
was restored by administration of exogenous LIF [48]. Administration of LIF to the site of
crush injury in wild-type mice increased the area occupied by regenerating fibers with an
associated increase in average myofiber diameter [48, 49]. These original studies suggested
that enhanced regeneration and increases in fiber size occurred, at least in part via
stimulation of the proliferation of muscle-forming myoblast cells, thus providing more cells
to fuse to and increase the size of regenerating fibers.
Molecular and Cellular Mechanism of Muscle Regeneration 7
In 1991, Austin and co-workers demonstrated that LIF stimulated myoblast proliferation in
culture [50], thereby showing that LIF functions as a mitogenic growth factor when added to
muscle precursor cells in vitro. To date, different groups have confirmed this finding and
shown that LIF induces satellite cell and myoblast proliferation, while preventing premature
differentiation, by activating a signaling cascade involving Janus kinase 1 (JAK1), signal
transducer and activator of transcription (STAT) 1, and STAT3 [51, 52]. In line with this, the
specific LIF receptor is primarily expressed by satellite cells and not by mature muscle fibers
[53]. Thus, it seems that LIF has the potential to affect satellite cells rather than mature
muscle fibers.
Earliest descriptions of LIF as a possible mitogen for myoblasts suggested that LIF treatment
increased the number of human and mouse-derived primary myoblast cells in a dosedependent manner after several days of culture, with the earliest increases noticeable after 6
days [50, 54]. There is evidence to suggest that LIF promotes survival of myoblasts and other
cell types [55, 56]. Hunt et al. [57] found that LIF treatment significantly reduced
staurosporine-induced apoptotic DNA fragmentation by 37% and also reduced the
proteolytic activation of caspase-3 by 40% compared to controls. This apoptosis-inhibiting
role of LIF was completely abolished by a PI3-K (phosphatidylinositol 3-kinase) inhibitor
(wortmannin). Therefore, LIF appears to increase the number of satellite cells by promoting
proliferation and blocking apoptosis.
4.2. Insulin-like growth factor-I and MAPK (proliferation phase)
The anabolic effects of IGF-I have been demonstrated in both muscle cell lines and animal
models [58-60]. For example, the addition of IGF-I to cultured myotubes results in an
enlargement of myotube diameters and a higher protein content, while the delivery of IGF-I
either through osmotic pumps or genetic overexpression results in increased muscular mass
in rodents [24, 58]. Mechanical loading also results in skeletal muscle synthesis of IGF-I [61,
62] in vivo, which stimulates gene expression, DNA and protein synthesis, different
transport mechanisms, migration, proliferation, and differentiation [63]. Therefore,
investigators conclude that IGF-I is a critical factor involved in skeletal muscle hypertrophy
in vivo as well as in cultured myotube enlargement in vitro.
IGF-I is thought to induce muscle growth through the increased proliferation of satellite
cells and the enhancement of protein translation resulting in an increase in the rate of
protein synthesis [63, 64]. In addition to stimulating myoblast proliferation, IGF-I stimulates
myoblast differentiation [65]. For example, IGF-I inhibits production of myogenin, a protein
that stimulates muscle cell differentiation, thus allowing increased myoblast proliferation. It
is known that the binding of IGF-I to its receptor, after tyrosine (auto)phosphorylation of the
receptor, results in the initiation of intracellular cascades of various kinase systems.
However, the interplay between the elements of these intracellular signaling pathways has
been described based on results of experiments with skeletal muscle cell types of different
species and under various conditions. Namely, in mouse and rat skeletal muscle
preparations, the involvement of both the MAPK (mitogen-activated protein kinase)
8 Skeletal Muscle – From Myogenesis to Clinical Relations
pathway and MAPK-independent signaling mechanisms, including PI3-K/Akt and protein
kinase C (PKC), was equally documented [66-68]. In primary cultured human skeletal
muscle cells, Czifra et al. [69] demonstrated that the proliferation-enhancing effect of IGF-I
was completely inhibited by the PKCδ-specific inhibitor Rottlerin but not by inhibitors of
the "conventional" PKCα and γ isoforms or by inhibitors of the MAPK or PI3-K pathway. In
addition, overexpression of a kinase inactive mutant of PKCδ prevented the proliferating
action of IGF-I. Furthermore, they showed, in mouse C2C12 cells, that the MAPK inhibitor
PD098059 partially inhibited the action of IGF-I. Taken together, these results demonstrate a
novel, central and exclusive involvement of PKCδ in mediating the action of IGF-I in human
skeletal muscle cells, with an additional yet PKCδ-dependent contribution of the MAPK
pathway in C2C12 myoblasts.
4.3. Notch-dependent signaling
The proliferating process in satellite cells appears to be controlled by Notch signaling during
muscle regeneration [70]. Within hours to days following muscle injury, there is increased
expression of Notch signaling components (Delta-1, Notch-1 and active Notch) in activated
satellite cells and neighboring muscle fibers [8, 70]. Up-regulation of Notch signaling
promotes the transition from activated satellite cells to highly proliferative myogenic
precursor cells and myoblasts, as well as prevents differentiation to form myotubes [8, 71,
72]. Proliferation was decreased and differentiation was promoted when Notch activity was
inhibited in myoblasts with a Notch antagonist, Numb, a gamma-secretase inhibitor, or with
small-interfering RNA (siRNA) knockdown of presenilin-1 [70, 71, 73]. In addition,
mutations in Delta-like 1 or CSL result in excessive premature muscle differentiation and
defective muscle growth [74]. Apparent impairment of Notch signaling occurs in aged
muscle, because expression of the Notch ligand, Delta, is not upregulated following injury
in this muscle. Forced activation of this pathway with a Notch-activating antibody can
restore the regenerative potential by inducing the expression of several positive regulators
(PCNA, Cyclin D1) of cell cycle progression [8, 9].
A recent study revealed that levels of TGF- are higher in aged than young satellite cell
niches [10]. Further analysis showed greater activation of the TGF- pathway in old satellite
cells, and physical competition between Notch and pSmad3 at the promoters of multiple
CDK inhibitors [10, 75]. Furthermore, the decline of Notch1 signaling with age is thought to
be another cause of the decreased regenerative potential of aged skeletal muscle. Indeed,
enhancement of Notch-1 signaling promotes muscle regeneration in old skeletal muscle [8,
9]. Although these experiments suggest a crucial role for Notch1 signaling in satellite cell
function, much remains to be determined, especially regarding the role of Notch3 signaling
during muscle regeneration. Notch3 was expressed in satellite cells, and various structural
and functional differences between Notch3 and Notch1/Notch2 have been reported [76].
More recently, Kitamoto and Hanaoka [77] conducted two very intriguing experiments.
They analyzed muscle after repeated injuries, by generating mice deficit in Notch3 and also
by repetitive intramuscular injections of cardiotoxin (CTX) into the Notch3-deficient mice.
Molecular and Cellular Mechanism of Muscle Regeneration 9
They found a remarkable overgrowth of muscle mass in the Notch3-deficient mice but only
when they suffered repetitive muscle injuries. Analysis of cultured myofibers revealed that
the number of self-renewing Pax7-positive satellite cells attached to myofibers was increased
in the Notch3-deficient mice compared to control mice. Given these findings, the Notch3
pathway might act as a Notch1 repressor by activating Nrarp, a negative feedback regulator
of Notch signaling.
5. The differentiation of satellite cells
5.1. MyoD family
Satellite cell myogenic potential mostly relies on the expression of Pax genes and myogenic
regulatory factors (MRFs: MyoD, Myf5, myogenin, and MRF4). Sequential activation and
expression of Pax3/7 and MRFs is required for the progression of skeletal myoblasts through
myogenesis. Pax7 is expressed by all satellite cells and essential to their postnatal
maintenance and self-renewal [78]. Pax7 induces myoblast proliferation and delays their
differentiation not by blocking myogenin expression [79] but by regulating MyoD [80]. In
parallel, myogenin directly down-regulates Pax7 protein expression during differentiation
[80]. MyoD is required for the differentiation of skeletal myoblasts [81, 82]. In addition,
MyoD null satellite cells showed reduced myogenin expression and absolutely no MRF4
expresion, and displayed a dramatic differentiation deficit [82]. Indeed, muscle regeneration
in vivo is markedly impaired in MyoD null mice [83]. In contrast, Myf5 regulates the
proliferation rate and homeostasis [84]. MyoD can compensate for Myf5 in adults. Myf5
deficiency leading to a lack of myoblast amplification and loss of MyoD induced an
increased propensity for self-renewal rather than progression through myogenic
differentiation. The differentiation factors myogenin and MRF4 are not involved in satellite
cell development or maintenance [84] but induction of myogenin is necessary and sufficient
for the formation of myotubes and fibers.
5.2. IGF-I and calcineurin-dependent signaling
IGF-I positively regulated not only the proliferation but also the differentiation of satellite
cells/myoblasts in vitro possibly through a calcineurin-dependent pathway. Since activated
calcineurin promotes the transcription and activation of myocyte enhance factor 2 (MEF2),
myogenin, and MyoD [85-87], calcineurin seems to control satellite cell differentiation and
myofiber growth and maturation, all of which are involved in muscle regeneration [88, 89].
In fact, our previous study [88] showed a marked increase in the amount of calcineurin
protein and the clear colocalization of calcineurin and MyoD or myogenin in many
myoblasts and myotubes during muscle regeneration. In addition, we showed that the
inhibition of calcineurin by cyclosporine A (CsA) induced extensive inflammation, marked
fiber atrophy, and the appearance of immature myotubes in regenerating muscle compared
with placebo-treated mice [88]. Several other studies indicated such defects in skeletal
muscle regeneration when calcineurin was inhibited [90, 91], whereas transgenic activation
10 Skeletal Muscle – From Myogenesis to Clinical Relations
of calcineurin is known to markedly promote the remodeling of muscle fibers after damage
[92, 93].
Many researchers have utilized CsA, though in different amounts, to determine the
downstream modulators of calcineurin signaling. We found that intraperitoneal CsA
treatment daily at 25 mg/Kg/day enhanced the expression of myostatin and Smad3 mRNA
in regeneration-defective tibialis anterior muscle after an injection of bupivacaine [89]. The
possibility that myostatin is a downstream mediator of calcineurin signaling has been
indicated by experiments with two different transgenic mice [94]. In addition, calcineurin's
pharmacological inhibition caused a decline in the transcription and activation of myogenin
and MyoD during myogenic differentiation by a downregulation of MyoD expression [95].
Considering these findings, calcineurin seems to block the myostatin-Smad3 pathway to
enhance the expression of myogenic differentiation factor (MyoD) during muscle
regeneration in vivo. Using CsA treatment in vivo, recent evidence including that obtained by
our group has also identified Id1 [87, 89], Id3 [87], and Egr-1 [87] as a possible downstream
negative hypertrophic effector target of the calcineurin-NFAT (nuclear factor of activated Tcells) pathway.
FOXO (forkhead box O)-induced expression of Atrogin-1 has been shown to inhibit
calcineurin activity [96]. More recently, the calcineurin variant CnAβ1 was suggested to
block the nuclear localization of the FOXO protein and the expression of several genes
targeted by FOXO [the muscle ring finger-1 (MuRF1), Gadd45a, Pmaip1, and atrogin genes]
in C2C12 myoblasts [93]. In addition, transgenic up-regulation of CnAβ1 expression
promotes the remodeling of cardiotoxin-treated muscle fibers [93]. In cardiomyocytes,
calcineurin directly binds and dephosphorylates (inactivates) Akt; FOXO indirectly activates
Akt by inhibiting calcineurin phosphatase activity [97]. In murine C2C12 myotubes, Akt was
shown to antagonize calcineurin signaling by causing hyperphosphorylation of NFATc1
[60]. Interaction between CnAβ1 and FOXO during muscle regeneration is a very attractive
idea, although it has not been demonstrated in adult skeletal muscle in vivo.
5.3. Serum response factor
SRF is an ubiquitously expressed member of the MADS box transcription factor family,
sharing a highly conserved DNA-binding/dimerization domain, which binds the core
sequence of SRE/CArG boxes [CC (A/T)6 GG] as homodimers [98]. Functional CArG boxes
have been found in the cis-regulatory regions of various muscle-specific genes, such as the
skeletal α-actin [99], muscle creatine kinase, dystrophin, tropomyosin, and myosin light
chain 1/3 genes. The majority of SRF's targets are genes involved in cell growth, migration,
cytoskeletal organization, and myogenesis [100, 101]. SRF was first shown to be essential for
both skeletal muscle cell growth and differentiation in experiments performed with C2C12
myogenic cells. In this model, SRF inactivation abolished MyoD and myogenin expression,
preventing cell fusion in differentiated myotubes [102]. SRF also enhances the hypertrophic
process in muscle fibers after mechanical overloading [103]. For example, we showed that,
in mechanically overloaded muscles of rats, SRF protein is co-localized with MyoD and
Molecular and Cellular Mechanism of Muscle Regeneration 11
myogenin in myoblast-like cells during the active differentiation phase [104]. Recent results
obtained with specific SRF knock-out models, by the Cre-LoxP system, emphasize a crucial
role for SRF in postnatal skeletal muscle growth and regeneration [105], by direct binding of
IL-4 and IGF-I promoters in vivo. These lines of evidence appear to indicate that SRF
modulates the differentiating process of satellite cells in adult mature muscle.
The expression and cellular localization of SRF and myocardin-related transcription factor-A
(MRTF-A) appear to be regulated by several upstream factors including 1-integrin, RhoA,
striated muscle activators of Rho signaling (STARS) [106], and MuRF2 [107]. For instance,
Lange et al. [107] demonstrated that SRF is blocked and relocalized by the nuclear
translocation of MuRF2, which regulates a signaling pathway composed of titin-Nbr1p62/SQSTM1 at the position of the sarcomere depending on mechanical activity. To date,
there has been no attempt to investigate whether titin-Nbr1-p62/SQSTM1 and MuRF2 affect
muscle regeneration. In addition, the mutation of SRF delineated the translocational action
of MRTF-A induced in vitro by STARS, a muscle-specific actin-binding protein [106].
5.4. Wnt-dependent signaling
Similar to Notch signaling, canonical Wnt signaling is critical for muscle repair [108-111]. The
canonical Wnt signaling cascade requires soluble Wnt ligands to interact with Frizzled
receptors and low-density lipoprotein receptor-related protein co-receptors (LRP). This
coordination stimulates phosphorylation of Disheveled and inactivates GSK3's
phosphorylation of -catenin. In the nucleus, the de-phosphorylated -catenin binds to T-cell
factor/Lymphoid enhancer factor-1 transcription factors [112], which may directly activate
Myf5 and MyoD or may upregualte MRF co-activators such as c-Jun N-terminal kinases [113,
114]. It is suggested that Notch activity presides during myoblast proliferation after which
there is a temporal switch to Wnt signaling and subsequent myoblast differentiation and
fusion into myotubes [108]. Inhibiting Notch (with soluble Jagged ligand or with a γ-secretase
inhibitor) or activating Wnt (by inhibiting GSK3 or adding Wnt3a) decreases Myf5
expression and promotes muscle differentiation providing evidence that Notch signaling
needs to be turned off and Wnt turned on for differentiation to ensue [108, 115].
This hypothesis was supported by the finding that aberrant activation of the Wnt pathway
can lead to fibrogenic conversion of cells in different lineages [116-118]. In fact, Wnt
signaling was shown to be enhanced in aged muscle and in myogenic progenitors exposed
to aged serum [116]. To directly test the effects of Wnt on cell fate and muscle regeneration,
Brack et al. [116] altered Wnt signaling in vitro and in vivo. Addition of Wnt3A protein to
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[116]. Conversely, the myogenic-to-fibrogenic conversion of aged serum was abrogated by
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after injury resulted in increased connective tissue deposition and a reduction in satellite cell
proliferation [116]. The authors therefore tested whether inhibiting Wnt signaling in aged
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Sweet Mustard Pickles - an old fashioned recipe - but an amazing taste!
Seeing puffins in Elliston, Canada is unlike anything I've ever experienced. I highly recommend taking a trip here when visiting Newfoundland!
Fasten your seatbelts! This summer, we took an Epic Canadian Road Trip, driving from our home in Kalamazoo, Michigan all the way out to St. John's, Newfoundland (yes, a ferry was involved). Why? Well, I'm the co-founder of Writing Walking Women, and we chose St. John's as the site of our conference this year – MUCH more on that to come! And, our daughter, Lillie (13), said
Marianne Parsons' "Colors to Live By" ~ Exhibitions of the Fibre Art Network
Sweet Mustard Pickles
Did you know that, in some countries, the local inhabitants actually eat the famous, highly photogenic Atlantic Puffin? We don't think that's the case in Newfoundland, Canada, at least we hope not! These adorable birds are elusive and shy, yet some have been able to catch a glimpse of them fleeting along the rocky coastline of Canada's eastern shore. Our illustrators at Anderson Design Group crafted this vintage rendition of the birds, a pleasant nod in our own way to one of Canada's most…
2,790 Likes, 38 Comments - Tour Canada 🇨🇦 (@tourcanada) on Instagram: "🇨🇦Petty Harbour, Newfoundland! Photographer | @_samoscientific_ #TourCanada @TourCanada"
Would get this as a tattoo but without the words
Of crepes and cod.
The East Coast Trail consists of 26 developed wilderness paths along 300km of coastline on the East Coast of Newfoundland, Canada. Bay Bulls to the lighthouse is the first 3.5km of the Spout Path.
Traditional music represents the province's history and culture, and forms a vital link between the past and present.
About quilts and quilting by an avid quilter
A History of Newfoundland [Prowse, Daniel] on Amazon.com. *FREE* shipping on qualifying offers. A History of Newfoundland


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24 piece Disney Flatware/Silverware/Cutlery Set
24 piece Disney Flatware/Silverware/Cutlery Set. 6 each of Fork, Knife, Spoon & Teaspoon. The item "24 piece Disney Flatware/Silverware/Cutlery Set" is in sale since Saturday, July 4, 2020. This item is in the category "Home, Furniture & DIY\Cookware, Dining & Bar\Cutlery\Cutlery Sets & Canteens". The seller is "ebc280800" and is located in Camberley. This item can be shipped to United Kingdom, Antigua and barbuda, Austria, Belgium, Bulgaria, Croatia, Cyprus, Czech republic, Denmark, Estonia, Finland, France, Germany, Greece, Hungary, Ireland, Italy, Latvia, Lithuania, Luxembourg, Malta, Netherlands, Poland, Portugal, Romania, Slovakia, Slovenia, Spain, Sweden, Australia, United States, Bahrain, Canada, Brazil, Japan, New Zealand, China, Israel, Hong Kong, Norway, Indonesia, Malaysia, Mexico, Singapore, South Korea, Switzerland, Taiwan, Thailand, Bangladesh, Belize, Bermuda, Bolivia, Barbados, Brunei darussalam, Cayman islands, Dominica, Ecuador, Egypt, Guernsey, Gibraltar, Guadeloupe, Grenada, French guiana, Iceland, Jersey, Jordan, Cambodia, Saint kitts and nevis, Saint lucia, Liechtenstein, Sri lanka, Macao, Monaco, Maldives, Martinique, Nicaragua, Oman, Pakistan, Peru, Paraguay, Reunion, Turks and caicos islands, Aruba, Saudi arabia, South africa, United arab emirates, Ukraine, Chile, Bahamas, Colombia, Costa rica, Guatemala, Honduras, Jamaica, Kuwait, Panama, Philippines, Qatar, Trinidad and tobago, Uruguay, Viet nam.
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For entreprenuer-turned-artist Sapna Dube, trips to the Amber fort in Jaipur and locations in Kerala, where she saw elephants in captivity being mistreated, provided the spark for a series of paintings, 'Elephant as God'.
Sapna says, "We worship Ganesha as the Elephant god, yet we mistreat the animal by keeping it chained and injuring it. I felt it more acutely when I travelled with friends from Belgium. That gave me a new perceptive on this issue. I decided to work on a project showcasing Ganesha in various forms. My series is created to bring awareness about elephants in captivity, especially those used in temples and processions. It has 21 paintings."
The paintings are titled 'Mouserider', 'Benevolent One', 'Cosmos', 'Lord of Fortune', 'First Among All', 'Pathmaker', 'Patron of the Arts', 'Listener', 'Trunk', 'Single Tusk', and 'Mr. Big'.
Sapna says, "Ganesha is my favourite god. I have enjoyed drawing him in my own manner. I have used colours and artistic influences from across the world. It has taken me six years to complete the project. I have used acrylic paint makers."
Talking about some of the paintings, Sapna says, "They showcase different attributes of Ganesha. For instance, in the 'Doorkeeper' painting, the elephant's head symbolises wisdom and knowledge and he is shown facing doorways to keep out unwanted elements."
'The Pathmaker', that showcases the ability of Ganesha to defeat all obstacles, is one of Sapna's favourites from the collection. She points out, "It's one of the first I painted in the series and embodies the role of the path maker that the elephant essays in the forest. We have assigned the same attributes to Ganesha as well."
The exhibition is on at the Taj Westend till the end of October.


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Governor Cuomo Directs Flags to Half-Staff - September 15, 2014
Governor Andrew M. Cuomo today directed flags on state government buildings to be flown at half-staff on Monday, September 15, 2014 in honor of a World War II soldier from Brooklyn whose remains were recently discovered and transported back to the United States. U.S. Army PFC Bernard Gavrin, 29, was last heard from in 1944 when he was reported missing in action during the battle for Saipan. His official burial will be on September 12 at Arlington National Cemetery in Washington, D.C.
"PFC Gavrin put his life on the line and paid the ultimate price to defend our nation and fight for the freedoms that it is built upon," Governor Cuomo said. "After far too many years, he has returned home and has been granted a proper burial alongside the many other heroes who answered the call. I ask all New Yorkers to join me in honoring PFC Gavrin's memory, his service and his sacrifice."
On June 15, 1944, as part of an Allied strategic goal to secure the Mariana Islands, U.S. forces were ordered to occupy Saipan. After a month of intense fighting, enemy forces conducted a suicide assault, known as a banzai attack. This was designed to inflict as many casualties as possible against the 105th Infantry Regiment, 27th Infantry Division. During these attacks, elements of the 105th Infantry Regiment sustained heavy losses, with more than 900 soldiers killed or injured.
PFC Gavrin was reported missing in action on July 7, 1944.
Governor Cuomo has directed that the flags on all state buildings be lowered to half-staff in honor of and in tribute to our state's service members and those stationed in New York who are killed in action or die in a combat zone.
Effective: Beginning of Business, Monday, September 15, 2014
Return to Full Staff: Sunset, Monday, September 15, 2014
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UNICEF Vacancy for a Maternal Newborn Child Health and Nutrition (MNCHN) Consultant
Nov 14, 2018, 7:43 PM
UNICEF works in some of the world's toughest places, to reach the world's most disadvantaged children. To save their lives. To defend their rights. To help them fulfill their potential. Across 190 countries and territories, we work for every child, everywhere, every day, to build a better world for everyone.
We are recruiting to fill the position below:
Job Title: Maternal Newborn Child Health and Nutrition (MNCHN) Consultant
Job Number: 517889
Locations: Adamawa, Bauchi and Kebbi
Work Type: Consultancy
Duration: The duration of this assignment is 11.5 months
- Under the guidance and supervision of the State Coordinators, FO Team Leader and the Chief of Field Office (CFO), the State Coordinator will work in close collaboration with the Field Office's steering committee to perform his/her job duties and roles
Key Expected Results/Major Responsibilities
State and LGA:
- Work with the SMOH, SPHCDA and other partners to support dissemination of national policy, strategy, guidelines and quality of care model for MNCHN at state and LGA level.
- Support SMOH and SPHCDA by providing technical assistance to the development of good quality state and LGA annual operational plans and their review using the Primary Health Care Mechanism.
- Support dissemination and use of approved training modules and guidelines on selected high impact Integrated Maternal New-born Child Health and Nutrition continuum of care package as defined ( such as- Focused AnteNatal Care (FANC); Skilled Birth Attendance (SBA); Emergency Obstetric and New-born Care (EmONC); Helping Babies Breath (HBB); Post Natal Care (PNC); Integrated Management of acute malnutrition, Infant and Young Child Feeding (IYCF), Micronutrient Prevention and Control, Community Based intervention including  New-born Care and Integrated Community Case Management (iCCM) for diarrhoea, malaria and pneumonia, etc.)
- Support state and LGA level supportive supervision, monitoring of implementation and evaluation of MNCHN programmes.
Health Facilities:
- Priority areas of work to be supported at this level and strategy based on TSS model (Training, Supplies, Supportive Supervision) including but not limited to the following:
- Set up and implement a quality of care model for MNCHN services in health facilities based on integrated selected high impact interventions.
- In collaboration with state and LGA Trainers, organize and conduct trainings for health workers using approved training modules and guidelines on selected integrated high impact interventions.
- Support end user monitoring of equipment and supplies provided to health facilities.
- Participate in on the Job Training, Supportive Supervision and mentoring of health workers and ensure report is timely done and submitted.
Communities, Villages, Households:
- Support setting up/organization of structures for implementation of Community based MNCHN interventions including (but not limited) WDCs, VDCs, training of VHWs, CHEWs, CHWs, CLTS, WASHCOM, MSG, etc.…and monitor respective interventions
- Work in close collaboration with C4D Consultant to promote uptake of selected high impact family care practices jointly with all sections namely Health, Nutrition, Child Protection, Media, Social Planning, WASH, advocacy and communications and WASH.
- Promote uptake of integrated Community Case Management for diarrhoea, pneumonia and malaria as well as other MNCHN community based interventions.
- Support and conduct the end user monitoring of supplies and demand for MNCHN services in communities and households.
- Use and monitor implementation of updated Mama/CHEW/CBNC kits as entry point for promotion of desired health seeking behaviour and link with immunization/polio related activities.
Qualifications & Experience
- University degree in Social Sciences, Public Health and/or related Technical field.
- Two years experience working on child survival issues or health projects.
- Computer skills, including internet navigation and various office applications.
- Demonstrated ability to work in a multi-cultural environment and establish harmonious and effective working relationships, both within and outside the organization.
- Willingness to travel to remote regions of the country
- Initiative, passion and commitment to UNICEF's mission and professional values.
- Fluency in oral and written English is required. Knowledge of another UN working language an asset. Knowledge of local working language of the duty station an asset.
- Knowledge of the latest developments and technology in related fields.
- Ability to make timely and quality judgments and decisions and very good training skills.
How to Apply
Interested and qualified candidates should:
Click here to apply
- Applicants should quote an all inclusive fee
- Only shortlisted candidates will be contacted and advance to the next stage of the selection process
Application Deadline Wednesday 28th November, 2018 by 23:55:00 GMT+0100 (West Africa Standard Time).
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Event: Portland 2020
| Team Leader:
Team Fundraising Goal:
| Amount Raised: $500.00
Ella's Elite has been going strong for fifteen years now... with teams all across the country... everyone working together to raise the much-needed funds for critical Cystic Fibrosis research. With Ella as our leader and inspiration... and with determination in our hearts we're at it again.
Please join us as we work to support the Cystic Fibrosis Foundation's search for better answers and more options.
Our Portland Walk is coming up on Saturday, May 4th at Oaks Amusement Park. We're still on the lookout for dedicated Team Members to fundraise and walk with us. And of course we're busy scouring our world for donations. We'd love to have you be a part of our Ella's Elite... either as a Team Member or contributor.
Please click one of the links above...
Until there's A CURE!!!
Due to the COVID-19 pandemic, indoor events sponsored by the CF Foundation are strictly prohibited at this time. Scheduled outdoor events must adhere to strict requirements to minimize the risk of COVID-19 infection. Events may be subject to change at any time based on guidance from the Centers for Disease Control and Prevention and local health officials.
To minimize the risk of COVID-19 infection, attendees at CF Foundation events must adhere to the following requirements:
Persons with CF should consult their physician before participation in any in-person event as they may be at an increased risk for severe illness from COVID-19. People with CF should maintain a safe 6-foot distance from persons outside of their household at all times.
© 2020 Cystic Fibrosis Foundation. Legal Terms & Conditions | Privacy Policy


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Welcome to My New Hard Cut Concrete Sawing & Drilling Blog
Importance of Hiring the Best Concrete Cutting Contractors
A technique or process of drilling, sawing, and removal of concrete performed by professionals or skilled operators is called Concrete cutting. It is several procedures or services through which concrete is manipulated into different shapes and sizes to better suit various applications.Sydney is one of the most beautiful cities in the entire world. To know… Read more
All You Need To Know About Core Drilling
In the world of construction, there are many instruments being used for various functions nowadays. Think about the different tools used for cutting, drilling, boring, plastering, holding-in-place and likes. Amongst these equipment mentioned above is the core driller. A core driller is a machine or equipment which is equipped with special types and grades of… Read more
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They say that jacks-of-all-trade are masters of none, but the history of Benjamin Franklin dares to say otherwise. A man considered to be a leading visionary of his time, it was no wonder he rose to the ranks and, through…Read More
64 Muhammad Ali Quotes on Life and Success
Inspirational and entertaining, Muhammad Ali is celebrated as one of the best sporting figures of all time not only for his exceptional skills inside the boxing ring but also for his wit. His life was riddled with pain and struggle.…Read More


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Body and Movement Plan
Project Title MOVE 2 DANCE Krumping is a street dance that is characterized by free expressive, exaggerated and highly energetic movement involving the arms, head, legs, chest and feet. In response to the brief I am going to create a photo montage … Continue reading
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Task 6 point of view shot report
A point of view shot is a shot used in film to show what the character is looking at.the point of view hot is usually positioned between a shot of a character looking at something and a shot showing the … Continue reading
Task 5 shot and reverse
shot and reverse Shot and reverse shot is a film technique where two or more shots are edited together, sometimes one character is would be shown starring at another character in off screen space and then the other character would … Continue reading
task 4 cuts examples
match cut a space odyssey-1968- stanley kubrick in this scene we see the ape throwing the bone in air after discovering the use of the bone as tool and then we see another shot of the space station in space … Continue reading
Task 3 continuity editing report
Continuity editing A predominant style of editing in narrative cinema and television. Continuity editing smooths the inherent discontinuity of the editing process. Continuity editing also establish a logical coherence between shots. The 180 degree is a rule filmmakers use to shoot from one … Continue reading
Task 2, 3 point editing system
The three point editing is an editing method which is done by setting three edit points. The first and second point are the in and out points of the clip. In a timeline were the clip fits into they are … Continue reading
Task one non linear and linear editing
Non Linear editing and Linear editing Non linear editing is a method that allows editiors to perform their film and video editing on a computer. In digital editing non linear also allows the editors to access frames in a digital … Continue reading
Case study
The piece of art work I chose is the work of William Hofhuizen the crucifixion(1965). In this painting William uses good color and compositions. The color scheme of this piece of art are the yellow, Ultramarine Blue and Cadium Red. The crucifixion 1965 … Continue reading
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Networks A social network is made up of individuals called nodes. social networks can focus on building and reflecting of social networks or social relations among people. As an online service these can lead to people meeting people who share … Continue reading
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Tiger Woods' mugshot released after arrest in Florida on DUI suspicion
The Tiger Woods mug shot was released shortly after he left the Palm Beach County Jail.
Tiger Woods' mug shot was released shortly after news of his arrest in Florida on suspicion of driving under the influence was released by the Palm Beach County Jail.
Woods, 41, was booked on Monday morning and released on his own recognizance at 10:50 a.m.
• A timeline of Tiger Woods' troubles since 2009
Woods has not played since he withdrew from the Dubai Dessert Classic on Feb. 3. He underwent fusion surgery on his back on April 19. It is his fourth procedure on his back since April 2014. He is expected to miss the rest of the PGA Tour season.


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Old Purple MIDI File
MIDI File Download Old Purple - David Kirkpatrick
Old Purple Karaoke MIDI File in the style of David Kirkpatrick
Old Purple MIDI File is a popular song recorded by David Kirkpatrick and written by David Kirkpatrick. Old Purple is produced and distributed by the team at Hit Trax and available in the following formats: MIDI Karaoke File and MP3 Backing Track. AMCOS Licensed and royalty paid. © 2000 - 2019
#oldpurplemidi #davidkirkpatrickmidi #midi #midifile #midikaraoke #backingtrack


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15 Best Texas Beaches
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Oahu Singles Vacations Written by Stephanie Dula Oahu singles vacations are a great way for singles to get to know other singles in a fun and friendly atmosphere. Singles vacations are planned along several guidelines. For instance, there are single parent vacations, singles vacations according to age range, and singles vacations according to activity preference, just to name a few. Oahu is the perfect place in which to experience a singles vacation, and you are certain to find one that suits you well. Many all-inclusive Oahu vacation packages are tailor-made for singles who are looking to meet other singles. Singles vacations are usually offered in a package form with accommodations and airfare coordinated together so that all attendees are in the same place at the same time. An individual can choose his or her desired location, activities, and dates, and more than likely one of many Oahu singles vacations will be a perfect fit. So many people are looking for new ways to meet others in the busy age of speed dating and computer matchmaking. Oahu just happens to be the perfect place to make a romantic connection given the extraordinary scenery. Singles really have the run of the island when choosing what kind of atmosphere in which to meet someone new.
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Understand the different levels of care you can expect from each facility. Detox facilities provide a safe, medicinally aided and monitored program for addressing the physical aspects of addiction. Residential rehabilitation programs provide care in a controlled environment with extra support to help mitigate the risk of relapse during recovery.
Partial hospitalization treatment options are most beneficial for those who with a dual diagnosis of a mental health disorder and substance abuse disorder.
Remember, this Hawaii dating site is absolutely free. Most other online dating services pretend to be free and tell you that registration is free. In fact, only registration will be for free, and you will be made to pay on most dating sites in order to be able to contact other singles.
Will you have me for dinner? In the end, they always How about I have you for dessert? Put your snake in my garden of eden! No one does bad like I do! I was a contender for Ms Hawaii, but I slept with one of the judges and got kicked out. I'm just wild and I can't help Why not have fun? Tall Blonde looking for interacial Message: I have blonde hair blue eyes. I like to look at asian, and women of color.
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Frameless Bathe Doorways could be geared up with quite a few various options. Developers, try our new API that lets you combine album artwork and content material from AllCDCovers into your web site, weblog, widget and iTunes software program. They allow asides to turn into entire episodes, experiment with style and Vikings box set Seasons aesthetic, often nodding to artwork-house cinema of previous.
These buildings have been all designed to harmonize with the rugged nature of the park by way of a style that became often called National Park Service Rustic. Junjie took control, banishing the Five and Po and capturing Shifu upon his return, intending to destroy him. Generally, Shifu was devoted to his grasp and hardly ever argued together with his wisdom, no matter how eccentric it may need seemed to him.
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Search 15 Best Texas Beaches Texas has huge variety of beaches , most long and hard packed with fine white sand, on the mainland and on the barrier islands. There are great party beaches such as Galveston Island East Beach and quiet deserted ones like Boca Chica on South Padre Island , where people go for solitude, romance and to enjoy unspoiled nature. Here are the best Texas beaches for your next weekend getaway from Houston , Dallas , San Antonio , Austin and other cities.
Boca Chica There are many spectacularly beautiful beaches on South Padre Island , some lively, others quiet and deserted, but Boca Chica Beach manages to stay empty of crowds, clean, and beautiful. It has completely escaped development and has only a few stilt houses and a jetty sticking out into the Gulf of Mexico.
Although Boca Chica is a state park Texas beach, it has no park facilities at all. What it does have is miles of fine sand beaches, clear water, and great swimming, snorkeling, fishing, kiteboarding, and birdwatching. It is a perfect spot to get away from the crowds and noise.
Sugar Daddy dating in HI, United States
If you misplace or the waiver letter has been lost, you must meet with your case worker and have them issue a replacement waiver letter. The case worker will manually fill out the letter with all the eligible children's names on it. This original replacement waiver must then be submitted with the summer fun registration form at the time of registration. It provides a free brunch to all summer fun participants under the age of 18 years old.
The brunch is a 3-item meal.
Use this dating site and become dating expert, chat with beautiful people or find the person of your soul. Online dating can help you find relationship. Single Women Hawaii – Use this dating site and become dating expert, chat with beautiful people or find the person of your soul.
Chronology Radiocarbon dating shows that Easter Island was inhabited by AD, and possibly by the 4th century. This fits in with the tradition that there had been 57 generations of kings since Hotu Matua; allowing an average of 25 years per generation, this takes us back to AD. Some archaeologists suspect that the island must have been settled several centuries earlier.
There is of course no evidence — only theories and assumptions — to rule out the possibility that the island was inhabited millennia before this; as mentioned in section 2, some native traditions point to pre-Polynesian settlement. As already explained, the standard view is that Polynesians discovered Easter Island by chance and that after its initial colonization it was not visited by anyone else until the Europeans began to arrive in the early 18th century.
As shown in section 3, the evidence is ambiguous but is certainly consistent with some sort of South American influence alongside the prevalent Polynesian influence. The island could have received settlers or visitors from both east and west on many occasions. There is clear evidence of different phases of development in statue carving and platform construction, and the insistence that all the archaeological remains must be crammed into a history spanning just years is theory-driven.
The rongorongo phenomenon is also difficult to fit into conventional theories about Easter Island. A great deal of excavation work still needs to be done. At Anakena the present surface of the sandy plains lies 4 m above the bedrock. At Rano Raraku the ground on which the giant statues were set up is often 6 m below the present surface.
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The culture of West Polynesia is conditioned to high populations. It has strong institutions of marriage and well-developed judicial, monetary and trading traditions. It comprises the groups of Tonga , Niue , Samoa and extends to the atolls of Tuvalu to the north. The pattern of settlement that is believed to have occurred is that the Polynesians spread out from the Samoan Islands into the Tuvaluan atolls, with Tuvalu providing a stepping stone to migration into the Polynesian Outlier communities in Melanesia and Micronesia.
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By Keith Wagstaff With plumes of ash and glowing lava, the area around Mount Kilauea looks more like Mordor than a tropical paradise. That hasn't been great for tourism. Can I roast a marshmallow over lava? Still, if you've got a plane ticket to Hawaii, there's no need to cancel your trip — you'll probably be fine. First of all, the Big Island of Hawaii is, well, big. The volcano is erupting on a tiny corner of an island that measures more than 4, square miles, which is more than double the area of the entire state of Delaware.
Most of the big resorts are in Kona, on the island's west side. That's about miles from the volcano. Concord University volcanologist Janine Krippner put the danger in perspective in this tweet: The Kilauea eruption is affecting people.
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Green sea turtle near Marsa Alam , Egypt Green sea turtle Scalation of carapace and plastron Taxidermied shell a Chelonia mydas Its appearance is that of a typical sea turtle. Unlike the closely related hawksbill turtle , the green turtle's snout is very short and its beak is unhooked. The neck cannot be pulled into the shell. The dorsal surface of the turtle's head has a single pair of prefrontal scales.
Its carapace is composed of five central scutes flanked by four pairs of lateral scutes.
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WA Delegate: None.
Founder: The Queen's State of Dawson Park
Last WA Update:
Embassies: Richard, Pacific Union, Brazzaville, Ecuador, Toronto, World, Nebraska, Central African Republic, London, Mackenzie, Prince Edward Island, Senegal, Samoa, Mongolia, Malawi, Missouri, and 330 others.Benin, Rhode Island, Laos, Pennsylvania, South Australia, Palau, Iowa, Massachusetts, Togo, Mexico City, Vermont Republic, Quito, New Brunswick, United Kingdom Of Great Britain, Kiribati, Botswana, Barbados, Great Britain and Northern Ireland, Tasmania, Mauritius, 0000, Brisbane, United Arab Emirates, Dominican Republic, Australian Capital Territory, Virginia, Liechtenstein, Sao Paulo, Okanagan, Sao Tome e Principe, New Hampshire, Colorado, Kuwait, Newfoundland and Labrador, Mississippi, Cameroon, El Salvador, Queensland, Costa Rica, Montserrat, Cote dIvoire, Wyoming, Seattle, Madagascar, Rio de Janeiro, British Columbia, Cape Verde, Solomon Islands, Dominica, Ceylon, New Mexico, Nunavut, Illinois, Crematorium, the Bahamas, Somalia, Brunei Darussalam, Zimbabwe, Saint Lucia, Uzbekistan, Burundi, Timor Leste, Idaho, Ethiopia, Detroit, Columbia, Malaysia, Guinea, Jamaica, Honduras, South China Sea, Knights of the Round Table, Edinburgh, Australia Commonwealth, Algeria, Qatar, Montreal, United States of America, Gabon, Saint Kitts and Nevis, Stornoway, Bonn, Uganda, Guatemala, Morocco, Djibouti, Northwest Territories, Belize, Delaware, Luxembourg, Ghana, North Dakota, South Dakota, Rwanda, Washington, The Central Pacific, Western Australia, Northern Territory, Guinea Bissau, South Carolina, Sierra Leone, Tanzania, Arizona, Pitcairn Island, Lesotho, Saint Vincent and the Grenadines, Utah, Ancient Lands, Washington DC, Republic of Russia, The Philippines, Maldives, Mozambique, Sakartvelo, Trinidad and Tobago, Maryland, Copenhagen, Seychelles, Tuvalu, North Carolina, Victoria, Kentucky, Crimea, Saskatchewan, Namibia, Calgary, Swaziland, Ottawa, Tonga, Minnesota, Nova Scotia, Democratic Republic of the Congo, Republic of Congo, Indonesia, Yukon, Sudan, Monaco, Republica del Paraguay, Michigan, Beijing, Kansas, Turkiye, Purgatory, Galapagos, Georgia, Nevada, Fiji, Antigua and Barbuda, Comoros, San Marino, Kenya, Vanuatu, Wales, The Gambia, Nauru, Two Letter, Commonwealth States, The Assembly of Gentlemen, The Awakening, Rhea, Vietnam, Angola, Kanada, Connaught, Leinster, Atlantic New Lands, Imperial Prussia, Kingdom Of England And Wales, United Assembly, Orient, Capital Heights, Sydney, The Six Counties of Ulster, Marshall Islands, Federated States of Micronesia, Northern Mariana Islands, Greater London, East Timor, Manchester, Birmingham, DBA, Palmyra Tudmur, Cotonou, Parkland, Iranian Islamic Republic, Federal Republic of Russia, Norsewood, Kigali, Plague, Queen Elizabeth Islands, Empire of Brazil, Indianapolis, Azerbaycan Respublikasi, Republic of Myanmar, Kandahar, Democratic Republic of Ecuador, Chan Nations, Salt Lake, The Republic of California, Russian Federation, RAMS, Heavenly Palace, Northern Africa, Kitai, Queens, Lusaka, Belfast, Long live the British Empire, Old Spirit, Union of South Africa, Zambia, Western Sahara, New Caledonia, Algiers, Republic of China, New Delhi, Wanderlust, 404 Not Found, Dhaka, Abuja, Jakarta, Islamabad, Brasilia, Addis Ababa, Mockba, Manila, Society of Commissioned Gentlemen, Kinshasa, Bangkok, Cape Town, Bloemfontein, Dodoma, Caracas, The Southern Pacific, Hsiao Han, Naypyidaw, Federation of Bosnia and Herzegovina, Kyiv, Kampala, Bogota, Buenos Aires, Rabat, Dao, Black Dragons, Joint Command, Riyadh, Imperio do Brasil, Bangui, Damascus, Warsaw, Canberra, Maputo, Tashkent, Taipei, God Complex, Kuala Lumpur, Khartoum, The Congo, Oslo, Nassau, Havana, Accra, Sanaa, Nuku Alofa, Lisbon, Santo Domingo, Republic of Korea, Antananarivo, Stockholm, Cemetery, Port Moresby, Santiago, Colombo, Superior, Dili, Malabo, Bern, Bishkek, Juba, Port Vila, Budapest, Sarajevo, Tifariti, Cardiff, Vaduz, Port of Victoria, Bamako, Bissau, Georgetown, Wellington, Windhoek, Saint Georges, Douala, Suva, Yamoussoukro, Dublin, Minsk, Saint Johns, Yaounde, Abu Dhabi, Niamey, Bratislava, San Salvador, Jervis Bay Territory, Apia, Fredonia, United States of Femdom, Western Africa, Maya, the Alliance Headquarters, Freedom Front, Nouakchott, Vientiane, South Bentley, Westminster, Conakry, Maseru, Halifax, Nursultan, Amman, Freetown, Lome, Lobamba, Port Louis, Great Europe, Kuwait City, Mbabane, Miami, Singapore City, and Little America.
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Edmonton contains 8 nations, the 1,935th most in the world.
Activity • History • Administration
Today's World Census Report
The Highest Crime Rates in Edmonton
World Census interns were dispatched to seedy back alleys in order to determine which nations have the highest crime rates.
As a region, Edmonton is ranked 3,080th in the world for Highest Crime Rates.
|WA Category
|The Queen's State of Margiana
|The Queen's State of Edmonton City
|Capitalist Paradise
|"From this beginning"
|The Queen's State of WhoLocks
|The Queen's State of Fort Edmonton
|Civil Rights Lovefest
|"Industry, Integrity, Progress"
|The Queendom of South-West
|Inoffensive Centrist Democracy
|"We Will Endure"
|The Queen's State of Oil Gateway
|Moralistic Democracy
|The Queen's State of Dawson Park
|Inoffensive Centrist Democracy
|The Queendom of Nathell
|Free-Market Paradise
|"Pride and Industry"
Regional Happenings
- : Embassy established between Little America and Edmonton.
- : Embassy cancelled between Zero Zero Zero Zero and Edmonton.
- : The Allied States of American Refugee of the region Little America agreed to construct embassies.
- : The Queen's State of Dawson Park proposed constructing embassies with Little America.
- : The Republic of Switz got Lazy 397 of the region Zero Zero Zero Zero ordered the closure of its embassy in Edmonton.
- : Embassy established between Edmonton and Singapore City.
- : Embassy established between Mbabane and Edmonton.
- : Embassy established between Kuwait City and Edmonton.
- : Embassy established between Miami and Edmonton.
- : The Airborne Flock of Migrating Geese departed this region for Edlhus.
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J.K. Rowling This is one of those books that no matter how many times I read it the material doesn't sink it. That probably has to do with the fact that this is about a sport! My discovery when reading this book (for the third time) was the mention of a Brutus Scrimgeour, this made … Continue reading Quidditch Through the Ages
Month: September 2012
Stephen Chbosky I am a bit behind with making a post for this book because I finished the book last week, but here it goes. I LOVED this book. It was written in a simple language that was moving, relate-able. and easy to understand. This is one of those books that you do not want … Continue reading The Perks of Being a Wallflower
Plato Personally this dialogue was a bit more difficult to get into than "Symposium" to me, and not because it related to death. The manner in which Socrates evolves his argument was hard to follow and understand for me. Here is what I got out of the texts. A quick summary of Phaedo: On the day of … Continue reading Phaedo
J.K.Rowling This is about the ninth time I've read this book. I am what is commonly called a "Potter-head," or a Harry Potter Obsessed person. I have been reading the series since I was eleven and have even been known to make things in relation to the series. The last time I read the series … Continue reading Harry Potter and the Sorcerer's Stone
"Isn't it odd how much fatter a book gets when you've read it several times?" Mo had said..."As if something were left between the pages every time you read it. Feelings, thoughts, sounds, smells...and then, when you look at the book again many years later, you find yourself there, too, a slightly younger self, slightly … Continue reading Quote
Plato Plato is most likely an acquired taste, when I started reading this dialogue it was slow and dull, but the further I read the more intriguing it became.Below are some passages I deemed important. "Evil is the vulgar lover who loves the body rather than the soul, and who is inconstant because he is … Continue reading Symposium
"A good library will never be too neat, or too dusty, because somebody will always be in it, taking books off the shelves and staying up late reading them." -Lemony Snicket (Horseradish: Bitter Truths You Can't Avoid)
Friedrich Nietzsche When reading sections 6-8 I absorbed nothing. The author is too obscure with his writing that I cannot understand what he was trying to get across in the slightest. The only thing I think he mentioned in these sections was the purpose of the chorus in Greek tragedy. Personally I think the Chorus … Continue reading The Birth of Tragedy (pt. 4)
Euripides Medea was a tale I have never read before. It was not as intriguing as Oedipus Rex or Agamemnon, but wasn't a difficult read in the least. I only found a couple quotes within the text that intrigued me. "The fiercest anger of all, the most incurable, is that which rages in the place … Continue reading Medea
"If one cannot enjoy reading a book over and over again, there is no use in reading it at all." - Oscar Wilde


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Tolpis pallidiflora Jord. & Fourr.
This is a synonym of Tolpis barbata (L.) Gaertn.
Other Data
Tolpis pallidiflora Jord. & Fourr. appears in other Kew resources:
First published in Brev. Pl. Nov. ii. 85 (1868)
Not accepted by
- Roskov Y. & al. (eds.) (2018). Species 2000 & ITIS Catalogue of Life Naturalis, Leiden, the Netherlands. [Cited as Tolpis barbata.]
Kew Names and Taxonomic Backbone
The International Plant Names Index and World Checklist of Selected Plant Families 2020. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0


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Where Can I Sell My Baseball Card Collection
If you fail to identify all of the superstars in your collection you might accidentally sell your cards for much less than you should have.
Where can i sell my baseball card collection. Whether you need money for financial reasons such as a wedding a down payment on a car or house or if your heart just isnt in it anymore it can be a tough decision. After all is said and done we are confident you will be satisfied with our quote. Again if you are selling cards printed from 1879 to 1979 and you need more guidance please feel free to contact me using this form. If you need information on how to sell your vintage baseball card collection you have come to the right source.
In my experience determining where to sell your baseball card collection usually depends on two factors. What is the best way to sell my large baseball card collection which consists of cards from 1959 late 2000. Our goal is to make you the customer happy. How to sell baseball cards.
What type of cards youre selling pre war vintage modern how quickly you need to sell them. This guide was created to help you through the process and equip you with the knowledge to sell your collection with confidence and ease. There is a large market out there for baseball cards mostly for. A collection of 10 baseball cards with three superstars is usually more valuable than a collection of 100 cards with only one superstar.
First if youre trying to find out the value of your cards you may want to visit your local sports card shop to buy a price guide. Here are some ways to sell your sports cards whether you have baseball cards basketball cards football cards hockey cards or all of the above. When you bring your items to us you will meet face to face with someone who will help you make the right choice on what to sell. Where can i sell my baseball cards whether you need cash or are just tired of the space they take up we hope to answer that for you through this page.
What is the best way to get a baseball card collection appraised. Sometimes the cards just arent worth parting with because of sentimental reasons. There are of course exceptions including old judges and t206s. What is the best way to sell an entire baseball card collection.
Its a popular question. Selling your sports card collection can be difficult for any collector. Selling your old baseball collection can become a fun endeavor. We buy baseball cards football cards basketball cards hockey cards miscellaneous sports collectibles and sports memorabilia as well as entertainment related trading cards collectibles and memorabilia.
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