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Epigenetics refers to heritable changes in gene expression that arise from changes in
chromosomes without alteration of DNA sequence. These changes occur throughout all
stages of development or in response to environmental factors such as exposure to toxins
or chronic stress and are implicated in diseases such as cancer. Epigenetic mechanisms
of gene regulation, which collectively make up the epigenome, include modifications to
DNA and histone components of nucleosomes as well as expression of noncoding RNAs
(ncRNAs). These modifications can affect gene accessibility to DNA-binding and
regulatory proteins such as methyl-CpG-binding proteins, transcription factors, RNA
polymerase II and other components of the transcriptional machinery, ultimately altering
transcription patterns, often in tissue- and cell-specific ways. A schematic diagram
showing the most well characterized epigenetic modifications are shown in Figure 16.1.
return to top of page
In vertebrates, DNA methylation occurs on the 5C position of cytosine residues to
yield 5-methylcytidine. This occurs almost exclusively within CpG dinucleotides,
although nonCpG methylation does occur in plants (primarily CpNpG and CpHpH methylation,
where H = A,T,C) and to a lesser extent, mammals. Other forms of cytosine exist,
including 5-hydroxymethylcytosine, 5-formylcytosine and 5-carboxylcytosine (Kriaucionis
and Heintz, 2009; Ito et al. 2011; Pfaffeneder et
al. 2011), which may be intermediates in a pathway for DNA demethylation.
In mammalian genomes, approximately 70–80% of CpG dinucleotides are methylated.
However, stretches of CpG-rich sequences with low levels of DNA methylation, known as
CpG islands, exist (reviewed in Blackledge and Klose, 2011; Deaton and Bird, 2011). DNA
methylation is typically associated with epigenetic gene repression, and many targets of
de novo DNA methylation during differentiation are promoters of stem cell- and
germline-specific genes (Weber et al. 2007; Mohn et
al. 2008; Farthing et al. 2008). DNA methylation also
recruits methyl-CpG-binding proteins, which recruit additional proteins that add
silencing modifications to neighboring histones. This coordination between DNA
methylation and silencing histone marks leads to compaction of chromatin and gene
CpG islands (CGIs) make up only 0.7% of the human genome but contain 7% of the CpG
dinucleotides. CpG islands often are highly enriched at gene promoters, and
approximately 60% of all mammalian gene promoters are CpG-rich. CpG islands are
typically unmethylated, open regions of DNA with low nucleosome occupancy. As such,
CpG islands promote relaxed chromatin structure that favors active transcription,
known as euchromatin, and increases accessibility of RNA polymerase II and other
components of the basal transcription machinery to the transcription start site. Most
CGI promoters have heterogeneous transcription start sites and lack TATA boxes, so
transcription factors with CpG in their recognition sites, such as SP1, can help
recruit TATA-binding protein to promoters without TATA boxes. Without additional
regulatory signals, transcription from CGI promoters results in nonproductive,
bidirectional cycles of initiation and premature termination. The regulatory signals
required for the transition from this nonproductive state to productive, directional
synthesis of full-length transcripts are not yet well characterized.
The mechanisms that keep CpG islands free of methylation appear to involve binding
of transcription factors and other transcriptional machinery or the act of
transcription itself. However, CpG islands can become hypermethylated (Meissner
et al. 2008; Mohn et al. 2008) to
silence specific genes during cellular differentiation, genomic imprinting and X
chromosome inactivation.
DNA methylation is catalyzed by DNA methyltransferases (DNMTs; reviewed in Carey
et al. 2011). DNMT3A and DNMT3B are involved in de novo
methylation (Okano et al. 1999) and are targeted to particular
genomic regions by specific histone modifications. During DNA replication, the
protein Np95 recognizes hemimethylated DNA and directs DNMT1 to the replication fork
to maintain patterns of DNA methylation (Pradhan et al.
The methylation status of a DNA sequence can be determined using a variety of
techniques such as the use of restriction enzymes (REs), which recognize short DNA
sequences and cleave double-stranded DNA at specific sites within or adjacent to
these sequences. Some REs are sensitive to methylation and will not cleave DNA if a
cytosine in their recognition sites is methylated, while other REs are insensitive to
methylation. The methylation-sensitive RE HpaII was used in early epigenetics studies
to determine that 55–70% of all HpaII sites (5'-CCGG-3') are
methylated in the mammalian genome (Bird, 1980; Bestor et al.
1984) and to identify CpG-rich, hypomethylated DNA regions [known as HpaII tiny
fragments (HTFs); Bird, 1986].
A list of REs that are sensitive to CpG and CpNpGp methylation can be found in the
Technical Reference section of the Promega web site
Pairs of isoschizomers where one RE is insensitive to methylation and the other is
sensitive (Table 16.1) are often used to query methylation status. DNA fragments
generated by a methylation-sensitive isoschizomer will differ in size from fragments
generated by a methylation-insensitive isoschizomer. The extent of cytosine
methylation can be estimated by calculating the ratio of the different DNA fragments.
For more information about restriction enzymes, visit the Promega Restriction
Enzyme Guide.
Bisulfite sequencing refers to techniques that assess DNA methylation through
bisulfite conversion, which converts unmethylated cytosine residues to uracil
residues. Methylated cytosine residues remain unmodified (Frommer et
al. 1992). The target DNA is purified, alkaline- or heat-denatured,
treated with sodium bisulfite, cleaned up, treated with alkaline, then cleaned up
again to remove salts and other components that can inhibit downstream applications.
DNA purification kits, such as the Wizard
SV Gel
and PCR Clean-Up System (Cat.# A9281), are commonly
used for this purpose. After bisulfite conversion and DNA cleanup, the DNA is
amplified by whole genome PCR, and the amplified products are analyzed using a
technique that distinguishes products derived from unmethyled DNA, which contain
thymine residues, from products derived from methylated DNA, which contain cytosine
residues. These techniques include pyrosequencing, methylation-specific PCR,
methylation-sensitive single-strand conformation analysis (MS-SSCA; Bianco
et al. 1999), high-resolution melting analysis (Wojdacz and
Dobrovic, 2007), methyl cytosine immunoprecipitation (mCIP; Zhang et
al. 2006), bisulfite methylation profiling (BiMP; Reinders et
al. 2008) and MALDI-TOF mass spectrometry (Schatz et
al. 2006). For high-throughput analysis, bisulfite-treated DNA can be
analyzed using microarrays with two sets of oligonucleotide probes, one of which is
complementary to cytosine-containing DNA and the other complementary to
thymine-containing DNA.
Typical bisulfite conversion protocols involve long incubation times under harsh
conditions, resulting in highly fragmented DNA. Promega offers the
MethylEdge™ Bisulfite Conversion System (Cat.#
N1301), which results in efficient DNA conversion and recovery with
reduced template fragmentation using a protocol that can be completed in less than
two hours, including desulphonation and cleanup. The MethylEdge™ Bisulfite
Conversion System does not require an additional cleanup kit.
MethylEdge™ Bisulfite Conversion System Technical
SV Gel and PCR Clean-Up
System Technical Bulletin
The firefly luciferase reporter protein (Fluc) can be used to assess DNA
methylation at the genome level or at specific DNA sequences. Researchers have
developed split-luciferase biosensors composed of two fusion proteins: a DNA-binding
domain fused to the N-terminal portion of Fluc, and a second DNA-binding domain fused
to the Fluc C-terminus (Badran et al. 2011). To assess levels of
global DNA methylation, both fusion proteins are constructed using the DNA-binding
domain of a methyl-CpG-binding domain protein such as MBD2, which has a 100-fold
preference for methylated CpG sites over unmethylated CpG sites. The fusion proteins
are expressed in a cell-free expression system, then incubated with the target DNA to
allow DNA binding. If multiple methylated CpG sites exist in proximity, the
N-terminal and C-terminal portions of Fluc will interact (Figure 16.2). The level of
restored Fluc activity is measured using a firefly luciferase assay, such as the
Luciferase Assay System, and
luminescence levels are indicative of DNA methylation levels throughout the genome.
To measure site-specific DNA methylation levels, the N-terminus of Fluc is coupled to
the MBD DNA-binding domain, but the C-terminus is coupled to a sequence-specific
DNA-binding domain (Porter et al. 2008).
Epigenetic gene regulation also is controlled by changes in histones that make up the
nucleosome and histone modification. Canonical nucleosomes are octamers that consist of
H2A, H2B, H3 and H4 proteins. However, there are several histone variants that can vary
by a small number of amino acids or include large insertions (reviewed in Sarma and
Reinberg, 2005). Often these histone variants are found at specific locations within the
chromatin or are used to demarcate boundaries between heterochromatin and euchromatin
The majority of histone-mediated regulation stems from histone modification, most
often modification of the exposed amino termini of histones protruding from the
nucleosome core. The predominant histone modifications include acetylation, methylation,
phosphorylation, ubiquitination and sumoylation, with thousands of potential
combinations of modifications within a single nucleosome. Of these, histone acetylation
and methylation are the best understood, and some general trends have been observed.
Trimethylation of histone H3, specifically the lysine at position 4 (H3K4me3), is a mark
associated with transcriptionally active chromatin, whereas H3K27me3 leads to compact
chromatin, which represses gene expression. The term "histone code" is used to describe
how different combinations of histone modifications affect transcription levels.
Identification of proteins that read, write or erase these marks is critical to help
unravel the complexities of epigenetic regulation. Chromatin immunoprecipitation (ChIP)
is a powerful assay to identify proteins that bind to chromatin and map protein binding
throughout the genome using techniques such as microarray analysis or high-throughput
In ChIP analysis, protein:protein and protein:DNA complexes are crosslinked,
immunoprecipitated using an antibody against the protein of interest and purified. The
DNA sequence of interest then is amplified from the immunoprecipitated material using
PCR. Alternatively, the immunoprecipitated DNA can be sequenced (ChIP-seq) or analyzed
using microarrays (ChIP-chip) to identify target sequences.
One challenge of the traditional ChIP method is the availability of specific
antibodies that recognize crosslinked epitopes. To overcome the need for suitable
antibodies, Promega scientists developed the HaloCHIP™ System (Figure 16.3).
This system takes advantage of the HaloTag
which is a mutated hydrolase (Los et al. 2005; Los and Wood, 2007;
Los et al. 2008; Hartzell et al. 2009) that
catalyzes a covalent attachment to a variety of ligands, including a resin-based ligand
for immobilization. This tag can be fused to any protein; for ChIP, the DNA-binding
protein of interest is fused to the HaloTag
by cloning the protein-coding region into a HaloTag
vector. The recombinant construct is transfected into cells for stable or transient
expression, then cells are treated with formaldehyde to induce covalent protein:DNA and
protein:protein crosslinks, lysed and sonicated to shear the DNA into smaller fragments.
The crosslinked complexes are captured directly from the lysate through covalent binding
of the HaloTag
moiety to the HaloLink™
Resin. Covalent binding allows more extensive and stringent washing than is possible
with noncovalent interactions, resulting in reduced background and increased
signal-to-noise ratio. Subsequent heating of the purified complexes reverses the
crosslinks and releases captured DNA fragments, which can be purified and analyzed using
PCR, sequencing or microarray analysis. For more information, see the
HaloCHIP™ System Technical Manual
Acetylation of a lysine residue neutralizes a positive charge on a histone
protein, reducing the electrostatic interaction with negatively charged DNA. This
reduction in affinity leads to increased accessibility of the DNA to protein
complexes, which can lead to increased gene expression. In addition, lysine
acetylation can recruit nucleosome-remodeling complexes, such as Swi2/Snf2, via their
bromodomains to promote and maintain euchromatin structure (reviewed in Bernstein
et al. 2007). However, the factors controlling gene
expression are complex, and histone acetylation also can lead to reduced gene
expression through indirect mechanisms.
Lysine acetylation occurs on the N-terminal tails of core histones and is
controlled primarily by two enzyme families: histone acetyl transferases (HATs) and
histone deactylases (HDACs). HATs use acetyl CoA as a coenzyme to transfer an acetyl
group to the epsilon amino group of the lysine side chain. These enzymes are grouped
into three families: GNAT, p300/CBP and MYST. HDACs reverse histone acetylation and
promote gene silencing. HDACs are often components of large protein complexes and are
recruited to sites of DNA methylation by methyl DNA-binding proteins. HDACs fall into
four categories: Class I, which includes HDAC1, 2, 3 and 8; Class, II, which includes
HDAC4, 5, 6, 7, 9 and 10; Class III, which includes the NAD+-dependent sirtuins
(SIRTs); and Class IV, which includes HDAC11 (reviewed in Sun et
al. 2012).
Misregulation of HATs and HDACs often is associated with development and
progression of cancer and other diseases such as neurodegenerative disorders and
cardiovascular diseases, making these enzymes attractive therapeutic drug targets.
Many HDAC inhibitors promote cell cycle arrest at the G1/S phase, and studies have
shown that tumor cells generally are more sensitive to HDAC inhibitors than normal
cells (Johnstone, 2002). Also, HDAC inhibitors can restore the ability of animals to
recall memory that had been lost in Alzheimer's and Parkinson's disease models,
possibly by changing chromatin structure in neurons (Fischer et
al. 2007).
To facilitate screening of potential HDAC inhibitors, Promega offers the HDAC-Glo™ I/II Assays and Screening Systems and SIRT-Glo™ Assays and Screening Systems. The HDAC-Glo™
I/II and SIRT-Glo™ Assays are single-reagent-addition, homogeneous,
luminescent assays that measure relative activities of HDAC class I and II enzymes
and sirtuins, respectively. The HDAC-Glo™ I/II Assays use an acetylated,
live-cell-permeant, luminogenic peptide substrate that is deacetylated by HDAC
activities from cells, extracts or purified enzyme sources (Figure 16.4). The
SIRT-Glo™ Assay uses a similar substrate to detect SIRT activities from
purified enzyme sources (Figure 16.5). Deacetylation of the peptide substrate is
measured using a coupled enzymatic system in which a protease in the Developer
Reagent cleaves the deacetylated peptide from aminoluciferin, which is quantified in
a luciferase-based reaction. The HDAC-mediated luminescent signal is proportional to
enzyme activity and persistent, allowing batch processing of multiwell plates in
high-throughput screening.
Materials Required:
HDAC-Glo™ I/II Assay and Screening System Technical Manual
SIRT-Glo™ Assay and Screening System Technical Manual
Histone methylation occurs at lysine residues, which can be mono-, di- or
trimethylated, and arginine residues, which can be mono- or dimethylated. Histone
methylation is catalyzed by protein lysine methyltransferases (PKMTs) and protein
arginine methyltransferases (PRMTs) but can be reversed by protein demethylases. To
date, researchers have identified >30 demethylating enzymes, >50
protein lysine methyltransferases and >10 protein arginine methyltransferases,
suggesting that protein methylation is a dynamic and complex process (Janzen
et al. 2010). Histone methylation has different effects on
transcriptional activity, depending on the number of methyl groups and position of
the amino acid being modified. In general, the H3K9me1 mark is activating, whereas
H3K9me2 and H3K9me3 are repressive; H3K4me3 and H3K36me3 are associated with active
chromatin, whereas H3K9me3, H3K27me3, H3K36me2 and H4K20me1 often are found in
transcriptionally repressed heterochromatin.
The downstream effects of histone methylation are largely determined by proteins
that bind to the modified histones. For example, H3K9me3 acts as a binding site for
heterochromatin protein 1 (HP1), which then can recruit histone methyltransferases,
histone deacetylases and other proteins that affect chromatin structure. H3K4me3
recruits proteins that promote euchromatin, whereas H3K9me1, H3K9me2 and H3K27me3
interact with proteins that promote heterochromatin. Two such groups of proteins are
the polycomb group (PcG) proteins and their antagonists, the trithorax (trxG) group
proteins, which were first identified as regulators of hox gene
expression in Drosophila (Schwartz and Pirrotta, 2008). More
recent studies have shown that related proteins exist in mammals and plants. PcG
proteins repress transcription; trxG proteins activate transcription. Some PcG and
trxG proteins possess histone methyltransferase activity and can modify histones
directly, while others bind to and interpret histone modifications.
In embryonic stem (ES) cells, CpG islands that are regulated by PcG proteins often
are "bivalent" in that they retain the permissive H3K36me2-depleted and
H3K4me3-enriched environment but also exhibit H3K27me3. Genes with bivalent promoters
often are actively silenced in ES cells but lose the repressive H3K27me3 mark while
retaining the activating H3K4me3 mark later during differentiation.
Histones can be phosphorylated on serine, threonine and tyrosine residues. Many of
the serine and threonine phosphorylation events play a role in DNA repair or DNA
condensation, segregation and decondensation during mitosis, but some are involved in
epigenetic regulation of transcription, including H3T3ph, H3T6ph, H3T11ph, H2.AS1ph,
H3S10ph and H4S41ph (reviewed in Pérez-Cadahía et al. 2010).
H3S10ph is one of the best characterized of these histone modifications. In addition
to its DNA-restructuring responsibilities during mitosis, H3S10ph seems important for
chromatin decondensation associated with transcriptional activation of target genes.
H3S10ph recruits chromatin-modifying enzymes and chromatin-remodeling complexes and
prevents binding of HP1 to neighboring H3K9me3 marks at the onset of mitosis.
H3S10ph, as well as H3T3ph and H3T11ph, can block binding of DNMT3a to H3, reducing
methylation of nearby chromatin (Zhang et al. 2010).
Several kinases are involved in phosphorylation of H3S10, including IkB kinase
a (IKKa) (Yamamoto et al. 2003; Anest
et al. 2003), proviral integration site for Moloney murine
leukemia virus 1 (PIM1) (Zippo et al. 2007) and ribosomal S6
kinase 2 (RSK2) (Sassone-Corsi et al. 1999). Addition of the
H3S10ph mark to H3K9me3 is catalyzed by Aurora B kinase (Sabbattini et
al. 2007), which also modulates chromosome structure during mitosis and
mediates chromosome alignment and attachment to microtubules of the mitotic spindle.
Histones contain many highly conserved tyrosine residues, many of which can be
phosphorylated. Phosphorylation of H3Y99 is critical for polyubiquitination and
subsequent proteolysis of excess histones, which can increase a cell's sensitivity to
DNA-damaging agents, cause genomic instability and induce apoptosis. Another tyrosine
residue, H3Y41, is important in chromatin structure and oncogenesis. In human
hematopoietic cell lines, phosphorylation of H3Y41 by Janus kinase 2 (JAK2)
destabilizes binding of HP1a to histone H3 (Dawson et
al. 2009), leading to a more open chromatin structure around certain gene
promoters such as leukemia oncogene LMO2, which can trigger
oncogenesis in hematopoietic cells. Overexpression or aberrant activation of JAK2
activity leads to higher levels of H3Y41, loss of HP1a binding and higher
expression of LMO2.
Promega offers a number of kinase enzyme systems to monitor the activity or
identify inhibitors of different kinases involved in histone phosphorylation,
including IKKa, PIM1, RSK2 and several cyclin-dependent kinases (CDKs) such
as CDK1, CDK2 and CDK5. These luminescent assays convert ADP produced by these
kinases to ATP, which is then converted to light by Ultra-Glo™ Luciferase.
The resulting luminescent signal positively correlates with ADP amount and kinase
activity. An example protocol is provided below. For a list of available Kinase
Enzyme Systems, refer to the Human Kinome
Reagent Preparation
Kinase Assay and Detection Protocol
ADP-Glo™ Kinase Assay Technical Manual
Kinase Enzyme Systems
Conjugation of ubiquitin, a 76-amino acid protein, to lysine residues of histone
proteins can affect transcription activity as well as nucleosome stability and, as a
result, gene accessibility. The consequences of histone ubiquitination depend on the
histone substrate and degree of ubiquitination (reviewed by Weake and Workman, 2008).
Mono-ubiquitination of histone H2A (H2Aub1) is often considered a repressive mark,
while H2B mono-ubiquitination can play a role in both transcriptional activation and
silencing. In addition, there is evidence of cross-talk between histone
ubiquitination and other forms of histone modification. For example, ubiquitinated
H2B has been identified as a docking site for the COMPASS protein complex
(Chandrasekharan et al. 2010), which includes the histone
methyltransferase responsible for H3K4 methylation. Also, H2Aub, but not H2A,
specifically represses di- and trimethylation of H3K4, and ubiquitin-specific
protease 21 (USP21) relieves this repression (Nakagawa et al.
Ubiquitination of histones can be reversed by cleaving the peptide bond between
ubiquitin and the ubiquitinated protein. Several deubiquitinases (DUBs) have been
reported to deubiquitylate histones 2A, 2A.Z and 2B, including USP3, USP10, USP21,
USP22 and Bap1. Histone deubiquination has been associated with both transcription
activation (Nakagawa et al. 2008; Draker et
al. 2011; Gutiérrez et al. 2012) and repression (van
der Knaap et al. 2005; van der Knaap et al.
Another post-translational modification that plays an important role in epigenetic
regulation is sumoylation, the addition of the small ubiquitin-related modifier SUMO
(reviewed in Ouyang and Gill, 2009). This modification can stabilize proteins, alter
subcellular localization, affect enzyme activity and mediate interactions with other
proteins. Many transcription factors and cofactors can be sumoylated, which is
generally indicative of transcription repression. In Drosophila,
the sumoylated form of Sp3 recruits the polycomb protein Sfmbt (Steilow et
al. 2008a) and HP1a, ß and ? (Steilow
et al. 2008b; Seeler et al. 2001) to
repress transcription.
Many histone-modifying enzymes, nucleosome-remodeling complexes and their
associated enzyme cofactors contain one or more SUMO interaction motifs (SIMs). This
motif allows these proteins to interact with sumoylated transcription factors and
cofactors, which can direct these enzymes to specific promoters. Two such groups of
histone-modifying enzymes recruited by SUMO are histone deacetylases, which decrease
histone acetylation at the target promoter, and histone demethylases such as
lysine-specific demethylase 1, which catalyzes the removal of methyl groups from
Many important regulatory proteins contain domains that bind to modified residues,
including plant homeobox domain (PHD) fingers, bromodomains, chromatin organization
modifier (chromo) domains, WD40 repeat and tudor domains. Transcription-friendly H3K4me3
acts as a binding site for effector proteins that contain a PHD finger, such as
nucleosome remodeling factor (NURF) and the ING4-containing histone acetyltransferase
complex. The H3K36me2 modification, which interferes with transcription initiation, acts
as a binding site for the chromodomain of the RPD3S histone deacetylase complex.
Histone modifications also can act in cooperation. The specific combination of
histone modifications at a particular site often determines which protein complexes and
accessory proteins are recruited to activate or repress transcription directly, catalyze
additional histone modifications or recruit other histone-modifying proteins. This
cooperativity can be explained, at least in part, by the fact that these proteins can
contain one or more modified-histone-binding domains. For example, the TFIID protein
complex contains both a PHD finger and bromodomain and so binds more strongly to H3K4me3
marks near acetylated H3K9 and H3K14 residues.
The absence of one of these domains through gene mutation or rearrangement can cause
serious lapses in gene regulation and diseases such as cancer. Recently, researchers
characterized a translocation involving histone demethylase KDM5A that resulted in
fusion of the H3K4me3-binding PHD finger of KDM5A to the transcriptional activator
NUP98, a common leukemia translocation partner, in an acute myeloid leukemia patient
(Islam et al. 2011). Similarly, mixed lineage leukemia (MLL) family
members, which act as histone methyltransferases, are involved in translocations in
There is increasing evidence that expression of noncoding RNAs, such as microRNAs,
small RNAs and large RNAs, play a role in epigenetic gene regulation (reviewed in Costa,
2008; Chuang and Jones, 2007). Noncoding RNAs can direct both cytosine methylation and
histone modification to silence DNA repeats in the genome. For example, a class of
29-nucleotide RNAs that was first discovered through their interaction with the
spermatogenesis-specific PIWI protein (piwi-interacting RNAs; piRNAs) map to repetitive
DNA sequences and are important for silencing short interspersed elements (SINEs), long
interspersed elements (LINEs) and long terminal repeat (LTR) retrotransposons.
Several noncoding RNAs also are implicated in X chromosome inactivation. The 17kb
X-inactive specific transcript (XIST) RNA binds and coats the inactive X chromosome and
forms complexes that modify chromatin structure to suppress transcription. XIST levels
are controlled by another large noncoding RNA, TSIX, which is transcribed from the
strand opposite of XIST. Noncoding RNAs are involved in genomic imprinting via a similar
mechanism (reviewed by Koerner et al. 2009).
Maintenance and inheritance of epigenetic marks during cell division is critical to
maintain a committed cell lineage and cellular phenotype in progeny cells, and set a
memory of transcriptional status. The transmission of epigenetic information through
multiple cell divisions involves many of the mechanisms discussed in this chapter: DNA
methylation, histone modification, histone variants and expression of noncoding RNAs
(reviewed in Zaidi et al. 2011). These same mechanisms govern the
inheritance of epimutations, which can lead to changes in chromatin structure and
transcription levels of genes important to diseases such as cancer and imprinting
Aberrant regulation of epigenetic mechanisms can result in genomic imprinting
disorders, such as Angelman syndrome and Prader-Willi syndrome, and may contribute to
the heritability of many forms of cancer, asthma, Alzheimer's disease and autoimmune
diseases such as systemic lupus erythematosus, rheumatoid arthritis and multiple
sclerosis (reviewed in Hirst and Marra, 2009; Hewagama and Richardson, 2009; Handel
et al. 2010). Epimutations can interfere with epigenetic
regulation at many levels, including DNA methylation, histone modification and noncoding
RNAs. Some epimutations are inherited, but many accumulate due to environmental factors
or with age. For example, even though monozygotic twins are epigenetically
indistinguishable at birth, their patterns of DNA methylation and histone acetylation
can differ dramatically as they age (Fraga et al. 2005).
Dual-Glo, Flexi, HaloTag, Nano-Glo, Steady-Glo and Wizard are registered
trademarks of Promega Corporation. ADP-Glo, DUB-Glo, Find My Gene, HaloCHIP,
HaloLink, HDAC-Glo, MethylEdge, SIRT-Glo and Ultra-Glo are trademarks of Promega
Products may be covered by pending or issued patents or may have certain limitations. Please visit our web
site for more information.
All prices and specifications are subject to change without
prior notice.
Product claims are subject to change. Please contact
Promega Technical Services or access the Promega online catalog for the
most up-to-date information on Promega products.


text:
Portland, OR –- The Office of University Communications will proudly host its first summer Portland State Tweetup at the University Pointe Leasing Office, 1955 SW Fifth Avenue, from 4:30 p.m. until 6:30 p.m. on July 19. Sponsored by University Pointe, Google, TwinStar Credit Union, Unitus Community Credit Union, and 7-Eleven, the Portland State Tweetup is a social media event for PSU social media fans as well as students, staff, faculty, alumni, and friends to meet each other in real life and connect. Like previous Tweetups, the event hopes to build campus community, pride, tradition and school spirit.
The Portland State Tweetup will feature free admission, BBQ and refreshments, and a photobooth. KPSU, PSU's college radio, will provide entertainment during the event. Attendees will receive passports and raffle tickets for the chance to win grand prizes including an Apple iPad, gift certificates, and gift baskets. The Ben & Jerry's national touring truck will also be at the event to sample out their new Greek Frozen Yogurt.
PSU's official social media presence can be found on Facebook, Twitter, YouTube, Flickr, Foursquare and Google+. Through social networking sites, PSU hopes to build an online community with the PSU campus community.
About Portland State University:
Located in Portland, Oregon, PSU has about 30,000 undergraduate and graduate students. PSU's motto is "Let Knowledge Serve the City," and we provide every student with opportunities to work with businesses, schools and organizations on real-world projects. Our downtown campus exhibits PSU's commitment to sustainability with green buildings, while sustainability is incorporated into much of the curriculum.
About University Pointe:
University Pointe at College Station is an American Campus community. American Campus Communities is the nation's premier owner and manager of luxurious, academically-oriented student housing.
Take advantage of our convenient location and unmatched amenities—everything you want and need in one place makes University Pointe the best in downtown apartments! University Pointe is located across the street from Portland State University in the heart of downtown Portland, OR, and has an on-site TriMet MAX light rail stop. You will also enjoy our amenities including a fitness center, computer center, multiple study lounges, social lounge with X-Box and PS3, wi-fi hot spots and much more
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subset: commoncrawl


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Helping police officers struck by the Joplin tornado
The cop struck by lightning in Joplin remains in critical condition — meanwhile, an 'Adopt-A-Cop Program' has been set up to assist local officers from the affected area who lost their homes and belongings
Two police officers were struck by lightning during the search for survivors which followed the massive tornado that killed scores in Joplin, Missouri. A local TV station indicated that "one of the officers was taken to the hospital, the other went back to work." The Wichita Eagle reported that the hospitalized officer had suffered "significant injuries" but that report offered no additional details.
Today I spoke with Captain Chris Skinrood of the Riverside (Mo.) Police Department, whose officer was the one critically injured in the lightning strike. Skinrood told me that the injured officer — who was identified as Officer Jeff Taylor in a press release issued this afternoon by the Riverside Police Department — remains listed in critical condition.
"He's doing better than what he was," Skinrood told me. "He has made some very, very slight improvements, but none the less they're improvements. He still has an extremely long road ahead of him..."
According to the abovementioned department press announcement, Officer Taylor was standing next to an all-terrain vehicle after returning to a command post when lightning struck the ground beside him. He was transported immediately to an area hospital and was reported to be in critical condition. Taylor was part of Riverside's 12-member contingent of police officers, firefighters and public works staff members who responded to the Joplin tornado disaster.
Family members, who have asked for privacy at this time, are accompanied by Riverside officers at Taylor's side. PoliceOne members looking for regularly-updated information about Officer Taylor, a website hosted by CaringBridge.org has been established here. There has also been an account set up — the Taylor Benefit Account — to accept donations to assist Jeff and his family with expenses.
Donations can be sent to:
K.C. Police Credit Union
2800 East 14th Street
Kansas City, MO 64127
Main Phone: 816-504-2800
Toll free: 800-338-8567
Fax: 816-504-2888
Internet: www.kcpcu.org
The thoughts and prayers of everyone here at PoliceOne go out to the family, friends, and fellow-LEOs of this injured hero.
Speaking of Thoughts and Prayers...
The tornado — now classified as an EF-5, the highest possible rating on the Enhanced Fujita scale of tornado power and intensity — that leveled a six-mile swath of Joplin had also struck police officers in other ways. Today I received an email from Scott Barthelmass, founder of the Missouri Law Enforcement Funeral Assistance Team, and a Corporal with the Overland (Mo.) Police Department.
Barthelmass told me about a joint effort between Ten Four Ministries, an organization based in Tulsa (Okla.) and whose Director is my good friend and PoliceOne Colleague, Travis Yates, and the Missouri Law Enforcement Funeral Assistance Team to create an 'Adopt-A-Cop Program' for the officers of Joplin Police Department and the Jasper County Sheriff's Department.
Barthelmass said that they've received a "list of seven officers who lost everything" in the disaster.
"We are in the process of contacting these officers so that we can meet their individual needs. If we are notified of more officers, we will most definitely add them to our list. We will also soon begin working on a list of officers whose homes suffered significant damage ... with the cooperation of many local, state, and national law enforcement organizations, we will spread the needs of our Adopt-A-Cop Program," he said.
The two lead partners in the effort are working with officers from the FOP Lodge (Lodge 27) to identify the needs of law enforcement officers there. Barthelmass said that he'd spoken with Lodge 27's William Davis, who said that the affected officers are "going through what is left of their homes."
Davis continued, "Financial support is primarily what they are needing at this time as they all looking for new places to live and places to store their belongings."
Donations can be sent to:
10-33 Fund
Adopt A Cop Program
Joplin, Missouri Area Tornado Victims
Ten-Four Ministries
PO Box 52005
Tulsa, OK 74152
Donations can also be made via the Internet. Because Ten Four Ministries and the Missouri Law Enforcement Funeral Assistance Teams are both 501c3 organizations, all donations at tax deductible.
According to Barthelmass, the specific needs of the officers and their families will be posted on the Missouri Law Enforcement Funeral Assistance Team and Ten Four Ministries websites, and I will update our coverage on PoliceOne as additional information becomes available. To wit, Lodge 27 late this afternoon sent an update to PoliceOne via our friends at Missouri Law Enforcement Funeral Assistance Team. The specific needs have now been identified for one officer, who has a two-year-old daughter who desperately needs clothes (Size 2T). She also lost all her toys.
I, for one, will tonight be gathering up as many toys as possible from my own son's playthings he's outgrown, and mailing them tomorrow in a box to the below address.
Joplin Police Dept.
Attn: William Davis
303 E 3 rd St.
Joplin, MO 64801
The officer himself, according to the latest update from Barthelmass, has just one pair of clothes. He wears a size 38x32 pant and XL shirts and size-14 boot. Lodge leadership reports, "If we could get him some BDUs and shirts and a pair of boots that would be great."
Death Toll Mounts, Search for Survivors Continues
The death toll of the mammoth tornado was recently estimated to be 125 souls, with some 1,500 people reported missing and more than 750 injured. Police officers from around the country continue to assist in the search for survivors, with 32 Kansas City Missouri Police Department employees reportedly still in the tornado-stricken region, and cops from other agencies are reportedly still arriving to assist.
The Associated Press reported this morning that Missouri Gov. Jay Nixon hopes more survivors will be found. "I still believe that because of the size of the debris and the number of people involved that there are lives to be saved," Nixon said.
In fact, according to several reports, search and rescue teams found two more survivors just this morning. The search, of course, will continue in earnest for days. Meanwhile, Nixon also conceded that he fears the death toll will rise. "I don't think we're done counting," he reportedly said.
The twister that struck Joplin is thought to be the deadliest single tornado to hit the United States since 1950. Barthelmass told me, "For those who have not seen the images of the destruction, it is truly almost unreal."
One civilian volunteer rescue worker — who has also worked at Ground Zero in New York following the 9/11 terrorist attacks and in New Orleans following Hurricane Katrina — described the destruction as "Katrina without the water."
As I mentioned above, the Joplin tornado has been classified as an EF-5 (upgraded from its original classification of EF-4). Such a storm has winds greater than 200 miles per hour, which compares to a Category Three hurricane. Katrina made its second landfall as a Category Three storm on the morning of Monday, August 29, 2005 in southeast Louisiana. Katrina killed more than 1,800 people and caused more than $81 billion in property damage.


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Emelisse Hotel
The Emelisse Hotel is situated along the rugged shoreline of the emerald Ionion Sea. It sits in a spot of supreme serenity amongst acres of native cypress and cedar trees on the beautiful island of Kefalonia. The pools and outdoor restaurants and bars provide front row seating for an unparalleled display of nature's splendour.
The sunset is mesmerising and has the power to bring you back to an inner stillness that is so rare in modern life. At night the hotel is lit by soft lamps and candlelight, transforming it into a romantic, whimsical wonderland. Snuggle up on a plush sofa on one of the numerous expansive terraces from which you can gaze up endlessly at a stellar canopy of velvety, indigo sky. The hotel provides a wide choice of accommodation ranging from double rooms to maisonettes and executive suites.


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Nutrition Facts
Ingredients and Preparation
- Remove mushroom stems (if present) and scrape out the brown gills from the undersides of the caps using a spoon.
- Mix together 1 teaspoon oil and lemon juice, and then brush all over the mushroom caps.
- Heat 2 teaspoons oil in a medium non-stick skillet over medium heat. Add garlic, shallot, bell pepper, and zucchini, and sauté until tender, about 3-4 minutes. Stir in tomatoes, 1 tablespoon basil, and marjoram, and cook for another 1-2 minutes. Remove from heat and season with salt and pepper.
- Spray grill rack or indoor grill with non-stick cooking spray and then place the mushroom caps stem-side down on the grill. Grill mushrooms for about 5 minutes on each side, until soft.
- Remove mushrooms from grill, fill with the vegetable mixture, and top with mozzarella cheese. Return mushrooms, filled-side up, to the grill. Cover and cook for about 3 minutes, until cheese is melted. Garnish with the remaining 2 tablespoons of chopped basil.


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Watch: Peyton, Eli Manning star in DirecTV Fantasy Football video
For the second consecutive year, DirecTV induced Peyton Manning and his younger brother Eli Manning into performing a promotional music video. This one is for the DirecTV NFL Sunday Ticket's new Fantasy Zone Channel.
Mobile Users: Click here to watch the video.
My only comment for now is this one is strange.
There are multiple cameos by past and current NFL players including members of the Broncos offensive line, Archie Manning, Chris Johnson and Joe Namath.
The brothers appeared in "Football on your Phone" last year.


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In a heated second straight battle against FC Dallas, Houston pulled out a 1-1 road draw despite missing Brian Ching to the World Cup and Dwayne De Rosario to a first-half injury. Carlos Ruiz scored in the first half to give Dallas the lead, but a late own goal from Clarence Goodson, under pressure from Eddie Robinson, gave Houston the tie.
Houston Dynamo (4-2-1) vs. FC Dallas (4-1-3)
May 13, 2006 -- Pizza Hut Park, Frisco TX
Scoring Summary:
FCD -- Carlos Ruiz 3 (Ramon Nunez 1, Drew Moore 1) 19
HOU -- Own goal (Clarence Goodson) 81
Houston Dynamo -- Pat Onstad, Wade Barrett, Eddie Robinson, Adrian Serioux (Julian Nash 70), Craig Waibel, Ricardo Clark, Brad Davis (Kevin Goldthwaite 62), Dwayne De Rosario (Ryan Cochrane 39), Brian Mullan, Ronald Cerritos, Alejandro Moreno.
Substitutes Not Used: Zach Wells, Kelly Gray, Stuart Holden, Chris Wondolowski.
FC Dallas -- Dario Sala, Clarence Goodson (David Wagenfuhr 82), Drew Moore, Greg Vanney, Alex Yi, Richard Mulrooney, Ronnie O'Brien, Simo Valakari, Kenny Cooper, Ramon Nunez (Aaaron Pitchkolan 78), Carlos Ruiz (Roberto Mina 72).
Substitutes Not Used: Ray Burse, Arturo Alvarez, Justin Moore, Mark Wilson.
Misconduct Summary:
FCD -- Carlos Ruiz (caution; Reckless Foul) 17
HOU -- Eddie Robinson (caution; Reckless Foul) 48+
Referee: Brian Hall
Referee's Assistants: Steven Davidson; Michael Salyers
4th official: Richard Heron
Attendance: 12,756
Weather: Sunny-and-84 degrees
All statistics contained in this boxscore are unofficial


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Promising results from Alzheimer's and BCI research raise hope of their transition into mainstream clinical foray, boosting prospects for growth, says RNCOS.
Noida, UP -- (SBWIRE) -- 02/03/2014 -- As per the new research report by RNCOS, "Brain Signal Analysis Software Market Outlook to 2017", brain signal analysis software represents a niche product category that caters to a very specific segment of the global medical device industry. Our study reveals that the market for brain signal analysis software stood at around US$ 30 Million in 2012 and is expected to grow at a double-digit growth rate; higher than the traditional medical device market.
Our research reveals that players in the market are mostly small with limited geographic presence. This presents an opportunity for new market entrants who can identify the regions with high market potential and low competition to launch their product. Besides, our study indicates that the market is not price sensitive and it is the range and flexibility of available analyses along with the speed of data processing that a software user looks at before purchasing the software. It is a market which focuses more on product quality rather than the price. Such a market is always open for new market entrants that can offer innovative products with unique differentiating product features.
The main factor propelling the growth of the market will be increase in its demand for epilepsy diagnosis and monitoring. Besides, rapidly expanding sleep labs in the world will also drive this growth. Also, this growth can really offshoot beyond expected limits if researchers successfully recognize EEG based biomarkers for early detection of Alzheimer's disease. This will bring EEG into the mainstay of clinical diagnosis of Alzheimer's disease, a field which has immense potential. Moreover, researchers are also reporting promising results from BCI research. If BCI picks up, as is expected from it, growth is expected to shoot up with it.
The research contains exhaustive information regarding the market dynamics, its current status and future outlook, opportunity assessment by application and analysis of key market players that will help clients in formulating apt market strategies and assess opportunity areas in the global brain signal analysis software market.
For FREE SAMPLE of this report visit: http://www.rncos.com/Report/IM647.htm
Check Related REPORTS on: http://www.rncos.com/Healthcare_Industry.htm
RNCOS is a leading industry research and consultancy firm incorporated in 2002. As a pioneer in syndicate market research, our vision is to be a global leader in the industry research space by providing research reports and actionable insights to companies across a range of industries such as Healthcare, IT and Telecom and Retail etc. We offer comprehensive industry research studies, bespoke research and consultancy services to Fortune 1000, Trade associations, and Government agencies worldwide.


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NTS and Other Entry Tests Number Series Quizzes
This quiz comprises of Number series which are an essential part of NTS tests, CSS/FPSC and other entry test exams. Please select the quiz number from below and start preparation. Please note that you can end the quiz any time by clicking the "submit all" button. This will end the quiz and you will be able to review the quiz with answers and explanations. Mobile users can end the quiz anytime by selecting the flag in the top right corner and then selecting and answering the last question. More Quizzes are in pipeline.


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TLC filed a lawsuit in Maryland court against Jon Gosselin for breaching his contract.
The cable network alleges the ex-reality TV dad hasn't stuck to the deal he struck when he signed on to "Jon & Kate Plus 8" by getting paid for appearances on other shows and making unauthorized disclosures about the show, according to court documents.
The news comes as Gosselin and estranged wife Kate have found themselves in a kind of nanny state.
The reality TV parents are said to be fuming after ex-nanny Stephanie Santorotold RadarOnline.com that Jon once bragged to her about hacking into estranged wife Kate's emails, phone records and bank accounts, according to reports.
Santoro previously said she had a fling with Gosselin while employed to look after his brood of eight, a claim Jon denies.
Kate Gosselin's law firm said Thursday that the octomom was "profoundly disturbed" by Santoro's latest allegations.
"Under the circumstances, Ms. Gosselin is carefully considering all of her legal options regarding this matter, and she will pursue them if and when the time is right," the firm, Schnader Harrison Segal & Lewis said in a statement.
Jon's lawyer also fired back, telling People that Gosselin "unequivocally told me that he's never illegally invaded Kate's electronic privacy in any way."
"He also finds it a little disappointing that Kate would give credence to an uncorroborated statement made by an individual who clearly has a motivation to tell stories about Jon that might result in financial (compensation)," lawyer Mark Heller said.
The nanny diaries are the latest twist in the celebrity couple's very public fall from grace since announcing in June they would divorce and put "Jon and Kate, Plus 8" on a brief hiatus.
TLC has reportedly decided to end filming new episodes of the reality show next month amid declining ratings. The network had earlier announced it would rename the show "Kate Plus 8" to reflect a reduced role for Jon, who then scuttled the arrangement by banning TLC from filming his children.
After trading repeated jabs through the media, Jon and Kate called a temporary truce earlier this week.
On Tuesday, a judge ordered Jon Gosselin to return $155,000 to a joint account he shares with Kate. He said he now plans to do so.
The couple's next court date is on Oct. 26.


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People either love Williamsburg or they hate it. In this rapidly developing artistic and eclectic neighborhood in the borough of Brooklyn where real estate continues to skyrocket as luxury lofts pop up, hipsters abound, and the original ethnic enclaves dwindle, it's easy to see why there are such contrasting viewpoints on the neighborhood that was once mostly inhabited by immigrants and Hasidic Jews. Now, having a glass-half-full outlook, with gentrification comes new sub-cultures and establishments that bring people (even Manhattanites) to an area rich in history, art, music and culture. And that, my friends, it turns out, is a pretty nice formula for a date! Take Wythe Avenue, for instance, it's got it all.
So, gentleman, hipsters or not, while Manhattan has a lot to offer (it can almost be overwhelming), I'm giving you this fail-proof guide to the perfect date… in a one-block radius.
Sunset Drinks on the Sixth floor of Wythe Hotel
The Wythe Hotel, located on 80 Wythe Avenue, is a cozy spot to grab pre-dinner drinks as the afternoon sun sets on a breathtaking view of Manhattan (The cliched description used is pretty accurate--here's proof). The new hotel opened this summer in a historic industrial building by the water in this non-hotel-scene 'hood. The view itself and some people watching will at least give you something to talk about at first--why not let Manhattan be your wingman.
Dinner Downstairs at Reynards or Down the Street at MyMoon
When it's time for dinner, saunter downstairs to Reynards, the restaurant in the lobby of the Wythe Hotel. Reynards is a the newest addition to Andrew Tarlow's culinary resume, as the owner of Williamsburg favorites Diner and Marlow & Sons. A slight diversion from the typical Williamsburg food scene, this brasserie's American Noveau menu lists dishes made of ingredients handpicked by the chef himself, fishes brought in by their fishmonger, meats cut by their in-house butcher, and desserts made with chocolates from the Mast Brothers, a couple well-known locals, which makes it very Williamsburg-y.
What the hell is "American Nouveau"? If you're thinking that, maybe MyMoon (184 N. 10th St.), a Mediterranean tapas spot around the corner, is more your speed with basic fare that you can share. The spacious dimly lit restaurant has a huge outdoor deck with both cabana-style booth seating and regular garden seating. Flowers in mason jars hanging from the garden lights set the mood. However, I must admit, the food alone should not be the reason to go--the romantic ambience makes up for it.
Post-Dinner Cocktails at Kinfolk Studios
If you've gotten this far, the night is going well, buddy! High five to you and treat yourself, and your date, to a cocktail at Kinfolk Studios (90 Wythe Ave.), a bar and restaurant, art and music venue, creative space, all in one. This hybrid destination is a cool hangout spot with extraordinary cocktails made by (quite good looking) mixology bartender types who are both attentive and friendly. Try the cucumber mint margarita.
Late-Night Music at Brooklyn Bowl
Still want to continue the night? Pop in (or uh, stumble over) to Brooklyn Bowl for some late-night music and/or bowling. Brooklyn Bowl posts their live music schedule weeks in advance, so if you're feeling pretty confident that the date will get to this point, buy your tickets in advance and plan your outing on the same night a rocking band is playing. Feeling peckish? Blue Ribbon bar snacks and Brooklyn Brewery brews are your options here. You really can't go wrong with that.
Finally, if you really don't want to leave that one amazing block, and you're ready for another date, but your wallet isn't... try a picnic at the East River State Park. Wine and cheese with Manhattan as your backdrop--she will love it. She won't even notice if you're broke. Win!


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Camp near Lewisburg Tennessee
Sunday morning May 3rd, 1863
Dearest Sallie, You will see from the heading of this letter that we have moved our camp, we moved out from Shelbyville last Monday some 4 or 5 miles and hadn't been there but a few days until we reed' orders to come here. We started Friday night about 9 or 10 o'clock and marched till about 2 and then slept till daylight and came on here yesterday about 10 or 11 o'clock We have stopped about 3 miles this side of Lewisburg - turned off the road to the right about half a mile and camped on the bank of a creek. We have a very pretty camp ground, but there is no telling how long we will stay here. They say we are sent here to support Van Dorn and I expect he will have us running all over this country in a few days. He is over at Columbia about 20 miles from here and whenever the Yanks get too hard for him, we will have to go to his assistance. Our brigade is all that came yesterday, I heard that 3 brigades were coming, but haven't seen any but ours yet. We may have some fighting to do pretty soon, I can't tell. They say the Feds have been


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Wavelet Design & Analysis App
The Wavelet Design & Analysis app is included in Wavelet Toolbox. You can find it, along with the rest of the apps for your installed products, by clicking the Apps tab in the MATLAB Toolstrip.
The Wavelet Design and Analysis app accesses the various graphical tools available in the Wavelet Toolbox.
Apps are interactive MATLAB programs you can use without writing any code. Most toolboxes come with apps, and you can download more for free from the MATLAB File Exchange. You can also package and share your own apps.


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By age 50, 5 out of 10 men have an enlarged prostate (BPH). By age 80, up to 9 out of 10 men have an
enlarged prostate. Having an enlarged prostate does
not always cause symptoms. But BPH symptoms also are age-related. By age 75, 5
out of 10 men complain of a decrease in the force of their urine
Lower urinary tract problems in
men under age 40 are most likely caused by other conditions, such as prostate
infection or failure of the smooth bladder muscles to relax during urination
(voiding dysfunction).
CitationsPresti JC, et al. (2008). Benign prostatic hyperplasia
(BPH) section of Neoplasms of the prostate gland. In EA Tanagho, JW McAninch,
eds., Smith's General Urology, 17th ed., pp. 348–370.
New York: McGraw-Hill.
Last Revised:
March 5, 2012
Healthwise Staff
Medical Review:
E. Gregory Thompson, MD - Internal Medicine & J. Curtis Nickel, MD, FRCSC - Urology
This information does not replace the advice of a doctor. Healthwise, Incorporated disclaims any warranty or liability for your use of this information. Your use of this information means that you agree to the Terms of Use. How this information was developed to help you make better health decisions.
To learn more visit Healthwise.org
© 1995-2012 Healthwise, Incorporated. Healthwise, Healthwise for every health decision, and the Healthwise logo are trademarks of Healthwise, Incorporated.


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When Congress returns from recess, House Republicans will begin a continuous assault on a series of health, environmental and labor regulations, which they say are hampering job creation. And they'll twin it with two tax cuts for both large and small businesses. One of those cuts will actually be aimed at preventing a scheduled tax increase — but it's not the payroll tax cut President Obama has asked Congress to extend.
In a memo to members, House Majority Leader Eric Cantor (R-VA) laid out a list of 10 rules, most of which have yet to be implemented, which they'll seek to prevent week by week. These include regulations that would limit the amount of mercury and other toxins boiler and incinerator operators can burn into the atmosphere; that could make it easier for workers to unionize; and that assure that employer insurance policies exempted from new health care law — so-called "grandfathered" plans — meet the law's basic requirements and aren't gamed by employers to reduce workers' existing benefits.The plan would also make two modest changes to the tax code. One, would be to eliminate a coming "three percent withholding rule." This policy, which is set to take effect in 2013, would require government at all levels to withhold three percent of their payments to large contractors until those contractors pay their taxes. The other would allow businesses with fewer than 500 employees to take a tax deduction equal to 20 percent of their income.
While obviously friendly to conservative interests, the plan broadly speaking views the slumping economy as a consequence of supply-side barriers. It aims money and laxity at employers, who, most economists agree, are sitting on trillions of dollars right now not because they may have to comply with certain new rules and laws but because they don't have customers.
Read the entire memo below.
TO: House Republicans
FR: Eric Cantor
DT: Monday, August 29, 2011
RE: Upcoming Jobs Agenda
As you know, we released The House Republican Plan for America's Job Creators earlier this year. While the debt crisis has demanded much of our attention, our new majority has passed over a dozen pro-growth measures to address the equally troubling jobs crisis, such as the Energy Tax Prevention Act and the Putting the Gulf of Mexico Back to Work Act. Aside from repeal of the 1099 reporting requirement in ObamaCare, however, each House Republican jobs bill now sits dormant in the Democrat-controlled Senate. You can view the progress of our jobs bills at MajorityLeader.gov/JobsTracker.
When we return next week, the Joint Select Committee on Deficit Reduction will begin meeting to take an additional incremental step towards addressing our debt crisis. During this time, it is essential that the House continue our focus on the jobs crisis. Below are two areas of our jobs agenda that I want to bring to your attention for our upcoming fall and winter legislative schedule.
Since passage of H.Res. 72 on February 11, our committee chairmen have been investigating and inventorying regulatory burdens to job creators. They've found many that have tied the hands of small business people and prevented job growth. By pursuing a steady repeal of job-destroying regulations, we can help lift the cloud of uncertainty hanging over small and large employers alike, empowering them to hire more workers.
Our regulatory relief agenda will include repeal of specific regulations, as well as fundamental and structural reform of the rule-making system through legislation like the REINS Act, the Regulatory Flexibility Improvements Act, and reform of the Administrative Procedures Act (all three bills are expected on the floor in late November and early December).
The following is a list of the 10 most harmful job-destroying regulations that our committee chairmen have identified, as well as a selective calendar for their repeal. These regulations are reflective of the types of costly bureaucratic handcuffs that Washington has imposed upon business people who want to create jobs.
Top 10 Job-Destroying Regulations:
· NLRB's Boeing Ruling (Week of September 12): On April 20, the National Labor Relations Board (NLRB) issued a complaint against The Boeing Company for the alleged transfer of an assembly line from Washington to South Carolina. Yet, not one union employee at Boeing's Puget Sound facility has lost his or her job as a result of the proposed South Carolina plant. Still, the NLRB is pursuing a "restoration order" against Boeing that would cost South Carolina thousands of jobs and deter future investment in the United States. H.R. 2587, the Protecting Jobs From Government Interference Act, sponsored by Rep. Tim Scott (SC), would take the common sense step of preventing the NLRB from restricting where an employer can create jobs in the United States.
· Utility MACT and CSAPR (Week of September 19): The Administration's new maximum achievable control technology (MACT) standards and cross-state air pollution rule (CSAPR) for utility plants will affect electricity prices for nearly all American consumers. In total, 1,000 power plants are expected to be affected. The result for middle class Americans? Annual electricity bill increases in many parts of the country of anywhere from 12 to 24 percent. H.R. 2401, the Transparency in Regulatory Analysis of Impacts on the Nation (TRAIN) Act, sponsored by Rep. John Sullivan (OK), would require a cumulative economic analysis for specific EPA rules, and specifically delay the final date for both the utility MACT and CSAPR rules until the full impact of the Obama Administration's regulatory agenda has been studied.
· Boiler MACT (Week of October 3): From hospitals to factories to colleges, thousands of major American employers use boilers that will be impacted by the EPA's new "boiler MACT" rules. These new stringent rules will impose billions of dollars in capital and compliance costs, increase the cost of many goods and services, and put over 200,000 jobs at risk. The American forest and paper industry, for example, will see an additional burden of at least $5-7 billion. H.R. 2250, the EPA Regulatory Relief Act, sponsored by Rep. Morgan Griffith (VA), would provide a legislative stay of four interrelated rules issued by the EPA in March of this year. The legislation would also provide the EPA with at least 15 months to re-propose and finalize new, achievable rules that do not destroy jobs, and provide employers with an extended compliance period.
· Cement MACT (Week of October 3): The "cement MACT" and two related rules are expected to affect approximately 100 cement plants in America, setting exceedingly stringent requirements that will be cost-prohibitive or technically infeasible to achieve. Increased costs and regulatory uncertainty for the American cement industry–the foundation of nearly all infrastructure projects–are likely to offshore thousands of American jobs. Ragland, Alabama, for example, recently saw the suspension of a $350 million cement production facility, putting 1,500 construction jobs on hold and additional permanent and high-paying plant operation jobs in limbo. H.R. 2681, the Cement Sector Regulatory Relief Act, sponsored by Rep. John Sullivan (OK), would provide a legislative stay of these three rules and provide EPA with at least 15 months to re-propose and finalize new, achievable rules that do not destroy jobs, and provide employers with an extended compliance period.
· Coal Ash (October/November): These anti-infrastructure regulations, commonly referred to as the "coal ash" rules, will cost hundreds of billions of dollars, affecting everything from concrete production to building products like wall board. The result is an estimated loss of well over 100,000 jobs. H.R. 2273, the Coals Residuals Reuse and Management Act, sponsored by Rep. David McKinley (WV), would create an enforceable minimum standard for the regulation of coal ash by the states, allowing their use in a safe manner that protects jobs.
· Grandfathered Health Plans (November/December): We all remember when President Obama promised Americans that if they liked their health care plan they could keep it. Now, the Obama Administration has been issuing further restrictions against those previously protected plans. The result, by the Administration's own estimates, will be a loss of 49 to 80 percent of small employer plans, 34 to 64 percent of large employer plans, and 40 to 67 percent of individual insurance plans. Meanwhile, employers losing their grandfathered status will face steep penalties, increasing their costs and negatively affecting wages and job growth. The Energy and Commerce, Ways and Means, and Education and Workforce committees will soon be working on legislation to repeal these ObamaCare restrictions.
· Ozone Rule (Winter): This effective ban or restriction on construction and industrial growth for much of America is possibly the most harmful of all the currently anticipated Obama Administration regulations. Consequences would reach far across the U.S. economy, resulting in an estimated cost of $1 trillion or more over a decade and millions of jobs. Unlike her predecessors, EPA Administrator Lisa Jackson is pushing for a premature readjustment of the current ozone standards, dramatically increasing the number of "nonattainment" areas. The new readjustment rule is expected early this fall and I expect the Energy and Commerce Committee to act swiftly to prevent its implementation, in order to protect American jobs.
· Farm Dust (Winter): The EPA is expected to issue revised standards for particulate matter (PM) in the near future. Any downward revision to PM standards will significantly impact economic growth and jobs for businesses and people throughout rural America that create dust, like the farmer in Atkinson, Illinois, who raised his concerns with the President at a town hall earlier this month. While the President may have sent him on a bureaucratic wild goose chase, the House will act promptly on H.R. 1633, the Farm Dust Regulation Prevention Act, sponsored by Rep. Kristi Noem (SD). H.R. 1633 would protect American farmers and jobs by establishing a one year prohibition against revising any national ambient air quality standard applicable to coarse PM and limiting federal regulation of dust where it is already regulated under state and local laws.
· Greenhouse Gas (Winter): The EPA's upcoming greenhouse gas new source performance standards (NSPS) will affect new and existing oil, natural gas, and coal-fired power plants, as well as oil refineries, nationwide. While the impact on the economy and jobs are likely to be severe, the rules are quickly moving forward, once again revealing the Administration's disregard for the consequences of their policies on our jobs crisis. Again, I expect Chairman Upton and the Energy and Commerce Committee to move swiftly in the coming months to protect American jobs and consumers.
· NLRB's Ambush Elections (Winter): This summer, the NLRB issued a notice of proposed rulemaking that could significantly alter current union representation election procedures, giving both employers and employees little time to react to union formations in the future. The result will increase labor costs and uncertainty for nearly all private employers in the U.S. The House will soon consider legislation that will bring common sense to union organizing procedures to protect the interests of both employers and their workers.
The second focus of our jobs agenda this fall and winter will pursue tax relief designed to help American employers create middle class jobs. Small businesses, in particular, are overburdened with taxes that destroy jobs and in a fragile economy, the worst thing government can do is impose tax increases that prevent small businesses from hiring. Instead, government should be focused on ways to enable businesses small and large to unlock more capital and create new jobs.
Therefore, under Chairman Dave Camp's leadership, the Ways and Means Committee will be exploring a number of ways to remove tax burdens for job creators and incentivize small businesses. These targeted efforts will not interfere with House Republicans' continued pursuit of fundamental tax reform, just as we proposed in our budget resolution this spring. Rather, the following proposals represent bipartisan and pro-growth steps that can be taken immediately to help businesses hire those struggling to find a job today.
3% Withholding Rule Repeal:
Beginning in 2013, federal, state, and local governments will be required to withhold three percent of all government payments made to contractors in excess of $100 million. While the law has been delayed multiple times, its effect once implemented will be massive–causing accounting burdens on governments and potentially harmful cash flow disruptions for contractors and subcontractors across all sectors. Therefore, we will move quickly this fall to repeal this burdensome requirement and relieve construction contractors, medical providers, manufacturers, farmers, and many others providing goods and services under government contracts of the uncertainty the impending law is creating.
20% Small Business Tax Deduction:
Small businesses employ about half of all Americans, yet President Obama is proposing federal tax rates that could take away more than 40 percent of their income. While the previous Democrat majority was pushing through their nearly $1 trillion stimulus, we began work on a proposal to allow small business people to take a tax deduction equal to 20% of their income. The goal was simple–immediately free up funds for small business people to retain and hire new employees, and reinvest in and grow their businesses. In light of the stimulus's failure, and our current position in the majority, I expect the House to move quickly in the coming months on this common sense and pro-growth small business proposal to create middle-class jobs.
In the weeks and months ahead, we will move aggressively on the items outlined above. But, our increased focus on the repeal of job-destroying regulations and pursuit of pro-growth tax relief will not distract from other equally important areas of our jobs agenda.
As he has promised in the past, we expect the President to transmit three vitally important trade agreements this fall. When he finally sends them our way, I will not hesitate to schedule them. In addition, the Senate has set up a vote on the House-passed patent reform bill next week. Increased movement in both of these areas on the part of the President and the Senate is welcomed. I hope the Senate and the President will join the House in acting on the other many pro-growth items we have already passed and will be passing in the coming months.
See you next week.
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Nokia has told TechRadar that there is no truth in the rumours it is set to join with the Open Handset Alliance and launch an Android phone.
The Guardian is citing industry insiders who claim that the new device will be unveiled at the Nokia World event in Germany in September.
But Nokia has told us: "There is no truth to this story whatsoever. It is a well known fact that Symbian is our platform of choice for smartphones."
It would have been quite a departure for Nokia to launch such a device given the company owns the Symbian Foundation (makers of the Symbian Mobile OS) and is a staunch stalwart of said platform (obviously).
Market vs Ovi
Were Nokia to have moved to Android, it would have placed Google Market in direct competition with the recently released Ovi Store, which does the same thing.
However, the current high end Symbian S60 platform Nokia (and Samsung) currently use on the likes of the N97 and i8910HD may still have a limited life span, although there's no word on when that could be.
This is because Nokia is still beavering away on the new Maemo platform, its OS for its internet tablets, which could make the jump to future mobile phones too, so we'll have to wait and see which path the Finnish bigwigs take.


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How to Configure Your Laptop for a Remote Desktop Connection
With Windows Vista, it's amazingly easy to set up the Remote Desktop feature so that you can work between your laptop and desktop computers. Keep in mind that you should use Remote Desktop only in a secure environment — say, over a local network and not over the Internet.
Set up a PC for Remote Desktop
You can configure either the desktop or laptop computer for remote access; either way works fine.
Choose Control Panel from the Start menu, and open the Control Panel's System icon.
In the list of tasks on the left side of the window, choose Remote Settings.
If a User Account Control (UAC) dialog box appears, click the Continue button.
In the System Properties dialog box, on the Remote tab in the Remote Desktop area, choose the item labeled Allow Connections from Computers Running Any Version of Remote Desktop.
Click OK, and close the Control Panel window.
That's it. The computer is now open to sharing its desktop remotely with another computer on the network.
Create an exception for the Windows Firewall
For the Remote Desktop to work, you have to create an exception for the Windows Firewall.
Choose Control Panel from the Start menu, and open the Control Panel's Windows Firewall icon.
In the Windows Firewall window, click the link that says Change Settings.
Click the Continue button if prompted by a User Account Control warning.
In the Windows Firewall Settings dialog box, click the Exceptions tab.
Scroll through the list to find the item called Remote Desktop, and place a check mark by that item.
Click the OK button, and then close the Windows Firewall and Control Panel windows.
What you can do with Remote Desktop
When the remote desktop is set up and connected, what you see on your laptop's screen is actually the display of another computer on the network. Moving the mouse on your laptop moves the mouse on the other computer, and ditto for the keyboard. It's just as though you're sitting at that computer, when you're actually working things from a remote location.
The remote desktop can be displayed on a full screen or in a window. In Full Screen mode, a strip appears across the top of the screen and acts as a sort of window control. In Window mode, the remote desktop appears in a window on your laptop's screen.
Sadly, you cannot copy files and folders between the remote system and your own computer by dragging things into and out of the remote desktop's window. The Remote Desktop Connection is more of a control and access feature than a file exchange utility.


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Starting MSRP from * $34,900
2020 Chevrolet Express Passenger Lexington, TN Jones Chevrolet Lexington
Jones Chevrolet Lexington has a large selection of 2020 Chevrolet Express Passenger for our Lexington, TN drivers. With miles of rigorous testing, the Chevrolet Express Passenger is one of the strongest, most intelligent and capable vehicles ever. The impressive engine provides power while making this one of the most fuel-efficient vehicles on the market. We encourage you to come test drive this vehicle to see for yourself.


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A family in a Toyota Sienna minivan were driving down a New Orleans street this weekend when a single-engined plane suddenly appeared in front of them. Unable to dodge the low-flying Piper's wings the two collided. Quick, someone call the Farmer's insurance university professor guy.
Police say 60-year-old Houston Hunter was flying over Lake Ponchartrain to drop off his grandson when the engine on his 50-year-old Piper suddenly starting cutting out on him. Unable to return to the airport, Hunter set down on West End Boulevard in New Orleans.
Unfortunately, a family of four and their grandmother were coming from the opposite direction in their minivan. We'll let the daughter's translation of the grandma explain what hat was like "She thought she would die… she never thought that she would see a plane in her face."
Both vehicles sustained major damage but neither the family in the van nor the pilot suffered serious injuries.
Photo Credit: AP/Gerald Herbert


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Baked chicory with chicken in a sage & mustard sauce
Nutrition and extra info
Nutrition: per serving
- kcal510
- fat32g
- saturates18g
- carbs11g
- sugars0g
- fibre1g
- protein42g
- salt1.16g
- 3 head of white chicory
Also known as endive, chicory is a forced crop, grown in complete darkness, which accounts for…
- 1 tbsp olive oil
Olive oilol-iv oyl
Probably the most widely-used oil in cooking, olive oil is pressed from fresh olives. It's…
- 3 tbsp dry white wine, chicken stock or water
- 4 skinless boneless chicken breasts
Chicken's many plus points - its versatility, as well as the ease and speed with which it…
- 25g butter
Butter is made when lactic-acid producing bacteria are added to cream and churned to make an…
- 1 shallot, finely chopped
Related to the onion (as opposed to being a younger version of it), shallots grow in clusters at…
- 1 garlic clove, chopped
For the sauce
- 25g butter
Butter is made when lactic-acid producing bacteria are added to cream and churned to make an…
- 1 tbsp plain flour
- 300ml milk
One of the most widely used ingredients, milk is often referred to as a complete food. While cow…
- 150ml crème fraîche or soured cream
- 4-6 sage leaves, chopped
Popular in both Italian and British cookery, sage has long, grey-green leaves with a slightly…
- 2-3 tsp smooth Dijon mustard
A condiment made by mixing the ground seeds of the mustard plant with a combination of…
- 2-3 tsp grainy Dijon mustard
A condiment made by mixing the ground seeds of the mustard plant with a combination of…
- 50g grated gruyère or cheshire cheese
Gruyère is an undoubted pinnacle of traditional Swiss cheese-making, a culinary masterpiece as…
Preheat your oven to fan 180C/conventional 200C/gas 6. Trim away the stalk ends of the chicory, discard any limp or tired outer leaves, then cut the vegetable lengthways into quarters. Pour the oil over the bottom of a large shallow baking dish and arrange the sections of chicory, cut-side up, in a single layer. Pour over the wine, stock or water and cover with foil. Bake for 10 minutes while you prepare the rest of the ingredients.
Slice the chicken into strips 1cm thick. Season lightly with salt and pepper. Melt the butter in a large frying pan until foaming, add sliced chicken and cook for 4-5 minutes or until the chicken is just turning golden brown. Stir in the shallot and garlic and cook for 1 minute. Remove the dish from the oven and add the chicken mixture to the chicory. Replace the foil and set the dish aside while you make the sauce.
Melt the butter for the sauce in the frying pan (no need to wash it) then stir in the flour. Gradually pour in the milk, stirring all the time until the mixture thickens. Remove from the heat and stir in the crème fraîche or soured cream and sage with 2 tsp each of both mustards. Taste and add extra mustard plus salt and pepper if you think they're needed.
Pour the sauce over the chicory and chicken and sprinkle cheese on top. Bake, uncovered, for a further 25-30 minutes until the sauce is bubbling and the top is golden.


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Viking Atla deck 4 plan (Sun)
Deck layout and review
Review of Viking Atla deck 4 (Sun)
- Navigation Bridge (called Wheelhouse on riverboats) is passenger accessible, and can be lowered when the ship passes under low bridges.
- Sun deck is enclosed by glass railings, with 360-degree scenery views, comfortable deck chairs, tables, loungers, canopy shaded sitting areas. The sundeck's relaxation area is encircled by the Walking / Jogging Track.
- Sky Bar is an enclosed bar with outdoor seating (small 4-seat tables and chairs).
- With aft location are two practice Putting Green areas, the ship's Herb Garden (organic herbal garden) and the deck games zone (Shuffleboard, giant Deck Chess set).
- Solar Panels at the rear are an alternative energy source for the Atla ship's hybrid engines.


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I knew a guy named Reed who turned out to be a scurvy dog, but I wouldn't say he was a wolf!
In all seriousness, the Marlin is a great rifle, too. I own a Model 60, one of the best rifles ever made.
Brownings are nice, but in terms of quality AND accuarcy AND ruggedness, it's hard to beat the Marlin.


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text: Lift chairs in the Heritage collection strike a balance between luxury features and price, giving you a lift chair that is an incredible value and will perform for many years to come. The GL-358L is a large-size model. It has 3 recline positions and comes with a full chaise pad and button back seat.


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Visit to Trogon Hall gallery for information about the conservation of Tabin Wildlife Reserve and briefing on birds, insects and animals that is found in this area. (For guest check in before 11:00am)
In the afternoon, visit the active Lipad mud volcano which provides a mineral salt lick for wildlife. After trekking through the rainforest for 20 minutes, arrive at the elevated bare area of the mud volcano which is frequented by wildlife and birds. Climb the observation tower and admire the contrasting landscape of the bare mud volcano and its surrounding thick forest, where birds are commonly sighted. Try to identify the multitude of animal footprints here. The adventurous can try the effects of the volcanic mud, said to be responsible for a fine complexion. Trek back to the resort along elephant trail.
After dinner, enjoy a CD/Slide presentation on Tabin's natural wonders, followed by a night safari to look out for nocturnal wildlife and birds. Remember to look up at the clear, dark sky of Tabin, identify the constellations above you and make a wish or two. (L,D)
Day 02
Join the early morning bird walk to discover some of Tabin's rich collection of almost 300 bird species. After breakfast, a drive to the core area for animal's spotting and for photo session before trekking through the lush green rainforest known for its rich diversity of plants and animal species to visit Lipad Waterfall. Take a dip in the natural pool and enjoy a natural and refreshing shower under the waterfall. Alternatively explore the surrounding vegetation or just relax.
In the afternoon, try out the Tabin Rainforest Foot Soak made with a natural blend of herbs, to relax tired muscles after long jungle treks.
In the evening, a dusk drive to look out for more nocturnal wildlife & birds coming out to look for food during this time of the day.
After dinner, go for a night walk along the trail nearby the resort in search of sleeping birds, frog and nocturnal wildlife. (B,L,D)
Day 03
After breakfast, free and easy until check out. (B,L)
You may opt to take up a dawn drive or jungle walk to look out for wildlife that is active at this time of the day.
[Not included in package/RM80.00 per person (dawn drive) , RM50.00 per person (Jungle walk)
By road:Kota Kinabalu to Lahad Datu (7 hours)
Sandakan to Tabin (4 hours)
Sukau Kinabatangan to Tabin (4 hours)
Semporna (Sipadan island) to Tabin (4 hours)
|Map Legend
|Borneo Rainforest Wildlife Adventure
Jalan Tengku, Lahad Datu, Sabah, Malaysia
|Sepilok Orang Utan Centre
|4 hours by road
|Sukau Kinabatangan
|4 hours by road
|4 hours by road
|Danum Valley
|4 hours by road


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The Nursing MSc degree confers both an academic award and a professional qualification in nursing. On completion of the programme, you will be recommended to the Nursing and Midwifery Council for professional registration.
Once qualified, you will have a range of employment opportunities in leading NHS and independent organisations. Staff development and continuing professional development programmes support and encourage life-long learning.
Healthcare is constantly developing and advancing as the needs of the population change, more research evidence emerges and new technologies are created. Once you've qualified, it's important to keep up to date with current healthcare knowledge and practice. This will be required by the Nursing and Midwifery Council (NMC) and encouraged by your employer.
Today's NHS is going through a period of unprecedented change. Nurses are learning new skills and taking on more responsibility. The number of nurse-led specialist services is growing and nurses are playing a more active role in managing people's health and wellbeing, both in hospital and the community.
There are many opportunities for nurses to advance their careers by developing specialist skills. These include clinical skills such as suturing, venepuncture, cannulation and prescribing, as well as running specialist nurse-led services in areas like diabetes and nutrition.
New roles such as nurse practitioner and nurse consultant demand more advanced clinical and academic skills and provide further career development opportunities with salaries that reflect these specialist skills.
Qualified nurses play a major role in educating nursing students and training their own colleagues. Many find mentoring very rewarding and most will be involved in teaching and supervising students on practice placements. The lecturer practitioner role also allows nurses to combine an interest in teaching with hands-on practice.
All modern nursing practice is evidence-based - which means applying clinical research findings to deliver the most suitable care to patients. Nurses are encouraged to reflect on their practice, undertake research, publish papers and speak at conferences and seminars. There are many examples of nurse-led research being used to change practice within clinical settings.
Professional nurses today have management and leadership responsibilities and make a real contribution to changing the way services are organised and delivered.
We offer a wide range of opportunities for continuing professional development and lifelong learning, including events and seminars, study days, short courses and qualifications at undergraduate and postgraduate level.
Find out more about our continuing professional development opportunities.
On successful completion of this programme students can apply to register as a nurse with the Nursing and Midwifery Council.
Admissions team, Faculty of Health and Social Care Sciences
Tel: +44 (0)20 8417 5735
Email us
This course is taught at Kingston Hill
Admissions team, Faculty of Health and Social Care Sciences
Tel: +44 (0)20 8417 5735
Email us
This course is taught at Kingston Hill