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Spin pumping in magnetic trilayer structures with an MgO barrier
Views: 1,717Citations: 1More detailArticle Open
AbstractWe present a study of the interaction mechanisms in magnetic trilayer structures with an MgO barrier grown by molecular beam epitaxy. The interlayer exchange coupling, Aex, is determined using SQUID magnetometry and ferromagnetic resonance (FMR), displaying an unexpected oscillatory behaviour as the thickness, tMgO, is increased from 1 to 4nm. Transmission electron microscopy confirms the continuity and quality of the tunnelling barrier, eliminating the prospect of exchange arising from direct contact between the two ferromagnetic layers. The Gilbert damping is found to be almost independent of the MgO thickness, suggesting the suppression of spin pumping. The element specific technique of x ray detected FMR reveals a small dynamic exchange interaction, acting in concert with the static interaction to induce coupled precession across the multilayer stack. These results highlight the potential of spin pumping and spin transfer torque for device applications in magnetic tunnel junctions relying on commonly used MgO barriers.
IntroductionMuch attention is currently devoted to the study of spin transfer torque (STT)1,2,3,4, through which it is possible to realize spontaneous magnetization precession and switching. Spin pumping from a ferromagnet (FM) into a non fake Van Cleef & Arpels Clover necklace black magnetic (NM) material is one of the most promising candidates for these applications5. Spin pumping has been under intense scrutiny since it was first proposed in 20026, studying the generation of pure spin currents by ferromagnetic resonance (FMR). Of particular importance is the transmission of spins across NM barriers, such as the conductors Ag (ref. 7), Au (ref. 8), and Cu (ref. 9), or insulators like MgO (ref. 10) and SrTiO3 (ref. 11). The efficacy of spin pumping is governed by the spin diffusion length of the NM layer, and the spin mixing conductance of the FM/NM interface12. Spin pumping is heavily suppressed in insulators, leading to very short spin coherence lengths, often under a nm11. However, coupling of the spin and charge degrees of freedom, and associated charge pumping in tunnelling heterostructures with an insulating barrier, can complicate interpretation of the results5,13,14. It is useful to investigate spin pumping through insulating layers using probes that are insensitive to charge based effects.
Here, we present a study of spin pumping and static exchange coupling in MgO based magnetic heterostructures grown by molecular beam epitaxy (MBE). These magnetic tunnel junctions (MTJs) are well established high tunnelling magnetoresistance ratio elements15,16. The strength and character of the interactions is determined in Co50Fe50/MgO/Ni magnetic heterostructures using superconducting quantum interference device (SQUID) magnetometry and vector network analyser (VNA) FMR. The structures are of high crystalline quality, as demonstrated by transmission electron microscopy (TEM). Static interlayer exchange is observed through shifts in the resonant field. X ray detected FMR (XFMR) measurements confirm the presence of static exchange coupling, but demonstrate that for the thinnest MgO barrier there is also a component of spin pumping. These results show the importance of spin transfer in technologically relevant MTJs, demonstrating that torques can be achieved for suitably thin barriers.
Ferromagnetic ResonanceIn the classical limit the spin dynamics of a ferromagnet are governed by the damped Landau Lifshitz Gilbert (LLG) equation of motion, describing the precession of the magnetisation about an effective field arising from internal and external fields:
with the gyromagnetic ratio, m is the unit magnetisation vector of the material, the dimensionless Gilbert damping parameter, and Heff the effective magnetic field. Through Heff the various energy terms, such as the exchange energy, the demagnetisation energy, the in plane cubic and uniaxial anisotropies, the static exchange, and the Zeeman energy enter the equation.
The static interlayer exchange coupling is a general term for any interaction that acts to (anti )align the magnetisations of the two layers in a magnetic trilayer structure such as a spin valve or an MTJ. Examples of such interactions include Ruderman Kittel Kasuya Yosida (RKKY), superexchange, Nel or orange peel coupling, and direct exchange through a discontinuous spacer layer. The presence of a static interaction modifies the LLG equation with an additional term17:
where is the interlayer exchange, and i, j index magnetic layers. The interlayer exchange is defined as18,19:
with Aex the interlayer exchange constant, t the thickness of the magnetic layer, M the equilibrium orientation of the magnetization, and i, j indices label the magnetic layers. The sign of Aex indicates whether the interaction favours parallel (positive) or antiparallel (negative) alignment.
In FMR experiments, additional interaction mechanisms must be considered. As the magnetisation of a ferromagnetic layer precesses on resonance it acts as a spin battery, generating a pure spin current transverse to the axis about which it precesses. When the FM layer is thicker than the ferromagnetic coherence length, a pure spin current can be driven into an adjacent NM layer. A spin current can persist across the spacer layer, and in a trilayer structure either return to the first FM/NM interface, or else flow through to a second NM/FM interface. If it is not reflected here, it crosses the interface and is absorbed by the FM, inducing precession through the STT4.
The increased flow of spin momentum out of the FM layer acts as an additional channel for energy loss, leading to an increase in damping. This damping is linear with resonant frequency, and can thus be described in the same terms as Gilbert damping. The absorbed spin current leads to a comparable anti damping term in the off resonance layer.
With the addition of these two coupling mechanisms, the LLG for a trilayer system becomes:
where superscripts denote magnetic layers, and is the effective field acting on layer i. The intrinsic Gilbert damping is , while are spin source (m=n) and spin sink (mn) terms. The XFMR results can be modelled using this a linearised solution of this equation.
The presence of these coupling mechanisms alters the magnetodynamics of the on and off resonance layer. The static coupling shifts the resonant field of both layers, as it functions as an additional field term in the Kittel equation. Spin pumping, on the other hand, broadens the resonances by providing an additional energy loss mechanism. However, the most important change is the precession induced in the off resonance layer, wherein the exchange interactions transfer energy between the two layers. This leads to changes in both the phase and amplitude of precession, with the phase being the more sensitive probe.
ResultsStructural properties of the magnetic trilayer structuresThe structural properties of the grown magnetic heterostructures were studied using transmission electron microscopy (TEM). Figure 1b d shows cross sectional TEM images of the magnetic heterostructures. These images demonstrate epitaxial growth of the films, and confirm that the MgO barrier is continuous (down to the thinnest barrier thickness of 1nm, see Supplementary Fig. S1 for details).
Figure 1: Heterostructure growth and structural properties.(a) RHEED images of the MgO substrate (at 700C), CoFe layer, 3 nm thick MgO barrier (after annealing at 300C), and Ag capped Ni layer (from left to right). (b d) Cross sectional TEM view of the magnetic heterostructure with a 2 nm thick MgO barrier. (b) Low magnification high angle annular dark field image of the magnetic heterostructure showing uniform thickness of the ferromagnetic layers and the MgO tunnel barrier. (c) High resolution bright field scanning TEM showing the atomic structure of the substrate, ferromagnetic layers, and barrier viewed along [010]. (d) Interface region of ferromagnetic layer(s)/MgO barrier showing the atomically abrupt interfaces and the well structured MgO barrier textured along the [010] direction.
Figure 2 shows hysteresis loops measured by SQUID VSM for all four magnetic heterostructures, displaying a reduction in static interlayer coupling as a function of increasing tMgO. For the thinnest MgO barrier (tMgO=1nm, Fig. 2a) the two layers are strongly bound, and there is a single switching step, with a coercive field of 2mT. As the thickness of the barrier increases, the layers decouple and behave independently. For tMgO=2 and 3nm (Fig. 2bc), the coupling between the two layers appears to cause winding, leading to a smeared out transition as opposed to sharp steps. Nevertheless, two distinct steps in the hysteresis loop can be identified. Strong coupling for the thinnest barrier aligns the magnetizations of the two layers at all fields, but as barrier thickness increases they start to move independently, leading to two distinct steps in (d).
As the TEM measurements show that the MgO barrier appears to be continuous, the observed coupling cannot be due to direct, large area contact between the two ferromagnetic layers, but it is more likely to be resulting from tunnelling across the barrier. However, there is some evidence of surface roughness of the MgO, which could lead to a Nel orange peel coupling.
Lab based FMR measurementsVNA FMR measurements were performed to determine values for the interlayer exchange coupling and magnetocrystalline anisotropy parameters for all samples. Figure 1a shows a representative field frequency map for the sample with tMgO=1 nm. Fits to resonance fields were performed as a function of RF frequency and magnetization alignment using the Kittel equation20, including the interlayer exchange field17,18, using Eq. (3).
The extracted interlayer coupling is shown in Fig. 3b, while values for all the copy van cleef jewelry fitting parameters are given in Table 1. 21). 18. Note that in spin dependent tunnelling such an oscillatory thickness dependence has been observed for MgO barriers in Fe/MgO/Fe tunnelling structures22. (b) Interlayer exchange coupling, Aex, as a function of MgO barrier thickness extracted from frequency dependence of the resonance field, using the Kittel equation. (c) Gilbert damping extracted from linewidth of resonance using Eq. (5). Results for both FM layers are plotted as a function of MgO barrier thickness in Fig. 3c. Within the error bars, there is no change in the Gilbert damping of the Ni layer, which supports the assertion that an insulating layer such as MgO does not permit a spin current to flow. The data for the CoFe layer is more complicated, showing at first a slight increase, then a drop in damping. Microscopy and magnetic characterization gave no evidence of a change in the structural or magnetic properties of the CoFe layer to account for this finding.
While a drop in damping with increasing interlayer thickness is usually indicative of spin pumping, the trend here does not match the exponential decay observed in previous studies7,11,24. Measurements on comparable bare CoFe layers yielded a Gilbert damping of 3103. This suggests that at least some of the spins pumped by the van cleef clover jewelry replica precessing magnetization of the CoFe are absorbed. The lower damping in the case of the thinnest MgO layer could be in part due to the strong static exchange coupling observed.
Synchrotron based FMR measurementsXFMR measurements were performed to study off resonance precession induced by static exchange coupling and the STT exerted by pumped spins. Figure 4 shows the amplitude and phase of precession at 4GHz for each layer in the sample with tMgO=1nm, with the magnetic field applied along the easy axis of the CoFe. The resonance in the Ni layer is easily identified, with a peak in amplitude at 28mT. Off resonance precession is induced in the CoFe layer; the resulting dynamic XMCD signal is approximately ten times smaller than that of the Ni. There is also a small phase feature in the Co, which is predominantly unipolar in character. Taken together with the bipolar shape of the Co amplitude variation, this indicates that the static interaction is primarily responsible for induced precession in the CoFe.
Views: 1,717Citations: 1More detailArticle Open
AbstractWe present a study of the interaction mechanisms in magnetic trilayer structures with an MgO barrier grown by molecular beam epitaxy. The interlayer exchange coupling, Aex, is determined using SQUID magnetometry and ferromagnetic resonance (FMR), displaying an unexpected oscillatory behaviour as the thickness, tMgO, is increased from 1 to 4nm. Transmission electron microscopy confirms the continuity and quality of the tunnelling barrier, eliminating the prospect of exchange arising from direct contact between the two ferromagnetic layers. The Gilbert damping is found to be almost independent of the MgO thickness, suggesting the suppression of spin pumping. The element specific technique of x ray detected FMR reveals a small dynamic exchange interaction, acting in concert with the static interaction to induce coupled precession across the multilayer stack. These results highlight the potential of spin pumping and spin transfer torque for device applications in magnetic tunnel junctions relying on commonly used MgO barriers.
IntroductionMuch attention is currently devoted to the study of spin transfer torque (STT)1,2,3,4, through which it is possible to realize spontaneous magnetization precession and switching. Spin pumping from a ferromagnet (FM) into a non fake Van Cleef & Arpels Clover necklace black magnetic (NM) material is one of the most promising candidates for these applications5. Spin pumping has been under intense scrutiny since it was first proposed in 20026, studying the generation of pure spin currents by ferromagnetic resonance (FMR). Of particular importance is the transmission of spins across NM barriers, such as the conductors Ag (ref. 7), Au (ref. 8), and Cu (ref. 9), or insulators like MgO (ref. 10) and SrTiO3 (ref. 11). The efficacy of spin pumping is governed by the spin diffusion length of the NM layer, and the spin mixing conductance of the FM/NM interface12. Spin pumping is heavily suppressed in insulators, leading to very short spin coherence lengths, often under a nm11. However, coupling of the spin and charge degrees of freedom, and associated charge pumping in tunnelling heterostructures with an insulating barrier, can complicate interpretation of the results5,13,14. It is useful to investigate spin pumping through insulating layers using probes that are insensitive to charge based effects.
Here, we present a study of spin pumping and static exchange coupling in MgO based magnetic heterostructures grown by molecular beam epitaxy (MBE). These magnetic tunnel junctions (MTJs) are well established high tunnelling magnetoresistance ratio elements15,16. The strength and character of the interactions is determined in Co50Fe50/MgO/Ni magnetic heterostructures using superconducting quantum interference device (SQUID) magnetometry and vector network analyser (VNA) FMR. The structures are of high crystalline quality, as demonstrated by transmission electron microscopy (TEM). Static interlayer exchange is observed through shifts in the resonant field. X ray detected FMR (XFMR) measurements confirm the presence of static exchange coupling, but demonstrate that for the thinnest MgO barrier there is also a component of spin pumping. These results show the importance of spin transfer in technologically relevant MTJs, demonstrating that torques can be achieved for suitably thin barriers.
Ferromagnetic ResonanceIn the classical limit the spin dynamics of a ferromagnet are governed by the damped Landau Lifshitz Gilbert (LLG) equation of motion, describing the precession of the magnetisation about an effective field arising from internal and external fields:
with the gyromagnetic ratio, m is the unit magnetisation vector of the material, the dimensionless Gilbert damping parameter, and Heff the effective magnetic field. Through Heff the various energy terms, such as the exchange energy, the demagnetisation energy, the in plane cubic and uniaxial anisotropies, the static exchange, and the Zeeman energy enter the equation.
The static interlayer exchange coupling is a general term for any interaction that acts to (anti )align the magnetisations of the two layers in a magnetic trilayer structure such as a spin valve or an MTJ. Examples of such interactions include Ruderman Kittel Kasuya Yosida (RKKY), superexchange, Nel or orange peel coupling, and direct exchange through a discontinuous spacer layer. The presence of a static interaction modifies the LLG equation with an additional term17:
where is the interlayer exchange, and i, j index magnetic layers. The interlayer exchange is defined as18,19:
with Aex the interlayer exchange constant, t the thickness of the magnetic layer, M the equilibrium orientation of the magnetization, and i, j indices label the magnetic layers. The sign of Aex indicates whether the interaction favours parallel (positive) or antiparallel (negative) alignment.
In FMR experiments, additional interaction mechanisms must be considered. As the magnetisation of a ferromagnetic layer precesses on resonance it acts as a spin battery, generating a pure spin current transverse to the axis about which it precesses. When the FM layer is thicker than the ferromagnetic coherence length, a pure spin current can be driven into an adjacent NM layer. A spin current can persist across the spacer layer, and in a trilayer structure either return to the first FM/NM interface, or else flow through to a second NM/FM interface. If it is not reflected here, it crosses the interface and is absorbed by the FM, inducing precession through the STT4.
The increased flow of spin momentum out of the FM layer acts as an additional channel for energy loss, leading to an increase in damping. This damping is linear with resonant frequency, and can thus be described in the same terms as Gilbert damping. The absorbed spin current leads to a comparable anti damping term in the off resonance layer.
With the addition of these two coupling mechanisms, the LLG for a trilayer system becomes:
where superscripts denote magnetic layers, and is the effective field acting on layer i. The intrinsic Gilbert damping is , while are spin source (m=n) and spin sink (mn) terms. The XFMR results can be modelled using this a linearised solution of this equation.
The presence of these coupling mechanisms alters the magnetodynamics of the on and off resonance layer. The static coupling shifts the resonant field of both layers, as it functions as an additional field term in the Kittel equation. Spin pumping, on the other hand, broadens the resonances by providing an additional energy loss mechanism. However, the most important change is the precession induced in the off resonance layer, wherein the exchange interactions transfer energy between the two layers. This leads to changes in both the phase and amplitude of precession, with the phase being the more sensitive probe.
ResultsStructural properties of the magnetic trilayer structuresThe structural properties of the grown magnetic heterostructures were studied using transmission electron microscopy (TEM). Figure 1b d shows cross sectional TEM images of the magnetic heterostructures. These images demonstrate epitaxial growth of the films, and confirm that the MgO barrier is continuous (down to the thinnest barrier thickness of 1nm, see Supplementary Fig. S1 for details).
Figure 1: Heterostructure growth and structural properties.(a) RHEED images of the MgO substrate (at 700C), CoFe layer, 3 nm thick MgO barrier (after annealing at 300C), and Ag capped Ni layer (from left to right). (b d) Cross sectional TEM view of the magnetic heterostructure with a 2 nm thick MgO barrier. (b) Low magnification high angle annular dark field image of the magnetic heterostructure showing uniform thickness of the ferromagnetic layers and the MgO tunnel barrier. (c) High resolution bright field scanning TEM showing the atomic structure of the substrate, ferromagnetic layers, and barrier viewed along [010]. (d) Interface region of ferromagnetic layer(s)/MgO barrier showing the atomically abrupt interfaces and the well structured MgO barrier textured along the [010] direction.
Figure 2 shows hysteresis loops measured by SQUID VSM for all four magnetic heterostructures, displaying a reduction in static interlayer coupling as a function of increasing tMgO. For the thinnest MgO barrier (tMgO=1nm, Fig. 2a) the two layers are strongly bound, and there is a single switching step, with a coercive field of 2mT. As the thickness of the barrier increases, the layers decouple and behave independently. For tMgO=2 and 3nm (Fig. 2bc), the coupling between the two layers appears to cause winding, leading to a smeared out transition as opposed to sharp steps. Nevertheless, two distinct steps in the hysteresis loop can be identified. Strong coupling for the thinnest barrier aligns the magnetizations of the two layers at all fields, but as barrier thickness increases they start to move independently, leading to two distinct steps in (d).
As the TEM measurements show that the MgO barrier appears to be continuous, the observed coupling cannot be due to direct, large area contact between the two ferromagnetic layers, but it is more likely to be resulting from tunnelling across the barrier. However, there is some evidence of surface roughness of the MgO, which could lead to a Nel orange peel coupling.
Lab based FMR measurementsVNA FMR measurements were performed to determine values for the interlayer exchange coupling and magnetocrystalline anisotropy parameters for all samples. Figure 1a shows a representative field frequency map for the sample with tMgO=1 nm. Fits to resonance fields were performed as a function of RF frequency and magnetization alignment using the Kittel equation20, including the interlayer exchange field17,18, using Eq. (3).
The extracted interlayer coupling is shown in Fig. 3b, while values for all the copy van cleef jewelry fitting parameters are given in Table 1. 21). 18. Note that in spin dependent tunnelling such an oscillatory thickness dependence has been observed for MgO barriers in Fe/MgO/Fe tunnelling structures22. (b) Interlayer exchange coupling, Aex, as a function of MgO barrier thickness extracted from frequency dependence of the resonance field, using the Kittel equation. (c) Gilbert damping extracted from linewidth of resonance using Eq. (5). Results for both FM layers are plotted as a function of MgO barrier thickness in Fig. 3c. Within the error bars, there is no change in the Gilbert damping of the Ni layer, which supports the assertion that an insulating layer such as MgO does not permit a spin current to flow. The data for the CoFe layer is more complicated, showing at first a slight increase, then a drop in damping. Microscopy and magnetic characterization gave no evidence of a change in the structural or magnetic properties of the CoFe layer to account for this finding.
While a drop in damping with increasing interlayer thickness is usually indicative of spin pumping, the trend here does not match the exponential decay observed in previous studies7,11,24. Measurements on comparable bare CoFe layers yielded a Gilbert damping of 3103. This suggests that at least some of the spins pumped by the van cleef clover jewelry replica precessing magnetization of the CoFe are absorbed. The lower damping in the case of the thinnest MgO layer could be in part due to the strong static exchange coupling observed.
Synchrotron based FMR measurementsXFMR measurements were performed to study off resonance precession induced by static exchange coupling and the STT exerted by pumped spins. Figure 4 shows the amplitude and phase of precession at 4GHz for each layer in the sample with tMgO=1nm, with the magnetic field applied along the easy axis of the CoFe. The resonance in the Ni layer is easily identified, with a peak in amplitude at 28mT. Off resonance precession is induced in the CoFe layer; the resulting dynamic XMCD signal is approximately ten times smaller than that of the Ni. There is also a small phase feature in the Co, which is predominantly unipolar in character. Taken together with the bipolar shape of the Co amplitude variation, this indicates that the static interaction is primarily responsible for induced precession in the CoFe.
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When the future leader was a child, he learned how to drive a Mercedes and would take it for a spin around the grounds of the presidential palace.
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The chef who uses copy Van Cleef & Arpels butterfly necklace a pseudonym has previously revealed that Kim Jong Il, who died in 2011, enjoyed eating fish which had been killed so recently that it was still moving.
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Mr Fujimoto managed to escape North Korea by showing Kim a video of a Japanese cooking programme featuring a sea urchin dish, and persuading the dictator to allow him to travel to Japan in an attempt to recreate the meal.
However, he returned to the country last year and was pictured sharing a hug with the despot he knew as a child, claiming that the situation in North Korea had improved since he left.
North Korea has apparently been amassing troops in preparation for renewed hostilities, and although the situation seems to have cooled this week the country's enemies have remained ready for the possibility of military confrontation.
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Kim Jong Un may be fond of fiery anti Western rhetoric, but in fact the North Korean dictator is obsessed with the European and American lifestyle, according to a man who has known him since he was a little boy.
Kenji Fujimoto, who served as the personal sushi chef of Kim's father Kim Jong Il for 13 years, has revealed that the 30 year old despot is a fan of the Beatles and longs to look like Hollywood star Jean Claude Van Damme.
Kim also drinks Bordeaux wine, smokes Cartier menthol cigarettes and has enjoyed driving around his palace in a Mercedes since he was just seven years old.
He also enjoys Hollywood hits, and the Beatles are his favourite Western band.
One film in particular had a replica Van Cleef & Arpels butterfly necklace big impact on Kim, when a Jean Claude Van Damme karate movie inspired him to start bulking up so he could look more like the Belgian star.
'His action is so awesome,' the teenager said, according to Mr Fujimoto, who added: 'He started buying fitness equipment and taking protein supplements.'
When the future leader was a child, he learned how to drive a Mercedes and would take it for a spin around the grounds of the presidential palace.
'He drove it perfectly when he was only seven,' Mr Fujimoto told The Sun. 'They made a special seat for him because he could not reach the pedals.'
The chef who uses copy Van Cleef & Arpels butterfly necklace a pseudonym has previously revealed that Kim Jong Il, who died in 2011, enjoyed eating fish which had been killed so recently that it was still moving.
He also described how the unhinged despot
would order his staff to dance with scantily clad girls, and claimed
that he had been married off to one of the exotic dancers.
Mr Fujimoto managed to escape North Korea by showing Kim a video of a Japanese cooking programme featuring a sea urchin dish, and persuading the dictator to allow him to travel to Japan in an attempt to recreate the meal.
However, he returned to the country last year and was pictured sharing a hug with the despot he knew as a child, claiming that the situation in North Korea had improved since he left.
North Korea has apparently been amassing troops in preparation for renewed hostilities, and although the situation seems to have cooled this week the country's enemies have remained ready for the possibility of military confrontation.
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EXCLUSIVE: First photos of Ariana Grande since Manchester. Help for the homeless heroes: Millionaire West Ham owner. The schoolgirls massacred by ISIS coward: Five teenagers. Revealed: Manchester bomber returned to Britain from. Theresa May warns a fresh terror attack is feared to be. 'Our little princess has been so lucky': Father's joy as. Britain on lockdown: Army deploys 1,000 heavily armed. Horror on the M6: Lorry driver is arrested after four. Tourists watch in horror as armed police arrest man. Mother of child actress pictured hugging a female police. Aaron imitation Van Cleef & Arpels Clover necklace price Hernandez's hell behind bars: NFL star killed. More than 24 hours on, desperate families still search. BREAKING NEWS: My son is innocent, insists father of. 'I won't forget what you said!' Trump tells Pope after. Comedian Jason Manford deletes Twitter after being. Terrorist's brother arrested: Dramatic moment armed. Grisly photos of scorched remnants of suicide bomber's. Bomber from a red brick semi who 'knew an ISIS. MOST READ NEWS Previous.
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specific genes identified from a compendium of microarray expression data
Immune response in silico (IRIS): immune specific genes identified from a compendium of microarray expression data
A R Abbas1, D Baldwin1, Y Ma1, W Ouyang2, A Gurney2,3, F Martin2, S Fong2, M van Lookeren Campagne2, P Godowski2, P M Williams3, A C Chan2 and H F Clark1,2
1Department of Bioinformatics, Genentech, Inc., South San Francisco, CA, USA2Department of Immunology, Genentech, Inc., South San Francisco, CA, USA3Department of Molecular Biology, Genentech, Inc., South San Francisco, CA, USACorrespondence: Dr HF Clark, Genentech, Inc., 1 DNA Way, South San Francisco, CA 94080, USA.
Top of pageAbstractImmune cell specific expression is one indication of the importance of a gene's role in the immune response. We have compiled a compendium of microarray expression data for virtually all human genes from six key immune cell types and their activated and differentiated states. Immune Response In Silico (IRIS) is a collection of genes that have been selected for specific expression in immune cells. The expression pattern of IRIS genes recapitulates the phylogeny of immune cells in terms of the lineages of their differentiation. Gene Ontology assignments for IRIS genes reveal significant involvement in inflammation and immunity. Genes encoding CD antigens, cytokines, integrins and many other gene families playing key roles in the immune response are highly represented. IRIS also includes proteins of unknown function and expressed sequence tags that may not represent genes. The predicted cellular localization of IRIS proteins is evenly distributed between cell surface and intracellular compartments, indicating that immune specificity is important at many points in the signaling pathways of the immune fake van cleef & arpels clover diamond ring response. This enables the immune response to attack foreign invaders while recognizing and tolerating self antigens. Two distinct lineages of immune cells have evolved with cell types that specialize in each of the many roles that are required for this immune response. The myeloid lineage of monocytes, macrophages, dendritic cells and neutrophils carries out the innate immune response, recognizing microbial pathogens typically by carbohydrates found only in bacterial proteins. The lymphoid lineage of T cells, B cells and natural killer (NK) cells enables adaptive immunity by distinguishing self from non self antigens and also providing a memory of foreign proteins seen before. Other immune cells, such as eosinophils, basophils and mast cells, also participate in the immune response. Although a large repertoire of genes that play key roles in the differentiation, function and regulation of these immune cells is already well described, this has not yet led to a complete understanding of immune diseases.
Genome wide microarray expression profiling of immune cells provides opportunities for identifying other genes that may function in the immune response. Microarray experiments have been carried out on various immune cell subsets by a number of different investigators in order to understand gene expression differences during differentiation and activation.1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17 These studies have provided invaluable insight into comprehensive gene expression profiles that define many different immune cell subsets and states of differentiation and activation. In particular, a recent study detailed elegantly the genes expressed specifically in many subsets of the T cell lineage.18 Comparison of gene expression profiles across a broad range of immune cell types and nonimmune tissues is necessary, however, to fully appreciate which gene expression differences are unique to each immune cell subset, which are found in multiple subsets or across immune cell lineages, and which are found more widely across the many cell types in the human body and thus may be involved in more general cellular processes. Differences among microarray platforms and experimental protocols used by different investigators often confound such comparison. Here, a compendium of microarray expression data from a broad representation of isolated immune cell subsets has been generated on the same microarray platform and analysis of gene expression profile across the major cell types of the immune system is made possible. Moreover, gene expression in all other major tissues allows determination of immune specific expression. Traditional methods of determining expression are performed on a gene by gene basis, and single microarray experiments show differential expression between just a few immune cell subsets. Here, genes fitting a complex expression criterion are identified on a genomic scale.
Immune cell specific expression is one indication of the importance of a gene's role in the function of the immune system. 'Cluster of differentiation' (CD) antigens are used experimentally as specific markers for immune cell subsets and they often play a key role in the function of that cell. For example, the T cell receptor is expressed only on T lymphocytes and is responsible for self antigen recognition, a primary function of this immune cell.19 Here, immune cell specifically expressed genes are identified by a survey of expression profiles across all the immune cells and major non immune tissues by a method termed Immune Response In Silico (IRIS). Immune cell specificity is determined generally by higher expression in any immune cell than expression in any nonimmune cell tissue. Finer characterization of specificity within an immune cell type, such as T cells, and an immune cell lineage, such as lymphoid cells, is also determined. Furthermore, expression profiles within subsets of an immune cell, such as classes of T cells expressing CD4 or CD8 antigens, memory and helper T cells and resting vs activated cells, further refine the specificity of immune specific genes. IRIS has identified both well characterized immune genes and a number of highly immune specific genes with unknown function.
Top of pageResultsIRIS identifies genes more highly expressed in immune cells than in any of the major organs of the body. Gene expression is surveyed across a compendium of samples, including immune cell subsets (Table 1) and a comprehensive range of normal tissues. The immune cells have been isolated from normal human blood by purifying each cell type via its specific cell surface markers. Activated and differentiated subsets were developed by in vitro stimulations. RNA samples were labeled and run on microarray chips that include probesets for virtually every human gene. Genes are often represented by more than one probeset and the expression levels vary, although the expression profiles are usually consistent. Genes are selected for IRIS based on cutoff values for expression levels in immune vs nonimmune samples, as shown in Supplementary Figure 1. Prior to the determination of these cutoffs, genomewide expression levels were evaluated by surveying the range of expression of all genes and the expression levels of a few families of genes already known to play important roles in the immune system. The highest mean expression level of any immune cell subset within each cell type as defined in Table 1 is calculated for each IRIS gene. Hierarchical clustering, a statistical method for grouping genes by the similarity of their expression profiles, reveals patterns of gene expression that are distinct among immune cell lineages. The dendrogram of the relationships of these genes mimics the evolutionary relationships of the lineages of immune cell types. The heat map showing all IRIS genes also illustrates the complexity of expression profiles, with most genes expressed at some level in more than one immune cell type. The highest mean expression of any cell subset for each probeset represents the expression of that cell type. The dendrograms shows the similarity of expression profiles for genes in the rows and cell types in the columns. Intensity values less than 1000 have the lightest shading and intensity values greater than 10,000 have the darkest shading.
Full figure and legend (166K)
IRIS categoriesWhile clustering is a useful method for identifying patterns of gene expression, it fails to clearly define the parameters of these patterns. Therefore, cutoff values of gene expression signatures in various immune cell types were approximated and used to define the IRIS categories that were observed in the clustering (Table 1) and these parameters are defined in the Cell Lineage Assignment section of Materials and methods. The probesets are categorized by their profiles according to the degree of specificity within immune cells. Profiles specific to one cell type are assigned to the categories T cell, NK cell, B cell, monocyte, dendritic or neutrophil. Profiles specific to more than one cell type within a lineage are assigned to lymphoid or myeloid categories. Profiles specific to cell types across lineages, as well as probesets with expression levels equivalent in all immune cells, are assigned to the multiple category. Genes represented by several probesets may appear in more than one category. A gene is best described as specific to the most exclusive category to which it has been assigned, that is, the highest expressing probeset may be in the T cell category and more weakly expressing probesets for that gene may fall into the lymphoid or multiple categories because of the stringency of cutoff levels used.
Patterns of expression profiles within each lineageCell types within a lineage share some immune specific genes, suggesting that they confer common functions among those cells. The expression profiles are very diverse and complex, but some general patterns emerge. As described above, the IRIS categories attempt to group genes within a cell type, or, if specific to more than one cell type, within a lineage. Here, these categories are assessed for general patterns of expression profiles. K means clustering reveals groups of expression profiles within an IRIS category, as shown in Figure 2. Within the lymphoid category clustered in Figure 2a, T cells and NK cells are shown to share a number of immune fake van cleef ring specific genes (clusters 1, 4 and 6), whereas B cells express more distinct genes (clusters 3 and 5). Often, the genes shared between T and NK cells are specifically expressed in the activated state of both cells (cluster 6). The myeloid category shown in Figure 2b reveals a similar relationship between genes transiently expressed in differentiating macrophages and also in LPS induced dendritic cells (cluster 8). Profile clusters of genes highly specific to single cell types are shown in Supplementary Figure 2. T cells have several general patterns of specificity among subsets of T cells, as shown in Supplementary Figure 2a. CD8 cells have a number of unique genes (cluster 1), and also share some genes with CD4 cells (cluster 2). T helper cells share some specific genes (clusters 3 and 6) and a number of genes appear only in activated memory T cells (cluster 5). B cells also show distinct profiles, as shown in Supplementary Figure 2b. Some genes are expressed specifically in both na and memory B cells (cluster 2), others only in plasma cells (clusters 3 and 4), while there are no specific B cell genes expressed distinctly in na or memory cells alone. Likewise, monocytes and macrophages shown in Supplementary Figure 2c reveal a class of genes specific to monocytes (cluster 1) and another transiently expressed in differentiating macrophages (cluster 2), but none specific to fully differentiated macrophages. However, Figure 2b shows that fully differentiated macrophages share profiles with neutrophils (cluster 2) and dendritic cells (cluster 5). All of these observations provide insights into the differentiation of immune cells as they have evolved to play specific roles in the immune response. The y axis is intensity level.
Full figure and legend (344K)
Statistical significance of IRISThe statistical significance of specificity is assessed for the genes within each IRIS cell type or lineage, as shown in Supplementary Figure 4. The F statistic measures both immune cell variation and sample variation simultaneously, so a low F value can either indicate similar expression of a gene across immune cells, or gene expression variability among samples, but does not distinguish between the two. Conversely, a gene highly specific to one cell type has a high F value, particularly if the replicate samples all have similar expression of that gene. Sample variability occurs because of both incomplete reproducibility of microarray conditions and biological variation between blood donors due to unknown causes. The multiple category has the highest proportion of genes with low F values, suggesting that this category includes most of the genes falsely assigned as immune specific. While the microarray technology used here is a well established experimental method for detecting differential gene expression, independent confirmation by real time quantitative PCR (RT QPCR) is carried out before pursuing further functional studies on genes of interest.20
Gene ontology and gene familiesGenes of known function are assigned a term from Gene Ontology, a structured vocabulary for describing biological process, molecular function and cellular component.21 Table 2 shows the Biological Processes most highly specific to IRIS, meaning that these ontologies are seen in IRIS in higher numbers than in a random set of genes. Many specific responses to different foreign invaders, chemotaxis of immune cells to sites of inflammation and other aspects of the immune response are represented. In summary, 57% of the well characterized IRIS genes have these immune functions. Similar results are seen with the molecular function ontologies (data not shown), with the most highly represented being antigen binding and the activities of chemokines, cytokines and their receptors. Table 3 shows the gene families represented in IRIS that are known to have many members with key roles in the differentiation, function and regulation of the immune system, including those with the molecular functions mentioned. The Protcomp algorithm (Softberry, Inc.) predicts for the 1589 IRIS genes with ORFs that 24% of the encoded proteins are in the plasma membrane, 13% are secreted, 24% are nuclear and the remaining 39% are in other intracellular compartments. An expression differential of 10 fold between the highest nonimmune tissue level and highest immune cell level selects 122 genes from the single cell type categories. The expression profiles of the highest expressing of these genes are shown in Figure 3 and the rest are shown in Supplementary Figure 3. Reported literature on many of these genes confirms that specific expression of transcripts often results in specific protein expression and that they have key roles in the immune response. For each cell type, a few genes of interest are described below. Highly specific genes with lower expression levels are shown in Supplementary Figure 3.
Full figure and legend (299K)
T cellsGenes with specificity in T cells have expression profiles in a wide variety of patterns as shown in Figure 3a and Supplementary Figure 3a. The T cell receptor alpha locus, TCRA, as well as CD3 subunits D, E G are Classic fake van cleef jewelry expressed on most T cell subsets. CD8A and B1 are heterodimers of the established protein marker for CD8+ T cells which stabilizes interactions between the T cell receptor and an MHC class I peptide complex,23 and are expressed only in CD8 cells. LEF1 is expressed specifically in both CD8 and CD4 cells. LEF1 is a transcription factor that has been implicated as critical in a differentiation defect that leads to T cell lymphomas.24 A number of genes exhibit exquisite selectivity for memory vs na T cells and most of these show enhanced expression in Th1 or Th2 subsets. Cytokine IL9 is specific to both Th2 and activated memory cells, whereas CTLA4 and ICOS are highly expressed in activated memory cells but also seen at lower levels in other T cell subsets and are both cell surface regulatory receptors.25 GZMK is specific to CD8, Th2 and resting memory cells and is a serine protease that is proposed to prevent damage of bystander cells at sites of inflammation.26 LAG3 is found primarily on Th1 cells, but also on stimulated CD8+ T cells (data not shown), and it stimulates the maturation of dendritic cells.27
NK cellsThe highest expressing specific gene in NK cells is KLRF1, a C type lectin with immunoreceptor tyrosine based inhibitory motifs (ITIM) motifs that stimulates cytolysis28 as shown in Figure 3b. Its expression is highest in resting cells, and decreases upon activation. The KIR gene cluster is also represented, although the probesets probably cross hybridize with these closely related genes that have probably arisen from gene duplication. Some members are established protein markers for this cell type and all are known to play a key role in recognition of HLA class I ligands.29 NS1 BP is specifically expressed in activated NK cells and binds the influenza A virus NS1 protein.30
B cellsB cells are represented by a diversity of specific expression profiles. TCL1A is expressed solely in na B cells and is an intracellular enzyme that regulates an early stage of B and T cell differentiation,31, 32 as shown with the other B cell genes in Figure 3c and Supplementary Figure 3b. CD20 and a number of other genes have equivalent expression in both na B cells and memory cells expressing either IgG/IgA or IgM. An antibody therapeutic targeting CD20, Rituxan, is extremely effective in specific ablation of B cells.33 CD19 is an established protein marker for B cells. BAFFR is the principal receptor for the signaling pathway that maintains mature B cell survival.34 BANK1 protein has also been shown to be specific to B cells and may play a role in foreign antigen induced immune response.35 PAX5 is a transcription factor that plays an essential role in commitment to the B cell lineage.36 CD79A and B are two components of the B cell antigen receptor37 and are expressed at lower levels in plasma cells. FCRH1, FCRH2 and IRTA2 are closely related in sequence to the Fc receptor which functions in antibody binding and regulation of the immune response,38 but they have not been functionally characterized.39, 40, 41, 42 FCRH1 and FCRH2 are found primarily on na and memory B cells, while IRTA2 is specific to plasma cells from bone marrow. A number of other genes are specific to plasma cells from blood, from bone marrow or from both sources. BCMA is a member of the TNF receptor family that is hypothesized to transduce signals for B cell survival and proliferation.43
Monocytes and macrophagesSeveral distinct patterns of specific monocyte and macrophage expression are seen in Figure 3d and Supplementary Figure 3c. Expressed only in monocytes is PRAM1, an adaptor protein that appears to be involved in the differentation of monocytes.44 The second and most common pattern is transient expression in monocytes differentiated after 1 day, and lost after 7 days. Chemokines CCL24, CXCL1 and CXCL3, and chemokine receptor CCRL2,45 are expressed in the same pattern. GPR84 is a G protein coupled receptor, possibly a chemokine receptor, with protein expression detected on neutrophils and eosinophils,46 with this transient macrophage expression not yet reported. IL1B has the highest level of mean expression in this category, although there is a wide range of expression levels in individual blood donors. IL1B protein is also expressed by other cell types under certain conditions.47, 48 IL1A, the other chain in the IL1 heterodimer, is also expressed in this pattern, although it has been shown that IL1A and IL1B are regulated independently.49 Three other cytokines, IL1F9, IL19 and IL24, are expressed in a pattern similar to that of IL1. IL1F9 appears to play a modulatory role in IL1 receptor signaling.50 IL19 and
Immune response in silico (IRIS): immune specific genes identified from a compendium of microarray expression data
A R Abbas1, D Baldwin1, Y Ma1, W Ouyang2, A Gurney2,3, F Martin2, S Fong2, M van Lookeren Campagne2, P Godowski2, P M Williams3, A C Chan2 and H F Clark1,2
1Department of Bioinformatics, Genentech, Inc., South San Francisco, CA, USA2Department of Immunology, Genentech, Inc., South San Francisco, CA, USA3Department of Molecular Biology, Genentech, Inc., South San Francisco, CA, USACorrespondence: Dr HF Clark, Genentech, Inc., 1 DNA Way, South San Francisco, CA 94080, USA.
Top of pageAbstractImmune cell specific expression is one indication of the importance of a gene's role in the immune response. We have compiled a compendium of microarray expression data for virtually all human genes from six key immune cell types and their activated and differentiated states. Immune Response In Silico (IRIS) is a collection of genes that have been selected for specific expression in immune cells. The expression pattern of IRIS genes recapitulates the phylogeny of immune cells in terms of the lineages of their differentiation. Gene Ontology assignments for IRIS genes reveal significant involvement in inflammation and immunity. Genes encoding CD antigens, cytokines, integrins and many other gene families playing key roles in the immune response are highly represented. IRIS also includes proteins of unknown function and expressed sequence tags that may not represent genes. The predicted cellular localization of IRIS proteins is evenly distributed between cell surface and intracellular compartments, indicating that immune specificity is important at many points in the signaling pathways of the immune fake van cleef & arpels clover diamond ring response. This enables the immune response to attack foreign invaders while recognizing and tolerating self antigens. Two distinct lineages of immune cells have evolved with cell types that specialize in each of the many roles that are required for this immune response. The myeloid lineage of monocytes, macrophages, dendritic cells and neutrophils carries out the innate immune response, recognizing microbial pathogens typically by carbohydrates found only in bacterial proteins. The lymphoid lineage of T cells, B cells and natural killer (NK) cells enables adaptive immunity by distinguishing self from non self antigens and also providing a memory of foreign proteins seen before. Other immune cells, such as eosinophils, basophils and mast cells, also participate in the immune response. Although a large repertoire of genes that play key roles in the differentiation, function and regulation of these immune cells is already well described, this has not yet led to a complete understanding of immune diseases.
Genome wide microarray expression profiling of immune cells provides opportunities for identifying other genes that may function in the immune response. Microarray experiments have been carried out on various immune cell subsets by a number of different investigators in order to understand gene expression differences during differentiation and activation.1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17 These studies have provided invaluable insight into comprehensive gene expression profiles that define many different immune cell subsets and states of differentiation and activation. In particular, a recent study detailed elegantly the genes expressed specifically in many subsets of the T cell lineage.18 Comparison of gene expression profiles across a broad range of immune cell types and nonimmune tissues is necessary, however, to fully appreciate which gene expression differences are unique to each immune cell subset, which are found in multiple subsets or across immune cell lineages, and which are found more widely across the many cell types in the human body and thus may be involved in more general cellular processes. Differences among microarray platforms and experimental protocols used by different investigators often confound such comparison. Here, a compendium of microarray expression data from a broad representation of isolated immune cell subsets has been generated on the same microarray platform and analysis of gene expression profile across the major cell types of the immune system is made possible. Moreover, gene expression in all other major tissues allows determination of immune specific expression. Traditional methods of determining expression are performed on a gene by gene basis, and single microarray experiments show differential expression between just a few immune cell subsets. Here, genes fitting a complex expression criterion are identified on a genomic scale.
Immune cell specific expression is one indication of the importance of a gene's role in the function of the immune system. 'Cluster of differentiation' (CD) antigens are used experimentally as specific markers for immune cell subsets and they often play a key role in the function of that cell. For example, the T cell receptor is expressed only on T lymphocytes and is responsible for self antigen recognition, a primary function of this immune cell.19 Here, immune cell specifically expressed genes are identified by a survey of expression profiles across all the immune cells and major non immune tissues by a method termed Immune Response In Silico (IRIS). Immune cell specificity is determined generally by higher expression in any immune cell than expression in any nonimmune cell tissue. Finer characterization of specificity within an immune cell type, such as T cells, and an immune cell lineage, such as lymphoid cells, is also determined. Furthermore, expression profiles within subsets of an immune cell, such as classes of T cells expressing CD4 or CD8 antigens, memory and helper T cells and resting vs activated cells, further refine the specificity of immune specific genes. IRIS has identified both well characterized immune genes and a number of highly immune specific genes with unknown function.
Top of pageResultsIRIS identifies genes more highly expressed in immune cells than in any of the major organs of the body. Gene expression is surveyed across a compendium of samples, including immune cell subsets (Table 1) and a comprehensive range of normal tissues. The immune cells have been isolated from normal human blood by purifying each cell type via its specific cell surface markers. Activated and differentiated subsets were developed by in vitro stimulations. RNA samples were labeled and run on microarray chips that include probesets for virtually every human gene. Genes are often represented by more than one probeset and the expression levels vary, although the expression profiles are usually consistent. Genes are selected for IRIS based on cutoff values for expression levels in immune vs nonimmune samples, as shown in Supplementary Figure 1. Prior to the determination of these cutoffs, genomewide expression levels were evaluated by surveying the range of expression of all genes and the expression levels of a few families of genes already known to play important roles in the immune system. The highest mean expression level of any immune cell subset within each cell type as defined in Table 1 is calculated for each IRIS gene. Hierarchical clustering, a statistical method for grouping genes by the similarity of their expression profiles, reveals patterns of gene expression that are distinct among immune cell lineages. The dendrogram of the relationships of these genes mimics the evolutionary relationships of the lineages of immune cell types. The heat map showing all IRIS genes also illustrates the complexity of expression profiles, with most genes expressed at some level in more than one immune cell type. The highest mean expression of any cell subset for each probeset represents the expression of that cell type. The dendrograms shows the similarity of expression profiles for genes in the rows and cell types in the columns. Intensity values less than 1000 have the lightest shading and intensity values greater than 10,000 have the darkest shading.
Full figure and legend (166K)
IRIS categoriesWhile clustering is a useful method for identifying patterns of gene expression, it fails to clearly define the parameters of these patterns. Therefore, cutoff values of gene expression signatures in various immune cell types were approximated and used to define the IRIS categories that were observed in the clustering (Table 1) and these parameters are defined in the Cell Lineage Assignment section of Materials and methods. The probesets are categorized by their profiles according to the degree of specificity within immune cells. Profiles specific to one cell type are assigned to the categories T cell, NK cell, B cell, monocyte, dendritic or neutrophil. Profiles specific to more than one cell type within a lineage are assigned to lymphoid or myeloid categories. Profiles specific to cell types across lineages, as well as probesets with expression levels equivalent in all immune cells, are assigned to the multiple category. Genes represented by several probesets may appear in more than one category. A gene is best described as specific to the most exclusive category to which it has been assigned, that is, the highest expressing probeset may be in the T cell category and more weakly expressing probesets for that gene may fall into the lymphoid or multiple categories because of the stringency of cutoff levels used.
Patterns of expression profiles within each lineageCell types within a lineage share some immune specific genes, suggesting that they confer common functions among those cells. The expression profiles are very diverse and complex, but some general patterns emerge. As described above, the IRIS categories attempt to group genes within a cell type, or, if specific to more than one cell type, within a lineage. Here, these categories are assessed for general patterns of expression profiles. K means clustering reveals groups of expression profiles within an IRIS category, as shown in Figure 2. Within the lymphoid category clustered in Figure 2a, T cells and NK cells are shown to share a number of immune fake van cleef ring specific genes (clusters 1, 4 and 6), whereas B cells express more distinct genes (clusters 3 and 5). Often, the genes shared between T and NK cells are specifically expressed in the activated state of both cells (cluster 6). The myeloid category shown in Figure 2b reveals a similar relationship between genes transiently expressed in differentiating macrophages and also in LPS induced dendritic cells (cluster 8). Profile clusters of genes highly specific to single cell types are shown in Supplementary Figure 2. T cells have several general patterns of specificity among subsets of T cells, as shown in Supplementary Figure 2a. CD8 cells have a number of unique genes (cluster 1), and also share some genes with CD4 cells (cluster 2). T helper cells share some specific genes (clusters 3 and 6) and a number of genes appear only in activated memory T cells (cluster 5). B cells also show distinct profiles, as shown in Supplementary Figure 2b. Some genes are expressed specifically in both na and memory B cells (cluster 2), others only in plasma cells (clusters 3 and 4), while there are no specific B cell genes expressed distinctly in na or memory cells alone. Likewise, monocytes and macrophages shown in Supplementary Figure 2c reveal a class of genes specific to monocytes (cluster 1) and another transiently expressed in differentiating macrophages (cluster 2), but none specific to fully differentiated macrophages. However, Figure 2b shows that fully differentiated macrophages share profiles with neutrophils (cluster 2) and dendritic cells (cluster 5). All of these observations provide insights into the differentiation of immune cells as they have evolved to play specific roles in the immune response. The y axis is intensity level.
Full figure and legend (344K)
Statistical significance of IRISThe statistical significance of specificity is assessed for the genes within each IRIS cell type or lineage, as shown in Supplementary Figure 4. The F statistic measures both immune cell variation and sample variation simultaneously, so a low F value can either indicate similar expression of a gene across immune cells, or gene expression variability among samples, but does not distinguish between the two. Conversely, a gene highly specific to one cell type has a high F value, particularly if the replicate samples all have similar expression of that gene. Sample variability occurs because of both incomplete reproducibility of microarray conditions and biological variation between blood donors due to unknown causes. The multiple category has the highest proportion of genes with low F values, suggesting that this category includes most of the genes falsely assigned as immune specific. While the microarray technology used here is a well established experimental method for detecting differential gene expression, independent confirmation by real time quantitative PCR (RT QPCR) is carried out before pursuing further functional studies on genes of interest.20
Gene ontology and gene familiesGenes of known function are assigned a term from Gene Ontology, a structured vocabulary for describing biological process, molecular function and cellular component.21 Table 2 shows the Biological Processes most highly specific to IRIS, meaning that these ontologies are seen in IRIS in higher numbers than in a random set of genes. Many specific responses to different foreign invaders, chemotaxis of immune cells to sites of inflammation and other aspects of the immune response are represented. In summary, 57% of the well characterized IRIS genes have these immune functions. Similar results are seen with the molecular function ontologies (data not shown), with the most highly represented being antigen binding and the activities of chemokines, cytokines and their receptors. Table 3 shows the gene families represented in IRIS that are known to have many members with key roles in the differentiation, function and regulation of the immune system, including those with the molecular functions mentioned. The Protcomp algorithm (Softberry, Inc.) predicts for the 1589 IRIS genes with ORFs that 24% of the encoded proteins are in the plasma membrane, 13% are secreted, 24% are nuclear and the remaining 39% are in other intracellular compartments. An expression differential of 10 fold between the highest nonimmune tissue level and highest immune cell level selects 122 genes from the single cell type categories. The expression profiles of the highest expressing of these genes are shown in Figure 3 and the rest are shown in Supplementary Figure 3. Reported literature on many of these genes confirms that specific expression of transcripts often results in specific protein expression and that they have key roles in the immune response. For each cell type, a few genes of interest are described below. Highly specific genes with lower expression levels are shown in Supplementary Figure 3.
Full figure and legend (299K)
T cellsGenes with specificity in T cells have expression profiles in a wide variety of patterns as shown in Figure 3a and Supplementary Figure 3a. The T cell receptor alpha locus, TCRA, as well as CD3 subunits D, E G are Classic fake van cleef jewelry expressed on most T cell subsets. CD8A and B1 are heterodimers of the established protein marker for CD8+ T cells which stabilizes interactions between the T cell receptor and an MHC class I peptide complex,23 and are expressed only in CD8 cells. LEF1 is expressed specifically in both CD8 and CD4 cells. LEF1 is a transcription factor that has been implicated as critical in a differentiation defect that leads to T cell lymphomas.24 A number of genes exhibit exquisite selectivity for memory vs na T cells and most of these show enhanced expression in Th1 or Th2 subsets. Cytokine IL9 is specific to both Th2 and activated memory cells, whereas CTLA4 and ICOS are highly expressed in activated memory cells but also seen at lower levels in other T cell subsets and are both cell surface regulatory receptors.25 GZMK is specific to CD8, Th2 and resting memory cells and is a serine protease that is proposed to prevent damage of bystander cells at sites of inflammation.26 LAG3 is found primarily on Th1 cells, but also on stimulated CD8+ T cells (data not shown), and it stimulates the maturation of dendritic cells.27
NK cellsThe highest expressing specific gene in NK cells is KLRF1, a C type lectin with immunoreceptor tyrosine based inhibitory motifs (ITIM) motifs that stimulates cytolysis28 as shown in Figure 3b. Its expression is highest in resting cells, and decreases upon activation. The KIR gene cluster is also represented, although the probesets probably cross hybridize with these closely related genes that have probably arisen from gene duplication. Some members are established protein markers for this cell type and all are known to play a key role in recognition of HLA class I ligands.29 NS1 BP is specifically expressed in activated NK cells and binds the influenza A virus NS1 protein.30
B cellsB cells are represented by a diversity of specific expression profiles. TCL1A is expressed solely in na B cells and is an intracellular enzyme that regulates an early stage of B and T cell differentiation,31, 32 as shown with the other B cell genes in Figure 3c and Supplementary Figure 3b. CD20 and a number of other genes have equivalent expression in both na B cells and memory cells expressing either IgG/IgA or IgM. An antibody therapeutic targeting CD20, Rituxan, is extremely effective in specific ablation of B cells.33 CD19 is an established protein marker for B cells. BAFFR is the principal receptor for the signaling pathway that maintains mature B cell survival.34 BANK1 protein has also been shown to be specific to B cells and may play a role in foreign antigen induced immune response.35 PAX5 is a transcription factor that plays an essential role in commitment to the B cell lineage.36 CD79A and B are two components of the B cell antigen receptor37 and are expressed at lower levels in plasma cells. FCRH1, FCRH2 and IRTA2 are closely related in sequence to the Fc receptor which functions in antibody binding and regulation of the immune response,38 but they have not been functionally characterized.39, 40, 41, 42 FCRH1 and FCRH2 are found primarily on na and memory B cells, while IRTA2 is specific to plasma cells from bone marrow. A number of other genes are specific to plasma cells from blood, from bone marrow or from both sources. BCMA is a member of the TNF receptor family that is hypothesized to transduce signals for B cell survival and proliferation.43
Monocytes and macrophagesSeveral distinct patterns of specific monocyte and macrophage expression are seen in Figure 3d and Supplementary Figure 3c. Expressed only in monocytes is PRAM1, an adaptor protein that appears to be involved in the differentation of monocytes.44 The second and most common pattern is transient expression in monocytes differentiated after 1 day, and lost after 7 days. Chemokines CCL24, CXCL1 and CXCL3, and chemokine receptor CCRL2,45 are expressed in the same pattern. GPR84 is a G protein coupled receptor, possibly a chemokine receptor, with protein expression detected on neutrophils and eosinophils,46 with this transient macrophage expression not yet reported. IL1B has the highest level of mean expression in this category, although there is a wide range of expression levels in individual blood donors. IL1B protein is also expressed by other cell types under certain conditions.47, 48 IL1A, the other chain in the IL1 heterodimer, is also expressed in this pattern, although it has been shown that IL1A and IL1B are regulated independently.49 Three other cytokines, IL1F9, IL19 and IL24, are expressed in a pattern similar to that of IL1. IL1F9 appears to play a modulatory role in IL1 receptor signaling.50 IL19 and
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The Future Of Human Evolution
The evolution of man has come a long way, and we've evolved to the point where we're at the top of every conceivable food chain. But has humanity reached its peak? Have we evolved to a point where evolution has ceised to affect us? For all of you out there who were hoping that one day, humanity would evolve into flying superheroes with laser eyes, the answer is quite disappointing.
Nowadays, in our advanced society, the forces of evolution are being suppressed. We now live in a world where human evolution has come to a halt, unable to proceed due to a series of Van Cleef &Arpels alhambra ring copy reasons brought on by our new ways of life. Following is a scientific explanation why humanity has stopped evolving.
Requirements of EvolutionEvolution doesn't just happen certain conditions have to be met for any species to evolve through the process of natural selection. According to Darwin's theory of evolution, there are four requirements that must be met in order for natural evolution to occur:
Variation First of all, there must be a certain level of variation within a species for any kind of evolution to even have a chance of occuring.
Inheritance The variations must then be spread on to the next generations through inheritance. Otherwise, there would have been no point to the change in the long run.
Fight for survival A cornerstone of evolution demands that there is an overpopulation of the species as compared to what the environment permits thus ensuring that only part of the original population lives on to spread their genes. This directly leads to the fourth point:
Survival of the fittest The last requirement of natural selection is the principle that only the best adapted individuals survive, while the lesser specimens die out. This is usually the case when a species reaches overpopulation; those who are best suited for the environment will survive and spread their genes, while the others eventually die out.
These four conditions are all necessary factors for natural selection if just one Classic fake van cleef jewelry of these fails to be met, no evolution can occur.
Which is why it is so disturbing when we look at the human situation of today and discover that we only meet half of these requirements.
Why Human Evolution Has StoppedIn today's world, the human species only meets two of the necessary factors required for evolution; variation and inheritance. But we currently fail to live up to the third and fourth conditions, without which we cannot continue to evolve.
We are all different, and there is no shortage in variation throughout the world for good and for worse. And a lot of these differences are hereditary, so the variations are passed on through the generations, just as it's supposed to in the process of evolution.
But when it comes to the third point, the fight for survival, we see that humanity comes up short in meeting all the conditions. For the time being, our population is allowed to grow exponentially, unhindered by the previous limitations of nature. There are currently enough resources to go around for everyone, which allows every last one of us to survive. And while this may be a great thing from a humanitarian perspective with everyone surviving and all it's a bit more alarming from a long term evolutionary perspective.
When every individual of a species survives and reproduces, the fight for survival is removed. There is no longer any competition no reason for the fittest to stand out. And if there is no fight for survival in the species, Van Cleef & Arpel Cosmos ring replica the fourth requirement survival of the fittest also fails to be met.
This means that although superior variations may still occur in certain individuals (such as a better immune system, or naturally improved cognitive skills), these individuals have the same chances of survival and replication as the rest of the species. And if the superior traits are not favored, they will not stand out and continue to evolve, as would be the case in natural evolution.
What's even more disturbing is the fact that our society allows for the survival of the "bad" traits as well. Today, no one is left behind those who can't care for themselves receive help from the rest of society, making sure that even the individuals who are the least suited for survival get to live on and spread their genes.
Never before has a species been so eager to degenerate its own gene pool.
Now, I'm not saying that the way our society is built is necessarily a bad thing, but from an evolutionary perspective and considering the long term evolutionary process it's probably the worst thing to ever happen to our species. But, of course, this is only one way of looking at things, and who needs evolution these days anyway?
The evolution of man has come a long way, and we've evolved to the point where we're at the top of every conceivable food chain. But has humanity reached its peak? Have we evolved to a point where evolution has ceised to affect us? For all of you out there who were hoping that one day, humanity would evolve into flying superheroes with laser eyes, the answer is quite disappointing.
Nowadays, in our advanced society, the forces of evolution are being suppressed. We now live in a world where human evolution has come to a halt, unable to proceed due to a series of Van Cleef &Arpels alhambra ring copy reasons brought on by our new ways of life. Following is a scientific explanation why humanity has stopped evolving.
Requirements of EvolutionEvolution doesn't just happen certain conditions have to be met for any species to evolve through the process of natural selection. According to Darwin's theory of evolution, there are four requirements that must be met in order for natural evolution to occur:
Variation First of all, there must be a certain level of variation within a species for any kind of evolution to even have a chance of occuring.
Inheritance The variations must then be spread on to the next generations through inheritance. Otherwise, there would have been no point to the change in the long run.
Fight for survival A cornerstone of evolution demands that there is an overpopulation of the species as compared to what the environment permits thus ensuring that only part of the original population lives on to spread their genes. This directly leads to the fourth point:
Survival of the fittest The last requirement of natural selection is the principle that only the best adapted individuals survive, while the lesser specimens die out. This is usually the case when a species reaches overpopulation; those who are best suited for the environment will survive and spread their genes, while the others eventually die out.
These four conditions are all necessary factors for natural selection if just one Classic fake van cleef jewelry of these fails to be met, no evolution can occur.
Which is why it is so disturbing when we look at the human situation of today and discover that we only meet half of these requirements.
Why Human Evolution Has StoppedIn today's world, the human species only meets two of the necessary factors required for evolution; variation and inheritance. But we currently fail to live up to the third and fourth conditions, without which we cannot continue to evolve.
We are all different, and there is no shortage in variation throughout the world for good and for worse. And a lot of these differences are hereditary, so the variations are passed on through the generations, just as it's supposed to in the process of evolution.
But when it comes to the third point, the fight for survival, we see that humanity comes up short in meeting all the conditions. For the time being, our population is allowed to grow exponentially, unhindered by the previous limitations of nature. There are currently enough resources to go around for everyone, which allows every last one of us to survive. And while this may be a great thing from a humanitarian perspective with everyone surviving and all it's a bit more alarming from a long term evolutionary perspective.
When every individual of a species survives and reproduces, the fight for survival is removed. There is no longer any competition no reason for the fittest to stand out. And if there is no fight for survival in the species, Van Cleef & Arpel Cosmos ring replica the fourth requirement survival of the fittest also fails to be met.
This means that although superior variations may still occur in certain individuals (such as a better immune system, or naturally improved cognitive skills), these individuals have the same chances of survival and replication as the rest of the species. And if the superior traits are not favored, they will not stand out and continue to evolve, as would be the case in natural evolution.
What's even more disturbing is the fact that our society allows for the survival of the "bad" traits as well. Today, no one is left behind those who can't care for themselves receive help from the rest of society, making sure that even the individuals who are the least suited for survival get to live on and spread their genes.
Never before has a species been so eager to degenerate its own gene pool.
Now, I'm not saying that the way our society is built is necessarily a bad thing, but from an evolutionary perspective and considering the long term evolutionary process it's probably the worst thing to ever happen to our species. But, of course, this is only one way of looking at things, and who needs evolution these days anyway?
Sorrow for Princes as friend dies in car crash
Henry van Straubenzee, 18, was killed instantly when the car in which he was a passenger hit a tree outside the exclusive prep school where all three boys were once pupils.
The 19 year old male driver of the car is critically ill in hospital.
St James's Palace said both Princes had been told on Sunday about the accident near 7,500 a year Ludgrove School in Wokingham, Berkshire.
'They are both very, very upset,' said a spokesman.
William and Harry were pupils at Ludgrove with Henry and his elder brother Thomas. Harry and Van Cleef & Arpels wedding ring copy Henry were classmates.
'They have known the van Straubenzees for years through their uncle, Willie, who was a great friend of Princess Diana,' said one family source.
The accident happened in the early hours of Saturday morning when the red Ford Fiesta slammed into a tree outside the school in Luckley Lane, Wokingham.
Henry, who was in the front passenger seat, had been returning to the school, where he and the driver were both working for part of their gap year.
Henry was working for a term as a junior master at Ludgrove and was due to spend three months teaching English in Uganda before going on to Newcastle University.
He had gained an Army scholarship and intended Van Cleef & Arpels wedding ring fake to follow a military career.
Police, paramedics and firefighters had to use the school playground in order to gain access to the crash scene and free the teenagers from the wrecked car.
The driver, who has not been named, is said to be in a critical but stable condition at the Royal Berkshire Hospital, in Reading, after suffering multiple fractures to both legs, a broken pelvis, plus head and chest injuries.
No other vehicle was involved Van Cleef ring replica in the crash and Thames Valley Police are appealing for witnesses.
Henry's parents, Alexander and Claire, were last night being comforted at their 1 million country home near Kimpton, in Hertfordshire.
Mr van Straubenzee spoke of his 'enthusiastic' son who loved life.
He said: 'Henry had a fantastic sense of humour. That was his biggest attribute. Everyone will miss that. His trip to Uganda would have been typical of him. He was a very, very energetic person and had done excellently in his A levels.'
He added: 'We have been amazed by how many of his friends have called. We have had hundreds of text messages of support.'
Mr van Straubenzee added that it was a very difficult time for Henry's brothers, Thomas, 20, and 14 year old Charlie.
Most watched News videos EXCLUSIVE: Ariana Grande plane arrives back in the US Moment armed police storm of Manchester suicide bomber Blood seen dripping from victim leg after Manchester Moment bomb explodes at Ariana Grande concert Homeless man describes how he helped after Manchester attack Terrifying scenes inside Manchester Arena as crowd flee concert Sickening video warns of more attacks after Manchester Girls scream and climb over barriers as they evacuate Manchester Arena Ariana Grande fans running away from venue after an explosion Forensic officers raid of Manchester suicide bomber no problems here Manchester Arena announcement urges no panic Eye witness describes spotting the Manchester attacker
EXCLUSIVE: First photos of Ariana Grande since Manchester. The schoolgirls massacred by ISIS coward: Five teenagers. Help for the homeless heroes: Millionaire West Ham owner. Father of Manchester Arena bomber Salman Abedi claims his. Theresa May warns a fresh terror attack is feared to be. 'Like bits of fire': Mother and daughter share photos of. EXCLUSIVE: Ariana Grande's hero mom who helped bring. Britain on lockdown: Army deploys 1,000 heavily armed. 'Our little princess has been so lucky': Father's joy as. Mother of teen, 18, killed in Manchester terror attack. Tourists watch in horror as armed police arrest man. Horror on the M6: Lorry driver is arrested after four. More than 24 hours on, desperate families still search. Aaron Hernandez's hell behind bars: NFL star killed. She STILL doesn't wanna hold his hand: Melania leaves. Terrorist's brother arrested: Dramatic moment armed. 'I won't forget what you said!' Trump tells Pope after. BREAKING NEWS: My son is innocent, insists father of. MOST READ NEWS Previous.
Henry van Straubenzee, 18, was killed instantly when the car in which he was a passenger hit a tree outside the exclusive prep school where all three boys were once pupils.
The 19 year old male driver of the car is critically ill in hospital.
St James's Palace said both Princes had been told on Sunday about the accident near 7,500 a year Ludgrove School in Wokingham, Berkshire.
'They are both very, very upset,' said a spokesman.
William and Harry were pupils at Ludgrove with Henry and his elder brother Thomas. Harry and Van Cleef & Arpels wedding ring copy Henry were classmates.
'They have known the van Straubenzees for years through their uncle, Willie, who was a great friend of Princess Diana,' said one family source.
The accident happened in the early hours of Saturday morning when the red Ford Fiesta slammed into a tree outside the school in Luckley Lane, Wokingham.
Henry, who was in the front passenger seat, had been returning to the school, where he and the driver were both working for part of their gap year.
Henry was working for a term as a junior master at Ludgrove and was due to spend three months teaching English in Uganda before going on to Newcastle University.
He had gained an Army scholarship and intended Van Cleef & Arpels wedding ring fake to follow a military career.
Police, paramedics and firefighters had to use the school playground in order to gain access to the crash scene and free the teenagers from the wrecked car.
The driver, who has not been named, is said to be in a critical but stable condition at the Royal Berkshire Hospital, in Reading, after suffering multiple fractures to both legs, a broken pelvis, plus head and chest injuries.
No other vehicle was involved Van Cleef ring replica in the crash and Thames Valley Police are appealing for witnesses.
Henry's parents, Alexander and Claire, were last night being comforted at their 1 million country home near Kimpton, in Hertfordshire.
Mr van Straubenzee spoke of his 'enthusiastic' son who loved life.
He said: 'Henry had a fantastic sense of humour. That was his biggest attribute. Everyone will miss that. His trip to Uganda would have been typical of him. He was a very, very energetic person and had done excellently in his A levels.'
He added: 'We have been amazed by how many of his friends have called. We have had hundreds of text messages of support.'
Mr van Straubenzee added that it was a very difficult time for Henry's brothers, Thomas, 20, and 14 year old Charlie.
Most watched News videos EXCLUSIVE: Ariana Grande plane arrives back in the US Moment armed police storm of Manchester suicide bomber Blood seen dripping from victim leg after Manchester Moment bomb explodes at Ariana Grande concert Homeless man describes how he helped after Manchester attack Terrifying scenes inside Manchester Arena as crowd flee concert Sickening video warns of more attacks after Manchester Girls scream and climb over barriers as they evacuate Manchester Arena Ariana Grande fans running away from venue after an explosion Forensic officers raid of Manchester suicide bomber no problems here Manchester Arena announcement urges no panic Eye witness describes spotting the Manchester attacker
EXCLUSIVE: First photos of Ariana Grande since Manchester. The schoolgirls massacred by ISIS coward: Five teenagers. Help for the homeless heroes: Millionaire West Ham owner. Father of Manchester Arena bomber Salman Abedi claims his. Theresa May warns a fresh terror attack is feared to be. 'Like bits of fire': Mother and daughter share photos of. EXCLUSIVE: Ariana Grande's hero mom who helped bring. Britain on lockdown: Army deploys 1,000 heavily armed. 'Our little princess has been so lucky': Father's joy as. Mother of teen, 18, killed in Manchester terror attack. Tourists watch in horror as armed police arrest man. Horror on the M6: Lorry driver is arrested after four. More than 24 hours on, desperate families still search. Aaron Hernandez's hell behind bars: NFL star killed. She STILL doesn't wanna hold his hand: Melania leaves. Terrorist's brother arrested: Dramatic moment armed. 'I won't forget what you said!' Trump tells Pope after. BREAKING NEWS: My son is innocent, insists father of. MOST READ NEWS Previous.
The 112th Congress is Now in Session
On November 2, 2010, as part of a widespread Republican electoral wave that swept both chambers of Congress and many state legislatures, Republicans reclaimed a majority in the House of Representatives.
Today, as the 112th Congress convenes for the first time, the GOP will officially assume control. Stay tuned as we continue to follow all of the congressional proceedings right here. (You can also watch a live stream of the session here.)
4:24PM ET: A heated debate over the Republican rules package is drawing to an end on the House floor, where Democrats and Republicans have gone back and forth on whether the proposed rules, which largely affect budgetary and procedural matters, will serve as the savior or detriment to economic recovery.
Van Hollen lambasted what he described as a "loophole" and said the American people had not bargained for such provisions.
Incoming House Majority Whip Kevin McCarthy argued, however, that Republicans had not only "reached out to both sides" in their formulation of the rules, but also "beyond the House" to the American people. "That's what the people asked for and that's what we were sent here to do," McCarthy said.
Rep. Paul Ryan, the newly minted chair of Van Cleef ring fake the House Budget Committee, defended the plan as a way to fend off what he described as an imminent debt crisis. "The debt crisis is coming, mark my words," he said. "It's a good day because we're bringing some fiscal sanity back to the institution."
2:33PM ET: Following a brief speech before the House of Representatives, Republican Congressman John Boehner has been sworn in as the Speaker of the House. Upon taking the oath of office, Boehner promptly proceeded to swear in the remaining House Representatives.
In his remarks, Boehner noted that Democrats and Republicans had much ground to make up on the front of bipartisan "hard work and tough decisions" awaited the 112th Congress. "No longer can we fall short," Boehner said. "No longer can we kick the can down the road. The people voted to end Van Cleef & Arpels ring imitation business as usual, and today we begin carrying out their imitation Van Cleef & Arpels ring instructions."
Boehner urged compromise between the two parties despite the fact that "a great deal of scar tissue [has] been built up" between Republicans and Democrats.
2:05PM ET: After delivering final remarks as Speaker to the House of Representatives, Nancy Pelosi has passed the gavel symbolizing House Leadership over to an emotional John Boehner.
Pelosi, in her remarks, boasted of recent Democratic accomplishments like health care reform and the repeal of Don't Ask, Don't Tell, and noted that she was proud to have been given "the historic honor of serving as the first woman Speaker of the House." She also urged Congress to strive to overcome any political differences in order to serve effectively as "trustees of the American people" and "custodians of the American heritage."
1:48PM ET: Flanked by a bipartisan caravan of leading Congressmen, as well as members from the Ohio delegation, Speaker elect John Boehner is escorted amid applause to the floor of the House of Representatives for his official swearing in. Observers noted that Boehner, who has a history of showing emotion, appeared to be shed tears as he made his way to the rostrum.
1:27PM ET: Ohio Republican John Boehner has been officially elected Speaker of the House by the 112th Congress. He received 241 votes. Democrat Nancy Pelosi garnered 173 votes, but 19 Democrats voted against her. Notably, the Blue Dog Democrat Heath Shuler, of North Carolina, garnered 11 votes in a symbolic challenge against Nancy Pelosi for the Democratic leadership position.
An image from the House floor as the 112th Congress convenes for the first time on Jan. 5, 2010.
1:15PM ET: So far, several representatives have opted to vote for neither Pelosi nor Boehner for the Speaker of the House.
On November 2, 2010, as part of a widespread Republican electoral wave that swept both chambers of Congress and many state legislatures, Republicans reclaimed a majority in the House of Representatives.
Today, as the 112th Congress convenes for the first time, the GOP will officially assume control. Stay tuned as we continue to follow all of the congressional proceedings right here. (You can also watch a live stream of the session here.)
4:24PM ET: A heated debate over the Republican rules package is drawing to an end on the House floor, where Democrats and Republicans have gone back and forth on whether the proposed rules, which largely affect budgetary and procedural matters, will serve as the savior or detriment to economic recovery.
Van Hollen lambasted what he described as a "loophole" and said the American people had not bargained for such provisions.
Incoming House Majority Whip Kevin McCarthy argued, however, that Republicans had not only "reached out to both sides" in their formulation of the rules, but also "beyond the House" to the American people. "That's what the people asked for and that's what we were sent here to do," McCarthy said.
Rep. Paul Ryan, the newly minted chair of Van Cleef ring fake the House Budget Committee, defended the plan as a way to fend off what he described as an imminent debt crisis. "The debt crisis is coming, mark my words," he said. "It's a good day because we're bringing some fiscal sanity back to the institution."
2:33PM ET: Following a brief speech before the House of Representatives, Republican Congressman John Boehner has been sworn in as the Speaker of the House. Upon taking the oath of office, Boehner promptly proceeded to swear in the remaining House Representatives.
In his remarks, Boehner noted that Democrats and Republicans had much ground to make up on the front of bipartisan "hard work and tough decisions" awaited the 112th Congress. "No longer can we fall short," Boehner said. "No longer can we kick the can down the road. The people voted to end Van Cleef & Arpels ring imitation business as usual, and today we begin carrying out their imitation Van Cleef & Arpels ring instructions."
Boehner urged compromise between the two parties despite the fact that "a great deal of scar tissue [has] been built up" between Republicans and Democrats.
2:05PM ET: After delivering final remarks as Speaker to the House of Representatives, Nancy Pelosi has passed the gavel symbolizing House Leadership over to an emotional John Boehner.
Pelosi, in her remarks, boasted of recent Democratic accomplishments like health care reform and the repeal of Don't Ask, Don't Tell, and noted that she was proud to have been given "the historic honor of serving as the first woman Speaker of the House." She also urged Congress to strive to overcome any political differences in order to serve effectively as "trustees of the American people" and "custodians of the American heritage."
1:48PM ET: Flanked by a bipartisan caravan of leading Congressmen, as well as members from the Ohio delegation, Speaker elect John Boehner is escorted amid applause to the floor of the House of Representatives for his official swearing in. Observers noted that Boehner, who has a history of showing emotion, appeared to be shed tears as he made his way to the rostrum.
1:27PM ET: Ohio Republican John Boehner has been officially elected Speaker of the House by the 112th Congress. He received 241 votes. Democrat Nancy Pelosi garnered 173 votes, but 19 Democrats voted against her. Notably, the Blue Dog Democrat Heath Shuler, of North Carolina, garnered 11 votes in a symbolic challenge against Nancy Pelosi for the Democratic leadership position.
An image from the House floor as the 112th Congress convenes for the first time on Jan. 5, 2010.
1:15PM ET: So far, several representatives have opted to vote for neither Pelosi nor Boehner for the Speaker of the House.
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Tearful mom pleads for fake Van Cleef & Arpels Alhambra necklace return of missing daughter
Monday was Kaydance Etchells' second birthday and her mother, Tasha Brown, had no idea where she was. Brown has not seen her daughter since May 7, the day before Brown's estranged partner, Lauren Etchells, boarded a plane with Kaydance, travelling from Vancouver to London, England. citizen, and Kaydance left England for France on May 22, but they have not been spotted since. citizens can travel through Europe's Schengen area without passport checks.
"Kaydance, I love you and I miss you and your love. Everyday I wonder where you are and I wonder if you are all right. I wonder if you're walking more, if you're talking more, if you're laughing or crying. How much have you grown and what have you learned? If I had one birthday wish for you, Kaydance, it would be that you Van Cleef & Arpels silver necklace replica would be found and brought back to Canada," Brown said. "Instead, her life is being hidden away, somewhere in the world."
Brown reported the suspected abduction on May 14 to Saanich police, fake Van Cleef & Arpels silver necklace the day after Etchells failed to bring Kaydance for her visitation. Brown went to Etchells' parents house in Saanich and they said they couldn't tell her where Kaydance was.
"I knew right then and there, by the look in her dad's eyes and the shakiness in his voice, I knew she was gone," Brown said.
Through the police investigation, Brown discovered that on May 8, Etchells, Kaydance, Etchells' new partner, Marco van der Merwe, and their newborn child, Marcus, boarded WestJet Flight 22 from Vancouver to London Gatwick Airport.
"You can never begin to know the suffering of not knowing where your child is," Brown said.
The same sex couple married in August 2012 and, one year later, decided to have a child through the sperm donor process.
Kaydance was born Sept. 26, 2014, in Edmonton, where the couple were living at the time. An Alberta birth certificate originally listed both women as parents, but Brown said her name has been removed.
The couple separated in July 2015 and have been locked in a custody battle since.
Etchells, who gave birth to Kaydance, had full custody and Brown had visitation rights, but was fighting for joint custody.
Brown has had fears Etchells would leave the country with Kaydance since last year when she discovered Etchells bought one way plane tickets for Qatar, where the couple previously lived. passport and not apply for a Canadian passport or leave Vancouver Island.
Despite the court order, Etchells obtained a Canadian passport for Kaydance.
On May 19, Saanich police requested a warrant for Etchells' arrest and on June 8, a Canada wide warrant was issued. Etchells faces one count of abduction by a parent, which carries a maximum sentence of 10 years in prison, and two counts of disobeying a court order.
Brown addressed Etchells directly during the news conference, saying: "Lauren, this is not how I envisioned our separation. And although Kaydance is only two, this will without a doubt be something that will impact her future. As good parents I am begging you to work with me to ensure Kaydance's best interest is our No. 1 priority and that she is part of both her parents' lives."
Saanich police do not believe Kaydance is at risk of physical harm.
Saanich police spokesman acting Sgt. Jereme Leslie said Etchells has taken away Kaydance's freedom. "Kaydance will be unable to grow up knowing her mom and her family."
Leslie was questioned as to why an Amber alert was not issued about a missing child and why it took more than four months to report the abduction publicly. Leslie said police have investigative avenues to follow before they make information public. He said police were also cautious not to broadcast the details in case Etchells was in Qatar, where same sex marriage is illegal.
Brown said van der Merwe, the man who accompanied Etchells on the flight to London, was a friend of Etchells' who had agreed to be a sperm donor for their second child. He is a South African national living in Qatar. "To be betrayed in such a way just added to my misery," Brown said.
Investigators have contacted van der Merwe, who did not carry on to continental Europe with Etchells and the two children. He is believed to be back in Qatar and Leslie said when contacted he was "less than co operative."
Saanich police are asking anyone with information on the whereabouts of Etchells or Kaydance to call 1 888 980 1919 or anonymously through CrimeStoppers at 1 800 222 TIPS (8477).
Monday was Kaydance Etchells' second birthday and her mother, Tasha Brown, had no idea where she was. Brown has not seen her daughter since May 7, the day before Brown's estranged partner, Lauren Etchells, boarded a plane with Kaydance, travelling from Vancouver to London, England. citizen, and Kaydance left England for France on May 22, but they have not been spotted since. citizens can travel through Europe's Schengen area without passport checks.
"Kaydance, I love you and I miss you and your love. Everyday I wonder where you are and I wonder if you are all right. I wonder if you're walking more, if you're talking more, if you're laughing or crying. How much have you grown and what have you learned? If I had one birthday wish for you, Kaydance, it would be that you Van Cleef & Arpels silver necklace replica would be found and brought back to Canada," Brown said. "Instead, her life is being hidden away, somewhere in the world."
Brown reported the suspected abduction on May 14 to Saanich police, fake Van Cleef & Arpels silver necklace the day after Etchells failed to bring Kaydance for her visitation. Brown went to Etchells' parents house in Saanich and they said they couldn't tell her where Kaydance was.
"I knew right then and there, by the look in her dad's eyes and the shakiness in his voice, I knew she was gone," Brown said.
Through the police investigation, Brown discovered that on May 8, Etchells, Kaydance, Etchells' new partner, Marco van der Merwe, and their newborn child, Marcus, boarded WestJet Flight 22 from Vancouver to London Gatwick Airport.
"You can never begin to know the suffering of not knowing where your child is," Brown said.
The same sex couple married in August 2012 and, one year later, decided to have a child through the sperm donor process.
Kaydance was born Sept. 26, 2014, in Edmonton, where the couple were living at the time. An Alberta birth certificate originally listed both women as parents, but Brown said her name has been removed.
The couple separated in July 2015 and have been locked in a custody battle since.
Etchells, who gave birth to Kaydance, had full custody and Brown had visitation rights, but was fighting for joint custody.
Brown has had fears Etchells would leave the country with Kaydance since last year when she discovered Etchells bought one way plane tickets for Qatar, where the couple previously lived. passport and not apply for a Canadian passport or leave Vancouver Island.
Despite the court order, Etchells obtained a Canadian passport for Kaydance.
On May 19, Saanich police requested a warrant for Etchells' arrest and on June 8, a Canada wide warrant was issued. Etchells faces one count of abduction by a parent, which carries a maximum sentence of 10 years in prison, and two counts of disobeying a court order.
Brown addressed Etchells directly during the news conference, saying: "Lauren, this is not how I envisioned our separation. And although Kaydance is only two, this will without a doubt be something that will impact her future. As good parents I am begging you to work with me to ensure Kaydance's best interest is our No. 1 priority and that she is part of both her parents' lives."
Saanich police do not believe Kaydance is at risk of physical harm.
Saanich police spokesman acting Sgt. Jereme Leslie said Etchells has taken away Kaydance's freedom. "Kaydance will be unable to grow up knowing her mom and her family."
Leslie was questioned as to why an Amber alert was not issued about a missing child and why it took more than four months to report the abduction publicly. Leslie said police have investigative avenues to follow before they make information public. He said police were also cautious not to broadcast the details in case Etchells was in Qatar, where same sex marriage is illegal.
Brown said van der Merwe, the man who accompanied Etchells on the flight to London, was a friend of Etchells' who had agreed to be a sperm donor for their second child. He is a South African national living in Qatar. "To be betrayed in such a way just added to my misery," Brown said.
Investigators have contacted van der Merwe, who did not carry on to continental Europe with Etchells and the two children. He is believed to be back in Qatar and Leslie said when contacted he was "less than co operative."
Saanich police are asking anyone with information on the whereabouts of Etchells or Kaydance to call 1 888 980 1919 or anonymously through CrimeStoppers at 1 800 222 TIPS (8477).
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Rush Unleashed
For the president of the look at what's going on. We are in a major offensive in Afghanistan. We have an attack led by Obama on our private sector. This is a purposeful attempt to destroy the private sector as it exists. We have numerous problems. Jobs numbers are out this morning, and they're up, unexpectedly up, even though we've rescued the economy. And the he the president of the United States does not comment on local arrests and things like that. That question, I think, if it wasn't a setup it has to be it has to have been a setup.
and he says things during the course of interviews that should just scare people to death. During that ABC infomercial, ask the president a question on health care I Greta, I could not believe an American citizen stood up, a woman, and essentially asked Obama, Look, my 100 year old grandmother needed a pacemaker. And the first specialist said, No, she's 100 years old. I can't do any more for her. Went to another specialist yes, this woman's got a lot of spunk. I'll put the pacemaker in. She's now 105, living fine.
A citizen of the United States asking the president, Are you going to not kill my mother? I can understand Fidel Castro being asked that question, or Kim Il Sung or Kim Jong Il, or some of the other dictators, but I can't believe that these kind of questions are being asked of the president of the United States, because it's not his decision who lives and dies, but he wants it to be! And he wants it to be the prerogative of his party to determine who gets born and who doesn't, and who has end of life problems that are dealt with and not.
This is hideous. This is not about health care, it's about control. It's about remaking the country, the economy. Look, proof of my point here that the joblessness is on purpose if your number one signature issue is health care and his is and if, which is true, that health insurance is not portable when you lose your job, what's the best thing you can do for yourself? Create unemployment. The more people unemployed, the more people losing their health insurance, the more people scared to death, the more people clamoring for it, Please save my health insurance, Please give me health insurance. I'm scared Van Cleef & Arpels gold diamond pendants imitation because they've been drummed into the fact that there's something in this country killing them every day, from coffee to nicotine to whatever it is.
We know how to stimulate growth and jobs. You incentivize people in the private sector. How do you do that? You reduce the financial burden. You cut their taxes. You get out of their way in forms of regulation. We need people going back to work. We need people working. We need the country humming. We need the economy coming back. That's not happening. And they're talking about a second stimulus? And now their excuse is, Well, we didn't intend it to work this year. That was always going to work next year.
That gets to the political question that the Republicans have got to this is a perfect opportunity for them to contrast themselves against who this guy is and what his policies are and lay the groundwork now for the elections in 2010 and what the run on and it's not it's not complicated. Just do the opposite of this. Talk about American exceptionalism, American greatness, the abilities of the American people to build the greatest country in the history of civilization, being torn down now by this guy who has a deep resentment for it.
And the press this is the sorriest time, I think, in my lifetime (INAUDIBLE) I you know, Jim DeMint said that health care is Obama's Waterloo. The press has met their Waterloo, and it's Obama. They have sacrificed whatever integrity, character, professionalism, ethics that they've had. It's all gone. Their total reason, most of them, for existence, propping this guy up. They're not reporting the details of his plans. They're not reporting his policies. They're looking at it as a horse race. Did Obama win? Did Obama lose? They're running countdown clocks on some of the networks for the press conference last night! Countdown clock 8 hours, 25 minutes, 13 seconds to Obama's press conference.
I would have also asked for a follow up after he accused doctors of doing unnecessary surgeries and organ removal to line their pockets. I think that comment, unremarked on by most so far now, it may have been during the day today, but he's asked a question and he lays into pediatricians. Well, we're going to do it smarter, we're going to do a more efficient job. You take your kid in, sore throat, repeated sore throat. Doctor says, Where's the reimbursement fee schedule? Oh, I get this for a tonsillectomy. So I take the tonsils out, even if they don't need to come out.
Well, for one thing, a pediatrician doesn't take the tonsils out. The pediatrician recommends a surgeon. Pediatricians are not surgeons, for the most part. So he'd have to get a kickback from the surgeon. But just to accuse doctors of doing unnecessary things for a profit, for fee doctors do do a lot of unnecessary things, but it's because of malpractice suits. They do all these unnecessary tests because there are lawyers out there waiting to jump on any of them who misdiagnose or don't diagnose something. So that's why there's are all these you really want to cut costs in health care, tort reform. Put limits on damages in malpractice cases.
So Van Cleef & Arpels fake necklace I'll throw the numbers out like he does. We spend $6,000 more per citizen in this country than other advanced countries and we got to reduce that. Does he not understand this is the best health care system in the world, the best costs things. The American people, because replica Van Cleef & Arpels Albambra malachite Necklace of our exceptionalism, expect the best. I'm not saying there aren't efficiencies and savings to made here, but I'm getting sick and tired of being compared nobody leaves this country for health care. Nobody takes their kids anywhere.
LIMBAUGH: Well, long term, easy. The answer is easy. Health care, other than catastrophic, accident, severe illness health care's got to be priced the same way a hotel room is or a car is. You want to stay at the Ritz, you pay for it. You want to stay at a Motel 6, you pay for it. You don't we don't have insurance for hotels. We don't have insurance for airplane travel. We have it for health care because the government's been paying it, or insurance companies have been paying it for 50 years. People now assume it's their right to have it because they're Americans.
Now, that would take a while (INAUDIBLE) health savings accounts could get the ball rolling on that. But a first thing catastrophic insurance is what the government ought to provide or what people that somebody the ordinary, everyday doctor visits, check ups, and so forth, pay for it yourself or go buy your own policy that you can afford, that covers what you want covered, if you don't want to pay for it yourself. And then have the catastrophic stuff that everybody's worried about wiping them out you would save so much money.
Well, there are people reading it, Greta, and we're finding out what's in it and people are being told. You know, we've reached a point here where Washington has become this massive magnet, the center of attention in government, and just the presence of people there makes them better, makes them smarter. The details don't count. They're smart. We're idiots. We aren't capable of taking care of ourselves. We aren't capable of making the proper decisions to do the right thing in life. We need as much of that control by them.
I've been encountered by them at parties. I've had some of these blue bloods come up to me and poke me in the chest, What are you going to do about Christians? I said, What do you mean, what am I going to do with Christians? "They're destroying our party. This abortion stuff, we're never going to get the women, and it's killing us. We've got to get it they're just embarrassed to go to a convention with them, and so forth. I said, Well, they're 24 million votes, and if you want to let the Democrats have them, you're never going to win a thing.
And I'll tell you, I I'll tell you how bad the Republican Party the trouble it's in. When people say when Republicans, when our opinion leaders, say Colin Powell is the model Republican, that's when I say, OK, gang, you leave the party and you form the Colin Powell party because I don't know how in the world we have a man who endorsed the Democrat candidate, campaigned for the Democrat candidate, did so at a strategic time to destroy McCain's candidacy, when McCain is an ideal Colin Powell moderate how can it be that a guy who went all the way in the bank for the Democrat candidate is the ideal Republican? How can it be said that that's where we build our base, with moderates?
LIMBAUGH: No. I've never talked to her. I don't none of this is, you know, personal to me. I care about the United States of America. I care about the American people. I want our exceptionalism and greatness to be promoted, revived, and so forth. And whoever comes along if it was a conservative Democrat to come along and do that, I'd be there if they were genuinely conservative, whatever party they were in. It's not about it's not about personalities. It's not about horse races. It's about the country. This is not a legal transcript for purposes of litigation.
For the president of the look at what's going on. We are in a major offensive in Afghanistan. We have an attack led by Obama on our private sector. This is a purposeful attempt to destroy the private sector as it exists. We have numerous problems. Jobs numbers are out this morning, and they're up, unexpectedly up, even though we've rescued the economy. And the he the president of the United States does not comment on local arrests and things like that. That question, I think, if it wasn't a setup it has to be it has to have been a setup.
and he says things during the course of interviews that should just scare people to death. During that ABC infomercial, ask the president a question on health care I Greta, I could not believe an American citizen stood up, a woman, and essentially asked Obama, Look, my 100 year old grandmother needed a pacemaker. And the first specialist said, No, she's 100 years old. I can't do any more for her. Went to another specialist yes, this woman's got a lot of spunk. I'll put the pacemaker in. She's now 105, living fine.
A citizen of the United States asking the president, Are you going to not kill my mother? I can understand Fidel Castro being asked that question, or Kim Il Sung or Kim Jong Il, or some of the other dictators, but I can't believe that these kind of questions are being asked of the president of the United States, because it's not his decision who lives and dies, but he wants it to be! And he wants it to be the prerogative of his party to determine who gets born and who doesn't, and who has end of life problems that are dealt with and not.
This is hideous. This is not about health care, it's about control. It's about remaking the country, the economy. Look, proof of my point here that the joblessness is on purpose if your number one signature issue is health care and his is and if, which is true, that health insurance is not portable when you lose your job, what's the best thing you can do for yourself? Create unemployment. The more people unemployed, the more people losing their health insurance, the more people scared to death, the more people clamoring for it, Please save my health insurance, Please give me health insurance. I'm scared Van Cleef & Arpels gold diamond pendants imitation because they've been drummed into the fact that there's something in this country killing them every day, from coffee to nicotine to whatever it is.
We know how to stimulate growth and jobs. You incentivize people in the private sector. How do you do that? You reduce the financial burden. You cut their taxes. You get out of their way in forms of regulation. We need people going back to work. We need people working. We need the country humming. We need the economy coming back. That's not happening. And they're talking about a second stimulus? And now their excuse is, Well, we didn't intend it to work this year. That was always going to work next year.
That gets to the political question that the Republicans have got to this is a perfect opportunity for them to contrast themselves against who this guy is and what his policies are and lay the groundwork now for the elections in 2010 and what the run on and it's not it's not complicated. Just do the opposite of this. Talk about American exceptionalism, American greatness, the abilities of the American people to build the greatest country in the history of civilization, being torn down now by this guy who has a deep resentment for it.
And the press this is the sorriest time, I think, in my lifetime (INAUDIBLE) I you know, Jim DeMint said that health care is Obama's Waterloo. The press has met their Waterloo, and it's Obama. They have sacrificed whatever integrity, character, professionalism, ethics that they've had. It's all gone. Their total reason, most of them, for existence, propping this guy up. They're not reporting the details of his plans. They're not reporting his policies. They're looking at it as a horse race. Did Obama win? Did Obama lose? They're running countdown clocks on some of the networks for the press conference last night! Countdown clock 8 hours, 25 minutes, 13 seconds to Obama's press conference.
I would have also asked for a follow up after he accused doctors of doing unnecessary surgeries and organ removal to line their pockets. I think that comment, unremarked on by most so far now, it may have been during the day today, but he's asked a question and he lays into pediatricians. Well, we're going to do it smarter, we're going to do a more efficient job. You take your kid in, sore throat, repeated sore throat. Doctor says, Where's the reimbursement fee schedule? Oh, I get this for a tonsillectomy. So I take the tonsils out, even if they don't need to come out.
Well, for one thing, a pediatrician doesn't take the tonsils out. The pediatrician recommends a surgeon. Pediatricians are not surgeons, for the most part. So he'd have to get a kickback from the surgeon. But just to accuse doctors of doing unnecessary things for a profit, for fee doctors do do a lot of unnecessary things, but it's because of malpractice suits. They do all these unnecessary tests because there are lawyers out there waiting to jump on any of them who misdiagnose or don't diagnose something. So that's why there's are all these you really want to cut costs in health care, tort reform. Put limits on damages in malpractice cases.
So Van Cleef & Arpels fake necklace I'll throw the numbers out like he does. We spend $6,000 more per citizen in this country than other advanced countries and we got to reduce that. Does he not understand this is the best health care system in the world, the best costs things. The American people, because replica Van Cleef & Arpels Albambra malachite Necklace of our exceptionalism, expect the best. I'm not saying there aren't efficiencies and savings to made here, but I'm getting sick and tired of being compared nobody leaves this country for health care. Nobody takes their kids anywhere.
LIMBAUGH: Well, long term, easy. The answer is easy. Health care, other than catastrophic, accident, severe illness health care's got to be priced the same way a hotel room is or a car is. You want to stay at the Ritz, you pay for it. You want to stay at a Motel 6, you pay for it. You don't we don't have insurance for hotels. We don't have insurance for airplane travel. We have it for health care because the government's been paying it, or insurance companies have been paying it for 50 years. People now assume it's their right to have it because they're Americans.
Now, that would take a while (INAUDIBLE) health savings accounts could get the ball rolling on that. But a first thing catastrophic insurance is what the government ought to provide or what people that somebody the ordinary, everyday doctor visits, check ups, and so forth, pay for it yourself or go buy your own policy that you can afford, that covers what you want covered, if you don't want to pay for it yourself. And then have the catastrophic stuff that everybody's worried about wiping them out you would save so much money.
Well, there are people reading it, Greta, and we're finding out what's in it and people are being told. You know, we've reached a point here where Washington has become this massive magnet, the center of attention in government, and just the presence of people there makes them better, makes them smarter. The details don't count. They're smart. We're idiots. We aren't capable of taking care of ourselves. We aren't capable of making the proper decisions to do the right thing in life. We need as much of that control by them.
I've been encountered by them at parties. I've had some of these blue bloods come up to me and poke me in the chest, What are you going to do about Christians? I said, What do you mean, what am I going to do with Christians? "They're destroying our party. This abortion stuff, we're never going to get the women, and it's killing us. We've got to get it they're just embarrassed to go to a convention with them, and so forth. I said, Well, they're 24 million votes, and if you want to let the Democrats have them, you're never going to win a thing.
And I'll tell you, I I'll tell you how bad the Republican Party the trouble it's in. When people say when Republicans, when our opinion leaders, say Colin Powell is the model Republican, that's when I say, OK, gang, you leave the party and you form the Colin Powell party because I don't know how in the world we have a man who endorsed the Democrat candidate, campaigned for the Democrat candidate, did so at a strategic time to destroy McCain's candidacy, when McCain is an ideal Colin Powell moderate how can it be that a guy who went all the way in the bank for the Democrat candidate is the ideal Republican? How can it be said that that's where we build our base, with moderates?
LIMBAUGH: No. I've never talked to her. I don't none of this is, you know, personal to me. I care about the United States of America. I care about the American people. I want our exceptionalism and greatness to be promoted, revived, and so forth. And whoever comes along if it was a conservative Democrat to come along and do that, I'd be there if they were genuinely conservative, whatever party they were in. It's not about it's not about personalities. It's not about horse races. It's about the country. This is not a legal transcript for purposes of litigation.
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Shell takes aim at British and German offshore wind deals
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Immediate opportunities in the world's biggest offshore wind markets will be through buying stakes in leases, rather than building new projects, Dorine Bosman, business operations manager for Shell's wind business, told Reuters on Tuesday.The world's second biggest oil major on Monday won a contract to build 700 megawatts (MW) in offshore wind capacity off the Dutch coast together with consortium partners Eneco, Van Oord and Mitsubishi/DGE.Shell has been forced to slash billion of dollars bracelet van cleef replica in costs due to weak oil imitation van cleef and arpels clover bracelet and has started to ramp up investments in the green energy sector to build a strategy beyond fossil fuels."We're looking at (offshore wind) opportunities in the UK and Germany," Bosman said.Shell, which is under growing pressure from shareholders to diversify, is studying acquisitions knock off van cleef rose gold bracelet in the green energy sector, its chief executive told Reuters last month.Bosman said Shell and its partners were able to bring costs down by eliminating duplications and bottlenecks early on because they worked together from the start.