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A role for Casein Kinase 2 in the mechanism underlying circadian
temperature compensation
Arun Mehra1,*, Mi Shi1,*, Christopher L. Baker1, Hildur V. Colot1, Jennifer J. Loros1,2, and
Jay C. Dunlap1,2,†
1Department of Genetics, Dartmouth Medical School, Hanover NH 03755
2Department of Biochemistry, Dartmouth Medical School, Hanover NH 03755
Most organisms employ biochemical clocks to keep daily time (Dunlap et al., 2004; Pittendrigh,
1993). Circadian clocks are oscillators defined by three key characteristics: they must free run
with an approximately 24 h period, are entrainable by relevant environmental stimuli, e.g.,
light; and exhibit temperature compensation (TC), the relative invariance of period lengths
over a physiologically relevant range of temperatures. Among these defining characteristics,
TC is the least well understood.
Temperature affects the clock in three ways. First, discrete temperature steps up or down can
reset the clock (e.g., Glaser and Stanewsky, 2007; Liu et al., 1998). Second, temperature can
impose physiological limits on rhythmicity (Kalmus, 1934; Liu et al., 1997; Njus et al.,
1977). Finally, while the phase of the clock responds to discrete temperature changes,
paradoxically, its period is buffered against changes in temperature. Although this relative
independence of period length had been noted (Kalmus, 1935; Pittendrigh, 1954; Wahl,
1932), Hastings and Sweeney (1957) first recognized that this buffering was not due to
temperature independence but rather reflected a TC mechanism. Because both the magnitude
and sign of the period change with temperature varied across the physiological range, they
reasoned that TC could be a network property perhaps arising from multiple reactions having
opposing contributions to period length, but unlikely to be based on a single temperature-
independent process as had been supposed. Notably, even homeotherms have temperature-
compensated clocks (e.g. Barrett and Takahashi, 1995; Tosini and Menaker, 1998), suggesting
that TC may not simply be a property appended to a temperature-dependent clock but may
instead be a property of the regulatory architecture.
Information obtained from Neurospora, Drosophila, and cultured mammalian cells is
consistent with a negative feedback loop of conserved architecture at the core of the clock. In
this loop two proteins that interact via PAS domains comprise a transcriptional activator that
drives expression of genes encoding protein(s) that, in turn, feed back to repress the activator
(see Allada et al., 2001; Schibler and Sassone-Corsi, 2002; Wijnen and Young, 2006). The
Neurospora clock provides an example of such a circuit. Briefly, WHITE COLLAR-1 (WC-1)
and WC-2 interact via PAS domains to make a transcriptional activator, the White Collar
†Corresponding Author: E-mail: email@example.com.
*These authors contributed equally to this work.
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Published in final edited form as:
Cell. 2009 May 15; 137(4): 749–760. doi:10.1016/j.cell.2009.03.019.
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Complex (WCC), which activates transcription of the frequency (frq) gene. FRQ protein in
turn, with the help of FRQ-interacting RNA helicase (FRH), depresses the activity of the WCC,
in part by promoting its phosphorylation. Over time, FRQ is progressively phosphorylated and
consequently targeted by the F-box and WD40 repeat-containing protein-1 (FWD-1) for
degradation by the proteasome (reviewed in, Brunner and Schafmeier, 2006; Dunlap and Loros,
2006; Liu and Bell-Pedersen, 2006).
frq and FRQ are implicated in a number of temperature-dependent responses (Nowrousian et
al., 2003). Temperature resetting has been explained by shifts in the level of FRQ without
underlying changes in frq expression (Liu et al., 1998), reflecting the fact that the steady state
level of FRQ rises as a function of constant temperature (Diernfellner et al., 2005; Liu et al.,
1997). Moreover, frq pre-mRNA is alternatively spliced to direct production of both long and
short FRQ, each of which can support compensated clocks (Diernfellner et al., 2007; Dunlap
et al., 2007b), and this splicing is temperature dependent (Colot et al., 2005; Diernfellner et
al., 2005). Augmented levels of long FRQ at high temperatures and increased amounts of short
FRQ at low temperatures extend the physiological range of rhythmic banding (Liu et al.,
1997). However, changes in the ratio of long to short FRQ do not lead to altered TC, nor is
this ratio relevant to TC as a general property (Diernfellner et al., 2007). These data implicate
frq/FRQ in a number of temperature responses; however, we emphasize that not all effects of
temperature on the clock contribute to TC processes.
Theory and experiment have suggested mechanisms underlying TC. The simplest model
invokes a pair of opposing biochemical reactions, each with normal and equal Q10s having
opposing effects on progress of the clock cycle (Hastings and Sweeney, 1957). As temperature
changes, cycle duration is kept constant as the rates of opposing reactions change by the same
magnitude. A similar solution proposed for Arabidopsis envisions reactions in two clock-
associated feedback loops with opposing responses to temperature changes so that the
composite is balanced (Gould et al., 2006). An attractive mathematical formulation for the idea
of opposing reactions, the balance equation (Ruoff et al., 2005; Ruoff et al., 1997), models the
rate of the circadian cycle as the sum of many reactions whose contributions to period must
equal 0. A mechanistically unspecified amplitude model (Lakin-Thomas et al., 1991) having
some empirical support (Lahiri et al., 2005; Liu et al., 1998) achieves TC by having the size
of the circadian limit cycle increase with temperature, thereby compensating for increases in
angular velocity. The possibility of competing intra-and intermolecular reactions involving
Drosophila PER (Huang et al., 1995; Price, 1997; Sawyer et al., 1997) led to a model in which
opposing effects of dimerization and nuclear entry lead to TC (Hong et al., 2007; Hong and
Tyson, 1997). This and additional mathematical models (Akman et al., 2008; Kurosawa and
Iwasa, 2005; Leloup and Goldbeter, 1997; Takeuchi et al., 2007) await experimental testing.
Genetic screens for TC mutants have suffered from the fact that circadian period is often
dependent on dosage of clock components (e.g., Smith and Konopka, 1982); thus, temperature-
sensitive alleles whose effective doses drop with increasing temperature can be mistaken for
TC mutants. In general, although TC is a universal aspect of circadian rhythms, mechanisms
that have been generalizable (e.g., Ruoff et al., 1997) have not been reducible to mechanistic
specifics, and mechanisms that are mechanistically specified (e.g., Huang et al., 1995) have
not been generalizable.
Two mutant strains, however, chrono (chr) and period-3 (prd-3), merited further study because
they displayed the unusual characteristic of enhanced or extended TC (Gardner and Feldman,
1981). The period length of the Neurospora clock is slightly undercompensated between 18°
C and 30°C beyond which the period shortens and TC is lost. In chr mutants, however, TC
extends beyond 30°C, and prd-3 strains are overcompensated such that the period increases as
a function of temperature (Gardner and Feldman, 1981). Reasoning that these types of TC
might be informative, we determined the molecular bases of the mutations in chr and prd-3
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and studied their roles in TC. The result is that these two independently derived period mutants,
later found to be defective in TC, identify separate subunits of the same enzyme, casein kinase
2 (CK2). Furthermore, genetic manipulation of CK2 activity, unique among kinases and
phosphatases examined, affects both period length and TC in a predictable manner. These and
additional data are consistent with a role for CK2 in the mechanism underlying TC.
chrono and period-3 bear mutations in the β1 and α subunits of CK2, respectively
First, we confirmed the phenotypes of chr and prd-3 and established an epistasis relationship
between these mutants. Consistent with previous data (Gardner and Feldman, 1981), chr shows
an extended range of TC (Fig 1A, black squares vs. white circles) and prd-3 overcompensates
compared to WT. In our hands, however, prd-3 shows poor rhythmic banding at 30°C and
above (Fig 1A, white triangles). Interestingly, prd-3; chr double mutants have longer periods
than either single mutant but overcompensate and band poorly above 30°C (Fig 1A, black
triangles), demonstrating additivity of period but clear epistasis of prd-3 over chr with respect
to TC.
chr and prd-3 were cloned using single nucleotide polymorphism (SNP) mapping. Genetic
mapping had placed chr on the right arm of LG VI and prd-3 near the centromere of LG I
(Gardner and Feldman, 1981) and these locations were refined by crossing onto a WT
Mauriceville background and following segregation of SNPs (Dunlap et al., 2007a; Lambreghts
et al., 2009). This localized chr (Fig 1B) and prd-3 (Fig 1C) on the physical map and suggested
candidate genes for both mutants in different subunits of CK2 (arrows, Fig 1B and 1C).
CK2, a multifunctional kinase (Litchfield, 2003), is a tetramer composed of two identical
catalytic α subunits and two, possibly different, regulatory β subunits (Sup Fig 1). Sequencing
of NCU05485.3, encoding the β1 subunit of CK2 (encoded by ckb-1) in the chr background,
revealed a C→T transition, resulting in an R265C change within an extended C-terminal tail
region of CK2β1 that is conserved among fungi (Sup Fig 1, top and middle). Since the β subunit
of CK2 mediates contacts between the catalytic α subunits and substrate (Litchfield, 2003), we
speculate that such functions may be perturbed in chr. Functional complementation of prd-3
(Fig 1C, bottom) identified NCU03124.3 on pLORIST H031 B12 as encoding the α subunit
of CK2. Sequencing revealed a T→C transition resulting in a T43H mutation near the highly
conserved phosphate anchor (Taylor et al., 1992) of the CK2 subdomain I (Sup Fig 1, bottom)
of CK2α (encoded by cka). Genetic rescue confirmed that these mutations in ckb-1 and cka
conferred the period length defects in chr and prd-3, respectively (see Sup Table 1).
Thus, these decades-old mutants, unique among the dozens of known clock gene alleles as
having extended-or overcompensation, identify separate subunits of the same holo-enzyme.
Together, the simplest interpretation of the data is that CK2 plays a role in facilitating TC.
Although not previously implicated in TC, CK2 has established roles in circadian clocks of
Arabidopsis (Sugano et al., 1998), Neurospora (Yang et al., 2003; Yang et al., 2002) and
Drosophila (Akten et al., 2003; Lin et al., 2002; Lin et al., 2005; Nawathean and Rosbash,
2004) that are consistent with the increased period lengths we see in the CK2 mutants. Since
a possibility was that these mutations were hypomorphic, we pursued the effect of CK2 dosage
on TC by altering dosage of both subunits to see whether the independent identification of two
CK2 subunits was more than a remarkable coincidence.
Reduction of CK2 subunit gene dosage alters TC
Phosphorylation-mediated turnover of FRQ is a major determinant of period length, and
reducing the activity of kinases acting on FRQ results in increased circadian period length
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(Liu et al., 2000). Thus, a trivial explanation for the chr and prd-3 mutant phenotypes would
be that period length increases with temperature simply because the mutant alleles chr and
prd-3 encode temperature-sensitive proteins. A more interesting result, however, would be if
chr and prd-3 were genuine hypomorphs having lower levels of activity but normal temperature
dependence; in this case reductions of the WT allele dosage should mimic the chr and prd-3
phenotypes. To achieve various gene dosages, we generated heterokaryons of either chr or
prd-3 bearing different proportions of knockout versus WT nuclei. In terms of protein
expression these approximate an allelic series. We confirmed this directly for CK2β1 by using
an antibody (Ab) against the CKB-1 protein and showed, while WT and chr strains have
comparable levels of CKB-1, Δckb-1 heterokaryons have lower levels of protein (Sup Fig 2).
Knockout heterokaryons bearing reduced dosage of cka (cka + Δcka, referred to as Δcka) or
ckb-1 (ckb-1 + Δckb-1, referred to as Δckb-1) mostly recapitulate the original homokaryotic
prd-3 and chr phenotypes, respectively. Like prd-3, many Δcka heterokaryons overcompensate
between 20°C and 25°C and all show increased periods (Fig 2A). These properties are
consistent with prd-3's being a hypomorph. Although prd-3 does not consistently express an
overt rhythm at 30°C, the knockdown strains show reduced period lengths at this temperature
(Fig 2A). Heterokaryon Δckb-1 knockdowns resemble the chr mutant, having increased period
lengths with extended-or overcompensation at the highest temperature (Fig 2B).
CKB-1 dosage qualitatively and quantitatively affects the type of TC
Heterokaryon knockdowns were crossed to WT to generate homokaryotic Δckb-1 knockout
strains. These strains appear overtly arrhythmic (Fig 2C) and provide a context in which to test
the hypothesis that CK2 plays an essential role in TC.
Because we suspected that chr might be hypomorphic, we hypothesized that as the level of
CK2 is reduced, the TC profile of the clock should move from WT slightly undercompensated
(negative slope), to extended TC (flat slope), and finally to overcompensated (positive slope).
We tested this by generating a heterokaryotic strain in which the sole source of ckb-1 mRNA
is driven by the quinic acid-2 promoter (qa-2, hereafter called qa) (Aronson et al., 1994; Dunlap
and Loros, 2005) in proportion to the concentration of quinic acid (QA) in the medium (Fig
3A, top panel); the strain bears the genotype (his-3 + his-3+∷qa-ckb-1; Δckb-1) and is referred
to as qa-ckb-1. We confirmed that addition of QA yields increased levels of CKB-1 in a dose-
dependent manner (Fig 3A, bottom panel). QA itself does not affect the TC curve of the WT
(data not shown).
Inducing ckb-1 gene expression in Δckb-1 has dramatic effects on period length and TC. Higher
levels of CKB-1 result in shorter periods matching WT period lengths with typical WT
undercompensation (Fig 3B, squares). By contrast, reduced levels of QA, yielding lower levels
of CKB-1, result in steadily longer periods progressing to overcompensation (positive slope)
(Fig 3B, circles). In confirmation of our working model, we see a consistent relationship
between ckb-1 dosage and the resulting slope of the TC profile.
In summary, as CKB-1 dose increases the TC profile changes from overcompensation to slight
undercompensation. It is important to note here that although it was the characterization of TC
mutants that initiated these studies, the experiment described in Fig 3B involves only WT
proteins whose level of expression is altered. By controlling the amount of WT CKB-1 we can
dictate the TC profile. Taken together, the data support a model in which CK2 modulates TC
under physiological conditions.
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Dosage changes in other kinases and phosphatases implicated in the Neurospora clock do
not affect TC
The phosphorylation state of FRQ and of all clock-associated proteins is affected by other
kinases and phosphatases. Given this, a trivial explanation of the preceding results would be
that any mutation affecting the activity of enzymes acting on clock components would affect
TC. We tested this by manipulating the levels of two kinases, casein kinase 1 (CK-1a) (Gorl
et al., 2001; He et al., 2006) and protein kinase A (PKAC-1) (Huang et al., 2007), as well as
two phosphatases, protein phosphatase 1 (PPP-1) and protein phosphatase 2A (PP2A) (Yang
et al., 2004).
We were particularly interested in whether changes in CK1 dosage might affect TC. Since
ck-1a (NCU00685.3) is an essential gene in Neurospora (Gorl et al., 2001), we used an
inducible-promoter knock-in approach (L. Larrondo et al., manuscript in preparation). Briefly,
we replaced ~0.6 kb of sequence upstream of the predicted CK-1a ORF with the QA-inducible
promoter similar to that described above (Fig 4A, top left). This allowed us to control and
reduce CK1 levels without completely eliminating the protein, thereby allowing cell survival.
We verified induction of message (data not shown) and protein expression using α-CK-1a
antibodies (Fig 4A, bottom left).
Like CK2, CK1 determines period length. Unlike CK2, however, changes in CK1 levels do
not significantly affect TC. In the absence of QA, the reduced dose of ck-1a in heterokaryons
bearing the knock-in construct increases period by approximately 3 h (Fig 4B left, top); this
phenotype is rescued upon addition of QA (Fig 4B left, bottom), consistent with previous
findings (Gorl et al., 2001;He et al., 2006). To complete the analysis, a heterokaryon bearing
the qa-ck-1a knock-in was crossed to obtain a homokaryon in which the only source of CK1
is the qa-driven construct. When the dosage of CK1 in this strain is restricted by limiting QA,
the effect on TC does not approach that seen with changing CK2 (Fig 4B, right).
The potential roles of PKA and the phosphatases were also examined by controlling their doses.
Endogenous promoters were replaced with ~1.1 kb of the qa promoter for both pkac-1
(NCU06240) and the catalytic subunit of PP2A (pph-1, NCU06630) and crossed to generate
homokaryons. Changes in steady state expression were measured by quantitative RT-PCR (Fig
4A, bottom right) on samples grown in the absence or presence of 10-2 M QA. Less pronounced
changes in period lengths were observed in homokaryons bearing qa-pkac-1 (Fig 4C, left) and
qa-pph-1 (Fig 4C, right) than were seen for CK1, but no significant differences were seen in
the TC profiles of these strains. Finally, heterokaryon knockouts of ppp-1 showed no significant
period reduction, consistent with RIP mutants (Yang et al., 2004), and showed no effect on TC
(Sup Fig 3).
Together, these data indicate that not all regulators of clock protein phosphorylation influence
TC. Rather, they suggest that CK2 plays a special role in TC.
CK2 directly phosphorylates FRQ
Since strains with low levels of CK2β1 show increasing periods as a function of temperature,
a trivial explanation would be that CK2 activity might be compromised with increasing
temperature, but only when CK2β1 levels are reduced to below normal physiological limits.
To address this, we assessed endogenous CK2 activity on synthetic peptides (Kuenzel and
Krebs, 1985) across a range of CK2β1 levels. Regardless of the amount of CK2β1, CK2 activity
approximately doubles over the indicated ten degree temperature (representative experiment,
Fig 5A); the trivial explanation does not hold. Additionally, CK2β1 levels do not vary, as
assessed by Western blot, between 20°C and 30°C (data not shown).
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As this analysis was performed on a synthetic substrate, we wondered how CK2 might
phosphorylate a bona fide substrate as a function of temperature. While CK2 has many targets
in Neurospora, one potential target likely to affect TC is the central clock component FRQ.
FRQ has a number of putative CK2 phosphosites and calmodulin kinase-free, partially purified
lysate fractions containing CK2 subunits phosphorylate FRQ in vitro while ckaRIP and
ckb-1RIP mutant extracts show reduced levels of FRQ phosphorylation (Yang et al., 2002).
To confirm direct activity of CK2 on FRQ, we performed in vitro kinase assays with full-length
GST-FRQ. CK2 is unique among kinases in that it can efficiently use GTP as a co-substrate
in phosphotransfer reactions; thus, phosphorylation by GTP is considered strong evidence that
a substrate is a direct CK2 target (Sugano et al., 1998; Yde et al., 2005). WT lysates incubated
with full-length GST-FRQ and GTP yielded efficient phosphorylation (Fig 5B), indicating that
CK2 can directly phosphorylate FRQ. However, since in vitro kinase assays can be
promiscuous, we also assayed for phosphorylation of endogenous levels of FRQ by CK2.
Briefly, whole cell extracts from cells grown at 30°C were incubated at 30°C with radioactive
GTP and were subsequently immunoprecipitated with antiserum against FRH, a protein that
interacts strongly and stoichiometrically with FRQ (Cheng et al., 2005). Immunoprecipitates
revealed bands the size of FRQ in WT, but not in frq10 or in qa-ckb-1 under non-inducing
conditions (Fig 5B, lower panels). FRQ is directly phosphorylated in a time-dependent manner;
however, as lysates contain both kinases and phosphatases and only a limited pool of
radioisotope, phosphorylation peaks at 30 min and then declines, presumably reflecting
phosphatase activity and depletion of radioisotope. In this assay, there appears to be no
detectable phosphorylation of FRQ when CK2β1 levels are low, so we could not assess Q10.
chrono and strains with reduced CKB-1 exhibit a temperature-dependent decrease in FRQ
phosphorylation in vivo
Since the rate of FRQ degradation is a key factor in determining period length (Liu et al.,
2000; Ruoff et al., 2005), we assessed the rate of FRQ degradation as a function of reduced
CK2β1 activity and temperature. We found FRQ to be degraded slightly more slowly in chr
than in WT at 25°C (Fig 5C, top panel), an effect that is exacerbated at higher temperature (Fig
5C, bottom panel). An artificially high temperature of 34°C was chosen to exaggerate the subtle
difference in FRQ stability seen between WT and chr. FRQ is similarly phosphorylated more
slowly in a chr ; frq7 background compared to frq7 (data not shown). On one level, these data
are simply consistent with expectations based on prior work, in that increased periods are due
to increased FRQ stability. However, Fig 5A shows that even with low CK2β1 activity,
consistent with chr, the rate of in vitro kinase activity appeared to increase with temperature.
Thus, WT levels of CK2 appear critical for modulating FRQ stability specifically at higher
To directly assess this, we tested FRQ stability in a strain expressing low levels of CK2β1. For
this, we used a standard light to dark transfer assay: steady state levels of frq/FRQ are high in
light, frq mRNA is rapidly degraded after the transfer to darkness, approximating the chase of
a pulse-chase. In WT, FRQ degradation does not vary between the two temperatures (Fig 5D).
In contrast, FRQ is roughly twice as stable at higher temperature relative to lower temperature
in the qa-ckb-1 strain in the absence of QA (Fig 5D, right). Taken together with the chr data,
it appears that FRQ is degraded more slowly as temperature increases when CK2β1 activity is
reduced. This is consistent with a model in which over-and extended TC in strains containing
hypomorphic CK2 or low dosage of WT CK2 results in a temperature-dependent misregulation
of FRQ turnover. As noted above, other kinases known to phosphorylate FRQ, including CK1,
do not show this effect. These data implicate CK2 in a special and perhaps unique role in
maintaining the rate of FRQ turnover specifically at higher temperatures and thereby sculpting
the TC profile.
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A putative CK2 phosphosite mutant in FRQ phenocopies chrono
Extended TC may arise at least in part due to misregulated CK2-mediated phosphorylation of
FRQ at high temperatures. Extended TC might thus also result from eliminating those CK2
target sites on FRQ less effectively phosphorylated at high temperature by low CK2 doses.
Hence, we analyzed FRQ strains bearing mutations in multiple putative CK2 phosphosites,
which increased period length (m3, m4 and m5 from Yang et al., 2003, generous gift of Y.
Liu). However, these strains did not show TC defects (data not shown). We then constructed
a set of S/T→A mutations in FRQ using three criteria to identify candidate sites; sites chosen
were predicted to be CK2 targets (score > 0.5 as assessed by NetPhosK 1.0
http:/www.cbs.dtu.dk/services/NetPhosK/), conserved in at least 3/5 related fungal species,
and known to be phosphorylated in vivo as assessed by mass spectrometry (C. Baker, J. Loros,
J.C. Dunlap, unpublished data). We made multiple rather than individual mutations as CK2
hypomorphs likely result in the coordinated hypophosphorylation of several sites. Thus, we
settled on eight possible CK2 sites and three cluster mutants were made, frqQ1, frqQ2 and
frqQ12 as well as a knock-in version of the WT, frqWT-KI (Fig 6A).
To assess the effects of these mutant frq alleles on TC, we used each to replace the endogenous
locus (Fig 6B). Homokaryotic strains bearing frqQ1 and frqQ12, but not frqwt-KI or frqQ2, show
defects in TC (Fig 6C). frqQ1 strains have a longer period, which persists as temperature
increases. Moreover, FRQQ1 has a longer half-life than FRQWT-KI in vivo at 28°C (Fig 6D).
This pattern is reminiscent of the chr mutant, albeit with a longer period. By contrast, frqQ2
strains show no significant difference from WT. Thus, we predicted and found phosphorylation
sites on FRQ that, when mutated, phenocopy the extended TC characteristic of chr. The
enhanced compensation characteristic of these strains is unique among existing frq missense
Finally, we wanted to ensure that CK2 was capable of directly phosphorylating those residues
that we had mutated in frqQ1. Thus, we made GST and GST-fusions of peptides bearing the
S/Ts of interest (Sup Fig 4A). These were purified (Sup Fig 4B) and radioactive kinase assays
were performed with either recombinant human CK2α/β (Sup Fig 4C, left side) or recombinant
N. crassa CKA (Sup Fig 4C, right side). All GST-peptide fusions were phosphorylated to
varying degrees (see bands at the level of the asterisks) while GST alone was not. The reduction
in phosphorylation intensity between GST-538 and GST-538A indicates that serine 538 is
indeed phosphorylated, while the other serines appear also to be phosphorylated. In the lane
with GST-554/558 either or both S/Ts are phosphorylated and we cannot distinguish between
these possibilities. Under these conditions, bands corresponding to the sizes of various CK2
proteins (arrows) are seen indicating the anticipated autophosphorylation of this kinase (Pagano
et al., 2005). Thus, this together with our previous data strongly suggests CK2 directly
phosphorylates these residues in vivo.
We suggest a mechanistic entrée to the cryptic processes underlying TC, a canonical
characteristic of circadian rhythms. Two classically isolated mutants were chosen because they
showed unusual characteristics, extended TC and overcompensation. Analysis of these mutants
identified CK2 as a key regulator of TC. The identities of chr and prd-3 are consistent with
mounting evidence for the role of CK2 in circadian clocks of many model organisms (for a
recent review see Mizoguchi et al., 2006). Moreover, the epistatic relationship we observed
between chr and prd-3 is consistent with their identities: any activity of the β1 kinase regulatory
subunit requires a functioning α catalytic subunit. The perhaps remarkable coincidence that
two strains selected for defects in period length, and only later shown to have defects in TC,
both bear mutations in subunits of the same kinase emphasizes that CK2 plays a key role in
establishing TC.
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Importantly, these analyses have defined a role for CK2 in TC without solely relying on the
use of missense mutant alleles. While many mutants affecting TC exist (e.g., frq7, perS, perL
etc.), it could not be excluded that their phenotypes were due to temperature-sensitive defects,
irrelevant to their WT function in TC control. The use here of controlled-expression strains
with only WT proteins, has ruled out this caveat. Moreover, these results demonstrate that not
all mutations that change period lead to defects in TC, consistent with other alleles e.g.,
timelessUltraLong (Rothenfluh et al., 2000) and frq1 and frq2 (Gardner and Feldman, 1981).
The data can be thought about in terms of a simple model (Fig 7). CK2 and CK1 are essential
for FRQ phosphorylation and period length determination; however, CK1 dose has little effect
on TC. We show that FRQ is a direct target of CK2 and that FRQ is poorly phosphorylated by
CK2 when CK2β1 is low. FRQ is degraded more slowly as temperature increases when
CK2β1 is at low levels. This is consistent with mutants in which putative CK2 sites are deleted
(frqQ1, frqQ12), thereby phenocopying CK2 hypomorphs. Thus, it appears that certain CK2
target sites are needed on FRQ for efficient FRQ degradation preferentially (though not
exclusively) at high temperature. Since FRQ steady state levels increase with temperature (Liu
et al., 1997), either CK2's differential phosphorylation of these sites on FRQ at high
temperature, or differing effects on degradation (e.g., FRQ might bind FWD-1 more efficiently
at high temperature when these sites are phosphorylated) would counteract the accumulation
of excess FRQ as temperature increases. In this way, we imagine that the TC curve is sculpted,
or to reiterate the conclusion of Hastings and Sweeney (1957), that the effect of ambient
temperature on period length is actively managed by the cell. By contrast, CK1 phosphorylation
of FRQ is not required differentially at different temperatures, as if CK1 is necessary but not
sufficient for timely FRQ turnover at elevated temperatures.
Finally, the data suggest that TC is an independently evolved layer that is not intrinsic to a core
oscillator. In this scenario, through evolution, proto-oscillators that may not have been initially
compensated may have co-opted the functions of utility kinases (such as CK2) to effect TC,
and perhaps for other advantages as well (see` Allada and Meissner, 2005). Since the Q10 of
CK2 activity seems to be within a thermodynamically predicted range, it seems that evolution
has not tinkered much with the catalytic arm of this enzyme to achieve TC. This is reasonable
given the many roles of CK2 in the cell; rather, evolution may have used the necessities arising
from the global involvement of CK2 to an advantage in developing a capacity for TC.
Interestingly, an observation in Arabidopsis is consistent with this view (Edwards et al.,
2005). It seems plausible that effective CK2 activity is held within limits to service the many
roles of the enzyme and that this homeostasis aids TC. We also note that TC is robust as regards
CK2 levels: frank overcompensation was seen only at inducer levels four log orders below
those that gave WT period lengths, levels that drove biochemically undetectable amounts of
CK2β1 synthesis. It is apparent that in this system or in similarly constructed oscillators, two
or even four-fold changes (as from loss of an allele in a diploid) would not affect TC. Instead,
we suspect that subtle changes in target clock proteins—e.g., in a constellation of CK2 sites in
FRQ—may allow the clock to fine-tune TC. This is akin to the idea of a region in
Drosophila PER that is polymorphic across latitudinal clines (Sawyer et al., 1997). A definitive
catalogue and time-history of FRQ phosphorylation would provide further insight into this
Strain constructions and culture conditions
WT Neurospora crassa strains used include 328-4 (ras-1bd A) and 87-74 (his-3; ras-1bd a).
Knockouts (Δcka, Δckb-1 and Δppp-1) were generated by using hph to replace the entire ORF
using the Neurospora Knockout Consortium's methods (Colot et al., 2006). Culture conditions,
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including race tubes and handling of Neurospora were as previously described (Dunlap and
Loros, 2005).
Antibodies and Western blotting
For α-CKB-1, a polyclonal Ab was raised in rabbits against two synthetic peptides,
MDTLKNAANYVGDKVC-amide and Ac-ISDKVSENKHDAKAC-amide, by 21st Century
Biochemicals, LLC (Marlboro, MA). Westerns were performed as previously described
(Garceau et al., 1997). The α-FRH Ab was made as previously described (Cheng et al.,
2005). The α-CK-1a Ab was a gift of M. Brunner.
In vitro kinase assays
Kinase assays with the synthetic peptide were performed with a modified protocol using a
commercial CK2 Assay Kit (Upstate, Lake Placid, NY).
FRQ degradation profiles
FRQ degradation was assessed by Western blot after transfer from light to dark and FRQ half-
lives were calculated by the following equation: t1/2 = ln(2) / m where m is determined from
an exponential fit in the form y = A e-mx. Half-lives were averaged from three independent
curves and standard deviations were determined.
For more details see Supplementary Experimental Procedures.
L. Larrondo kindly provided strains 1061-1 and 1066-6. We thank C. Hammell for technical assistance with GST-
FRQ purification, P. Ruoff for helpful discussions and R. Lambreghts and H. Benchabane for critical reading of the
manuscript. This work was supported by a Canadian Institutes of Health Research Postdoctoral Fellowship to A.M.
and by grants from the National Institutes of Health to J.C.D. (RO1 GM34985 and PO1 GM68087) and to J.J.L. and
Akman OE, Locke JC, Tang S, Carre I, Millar AJ, Rand DA. Isoform switching facilitates period control
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Figure 1. chrono and period-3 bear mutations in CK2 subunits
(A) The TC profiles of chr and prd-3 are shown. Epistasis between chr and prd-3 is observed
with respect to TC and presence or absence of clear banding at 30°C. Values are the mean of
N=3 race tubes ± SD. See also Fig. 2 in Gardner and Feldman (1981) for single mutant TC
(B) SNP mapping of chr reveals linkage to a SNP on supercontig 311 (Neurospora genome
release 3) of linkage group (LG) VI. Genetic data, shown as distance in cM between chr and
the indicated SNP (open circles), were consistent with a location for chr on the end of 311, a
position occupied by ckb-1.
(C) Mapping localizes prd-3 approximately equidistant between SNPs 164a and 164b. This
location was used to identify a series of pLORIST cosmids (horizontal lines) spanning the
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region. Transformation of these into prd-3 recipients revealed rescue only by pLORIST H031
B12. Predicted ORFs within this cosmid included cka (grey box).
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Figure 2. Heterokaryon knockdowns of cka and ckb-1 resemble prd-3 and chr respectively
(A) Reduction of cka dosage in heterokaryons increases period lengths and extends TC. TC
profiles of WT (circles), prd-3 (large triangles) and 7 independent Δcka heterokaryons (small
triangles) are depicted ± SD, reflecting analysis of of 3-6 bands in a single race tube.
(B) Reduction of ckb-1 dosage in heterokaryons reduces period length and results in enhanced
TC. TC profiles of WT (circles), chr (large squares) and 8 independent Δchr heterokaryons
(small squares) are depicted ± SD, reflecting analysis of 3-6 bands in a single race tube.
Heterokaryons that overcompensate across the temperature range are shown with dotted lines.
(C) Four independent Δckb-1 homokaryons display an arrhythmic phenotype whereas two WT
sibling controls have a normal phenotype.
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text: The South Carolina Higher Education Efficiency and Administrative Policies Act, signed into law on August 3 by Gov. Nikki Haley, is a big step for transparency in South Carolina's public institutions. The twofold law requires them to post all purchasing transactions online and eliminates portions of the timely and costly process for having new facilities or major purchases approved.

subset: commoncrawl


text:
Loka Yoga Spring Fling – This Sunday 3/22 8:00 AM – 4:00 PM
Join Loka Yoga in welcoming Spring this Sunday March 22 from 8:00 AM to 4:00 PM. Meet the awesome teachers and students at one of Oakland Magazine's Best of Oakland http://www.oaklandmagazine.com/Oakland-Magazine/2015-Best-of-Oakland-ballot/ nominees for the Best Yoga Studio 2015! Drop in for one or both of the free class with our very own Haley Ebersole http://www.lokayogaoakland.com/p/about.html, join in the fun of our clothing swap, share a meal with your fellow community members at our potluck, and don't forget to make an appointment to give blood from 1-4 pm with Loka's very own Blood Center's of the Pacific mobile donation coach! We look forward celebrating the arrival of spring and the wonderful Oakland community with you this Sunday.
With Love,
Dari and the Loka family
SUNDAY 3/22 Loka Yoga Spring Fling Schedule
8-8:45AM - FREE Meditation with Haley Ebersole
9-10:30AM - FREE Hatha Vinyasa 1-3 with Haley Ebersole
This is a general and intermediate class for most levels, including advancing beginners with general body awareness and/or good listening skills. It offers more variety for those in the class who wish to try something a little more challenging than an Intro class. Variations are offered all along the way so most levels will feel comfortable and accommodated to suit their needs.
1PM to 4:00 PM – Community Clothing Swap and Potluck
Bring clothes you are ready to pass on and score some new items for Spring! All items left behind will be donated. Bring a dish to share and enjoy a community meal with your fellow yogis and neighbors.
1:00 PM to 4:00 PM – Blood Drive with Blood Centers of the Pacific Mobile Coach
Please register yourself for the Blood Drive now
Appointments are available to help you plan your day of fun!
click on DONATE BLOOD, and enter sponsor code: Loka
Appointments are available to help you plan your day of fun!
Loka Yoga
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Feel good about feeling good
Try these exercises to feel good about feeling good:
Reframe the way you think about your choices.
If you have to make yourself work out regularly, write a list of all the things you enjoy about exercising.
Examples: I love the energy I have after running, I love being able to fit into my favorite clothes, I can balance on one foot thanks to yoga.
Do what's best for you — not just what you think you should do.
In one recent experiment, researchers offered people a choice between a gift certificate for groceries and one for spa services.
The people who made the "financially sensible" choice (groceries) were then angrier than the people who picked the spa certificate.
If you can afford to visit a spa, you shouldn't deny yourself the trip because someone else might consider it frivolous.
Allow yourself the occasional indulgence, and indulge in things that have some health benefits.
Upgrade to whole-wheat pasta, or choose dark chocolate over milk chocolate for the antioxidant boost.
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Search for mussels comes up empty
On Wednesday afternoon, members of the Douglas County Dive Team went in search of zebra mussels on Lake Victoria.
They were accompanied by two members of the Department of Natural Resources Invasive Species Program, who walked along the shoreline.
The divers and DNR spent at least two hours on Lake Victoria, mostly on the north end near the Lake Victoria / Lake Geneva channel. They also spent some time on Lake Darling.
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Photographer's Corner; Your Brand, As Unique as Your Fingerprints
February 08, 2013
In the photography industry the term "branding" is tossed around quite a bit. What is branding? And why is it important to us as photographers? A good place to start find...Read the Full Post »
Product Spotlight: Lensbaby Part I
January 31, 2013
Lensbaby Composer Pro with Sweet Optic 35 by Laura Tillinghast LensBaby's selective focus lenses have piqued my interest for sometime now. Back in college, when I was st...Read the Full Post »
Mpix: 25% Off All Gallery Wraps
January 31, 2013
Mpix has just released their first promotion of the year. Purchase any size canvas gallery wrap (sizes range from 8x10 to 24x36) for 25% off. If you've made a New Year's...Read the Full Post »
Package Deal: Balancing Customer Demands with Quality Control
January 18, 2013
Sunee is a professional family portrait photographer living in the San Francisco Bay area. She started photographing children and families when she worked as a nanny and...Read the Full Post »
Photographer's Corner: Tips for Tax Time
January 04, 2013
With tax season approaching we know that it can be somewhat daunting and even overwhelming to prepare your tax returns. As a small business owner there are a lot of thing...Read the Full Post »
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text: ArcGIS Jobs 2 were found based on your criteria Post a job. It's free! GIS Mapping and geological data collation Hourly – 3 to 6 months – 10-30 hrs/week – Posted Feb. 12 2015 Exciting opportunity to work with super switched on young Australian team in building and scoping tenement changes in Western Australia All systems and data are publicly available The applicant will need to utilise a high-end GIS such as MapInfo / Arc GIS or similar The task will require; - Building a 'base' map from available public data (geological characteristics with tenement overlays) - Daily updating of tenure changes through excel downloads and GIS updates - Identification of newly available tenure - Filtering of newly available ... arcgis data-analysis database-management data-entry data-mining digital-mapping gis internet-research mapinfo microsoft-excel web-scraping GIS project Fixed-Price – Est. Budget: $30.00 Posted Nov. 16 2014 I have a small course project on GIS, which involves the basics of ArcGIS. I am looking for someone to get it done. To give you an idea, what are the expectations of the project: 1. You are required to find out a project on your own such as: Using GIS to determine change of the forest levels for the last 5 years in a specific part of the world. Using GIS in determining water table decline in the world ... arcgis gis


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Photos That Will Tug On Your Heartstrings
New Diabetes Breakthrough?
5 Simple Rules for Giving Your Pet a Healthy Life
This 15-Year-Old Babysitter Is Making $500,000 a Year
A Baby Costs How Much in Those First Few Years?
10 Air Travel Tips for Dogs
Why You Need to Run Lemonade Kool-Aid In Your Dishwasher
No One Believes These Teen Girls Are Twins
The Chow Chow: Beautiful, Strong and Independent
20 Unforgettable Vintage Photos of Real Circus Freaks & Sideshow Acts
30 Really Good Examples of Why Japan is So Weird
21 Things That Could Only Happen in Russia
She Dyed Her Kitten Pink, and Then the Kitten Died (Or Did It?)
The Miracle 21-Week-Old Baby Who Survived Against All Odds
5 Hardest Yoga Positions to Practice
2-Year-Old Boy Meets with Hero Who Has the Same Rare Condition
The 10 Creepiest Things Ever Unearthed During Archaeological Digs
Woman Gets Ultimate Revenge On Childhood Bully
15 Idiotic Laws That Only Exist in America
Father Forces "Grown" Daughter to Dress Like a Princess
These Terrifying, Oversized Animals Will Haunt Your Dreams Forever
Hitler's Last Food Taster Feared Death with Every Bite
10 Foods Banned In The U.S.
30 Convenient Ideas You Never Knew You Wanted
10 Inexpensive Date Ideas for New Relationships
Do You Know That There Is A Devil Santa Clause Too?
The Worst Parents of 2014
Supermarkets & Toy Stores Welcome Ugly Fruit
28-Year-Old Creates App that Lets You Take Back Texts
Did Ray J Bail on Reality TV to Save His Romance?


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GreenPower Premium
50,000 - 100,000 downloadsAdd this app to your lists
+ By Binary Mango AS
★ Support for Mobile Data on Android 5.1: Only for rooted phones. ★
"Charge less by extending your battery life with this amazing app""!! (Sprint newsletter)
Unlike other battery savers like JuiceDefender that require regular user manual actions, GreenPower is an easy battery saver that is fully automatic: Once configured, it runs and saves your battery by itself. It does so by smartly managing Wifi, Mobile data, Bluetooth and Screen Brightness: Turning them off when you don't need them, but ensure minimum disturbance: Automatic turned ON when needed by you or by other apps (e.g retrieving mails). Easier, cleaner and faster than competitors like JuiceDefender , 2x battery saver and easy battery saver!
GreenPower battery saver: 1.5 million downloads & 2 years of experience in Android Battery Saving!
"GreenPower is a simple and easy battery saver, battery booster and power management software" (Engadget Chinese edition)
★ Adds up to 100% battery life depending on usage
★ Automatically manages your Wi-Fi, Mobile Data (2G, 3G, 4G), Bluetooth, GPS, display brightness
★ Battery saving works with almost all devices and carriers
★ Supports all Android versions from Froyo to Lollipop
★ Easy yet powerful battery saver and battery booster
"GreenPower battery saver mainly does one thing, but does it so well and so flexibly that the benefits reaped are considerable." (InfoWorld)
Or take time to configure the battery saving settings to get the most out of it and the best battery longevity!
Try Premium version for 24H and get a refund if you didn't like it by just sending us a mail.
"GreenPower battery saver is a remarkable tool. Battery booster, time saver, top notch!" (AndroidPit)
• Battery life indicator in notification area.
• Wifi management: Based on schedule, screen state, location, power connected, signal level...
• Mobile data management: Based on schedule, screen state, power connected...
• BLUETOOTH (premium) management: Based on devices connected or in range, screen state, power etc
• Traffic check: Configurable to prevent disturbing other apps
• Apps Whitelist & Blacklist (premium): To keep data on when using specific apps (like music streaming)
• Simple WIDGET (premium): To quickly pause or resume
• Night / Sleep mode (premium) fully configurable, Airplane mode...
• Tasker & Locale Plug-in (premium)
• Compatible with Cerberus 2.0
• Very lightweight and fast app
• Easy and clean interface
• Fully configurable settings
• Settings backup / restore
• 30 Languages
• And more coming all the time...
• Dedicated Forum for support issues
• Support from the developer. I'm answering each issue.
• Regular updates with bug fixes and new features
• Widget
• Night mode (sleep mode)
• Tasker & Locale Plug-in
• Location-based Wi-Fi
• Apps whitelist & blacklist
• No ads
• The good feeling to support an independent developer
Tags: green power premium, greenpower premium, green power premium apk, green power premium 5.0, green power battery saver, green power battery saver premium, paristo green power, green power premium review, green power battery saver premium v3.1, greenpower premium apk.
This new release brings several improvements and corrections.
The complete release notes are available online:
(69 stars)
Has got to be the best on the market. Dev always responsive! Great job!
(69 stars)
The app automatically manages data which helps conserve battery life so you don't think about it.
(69 stars)
It works good but settings are too complicated and poorly explained . it needs a complete UI renovation and settings explained a little better. But all in all it does the job.
(69 stars)
Even with set on permantly, data and WiFi always seem to go off and on randomly. Have to manually turn them back on.
(69 stars)
Used to love this app for saving my battery but since upgrading to android lollipop it will no longer turn off mobile data without rooting. Dont much see the point in it anymore!!
(69 stars)
I wish that all functions would work without rooting, especially the premium edition.
(69 stars)


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Use of Resolidified Melts in Infrared Spectroscopy: Application to Polymorphism and Solvation Studies
Applied Spectroscopy, Vol. 15, Issue 6, pp. 157-159 (1961)
Acrobat PDF (289 KB)
A sample melt technique has been developed for the rapid identification by infrared spectra of polymorphs and solvates. Excellent spectra are obtained; no decomposition or rearrangements have been detected in studies with 35 compounds, which included 25 steroids. Resolidified melt spectra of known polymorphs have been identical. The problems of low solubility and unusable regions of the spectra encountered in solution studies are avoided. In the case of solvates, the solvent may be identified by its spectrum in the gaseous or liquid state, and the spectrum of the solvent-free compound may be determined on the resolidified melt.
Marvin F. Grostic and George E. Bronson, "Use of Resolidified Melts in Infrared Spectroscopy: Application to Polymorphism and Solvation Studies," Appl. Spectrosc. 15, 157-159 (1961)
Cited By
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CWB DB9 138 cm. Pro Series Wakeboard Blank with FinsItem # HN-CNN230
- PVC foam core wakeboard
- Features poplar wood stringer tip to tail
- Smooth bottom design for clean ride
- Variable edge rail with bevel
- LLightweight, strong wakeboard with fins
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KitchenAid KSM150PSGC 5 Qt. Artisan Series Stand Mixer - Gloss Cinnamon
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Precision Cape Cod Chicken Coop
Hayneedle Celestron AstroMaster 114 EQ Reflector Telescope
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Description: Classic Comfort Everyday Design. Soft Touch® supreme pillow inserts have the look and feel of down without the price. These pillows are filled with 100% recycled polyester fiberfill and feature a 65% polyester 35% cotton 200 thread count high quality ...
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Description: Pellon® Pillow Forms are perfect for when you want to make a unique and wonderful pillow! They are soft, yet supportive and economically priced. They are ideal for the Home Crafter or Decorator. Cover is brushed microfiber. Fill is Siliconized ...
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Hank answers a caller who asks for an explanation of 1 Corinthians 15:1-2 in context.
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Advantages of Valuing Diversity in the Workplace
With a varied labor force, comes numerous various concepts, viewpoints and views that include strength to employees' capability to interact, provide and plan. Right here is how having employees with various backgrounds can benefit a business:
A varied and inviting office where every employee feels supported and equipped, is the very best technique to keep and bring in quality employees. With high-performing employees, come enhanced efficiency, quality and spirits. When employees see that their business values and promotes diversity, they are more probable to be pleased with their tasks, and this in turn makes them work even harder.
Employees with various backgrounds, offer an organization with lots of various viewpoints and concepts on ways to enhance its items, services or advertising techniques, which assists to much better record the marketplace section where they originate from. This, in turn, assists in making more ingenious choices focused on benefiting the business.
Enhancing performance can be as easy as giving everyone respect
Smart, growing business that wish to take the lead in development value the value and understand of imagination. As an outcome, they frequently motivate their employees to come forward with originalities that can assist enhance their business.
Companies offer their services and items to a progressively varied consumer base. Having personnels that comes from various market sectors and demographics can actually assist an organization comprehend its consumers much better, leading to enhanced services.
With a deep understanding of various cultures, an organization will certainly have the ability to customize its methods to item formulation, sales and services to match requirements of various customers. In addition, it will certainly be in a much better position to satisfy the numerous customers' requirements far better than their rivals.
Better development will come through improving your work's respect level
In a progressively around the world market, lots of business with abroad operations benefit considerably from employing multilingual employees, as they have the ability to interact with consumers from various ethnic backgrounds. Having a varied labor force gets rid of language obstacle issues, allowing a business to reach a larger customers base.
Terrific skill can be discovered by employing both global and domestic employees. That is why some business are prepared to sponsor the migration for global employees when the abilities they are looking for are in brief supply back house.
Diversity in the office makes a business more powerful, which assists to enhance development and success. Employees from various backgrounds will certainly supply a business with a much better viewpoint concerning the requirements of all its customers; not simply a couple of groups. They can provide important understanding on the finest client service suggestions that will certainly assist enhance sales amongst their particular groups.
There are numerous factors which could lead employees to do something about it that show no stability or respect at all. The majority of the time, these factors could be asserted sensible in one usual viewpoint called "completion validates the ways". The outcomes of some of these actions cost the business terrific loses.
Some reasons for lost work respect and stability consist of the following:
If you like this go to Http://www.frameAssociates.com/ for more.
Lacking of proper management causes more failures and slowdowns
A lot of high level employees sitting on managerial positions tend to utilize old clinical management concepts where lower level personnels are not included in choice making. The employees start to work simply for the sake of completing the appointed tasks and will certainly lie to cover up their faults and even toss away any proof that might jeopardize their word.
Employees discover it traumatic to approach management with their concerns and end up taking the incorrect choices. This constantly results in employees experiencing ill mindsets in the direction of management and might lead them to take things for given.
Language barriers will be a thing of the past with diversity
With a varied labor force, comes lots of various concepts, point of views and views that include strength to employees' capability to interact, provide and plan. Right here is how having employees with various backgrounds can benefit a business:
A varied and inviting work environment where every employee feels supported and equipped, is the finest approach to keep and draw in quality employees. When employees see that their business values and promotes diversity, they are more most likely to be pleased with their tasks, and this in turn makes them work even harder.
Employees from various backgrounds will certainly offer a business with a much better viewpoint relating to the requirements of all its customers; not simply a couple of groups.


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What's New
Virtual Reality Showcase
- Virtual Fashion demonstration, showing digital fabrics draped over models.
- 3D scanning technology, utilizing the Matterport camera.
- Synthetic 3D environments. Including Architectural Visualisations.
circusAR(Augmented Reality)
Scanning can do wonders.
Marvelous and sublime harmony of reality and virtual reality
●How to Use circus AR
1. Run the app and scan the object, directly facing it to the full screen of your device.
(For a practice, scan the front and back side of 1$ ~ 100$ bill)
* Japanese Yen: 1,000 Yen – 10,000 Yen
* Chinese Yuan: 1 Yuan – 100 Yuan
2. After recognition, contents will be brought on with vibrations (please wait for a moment)
(The downloading speed will be different according to your internet speed and content size)
3. Enjoy various augmented reality contents already registered.
(We are offering more than 4,500 types of contents – being updated continuously)
● Helpful Tips
- Not all images but the images registered at circusAR can be recognized by the app.
(You can scan the PC screen or the printout)
- Without the recognition process, you can directly enjoy the augmented reality from AR contents List, 3D Model View and AR Photo.
* Run the app> Menu > Contents List, Play More > Select the content > display the content
- For clear recognition, scan the image or objects directly facing them to the full screen of your mobile device.
- Internet connection is necessary. We recommend 4G or LTE rather than WiFi.
* Inquiry for Content Registration firstname.lastname@example.com / +82-2-6261-2330
●Application Fields of circusAR Service
We provide our augmented reality service in many different industries: education, exhibition, convention, MD products, entertainment, marketing promotion, game, publication, tourism, art, performance, printed publication, product packaging, etc.
●What's New? (2015.02.25)
- Faster recognition speed & 3D object recognition function
- Contents can be utilized without recognition-process (Contents List)
- My page (Bookmark, History and favorites list)
- 3D Sketchbook (AR coloring book), 3D Model View, AR Photo, Etc.
- Moving and re-sizing contents images function
- New contents platform (Audio gallery, Docent, Image show, Magazine and other 10 kinds.)
●About circus company
We make your ambitious visions come true.
We research to develop Augmented Reality (AR) and new technologies, and create an amazing experience and enjoyable contents; we are ICT+ creative partners.
When you meet us, technology becomes an exciting game and contents become more attractive.
We offer you something charming, beyond what you expect.
www.circusar.com/download/circusAR_proposal.pdf (circus company Introduction)
●Contact us
With circusAR, IT is fun and interesting.
firstname.lastname@example.com / +82-2-6261-2330
Homepage : www.circusAR.com
Facebook : www.facebook.com/circusAR
Youtube : www.youtube.com/circusAR1
Twitter : www.twitter.com/circusAR
Pinterest : www.pinterest.com/circusAR1
Instagram : www.instagram.com/circusAR_circuscompany
Vimeo : www.vimeo.com/circusAR
※ We love to collaborate with talented artists to create diverse AR contents.
We always welcome you."
Stairs Tools Free
This version includes some tools of the full version.
It serves to make you understand how the application works and to study the interface.
If you like you can buy the full version.
Remember that Google gives you 15 minutes to try the applications you buy.
Please do not give 1 star because the application is not full.
"Stairs Tools" is a fantastic tool that allows you to design various types of stairs in few seconds.
Choose the type of stair among those available, enter the height of the stair and calculate it.
The app will calculate all the parameters of the stair.
After, you have the ability to change the settings you want, and recalculate the scale to suit your needs.
Watch the video on youtube to find out how easily you can design a staircase.
*If you think this app can be usefull for you please rate it for support us.
Follow us on Google plus and follow our YouTube channel.
Download other product for engineering and architecture.
- Beam Tools
- Pillar Tools
- Balcony Tools
Amway Realidad Aumentada
Para usar la aplicación solo debes abrirla desde tu dispositivo, apuntar y escanear las imagenes con el logo de Amway Realidad Aumentada.
HOLO - Virtual Reality
Mode 1 : Stereoscopic 360° Video Demo - CITY TOUR
Mode 2 : Interactive Realtime 3D Walktrough
Mode 3 : Liveview with recording and social media sharing
1. Download and install the HOLO Virtual Reality App.
2. Press START in the main menu and select one of the available contents
(3D, Video, Live).
3. Insert your smartphone into your Virtual Reality viewer and secure it.
4. Enjoy!
In Virtual Reality mode look downwards to activate the menu at anytime.
The bigger center square is always a On/Off button to trigger
actions. In the 3D world this button activates/deactivates walk mode.
In the video example you can use it to start/stop the active sequence.
In the live example it allowes you to start/stop recording what you see and later share it on Facebook,
Youtube and other plattforms. By looking at the rectangles left or right of the big square
you can switch between the different contents (3d, Video, Live) - Or look at Exit to switch to the main menu,
where you can continue with functions outside of your Virtual Reality device.
SketchUp Mobile Viewer
View models in your own 3D Warehouse account, or explore the millions of free designs in the 3D Warehouse -- the world's largest collection of 3D models.
Explore and share 3D models:
• Easily navigate complex models using intuitive multi-touch gestures for orbit, pan, zoom.
• Use the Camera menu to select any of the standard camera views (top, side, bottom, etc.) or choose from the list of presaved scene-based camera angles (note: camera angles are the only supported scene property at this time).
• Zoom out to the extents of a model instantly to easily reorient your view.
• Search and browse through over 2 million, free 3D models on 3D Warehouse.
• Use the detailed search result info to preview model info before downloading to your device.
• Download models to your device from 3D Warehouse for seamless offline viewing.
For more information, visit: http://help.sketchup.com/en/article/3000042
This app is recommended for Android tablets running Ice Cream Sandwich(4.0) or higher with a minimum of 1024Mb of RAM. Signing in to 3D Warehouse requires a Google ID.
Visuar Augmented Reality
With "Visuar Augmented Reality" you can upload your own models and show in 5 minutes.
Visuar is a generic visor that you can use for whatever you want, publicity, medicine, education, tourism, architecture, marketing, etc.
Besides, with Visuar you can make advertising and marketing campaigns at very low cost.
With the augmented reality leader App have:
- Your own channel with your corporate image.
- You can move, rotate and escalate in all perspectives.
- Add video, sounds, animations and video textures to your models.
- Change the textures in real time.
- Add personal targets in real time.
- Take photos and send by social networks and email.
- You can center the view int the model and saw the interior.
… and it´s only the beginning.
You can have your own channel now in http://channels.visuar.es, for FREE.
Would you like to try it right now?
1-Download the app for free.
2-Print the marker that you will find at http://www.visuartech.com/target.pdf
3-Select the project you wish to view in Augmented Reality and download it.
4-That is it! Enjoy!.
Visuar is been developed by Visuar, leading company in the development of augmented reality applications to all needs for any productive sector.
Visuar brings you a huge and attractive potencial so far never seen.
Architecture, engineering, tourism, food, fashion, manufacturing, advertising, naval automotion, entertainment, marketing, medicine, education, and more...
Visuartech will advice you the best way to integrate your projects to Augmented Reality in a suitable and useful way for your business.
If you have any questions don't hesitate to contact us.
Phone +34902929233
Skype: visuar-ar
R U McSafe Augmented Reality
The R U McSafe app uses stunning Augmented Reality (AR) to showcase health and safety in the workplace. You can use your mobile device to view various different safety scenarios in realtime/space (using the printed target provided)
You can move the mobile device at any angle and the 3D simulation will stay in relation to space around you in real-time as long as the printed marker is constantly in view.
Get the marker:
What easy steps do I need to do before I can view the Amazing R U McSafe Augmented Reality?
Step 1 Print the image of the Marker (This is important as you will need the image printed and placed on the desk/table in front of you) - Why? The printed image is called a marker or target and will be the 'stage' where the 3D objects will magically appear from. Without the target the app will not function.
Install the APP ?
Step 1 Install the App - This will take a few minutes dependent on connection speed
Step 2 Once the App is installed - run the App
Step 3 Once open simply position your mobile device or tablet so that the printed marker fills the screen when viewed from above. Keep the printed marker onscreen at all times and watch the 3D appear as if they are right in front of you!
You can interact with the engaging 3D animation by using your finger to spin the icons and then select the scenario you wish to view.
Step 4: Updates - As part of a continual development programme we will from time to time improve and update Apps at any point in the future without prior written notice.
HP Realidad Aumentada
1. Descarga gratis la aplicación HP Realidad Aumentada desde Google Play o desde el App Store. Para esto tienes que escribir "HP Realidad Aumentada" en el buscador e instalar la aplicación.
2. Entra a tu nueva aplicación y dirige la cámara de tu dispositivo hacia las imágenes que encontraras en la siguiente liga:
3. Interactúa con las animaciones en Realidad Aumentada y descubre todo lo que ofrece hp y sus campañas al medio ambiente. ¡Vive la experiencia!
Esta aplicación es compatible con tu iPhone, iPod y iPad.
Virtual Environments
With a sleek, finger-friendly design, the VE app experience allows you to engage in fully immersive, interactive events. The intuitive user interface mirrors our desktop platform, allowing you to quickly and easily leverage the same great features and functionality.
• Live and VOD Webcasts complete with Video, Slides, Q&A, Polling
• Mobile eLearning Courses and Certification on Android tablets
• Search and discovery of content
• Interactivity with built-in chat and networking tools
• Engagement with subject matter experts rooms
• And much more…
Login Process:
• To attend a virtual event via your Android mobile device, please download this app first.
• After downloading and installing the VE app, go ahead and register for the event at the URL specified in your event invitation.
• On the confirmation page or within the confirmation email you've been sent, click the URL provided and you will be prompted to install the mobile app. Since you're already installed the VE app, simply confirm you have already installed it and you will enter the virtual event. *If you have not yet installed the app, please do so via the link on the mobile app page and start at the top of the Login Process.
• Prior to joining your first virtual environment, the VE app must be installed.
• A minimum Android version of 2.2 (Froyo) and above is required.
• Users must be on a WiFi or 3G/4G network connection with a recommended broadband connection of at least 2 Mbps.
• Presentation quality is dependent upon quality of the network connection.
• Android smartphones only support Conference Centers and Webcasts. Please use an Android tablet or iPad if attending an eLearning course or Virtual Event with booths.
3D Print: 3D Printing Ideas
Qubism 3D modeling
Qubism has been designed from scratch with a dedication on touch operation and performance. Focus on and Zoom in on areas that need work at higher precision and Qubism will accommodate.
Work is ongoing and takes a fair bit of my free time but constructive feedback is welcome. Please email me if you experience any problems.
Exporting and saving Images and Models:
Select the 'camera' tool from the leftmost menu and select the file type from the rightmost (tick) menu.
•PNG: Show off your model to you friends, should work with any social and photo apps installed
•JSON: Share you model with other Android devices running Qubism via email with this option
•DAE: Export your model in Collada format to develop it further with tools like Blender or Sketchup
All these files can be sent via gmail or saved to SD card if you have an app like 'Send to SD card' installed.
All tools available in the menu only need a single touch to operate but the following shortcuts can speed things up:
•Cancel operation in progress: Tap screen with a second finger while the first is still on the screen
•Pan, Zoom and rotate: Use two fingers on the screen (e.g. pinch screen)
•View: Place two fingers on the screen then remove one
•Focus: Place one finger on a qube and tap twice quickly with a second finger
Known limitations:
•Qubes that are too close or two far away from the centre of the red axes will be cropped by OpenGL. Keep the red axes near the area you want to work on using the focus and pan tools before zooming in.
•The line removal rendering options are not perfect, it trims outlines where two Qubes meet but this can have unexpected results.
•Transparency requires Qubes to be rendered back to front, this is fine when editing but in camera mode this order is only calculated when the rendering mode changes and may be incorrect once you change the view. The new refresh button will appear once you change view and will recalculate the order but artifacts are still possible.
•Out of Memory issues can unavoidably occur outside of my control. Qubism will save your model for you and it will have a ? for its thumbnail when you re-enter the application.
•Qubism will never be able to load existing 3D file formats (Obj and even DAE) because it would be extremely difficult and often impossible to convert triangles meshes into qubisms internal format.
Autodesk FormIt
Express design ideas with easy-to-use tools
• Create 3D forms quickly by using our innovative 3D sketching system
• Sketching is not constrained to one plane
• Use other objects in the scene as snap references, including levels
• Sketch lines, splines and circles with local and global axis alignments and snapping options
• Directly manipulate forms using gestures for easy model changes
• Move objects more accurately by first selecting the point you wish to snap with
Easy access to site information
• Set project location in a searchable maps interface
• Review historical climate data based on location
• Import satellite image of your site
• Design directly in the context of your proposed building location
• Explore the effects of the sun using the location-aware Shadow Study tool
Collaborate with team members
• Store and share designs in the cloud using Autodesk® 360 cloud services
• Easily transfer designs to desktop applications via RVT and OBJ file formats
• Explore design ideas and add detail to early design forms using award-winning BIM technology
Participate in the community
• We want to hear about how you use FormIt! Take our survey at: https://www.surveymonkey.com/r/formit_android
• Our support forum has moved to: http://forums.autodesk.com
• Visit us at http://formit.autodesk.com and share your images with the community
• Follow us on Twitter @FormIt3D
AutoCAD 360
AutoCAD 360 is a free*, easy-to-use drawing and drafting mobile app that allows you to view, edit, and share AutoCAD drawings—anytime, anywhere. Simplify site visits and fieldwork with powerful design review and markup tools—available online and offline. Seamlessly share drawings with others across desktop, web, and mobile devices. Easily access drawings from web browsers using the free companion AutoCAD 360 web app. AutoCAD 360 Pro subscription plans further expand the functionality of the app, bringing additional capabilities to the field.
• Open 2D and 3D DWG drawings uploaded to your free* AutoCAD 360 online account
• Work offline, then easily upload changes when you are back online
• Upload and open files directly from email on your device
• See all aspects of your DWG file, including external references, layers, and image underlays
• Use multi-touch zoom and pan to easily navigate large drawings
• Use your device's GPS to orient yourself within a drawing
• Draw and edit shapes with accuracy using Snap
• Select, move, rotate, and scale objects
• Add and edit text annotations directly on your drawing; no need for paper mark-ups
• Validate distance measurements in the drawing while you are onsite
• Save edits to your AutoCAD 360 online workspace, so your drawings stay up to date
• Share your designs with others directly from the mobile app
• Add comments and images and invite responses using the Design Feed
• Print remotely from your mobile device using HP's ePrint & Share service
• Plot your designs to PDF or DWF and share them via email
AutoCAD 360 Pro Subscription Plans
AutoCAD 360 is free* to download and use. AutoCAD 360 Pro subscription plans allow you to get much more out of the app, enabling:
• New drawing creation
• Larger file size support
• Connectivity to third-party storage, including Dropbox, Box, and more
• Advanced layer management
• Properties and attributes
• Block libraries
• Drawing coordinates
• Priority support
• Increased storage space
• New tools and much more
AutoCAD 360 Pro subscription plans are available in the following options:
• AutoCAD 360 Pro monthly for $4.99
• AutoCAD 360 Pro annually for $49.99 (save $9.89 vs. monthly membership)
• AutoCAD 360 Pro Plus annually for $99.99
Subscriptions will be charged to your credit card through your Google Play account. Your subscription will automatically renew unless cancelled at least 24-hours before the end of the current period. You will not be able to cancel a subscription during the active period. You can manage your subscriptions in your Account Settings after purchase.
* Free products and services are subject to the Autodesk, Inc. terms of use that accompany them.
Terms of use: http://usa.autodesk.com/adsk/servlet/item?siteID=123112&id=21310328
Privacy policy: http://usa.autodesk.com/privacy/
Floor Plan Creator
* Predefined room shapes as well as free drawing of a room shape with S-Pen, mouse or touch.
* Symbol library: doors, windows, furniture, electrical, fire survey.
* Cloud synchronization to automatically backup and share plans between devices (purchased, 3 days free trial)
* Export as image (free with watermark); DXF, SVG (purchased).
* Supports metric and imperial unit systems.
* Supports Leica Disto™ bluetooth laser meters: http://www.youtube.com/watch?v=xvuGwnt-8u4