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Aston Villa's players are a disgrace... you would not sign any of them
- Villa have been a disgrace and that 6-0 defeat to Liverpool was as bad a performance, in every aspect, as I have ever seen
- The Derby side of 2007-08 set the benchmark for the worst we have seen but, with seven games to go, Villa are in danger of wrestling that tag
- Six years ago they finished sixth under Martin O'Neill with 64 points, sandwiched between Manchester City and Liverpool
- Since his abrupt departure, five further managers have come and gone but the problems have spiralled
- Villa have got the biggest problem you can find in football which is never looking like scoring (their current total is 22) and conceding too many (58)
- To make matters worse, they have got poor players with a shocking attitude, who have failed to show any self-awareness
- Read our Premier League betting guide for this weekend
The worst team ever to play in the Premier League: those were the words I could not stop myself saying on February 14.
I'd just watched Liverpool tear Aston Villa to shreds and the 6-0 final score in no way flattered my old club. The home team were appalling and my instinct was to make that damning assessment. Immediately, though, plenty of people got in contact on social media to tell me I was wrong.
The same two examples kept being offered to say I'd been overly harsh on Aston Villa. The first one was Derby County, who finished rock bottom in 2007-08; the second was the Sunderland team of 2005-06 that was relegated with 15 points.
Derby hold the unwanted records of scoring the fewest goals (20) and conceding the most (89) in a 38-game season, not to mention the lowest points total (11). They set the benchmark for the worst we have seen but, with seven games to go, Aston Villa are in danger of wrestling that tag away from them.
Aston Villa's players have failed to show any fight this season, with Harry Kane scoring here
Daniel Sturridge opens the scoring during Liverpool's 6-0 rout against the Premier League's bottom side
Villa are bottom of the Premier League and without a manager after a woeful campaign
Everything I've seen since that capitulation against Liverpool makes me think I was right to be so critical. My adrenaline isn't running the way it would in a television studio after a live game but I stand by my statement. Here we are on April 2 and Villa haven't added to their points tally.
Sunderland, remember, were historically a yo-yo club. The squad they had 10 years ago had also only just won promotion and they signed players who were Championship standard.
Derby, meanwhile, were not ready to be in the Premier League. They came up via the play-offs with a squad of thirtysomethings; 12 months before they were promoted, they had finished 20th in the Championship, so it shows how big a leap they had taken.
But Villa? Six years ago they finished sixth under Martin O'Neill with 64 points, sandwiched between Manchester City and Liverpool. Since his abrupt departure, five further managers have come and gone but the problems have spiralled.
Normally when a side go down, three or four get cherry-picked and the same number are brought in with Championship experience but, given the chance, who would you want your club to buy from this Villa squad? My answer would be nobody.
There will be arguments about them losing Christian Benteke and Fabian Delph last summer, along with Tom Cleverley and Ron Vlaar. In total, Villa received more than £44million in transfer fees but they still spent £52m on players who have been totally inadequate.
The Sunderland team of 2005-06 went down to the Championship with just 15 points to their name
The Derby side of 2007-08 won once, amassed 11 points and went a record 32 games without a victory
Look at Southampton. They have lost a succession of top performers but they have invested well. Yes, there has been a lack of investment from owner Randy Lerner but, other than Benteke, can you name a good player Villa have signed in the past five years?
Since Liverpool demolished them six weeks ago, there has been no positive reaction. Five games have produced five defeats. Not a single point.
The disastrous sequence culminated with Remi Garde losing his job on Tuesday. The decision to sack him will have no influence on whether they stay up.
Garde has been heavily criticised with some justification, as too has Lerner. Garde looked totally ill-equipped to deal with a dogfight from the start and his quiet ways never looked like inspiring a group that Tim Sherwood had failed to connect with.
In fairness to Sherwood, he galvanised the players last year to keep them up. When he was sacked last October, why did they not go for the obvious candidate in Nigel Pearson, who was available? Had Sherwood gone 16 days earlier, Villa could have had Sam Allardyce. Yet for all the fury being aimed at Garde and Lerner, why is the squad getting off scot-free?
Remi Garde took over in November and never looked like saving the club from relegation
Tim Sherwood galvanised the club with his skills of motivation but was replaced by Garde in November
They have been a disgrace and that 6-0 defeat to Liverpool was as bad a performance, in every aspect, as I have ever seen.
The least you can do in such a position is run and tackle but they won't even do that — their average distance per game (66 miles) is the worst in the Premier League.
Villa have got the biggest problem you can find in football which is never looking like scoring (their current total is 22) and conceding too many (58) but, to make matters worse, they have got poor players with a shocking attitude, who have failed to show any self-awareness.
At least with the appointment of Brian Little at board level this week, moves are being made to correct that.
I wrote about Jack Grealish last November, when he'd been out partying in Manchester after a 4-0 defeat at Everton. Technically, he'd done nothing wrong but he gave out the wrong signals at that time.
You would think people would learn but now Gabby Agbonlahor has had a similar incident.
Again, Agbonlahor, who has been suspended, had permission to be in Dubai with club captain Micah Richards during the international break but, as someone who has been at Villa Park for his entire career, did he not realise how fans would react to seeing him photographed in a nightclub? It was embarrassing.
Gabby Agbonlahor (centre)was pictured in a Dubai nightclub with Dwight Gayle and Frazier Campbell
Jack Grealish was pictured in a nightclub in November following a 4-0 drubbing at the hands of Everton
Relegation has been inevitable since autumn but that won't lessen its impact.
Villa — one of only seven teams to keep their place in the Premier League since 1992 — will be the highest profile club to lose their top-flight status in the modern era, bigger than Leeds (2004) and Newcastle (2009).
Why? In my eyes, Villa are the biggest club in the Midlands. In terms of history, the trophies they have won — not least the 1982 European Cup — and their stadium, they outrank both Leeds and Newcastle.
When they do go, there is little chance of them coming back up with that squad. In fact, I would say at this stage that another relegation, rather than promotion, would be on the cards. How will a group who can't run and won't run cope with 46 ferocious games?
What makes their plight so bad is the fact this has been the season when strange things have happened and all clubs have been taking points off each other.
All clubs, that is, bar Aston Villa.
Cruyff reigns supreme
There is always a debate raging about who is the greatest player we have seen or who is the best manager. As much as we crave it, there will never be a definitive answer.
I have tended to look at things another way, though, and looked for who has been the greatest for football in general. My answer every time would be Johan Cruyff.
Pele and Diego Maradona may have been considered a shade better as individual players than Cruyff but neither could come close to him in terms of what he did as a manager with Ajax and Barcelona.
Franz Beckenbauer, perhaps, has outstanding credentials, given he won the World Cup as a player and a manager, but what sets Cruyff apart is his legacy. Ajax and Holland were nothing, really, before he started to shine for them and his ideas still glow in the glorious Barcelona team we see today.
Johan Cruyff won eight Dutch titles with Ajax over the course of two spells with the club
Johan Cruyff (right) helped Holland beat Uruguay 2-0 in the opening game of the 1974 World Cup finals
Ex-players are forever accused of living in the past but the great thing about Cruyff, regardless of what he had achieved and how good he had been, was the fact he was always looking to see how the game as a whole could be moved on.
I loved Cruyff. I loved the way he played, I loved the way his teams played and I loved listening to his words. His standout quote for me was when he said: 'Quality without results is pointless. Results without quality is boring.'
During the 2006 World Cup, I had the honour of meeting him — he stayed at the hotel where the England wives were based in Baden-Baden — and it was an ambition of mine to interview him. We tried to set it up several times in recent years but circumstances never allowed it to happen.
It would have been an absolute privilege to have an audience with Cruyff. I don't think we will ever see another like him.
Man of the week: Dimitri Payet
Another game, another stunning moment from West Ham's maestro — his goal against Russia for France in midweek was sensational.
As West Ham enter the defining period of the season, when they have the potential to qualify for Europe and possibly win a first piece of silverware since 1980, it will be hugely reassuring to Slaven Bilic that Dimitri Payet is in such sparkling form.
There is no doubt he will be on the shortlist for the PFA Player of the Year award. He and Harry Kane are the two candidates most likely to deny Leicester's Riyad Mahrez and Jamie Vardy, and it has been a joy to watch Payet play this season.
Dimitri Payet restored's France's two-goal cushion against Russia with a stunning free-kick
Payet (centre) is back with West Ham, who welcome Crystal Palace to the Boleyn Ground on Saturday
He has set the Premier League alight and after that majestic goal in the Stade de France, there is every chance he will do the same for the hosts when Euro 2016 begins in June. Didier Deschamps, the France coach, is finding it impossible to resist his claims.
Payet's form also shows the strength of the Premier League.
It says everything for the power our clubs have that a club who have not traditionally been in the fight for the top four have a player who is so important to one of the major nation's ambitions.
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Visual Overview
Despite a substantial appreciation for the critical role of macrophages in cardiometabolic diseases, understanding of human macrophage biology has been hampered by the lack of reliable and scalable models for cellular and genetic studies. Human induced pluripotent stem cell (iPSC)–derived macrophages (IPSDM), as an unlimited source of subject genotype-specific cells, will undoubtedly play an important role in advancing our understanding of the role of macrophages in human diseases. In this review, we summarize current literature in the differentiation and characterization of IPSDM at phenotypic, functional, and transcriptomic levels. We emphasize the progress in differentiating iPSC to tissue resident macrophages, and in understanding the ontogeny of in vitro differentiated IPSDM that resembles primitive hematopoiesis, rather than adult definitive hematopoiesis. We review the application of IPSDM in modeling both Mendelian genetic disorders and host–pathogen interactions. Finally, we highlighted the potential areas of research using IPSDM in functional validation of coronary artery disease loci in genome-wide association studies, functional genomic analyses, drug testing, and cell therapeutics in cardiovascular diseases.
- coronary artery disease
- genome-wide association study
- hematopoiesis
- induced pluripotent stem cells
- macrophages
Macrophages represent a critical cell type in the pathogenesis of chronic inflammatory cardiometabolic diseases, such as atherosclerosis, central obesity, and insulin resistance.1 The differentiation of human induced pluripotent stem cells (iPSCs) to macrophages (iPSC-derived macrophages [IPSDM]) offers a powerful platform to study human macrophage biology with enormous potential in disease modeling, functional genomic analyses, drug screening, and cell therapeutics. The goal of this brief review is to provide a concise but comprehensive overview of new advances in the differentiation and application of IPSDM, with a major focus on cardiovascular research, while also highlighting cutting edge applications across the vast landscape of macrophage biology in health and diseases (Figure 1).
Please see http://atvb.ahajournals.org/content/atvb-focus for all articles published in this series.
IPSDM Provide an Important Tool to Study Human Macrophage Biology
Primary murine macrophages, despite being widely used as a powerful model in the study of macrophage biology, have fundamental interspecies differences compared with human macrophages.2,3 Functionally, humans are physiologically more sensitive than mice to lipopolysaccharide stimulation.2 Human monocyte-derived macrophages (HMDM) have human-specific TLR4 target genes, for example, CXCL13,4 and relative to murine macrophages slower induction of several negative feedbacks of the toll-like receptor pathway.3 In HMDM, pattern recognition receptor stimulation drives release of endogenous ATP that triggers inflammasome activation,5 but murine macrophage inflammasome activation requires cholesterol crystals.6 Further, compared with mouse, acetylated low-density lipoprotein loading in HMDM induces much greater triglyceride enrichment yet many-fold lower levels of cholesteryl ester (≈30 versus >150 μg/mg protein).7 Transcriptomically, compared with mRNA, noncoding RNAs are mostly not conserved in mice, highlighting the need to study human macrophage-specific noncoding RNAs in human macrophages.
Experimental human macrophages are derived mostly from 2 sources: tumor-derived cell lines (eg, U937, THP-1 cells) and primary cells, including isolated tissue resident macrophages such as microglia and alveolar macrophages, and HMDM. THP-1 derived macrophages (THP-1Φ) have unlimited replicative potential but are karyotypically abnormal and functionally immature.8 Tissue macrophages, especially those from healthy subjects, have extremely limited availability. HMDM are relatively easy to obtain, but are difficult to be genetically manipulated, and do not self-renew limiting scalability. IPSDM provides a genotype-specific, scalable, and renewable source of human macrophage8 that are amenable to genetic manipulation by clustered regularly interspaced short palindromic repeats/clustered regularly interspaced short palindromic repeat–associated 9 (CRISPR/Cas9)–mediated gene editing. With continual improvement in differentiation efficiency and fidelity, IPSDM is emerging as a powerful tool in multiple areas of research for disease modeling and more recently, drug testing and cell therapeutics.
Generation, Functional Feature and Molecular Profiling of IPSDM
Multiple protocols have been described to differentiate iPSC into macrophages. The first 2 IPSDM differentiation protocols published by Choi et al9 and Senju et al10 in 2011 used coculture of iPSC with OP9 mouse stromal cell monolayers to induce hematopoietic differentiation, followed by expansion of multipotent myeloid progenitors and differentiation into macrophages with GM-CSF (granulocyte macrophage colony-stimulating factor)9 or M-CSF (macrophage colony-stimulating factor).9,10 In 2013, modified from a protocol published by Karlsson et al11 for human embryonic stem cell to macrophage differentiation, van Wilgenburg et al12 established an embryoid bodies (EBs)-based protocol for IPSDM differentiation in which hematopoiesis was induced under serum-free condition followed by directed differentiation to macrophages with M-CSF either in serum-free media or in media with serum. Yanagimachi et al13 developed a monolayer-based protocol by culturing iPSC on a layer of extracellular matrix component under serum-free condition and differentiated floating monocyte-like cells to macrophage in media with serum and M-CSF. In 2015, our group8 published an EB-based differentiation protocol and for the first time performed transcriptomic characterization using deep RNA-sequencing in HMDM and IPSDM lines derived from the same subjects. The results highlighted the transcriptome similarity between HMDM and IPSDM with a small percentage of genes (≈12%) differentially expressed.8 Further, more sophisticated functional characteristics, such as cholesterol efflux, cholesteryl ester hydrolysis,14 and cytokine secretion profile in macrophages with M1 (lipopolysaccharide+interferon-γ) and M2 (interleukin-4) activation, were compared between multiple HMDM and IPSDM lines and have demonstrated remarkable similarity.8 Two additional studies from our group further elaborated that IPSDM recapitulate important alternative splicing events15 and long noncoding RNA profiles (article under review) of HMDM during macrophage activation, identifying IPSDM as uniquely suited to study human macrophage-specific transcriptome regulation. These protocols8,12,13 used sequential exposure of iPSC to different cytokines that resemble hematopoietic specification and myeloid expansion, and myelomonocytic cells were harvested for directed differentiation to macrophages with M-CSF. The differentiation efficiency was evaluated differently across studies, but generally these protocols have demonstrated high efficiency and scalability with >10 million IPSDM generated from 1 6-well plate of cultured iPSCs.8,12
Although IPSDM have demonstrated similar phenotypic, functional, and transcriptomic characteristics to HMDM, their ontogeny and identity have not been fully defined and this recently has received further study. It has been recognized that in vitro hematopoietic differentiation of PSCs resembles in vivo primitive hematopoiesis rather than definitive hematopoiesis.16 Vanhee et al16 used a reporter human embryonic stem cell line with MYB-eGFP, a marker for hematopoietic stem cell-dependent hematopoiesis, and confirmed the absence of in vitro generation of MYB+ hematopoietic stem cell during EB-based IPSDM differentiation. The results were further validated in human iPSCs,17 showing that knockout of MYB in iPSC by CRISPR/Cas9 did not impact IPSDM differentiation.12 In contrast, when SPI1 and RUNX1, important transcription factors in yolk sac hematopoiesis, were knocked out, iPSCs were unable to produce CD14+ macrophages. Thus, IPSDM may developmentally relate to and be a good model for MYB-independent tissue resident macrophages. Indeed, several iPSC to microglia-like cell differentiation protocols were published this year using cytokines representing the main driver of microglia lineage,18,19 coculture with iPSC-derived neurons20,21 and astrocytes,22 or incubating with cytokines derived from those cell types23 to recapitulate organ-specific cues. Takata et al21 has established a protocol that specifically resembles primitive hematopoiesis and yolk sac macrophages for further differentiation into tissue macrophage-like cells by coculture or engraftment. The iPSC-derived microglia-like cells showed transcriptome profile highly similar to that of human adult and fetal microglia,23 and were clustered together with cultured human microglia and away from HMDM and cultured human hepatic macrophages,18 supporting their role as a more suitable cellular source of microglia than adult hematopoiesis-derived myeloid cells.
It is also noteworthy that the myelomonocytic cells that emerge from EB culture and are collected for subsequent differentiation to macrophages,8,12 indeed originally termed monocytes,12 are now recognized as MYB-independent primitive myeloid cells that clustered away from peripheral blood monocytes based on their transcriptome profile.20 We also have performed RNA-sequencing of myeloid progenitors (CD34+CD43+CD45+CD18+) harvested during progressive commitment and maturation to IPSDM8 and showed that the expression profile of those progenitors is highly correlated to that of IPSDM but not CD14+ monocytes (Figure 2). Thus, successful differentiation of circulating monocytes will rely on a more complete understanding of mechanisms regulating definitive hematopoietic specification24 and identification of factors governing the generation of hematopoietic stem cells from iPSC,25 knowledge gaps that remain major goals of regenerative medicine.
Future studies to further improve efficiency, reproducibility and scalability of IPSDM differentiation with potential for automated differentiation will facilitate the application of IPSDM in multiple areas of research. It has been suggested that the differentiation efficiency and yield are higher when using serum-free media to differentiate EBs into myeloid precursors as opposed to serum-containing media.26 Monolayer-directed differentiation as opposed to an EB-based differentiation that generates cells of all 3 germ layers may have advantage for differentiation in defined conditions,21 but direct comparison has not been performed. Novel protocols to differentiate iPSCs to tissue resident macrophages, for example, cardiac macrophages, will further raise the possibilities to delve into their specialized properties and tissue-specific functions. It is also important to consider whether IPSDM may resemble the heterogeneity of human tissue resident macrophages at single cell levels and the mechanisms driving the heterogeneity and downstream functional consequences.
Disease Modeling and Functional Genomic Analyses With IPSDM
IPSDM overcome the limitations of the lack of an adequate supply of disease-specific primary macrophages in studying rare diseases, patients-derived IPSDM are increasingly used to model Mendelian diseases, for example, Tangier disease (mutations in ABCA1), lysosomal storage diseases including neuronal ceroid lipofuscinoses27 and Gaucher disease (mutations in GBA),28,29 chronic granulomatous disease (defective NADPH [nicotinamide adenine dinucleotide phosphate] oxidase),30 Blau syndrome (mutation in NOD2),31 and chronic infantile neurological cutaneous and articular syndrome (mutation in NLRP3).32 Some of these studies compared the phenotypes of IPSDM and HMDM derived from the same subjects8,29 to further validate the fidelity of IPSDM in assessing the functionality of genetic mutations. CRISPR/Cas9 gene editing technologies further expanded the potential by introducing disease mutation in iPSC lines derived from healthy subjects and to correct mutations in the iPSC lines of affected patients to demonstrate causality and the functional impact of the disease mutations on a specific isogenic background.14,33 IPSDM also serves as a powerful tool to study macrophage host–pathogen interaction. IPSDM can be infected with various strains of HIV-1,34 and support Salmonella26 and Chlamydia infection.35 Knockout of IRF5/IL10RA by CRIPSR/Cas9 led to increased susceptibility to Chlamydia trachomatis infection in IPSDM. This approach can potentially be extended to study other pathogens to advance our understanding of mechanisms of host–pathogen interactions in macrophages and the role of human genetics in influencing the susceptibility to specific pathogens and outcome of infections.
IPSDM provides a unique platform for functional genomic analyses. To date, genome-wide association studies have identified 73 coronary artery diseases–associated regions.36 Genome editing in human iPSCs to introduce separately each risk allele, or a combination of alleles, and to correct risk alleles and then differentiate to macrophages allows the interrogation of the macrophage-specific function of the causal variants at genome-wide association study coronary artery disease loci. IPSDM are also powerful tools for modeling genetic interactions, alternative splicing, and macrophage-specific functional noncoding transcripts and regulatory elements. By establishing a biobank of iPSC lines derived from a large number of healthy subjects,37,38 one can study gene–environment interaction and map quantitative trait locus under various stimuli to determine how genetic differences between individuals may alter immune cells' response to environmental stimuli, and whether variation in the response may predict susceptibility or resilience to macrophage-related disorders in humans including atherosclerosis and coronary artery disease. The coculture or 3-dimensional systems, involving IPSDM and other iPSC-derived cell types relevant to atherosclerosis, including endothelial cells and smooth muscle cells, to form 3D tissue-engineered blood vessel organoids, will be crucial to advance in vitro disease modeling, for example, organs on a chip,39 and better recapitulate the complexity of the tissue-level in vivo environment. The scalability of the IPSDM platform can allow genome-wide CRISPR screening of human macrophage-specific regulatory element or knockout screening to identify novel regulators of functional phenotypes, which to date have only been performed in immortalized cell lines or PSC lines.40
Drug Testing and Cell Therapeutics
Patient-derived IPSDM-based in vitro assays are starting to impact the drug discovery process. IPSDM derived from Gaucher disease patients because of mutations in GBA, a gene encoding glucocerebrosidase, were used to test the efficacy of a noninhibitory small-molecule chaperone drug previously identified from high-throughput screening of compound libraries to enhance glucocerebrosidase activity and reverse the disease phenotype.29 IPSDM derived from chronic infantile neurological cutaneous and articular syndrome patients carrying mutations in NLRP3 were used to test existing anti-inflammatory compounds in inhibiting the abnormal IL-1β secretion,32 and can be potentially used to seek drug candidates that directly modulate NLRP3-inflammasome activation for chronic inflammatory conditions such as atherosclerosis.32 Beyond the use of patient-derived IPSDM, CRISPR/Cas9-edited IPSDM for key regulators in inflammation and lipid metabolism pathways can be exploited for the development of novel therapeutics and delivery systems.41 The development of organoid systems derived from isogenic iPSC lines will allow drug effects to be tested within the context of the symbiotic interactions between macrophages and their host tissue with the potential for development of individualized treatment strategies.
IPSDM-based cell therapeutics have been studied mainly in cancer42,43 and airway diseases44 using murine models particularly relating to the protective benefits of macrophage-mediated phagocytosis. The intraperitoneal injection of iPSC-derived myeloid cells10 genetically engineered to express interferon-β significantly inhibited the growth of human gastric and pancreatic cancers implanted in the peritoneal cavity,42 as well as inhibiting the growth of disseminated human melanoma cells in immunocompromised mice.43 iPSC-derived primitive macrophages intranasally transferred to the lung matured into alveolar macrophages and eliminated the surfactant protein accumulated as a result of Csf2ra deficiency in mice.44 Indeed, evidence is emerging that iPSC-derived myeloid cells may have specific advantages as in cell therapeutics. In mice, iPSC-derived primitive macrophages exhibit greater capacity to be differentiated into microglia-like cells by coculturing with neurons than monocytes and bone marrow-derived macrophages.21 Similarly, murine iPSC-derived alveolar macrophages engraft in lung better than bone marrow-derived macrophages.44
Adult mammalian myocardium contains diverse macrophage populations with distinct origins and function.45 At steady state, the majority of the resident macrophages is derived from yolk sac progenitors and primarily maintained through local proliferation. However, during injury, circulating monocytes are recruited, differentiated to macrophages and reside, whereas resident macrophages expand through proliferation. Transcriptional analyses reveal that monocyte-derived macrophages coordinate cardiac inflammation, while playing lesser roles in antigen sampling and efferocytosis.46 Similarly, embryonic and neonatal macrophages regulate tissue remodeling during development with a lower capacity to generate inflammatory response compared with monocyte-derived macrophages.47 Therefore, it has been proposed that preserving resident cardiac macrophages expansion, while targeting peripheral monocyte recruitment may lead to improved myocardial recovery after injury.46 Our RNA-sequencing data suggested that IPSDM may be less inflammatory, as genes expressed at lower levels in IPSDM versus HMDM were enriched in gene ontology terms immune response, antigen processing and presentation, and response to stimulus,8 but whether IPSDM could exert therapeutic benefits in cardiac injury remains an unexplored question. Deeper understanding of the functions of cardiac-resident macrophages in the steady state and injured heart and macrophage populations in the diseased vasculature, will reveal the potential of IPSDM as a cell therapeutic strategy in cardiovascular diseases. Yet significant challenges and barriers for clinical application of IPSDM and any PSC-based products remain to be addressed.
Limitations and Cautions
Application of iPSC-derived cells for functional genomics and translational studies has many strengths, but current limitations should be considered carefully. These include (1) clonal variability can occur during iPSC reprogramming and has been well recognized to cause variations in differentiation efficiency and the functional properties of differentiated cells. Somatic cell source and reprogramming methods also may affect differentiation efficiency, but the impact on IPSDM has not been systematically evaluated. It is therefore essential to use well-characterized iPSC lines, highly consistent differentiation protocols with quality assessment at multiple stages of the differentiation process, and multiple iPSC lines/clones per subject with independent replications to minimize variability and render statistically significant and meaningful experimental outcomes. (2) The gene of interest and functional phenotypes to be tested in IPSDM need to be carefully chosen and evaluated, for example, whether the target genes or noncoding transcripts show similar expression profile between IPSDM and HMDM, or whether the alternative splicing events or the regulatory features of HMDM are also represented by IPSDM. For initial screening, our studies characterizing the coding genes8 and lincRNA (long intergenic noncoding RNA) transcriptome (article under review), and alternative splicing events15 in isogenic IPSDM and HMDM lines provide a powerful resource for planning IPSDM studies to model such events, but specific validation for each chosen target is essential. (3) Some target genes may affect iPSC reprogramming or IPSDM differentiation efficiency, and cannot be reliably studied in IPSDM. Also IPSDM modeling of aging-associated cellular phenotypes has not been established. (4) Carefully designed controls and a combination of experimental strategies are essential for faithful interpretation of the functional consequences of target genes when using IPSDM, for example, patient-derived iPSC lines with correction of disease causing mutations in conjunction with wild-type lines with gene knockout or knockin of the mutations, and CRISPRi- and CRISPRa-mediated gene inhibition and activation add powerful complementary layers of information. (5) Use of IPSDM does not eliminate the need for traditional cellular models or distinct genetic manipulations, for example, siRNA or antisense oligonucleotides mediated knockdown, which are complementary and alternative approaches to further ensure validity and reproducibility.
IPSDM possess enormous value in advancing our understanding of diseases that involve human macrophages and already have demonstrated proof of principle utility in the development of novel therapies. Future studies will focus on automation of IPSDM differentiation, via both primitive and definitive hematopoiesis, with high scalability and reproducibility to produce the full range of myeloid cells that have tissue-specific properties and are applicable for tissue engineering. In such contexts, IPSDM will be instrumental in validating human genetic discoveries, advancing functional genomics, and facilitating clinical and therapeutic translation for diseases such as atherosclerosis and coronary artery disease in which macrophages play a modulatory role in the clinical outcome.
The differentiation of human induced pluripotent stem cells to macrophages (induced pluripotent stem cell–derived macrophages) offers a powerful platform to study human macrophage biology, understand human disease, and develop novel therapies.
Induced pluripotent stem cell–derived macrophages show similar phenotypic, functional and transcriptomic characteristics as human monocyte-derived macrophages.
In vitro hematopoietic differentiation of pluripotent stem cells resembles in vivo primitive hematopoiesis rather than adult definitive hematopoiesis. Protocols that specifically differentiate induced pluripotent stem cells to microglia-like cells have been reported.
Future studies will focus on automation of induced pluripotent stem cell–derived macrophages differentiation, via both primitive and definitive hematopoiesis, with high scalability and reproducibility to produce the full range of myeloid cells that have tissue-specific properties for use in tissue engineering, functional genomic analyses, drug testing, and cell therapeutics.
Sources of Funding
This study is supported by National Institutes of Health (NIH) grants R01-HL-113147 and K24-HL-107643 (to M.P. Reilly), and partially supported by American Heart Association Postdoctoral Fellowship 15POST25620017 and NIH grant K99-HL-130574 (to H. Zhang). M.P. Reilly is also supported by R01-HL-111694 and R01-HL-132561.
- Nonstandard Abbreviations and Acronyms
- Cas9
- CRISPR–associated 9
- clustered regularly interspaced short palindromic repeats
- embryoid body
- granulocyte macrophage colony-stimulating factor
- human monocyte-derived macrophages
- iPSC
- induced pluripotent stem cell
- iPSC-derived macrophages
- macrophage colony-stimulating factor
- Received August 18, 2017.
- Accepted September 12, 2017.
- © 2017 American Heart Association, Inc.
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This Issue
Jump to
- Article
- Visual Overview
- Abstract
- IPSDM Provide an Important Tool to Study Human Macrophage Biology
- Generation, Functional Feature and Molecular Profiling of IPSDM
- Disease Modeling and Functional Genomic Analyses With IPSDM
- Drug Testing and Cell Therapeutics
- Limitations and Cautions
- Conclusions
- Sources of Funding
- Disclosures
- Footnotes
- References
- Figures & Tables
- Info & Metrics
- eLetters
Article Tools
- Human Induced Pluripotent Stem Cell–Derived Macrophages for Unraveling Human Macrophage BiologyHanrui Zhang and Muredach P. ReillyArteriosclerosis, Thrombosis, and Vascular Biology. 2017;37:2000-2006, originally published October 5, 2017https://doi.org/10.1161/ATVBAHA.117.309195
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Via the Moonie Times: Missouri's Republican legislative leaders are seeking to appeal a court ruling requiring officials to recognize same-sex marriages performed in other states. Senate President Pro Tem Tom Dempsey and House Speaker Tim Jones have filed a motion asking to intervene in a Jackson County court case so they can appeal it to the state Supreme Court. Judge ...Read More »
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D: Don't Be A Douchebag
An "Ex-Gay" At Vatican Marriage Meeting
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Quote Of The Day: Pope Francis
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Tony Perkins Has The Sixth Circuit Gladz
"We applaud the U.S. Court of Appeals for the Sixth Circuit for upholding the freedom of the people to define marriage as the union of a man and woman. The Sixth Circuit rightly recognizes that the Constitution does not demand that this modern redefinition of marriage be forced on the States. The American people simply will not accept a nationwide ...Read More »
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Via press release from Tony Perkins: If you want to motivate voters, try trampling their right to self-govern. For more than a year, Americans have watched it happen in courtrooms across the country, as a handful of judges try to junk the marriage amendments passed by more than 41 million people. Yesterday, FRC did its part to stand with those ...Read More »
Tony Perkins: Our Houston Hate Rally Was "High-Water Mark For FRC"
"If the Mayor thought pulling the subpoenas would take some of the heat off, she quickly discovered that it only exposed the real controversy — which was invalidating the referendum to begin with. In the fierce back and forth over the measure, Parker said the bill was 'personal.' What she didn't realize was how personally Houston would take it. And ...Read More »
Perkins: Houston's Hate Rally Gives Hope To Anti-Gays Causes Around The World
HOUSTON: Hate Rally Claims 6000 Attended, 700 Churches Simulcast Event
Via the Moonie Times: Thousands showed their support Sunday the five Houston pastors who drew national attention for challenging the city's now-withdrawn effort to subpoena their sermons. I Stand Sunday, a combination church service and religious-freedom rally, drew an estimated 6,000 congregants to hear a high-profile line-up of speakers at Grace Community Church in Houston. The two-hour event was simulcast ...Read More »
Seen At The Houston Hate Rally
(Tipped by JMG reader Str8 Grandmother)Read More »
LIVE VIDEO: Houston Hate Rally
The Family Research Council's "I Stand Sunday" hate rally against Houston's LGBT rights law and Mayor Annise Parker begins at 7PM Eastern time at the Grace Community megachurch, whose largest hall seats 10,000. As you can see by the list above, the evening will provide a cavalcade of crackpottery. LGBT activists are encourage to take part in the already underway ...Read More »
Tony Perkins On Tim Cook
"Apple's old logo was colored by the rainbow — and it looks like their top management still is. Three years after taking the company over for an ailing Steve Jobs, Apple CEO Tim Cook confirmed what many people suspected: he identifies as a homosexual. Cook, the head of the tech world's colossus, said he made the announcement because his predecessor ...Read More »
New Zach Dasher Ads: Bibles And Guns Brought Us Here & They'll Keep Us Here
In three new campaign ads, the US House bid of Louisiana's Zach "My Platform Begins With God" Dasher is endorsed by Phil Robertson (Dasher's uncle), and Sen. Ted Cruz, who's apparently OK with vertical cellphone video ads. Via NOLA.com: Fifth Congressional District candidate Zach Dasher's new ad says he believes in guns, and he's just unloaded the biggest gun he ...Read More »
HOUSTON: Mayor Annise Parker Says She'll Withdraw Subpoenas Of Pastors
Via the Houston Chronicle: The City of Houston will withdraw its controversial subpoenas of five pastors tied to a lawsuit over the city's equal rights ordinance, Mayor Annise Parker announced at a news conference Wednesday. The announcement came amid a national firestorm about the subpoenas, which have prompted outrage among Christian conservatives. Parker said two meetings yesterday, one with local ...Read More »
FRC Hate Rally Draws Non-Crowd
I count exactly ten people visible under the trees. Notice the t-shirt sales tent in the back. That's Perkins on the stage. (Via Scott Wooledge)Read More »


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Music Videos
Marching to the Beat of Their Own Drum
Mesmerizing Harry Potter Theme Song Played on Glasses by Street Performer
Listen as this street performer plays the classic Harry Potter Theme Song on glasses. The haunting melody sounds incredible ringing from the glass harp surrounded by captivated strangers.
Enjoy the Sounds from the World's Newest Instrument
The Venova is the world's newest instrument designed by the folks at Yamaha. The wind instrument is a combination of a saxophone and recorder and looks like something out of a Star Wars cantina.
Musical Road's Aren't All They're Cracked Up to Be
There are certain stretches of California's highway that have special grooves cut into them that play music as you drive. The only problem is, they really suck.
View the World Through a Guitar
This mesmerizing video was shot with a camera in an acoustic guitar and captures the vibrations of the strings. The ever changing scenery and magical motion of the strings make for a captivating scene.
How Computers Know What Music You'll Like
If you've used any streaming services like Spotify you'll be familiar with their recommendation services. While some are way off there are those like Spotify's Discover Weekly that work pretty well and here's why.
Making Mad Music for Rick and Morty
If you've ever watched Rick and Morty you'll know there are some pretty out there ideas and at some point someone had to make music for those situations. That someone is composer Ryan Elder who has the task of making those strange requests a reality.
Italy is the Home of the Violin
Violins are one of the most well known of the classical music instruments but the very best of them all originate from one place. Cremona, Italy is known as the city of violins and is the place to go for quality instruments.
The Deaf Musician Helping to Visualize Sound
Deaf musician Myles de Bastion has been in love with music all his life and has found a way of sharing it with others. Myles has created the Audiolux visualization system to help other deaf people see sound.
Listening to the Same Song for a Thousand Years
Meet composer Jem Finer who has created a symphony that will play non stop for a thousand years. Needless to say you don't want to run into this piece on your iPod shuffle.
Hired Gun Rock Documentary Trailer
Hired Gun is a documentary about the world of Rock and the musical geniuses that have gone on to create iconic rock solos and more.
The Song Played at Every American Graduation
Ever wonder why the same song is played at every American college graduation? Turns out the song, Pomp & Circumstance, has its origin's in the British war for gold.
Creating Horror with The Apprehension Engine
The Apprehension Engine is a custom built instrument designed to create horror sound effects. The various elements can be used to create all manner of sinister and creepy sounds to make the hair on the back of your neck stand on end.
Who Knew Cows Loved Jazz?
A dairy farmer in Florida named Ed Henderson has taken his love of jazz outdoors and now shares it with his bovine brethren. What was once a backyard practice session has now turned into a regular session entertaining over 6,000 cows.
Tapping Along with the Typewriter Orchestra
Who needs traditional instruments when you have a typewriter? Meet the 6 men that make up the Boston Typewriter Orchestra and play various original and cover songs using old mechanical typewriters.
The Unique Sound of Funny from the 90's
Meet Fred Newman, known for his sound effects and voice overs in the Nickelodeon's classic "Doug". Fred has also lent his wide range of silly sounds to the likes of "Gremlins" and "Who Framed Roger Rabbit".


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HONG KONG, Aug 10 (Reuters) - Cheung Kong (Holdings) , the property developer founded by Asia's richest man, Li Ka-shing, said on Friday it had won a tender from subway operator MTR Corp to build the property above a rail station in Hong Kong, paying HK$9.6 billion ($1.24 billion) for the land.
Managing director and executive deputy chairman Victor Li, the elder son of the company's 83-year old founder, announced the tender win for the Tsuen Wan project at the company's headquarters, adding it would spend more than HK$20 billion ($2.6 billion) on the project.
The MTR had withdrawn the tender in January when bidders failed to meet its reserve price of HK$10 billion. Developers have grown cautious on land purchases with Hong Kong prices at some of the highest levels in the world. ($1 = 7.7556 Hong Kong dollars) (Reporting by Alex Frew McMillan; Editing by Muralikumar Anantharaman)


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Moscow main Department of the Ministry of internal affairs has placed on the portal of public procurement on the purchase of horses on two million rubles
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- In Egypt, a bomb exploded in a mosque, was injured about 150 people
- One person was injured in a car accident in the South-West of Moscow
- Break new ground in the space: "the core instruction" will ensure Russia's leadership
- Russia and Venezuela have agreed on the supply of wheat
- Prevent ex.: bobsledder Zubkov: I will defend his good name
5 recent posts for today:
- Mubarak's family is on the state of Egypt to return nearly $ 3 million, January 14, 2013
- Lavrov: US rejected the offer of Russia meeting on Syria, October 14, 2015
- Get rid of internal Islamists European countries, May 20, 2013
- The mother asked the pilot Yaroshenko power? Usland exchange for Snowden Son, July 16, 2013
- The last watching harry potter you will ever need, July 9, 2011


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Jimmie skinner guitar
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So this app is completely useless for my B string. When they sing You Are My Sunshine you realize this isn't just a song for Barney and friends - in fact, jimmie skinner guitar pretty depressing. Since I didn't test the unit with PC or Android devices, I can't be sure if the class-compliant rule still applies. When you find a format that fits your learning style best, check out other songs they have on their Jimmie skinner guitar channel. No part of this site or its content may be reproduced without the permission of the copyright holder. To demonstrate its skills, the firm has created a President Obama urn and says that it can even make wigs for the functional busts to create a more life-like effigy of people with long hair. For practice what you preach guitar pro left jimmie skinner guitar, Erik is running through some scales and light arpeggios. Today we will talk about the difference between these two types and talk about what each of them has to offer. 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So if you were to continue on past stephane laplante guitariste G, the next bar chord would be A minor because A comes after G as seen in the chromatic scale - A, Jimmie skinner guitar, Guitad, C, C, D, D, E, F, F, G, G and then back to A, then A and so on etc. The following sskinner are all advice I was given by musicians I trust and respect, and if jimmie skinner guitar worked for me odds are that jimmie skinner guitar help some of you as well. Christine, a consummate Yogi and beautiful musician, is cool tune guitar and bass tuner in Jimmoe on production tour - we look forward to having her back as guest musician. Hi, I'm glad my lessons helped you out. Register on the forum. By subscribing to the site you will receive my newsletter that I jimmie skinner guitar out every couple of weeks letting you know what the new jimmie skinner guitar are for guktar past couple jimmie skinner guitar weeks and where to find them. The staff are super sweet and polite. 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What Appliance Repair Is – and What it Is Not
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New Zealand Consul General Graham Sims works with Vietnam National University – Hochiminh City
On June 4th, 2009, New Zealand Consul General Graham Sims paid a visit and worked with VNU-HCM.
At the meeting, Assoc. Prof. Dr. Le Quang Minh, Vice-president of VNU-HCM, sent his congratulations to Mr. Graham Sims on the occasion of taking up his duties at New Zealand Consulate General in Hochiminh City, simultaneously introduced an overview of organization and operation of VNU-HCM, gave a summary report on education cooperation between VNU-HCM and New Zealand's partners, and expressed a desire to receive the enthusiastic support of the New Zealand Consulate General in the relationships with New Zealand's universities.
Speaking at the meeting, Mr. Graham expressed pleasure of the achieved results in the relationship between two countries and promised in his new position he will continue ativities to support education cooperations between
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- Over a thousand students attended the speech of Harvard's President
- Public lecture of the President of Harvard University at VNUHCM-USSH


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Square One's employees stayed loyal
Published 4:00 am, Wednesday, July 24, 1996
WHEN Joyce Goldstein announced eight weeks in advance that she planned to close her Square One in July, not a single employee left. Displaying the kind of two-way loyalty that's unfashionable these days, they all stayed through July 13 so that Square One could go out with the fully staffed style it had shown through 12 years of fame. Joyce, meanwhile, helped the employees, many of whom had been with the restaurant for years, find new positions at first-class places.
Said a happy Edwin Aguilar, a Square One dishwasher for four years: "I've got a new job at the Ritz Carlton." Stop cruelty to oysters!
I mentioned risotto to Janet Trefethen of Trefethen Vineyards and to my surprise she asked what kind of rice I used. I said arborio, which of course is traditional. Did I ever use California pearl rice? she pressed. By now, I began to have a tiny suspicion. Yes, I answered, it's good but I preferred arborio, although I've never blind tasted one against the other.
She gave me a winning smile and suggested I try just such a test, and then she copped. One of her many interests, in addition to fine wine and quarter horses, is raising California rice.. Frogs have feelings too!
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When the sophisticated Jaguar Club, 150 owners of Jaguar motor cars, zoomed to the hinterlands of Guenoc Valley, they found an old fashioned country welcome, roasted wild pig. Orville Magoon, owner of the 23,000 acre estate, said that all the pigs at the banquet, were government inspected. Henry VIII never had it so good. Halt crab torture!
Drew Nieporent, the New York restaurant operator who put together Rubicon here in San Francisco, picked a good weekend to visit the Bay Area. The same Friday that he was celebrating in the Napa Valley, The New York Times ran a glowing review of his latest New York venture, Zeppole, one fine Italian restaurant, according to reviewer Ruth Reichl, whom I admire.
Nieporent, who I saw a couple of times during the weekend, is an entertaining fast-talker, a former cruise-line waiter who knows food and restaurants. You can occasionally catch him here at Rubicon with investors like Francis Ford Coppola. Down with fresh!
And speaking of the Napa Valley, I stopped in for lunch at Mustards Grill in Yountville recently. I'm happy to report it's as good as ever, which is mighty good, indeed. I enjoyed a grilled salmon that went perfectly with a glass of 1994 Hess Chardonnay. The sauce was baby beans and a few kernels of sweet corn with thin-sliced potatoes. There were a few Southeast Asian red peppers which added zip to the sauce, but remember not to eat them lest you cauterize your tongue. Ban fish murder!
The S.F. chefs' shuffle: Mark Franz, formerly of Stars, is joining Lark Creek Inn as acting chef; Sylvain Portay of Le Cirque in New York is the new chef at The Dining Room of the Ritz Carlton, replacing Gary Danko, who is writing a book; and David Lawrence, formerly of 231 Ellsworth and Cityscape in San Francisco, will take over the kitchen at the Essex Supper Club, the successor to Ernie's.
While a chef in England, Lawrence prepared meals for the Prince and Princess of Wales, Princess Margaret and Prime Minister Margaret Thatcher, so he should be able to handle the Essex patrons OK. Regarding the allegations about sloppy selling of fresh seafood in Chinatown, stop and think a minute. Do you really believe customers would buy fish and seafood that looked sick? Sanitary selling practices are just good business.
The little bold-faced slogans between items are just to give equal space to the other side.<


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It has been clinically tested clinically, contains 50 mg of epihydroksytiolano-17-ester per dose. By building muscle mass in arms controlling the economy testosterone and estrogen through the preparation Probolan 50 natural processes of the body starts to burn fat and develop muscles. Probolan 50 and each of its components have been thoroughly tested for safety and the impact on consumption accelerate muscle growth. Muscles respond to training in three ways. When you're doing a lot of times repeated (more than 15), increases muscle strength, without an accompanying increase in their size and strength. 6 to 12 times repeated - the amount on which bodybuilders rely on - increase in both size and muscle strength. Lifters generally do little times repeated, 2 to 4 per set, which provides and low magnification. If, however, for some time, you will be lifted weights while doing little of times repeated, then - when then go back to their 6-12 times repeated - you'll be stronger. The principle is: greater force shall be at greater muscle tension equals more growth. This allows your muscles gain strength, become more durable and ready to take intense weight building muscle mass in legs training. The secret of the effectiveness of this Epihydroxetiolan-17-ester matrix - available in appropriate proportions, which are the key to excellent results. Many bodybuilders are not able to increase the weight, because they train constantly, so that their muscles can not fully recover in August. Taking a few days off can restore glycogen, increase anabolism and allow a return to an optimal level of hormones such as testosterone and cortisol. Aerobic exercise has a detrimental effect on mass building. Aerobics interfere with strength gains and recovery while burning up valuable glycogen and amino acids with branched łaącuchach. Adding mass is the best way to increase basal metabolic rate. The higher it is, the more building muscle mass in arms calories are burned and it is easier to stay lean. Muscle growth is the logical consequence of their contractions. Usually emphasize the role of the concentric contraction phase, when mięsieą getting shorter but stretching the muscle in the concentric phase (negative) when mięsieą lengthens while maintaining tension, or directly cause muscle hypertrophy. Emphasizing the negative is an easy technique to overload muscles and promote radical gains in mass. Creatine is associated with increased power and higher production ability to adezynotrójfosforanu (ATP) - which is a chemical fuel for training and growth. Supplement creatine allows bodybuilders to increase muscle creatine concentration and thus increase their power and the amount of ATP, without all the fat that you would if you were building muscle mass in legs burdened only fetch blood creatine. As the only commercially available formulation contains up to 500 mg Epihydroxetiolan-17-ester matrix. The best bid for the winter 2014Jest is the highest permitted dose of the active ingredient. Therefore, only Probolan 50 for maximum production of testosterone - the hormone needed to build a perfectly sculpted muscles. Never achieve a positive nitrogen balance with a low calorie diet. To build new muscle mass and muscle regeneration provide, the need for raw materials: carbohydrates, proteins and fats. Increasing your calories by 50% (eg. From 3000 to 4500 calories per day) for three days can spur growth without causing weight gain (or only slightly). The idea is to limit the increased calories to a designated three days; you'll be able to stimulate growth by improving muscle sensitivity to insulin and by providing more carbs for glycogen storage. If you are not overtraining muscles grow between you, try this method. Probably easier anabolism extra calories, fat hangs before. Therefore, you must limit the increased caloric intake by half to three days. After this period go back to normal calorie intake - has provoked an increase in his new weight without adding an unwanted fat.
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