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Breaking news
Business news and information from the European Union
- EU leaders strengthen resolve as 2nd Covid wave hits Europe
EU leaders at a summit Thursday resolved to strengthen their collective effort to fight the COVID-19 pandemic, with a focus on testing and tracing policies, and on vaccines.
- Harmonised procedures boost rail sector across EU
As of 31 October, Europe's rail sector will benefit from new harmonised procedures expected to significantly simplify procedures and reduce costs for railway undertakings operating across Europe.
- Covid crisis needs adequate minimum wages, says EC
The European Commission proposed Wednesday a new directive to ensure that EU workers are protected by adequate minimum wages allowing for a decent living during the COVID crisis.
- Brussels puts forward Atlantic and North Sea 2021 fish quotas
The EU Commission set out its proposal for catch limits for fish stocks in the Atlantic and North Sea Tuesday, the basis for EU fisheries ministers to set final catch limits at a 15-16 December Council.
- EU gets WTO green light for tariffs on US exports over Boeing
The WTO's dispute settlement body gave formal authorisation Monday for the EU to increase duties on U.S. exports worth up to $4 billion, in retaliation for illegal subsidies granted to U.S. aircraft maker Boeing.
- EU states agree informal deal to improve rail passenger rights
EU ambassadors endorsed a reform of rail passenger rights Wednesday with a view to strengthening the rights of all rail passengers, in particular those with disabilities or reduced mobility.
- Strong take-up of EUR 17 bn inaugural EU 'social bonds'
There was strong investor interest in the EUR 17 billion inaugural 'social bonds' issued Wednesday under the EU's 'SURE' instrument, set up to help protect jobs and keep people in work over the pandemic.
- Euro-Parliament awards 2020 Sakharov Prize to Belarus opposition
The European Parliament awarded its Sakharov Prize for Freedom of Thought Thursday to the democratic opposition in Belarus - in a political crisis since the disputed 9 August presidential elections.
- Eco-schemes feature in EU's greener common farm policy
Farmers will be afforded flexibility on the way to higher environmental standards in farming in the post-2020 common agricultural policy reform package agreed provisionally by EU ministers Wednesday.
- Cyber attacks threat grows under Covid-19
Cyber criminals have advanced their capabilities during the pandemic, adapting quickly and targeting victim groups more effectively, according to a report Tuesday from the EU cybersecurity agency.
- EU Baltic 2021 fish quotas to help stocks recover
The EU agreed on 2021 fishing catch limits for the Baltic Sea Tuesday, with an acceptance of the need to continue decreasing the fishing opportunities for several fish stocks to help them recover.
- Europe's nature under pressure
Intensive farming, urbanisation and unsustainable forestry are contributing to the decline of protected habitats and species in Europe, a review of vulnerable species and habitats showed Monday.
- 'Farm to fork' strategy commits EU to sustainable agri-food sector
The EU Council adopted Monday a set of conclusions on a 'farm to fork' strategy, endorsing the goal of developing a European sustainable food system, from production to consumption.
- Brussels points way to greener, more digital economy post-COVID
A more healthy and fairer Europe is promised in the EU Commission's 2021 work programme, published Monday with the aim of accelerating a long-term transformation to a greener economy, fit for the digital age.
- EU steps up 'no-deal' preparations as UK refuses to budge
With time running out for successful conclusion of an EU-UK trade deal and insufficient progress on key issues, EU leaders called on the UK at a summit Friday to make moves to make an agreement possible.
- EU imposes sanctions on Russian officials for Navalny poisoning
The EU imposed sanctions Thursday against six Russian officials involved in the assassination attempt on Russian opposition leader Alexey Navalny, poisoned in Russia with a Novichok toxic nerve agent.
- New EU chemicals strategy targets harmful consumer products
A new strategy on chemicals, unveiled Wednesday by the EU Commission, would prohibit the use of the most harmful chemicals in consumer products such as toys, cosmetics and detergents.
- Auditors look into effectiveness of EU funding for SMEs
The European Court of Auditors launched a new audit Wednesday to examine whether EU funding for small and medium sized enterprises (SMEs) is helping to make them more competitive and future-proof.
- EU puts in place rail safety measures for Channel Tunnel
The EU Council adopted legislation Wednesday to ensure safe and efficient operation of the Channel Tunnel rail link between continental Europe and the United Kingdom after the end of the Brexit transition period.
- MEPs want to increase 2030 emissions reduction target to 60 pct
All EU Member States must become climate neutral by 2050, the European Parliament said Thursday in a vote on EU climate law, with MEPs calling for ambitious 2030 and 2040 emissions reduction targets.
- Helping small businesses trade internationally
The European Commission launched the Access2Markets online portal Tuesday, to help small and medium-sized firms trade beyond the EU's borders and successfully navigate the world of international trade.
- EU states look to coordinate COVID travel measures
The EU adopted Tuesday a common framework on travel measures in response to the COVID-19 pandemic, with recommendations to help states take decisions based on the epidemiological situation region by region.
- Brussels settles antitrust chips dispute with Broadcom
The EU Commission accepted commitments by Broadcom Wednesday to suspend exclusivity agreements and/or leveraging provisions for Systems-on-a-Chip used in TV set-top boxes and Internet modems.
- EU to issue EUR 100bn social bonds to protect jobs
The EU Commission announced Wednesday that it will issue its forthcoming SURE bonds of up to EUR 100 billion - designed to help protect jobs and keep people in work through the pandemic - as social bonds.
- EU likely to miss targets for recycling plastic packaging
There is a significant risk that the EU will not meet its plastic packaging recycling targets for 2025 and 2030, according to a review published Tuesday by the European Court of Auditors.
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Opinion ARTICLE
"Of mice and men": arginine metabolism in macrophages
- 1Bristol Heart Institute, School of Clinical Sciences, University of Bristol, Bristol, UK
- 2Department of Microbiology and Molecular Genetics, University of Pittsburgh, Pittsburgh, PA, USA
Macrophages are involved in inflammation from induction to resolution. Polarization of macrophages along the M1 (classical) or M2 (alternative) axis occurs during inflammation and can be at least partly categorized by the route of arginine metabolism within the macrophage, balancing the activities of the arginase and nitric oxide synthase (NOS) enzyme families (1, 2). Arginase activity is associated with tissue repair responses (via ornithine production and pro-proliferative effects). In contrast, NOS2 generates nitric oxide (NO) species with anti-proliferative effects that is necessary for protection against pathogens and aberrant cells (2, 3). Other NOS enzymes produce NO that acts in the regulation of smooth muscle tone and other cellular processes (4). Macrophages preferentially expressing the arginase or NOS2 pathways enzymes also influence T-cell activation, proliferation, signaling, and apoptosis in different ways (1).
While arginase and NOS enzymes can be used to ascertain the pathway of macrophage activation in rodents, there has been debate as to whether they are present in macrophages from humans and other mammals. The arginase and NOS enzymes are extensively conserved, and the NOS forms found in mammals are similar to those in cnidarians, mollusks, and other chordates (5, 6). These arginine-metabolizing enzymes are present in some human leukocytes, and there is evidence that they are also present in macrophages from other vertebrates, including chickens, rabbits, cows, and primates (7–12). However, comparisons of tissue macrophages of different species are lacking, which limits our understanding (13). Many studies in humans have principally focused on blood monocytes, leading some researchers to question the suitability of rodents as model of macrophage activation, as there is not always a direct correlation with human cells. Was Robert Koch correct when he said "Gentlemen, never forget that mice are not humans," or can the differing results between species be explained, in part, by differences in the types of monocyte or macrophage studied? Our purpose here is to examine this question.
Arginine Metabolism in Mammalian Cells
Many mammalian cells, including neutrophils, granulocytes, erythrocytes, hepatocytes, cardiac myocytes, dendritic cells, myeloid-derived suppressor cells, foam cells, natural killer cells, endothelial cells, and smooth muscle cells, have arginase (12, 14–16) or NOS activity (8, 17–19), albeit to different degrees. Macrophages are the primary circulating cells that can express either of these enzymes, depending on the inflammatory circumstance. Experiments that detect NO, ornithine, or urea production (via NOS2 or arginase) have most often been performed on rodent macrophages. Macrophages from some mouse strains (e.g., the M1-biased C57BL/6 strain) can be stimulated by lipopolysaccharide (LPS) to produce considerable quantities of NO. Macrophages from others strains (e.g., M2-skewed BALB/c mice) produce much less NO (20) and produce more ornithine instead. Some researchers did not detect any NO production in macrophages from humans, pigs, and rabbits (8, 11, 14, 21–23), but others (including ourselves) have observed NOS or arginase activity in macrophages from rabbits, humans, and other primates (4, 7, 10, 12, 17, 24–26).
Why is There Controversy?
One main difference between the studies from laboratories is that some use monocyte-derived macrophages (MDM), while others study tissue macrophages directly. A number of groups have detected NOS or arginase activity in human monocytes or macrophages (3, 27–29); but others have not. Why is this so? Part of the explanation lies in the fact that in vitro-derived macrophages can generate different responses from macrophages obtained in vivo as discussed below (and shown in Table 1). Another explanation is that many groups use the identification of enzyme protein rather than detection of enzyme activity as evidence of enzyme expression. Failure to detect the presence of a protein is not definitive evidence for absence of expression (especially when considering potentially different detection thresholds of antibodies or the high Vmax of arginase, i.e., very little enzyme is required for ornithine production).
Table 1. The presence of arginine-metabolizing enzymes in human monocytes and macrophages varies with cell source, treatment and health status/stress level of the individual.
Macrophages Produced in vitro
Macrophages have been produced in vitro in a number of ways. Cells from bone marrow have been isolated and "differentiated" in culture medium containing high levels of cytokines (such as colony stimulating factors, CSFs) to produce bone marrow-derived macrophages (BMDM) (13, 23, 44–46). Macrophages have also been produced by isolating and culturing monocytes from blood, to produce MDM (10, 13, 22, 30, 37, 47, 48). Production of these in vitro-derived macrophages is cheap, simple, and reproducible, but they may not be a full representative of tissue macrophages, as the preparation and culture procedures may not be sufficient to induce cell activation (4). The differences between tissue macrophages and in vitro-derived macrophages are at least partly dependent on cell source, time in culture, and the degree of manipulation in culture. Each research group will use different types and sources of culture media and sera, which vary greatly in the concentrations of factors that influence NOS2 or arginase expression, such as transforming growth factor β (TGFβ) (4, 20, 49). Another confounding issue is that circulating monocytes and tissue macrophages arise from different stem cell populations (50), although some macrophages found at sites of infection or inflammation may derive from infiltrating monocytes (51). Together, these factors may account for many of the differences observed in NO and urea production in these macrophages (8, 20).
Monocyte-derived macrophages or BMDM from different strains of mice can differ in their response to interferon-γ (IFNγ), LPS, and tumor necrosis factor-α (TNFα) (4, 8), and differences in the rodent background can result in differences in macrophage gene expression (13, 20, 49). Human in vitro-derived macrophages also show variability in their responses to LPS (4, 22, 46). It may be that the same stimulus is able to generate quite different responses in genetically diverse individuals, as it does between mouse strains (38, 49, 52). In general, human macrophages are not as responsive to LPS as mouse macrophages, possibly because of the lower environmental exposure of humans to LPS. It is also possible that human monocytes may be more effectively stimulated to become M1-activated macrophages by cytokines other than IFNγ and LPS/TNFα (e.g., IFNα) (4, 18, 43). Human macrophages take longer time to respond to the stimulatory factors in vitro than mouse macrophages, and some experiments using human MDM may have ended before a response was detected (48). There are other indications that the timing and length of the exposure of the cells to varying cytokines in vitro are important. For example, when M1-polarizing cytokines were removed from the culture medium, NOS2 levels in mouse BMDM were reduced and NO production (measured as nitrite) ceased (45). In addition, whichever arginase or NOS enzyme was induced earliest, the alternative enzyme decreased in expression and activity, unless arginine was present in excess (15, 45, 53). Macrophages require the local environment to continuously give appropriate activation cues. Changes in environmental cues can stimulate macrophage populations in vitro to express varying percentages of M1 or M2 dominant activity (54). When activation cues are reduced or removed, macrophages may become deactivated (e.g., M2c) or indeterminate (e.g., have features of M1 and M2).
Macrophages Obtained in vivo
Macrophages can be identified in whole tissues and organs or isolated in large numbers from in vivo sources such as the peritoneum or granulomas, and either examined immediately or used ex vivo. Macrophages obtained in vivo or made from monocytes can respond differently to the same stimulus (35, 47). In one study, monocytes and tissue macrophages were obtained from patients with an inflammatory disease (either rheumatoid or psoriatic arthritis). Compared with tissue macrophages, the MDM had a blunted response to the M2 cytokines interleukin-4 (IL-4) and IL-13, at least partly due to a reduction in some of the receptor elements for these cytokines (47). These results suggest that the response of the macrophages to M2 cytokines may be source specific, but it is possible that these cytokines alone were not sufficient to fully stimulate the MDM (38). Several lines of evidence suggest that macrophages in vivo express functional NOS2. Blood monocytes and peritoneal macrophages obtained from women during laparoscopic procedures contained NOS2 mRNA and protein. The macrophages had higher NOS levels than the monocytes, and this could be increased by treatment with LPS. The monocytes and macrophages also produced neopterin, nitrite/nitrate, and citrulline (suggesting that the enzyme was active). Although the production of NO from these macrophages was low, it would probably have been sufficient to cause functional changes (35).
Macrophages can also be obtained from alveolar aspirates, skin, and the placenta (10, 16, 21, 38, 39, 55, 56). For example, sponges placed subcutaneously into mice, rats, or rabbits attract large numbers of macrophages. The sponges can be removed from the animal and the macrophages were isolated and purified (10, 55, 56). It is a little more difficult to obtain and purify macrophages from other tissues, such as atherosclerotic vessels (44), but intact biopsy, surgical, or cadaveric specimens can also be investigated. It should be noted that resident macrophages from different tissues observed at different times (and different health states) may not necessarily have identical properties (51, 57).
In order to perform their full range of functions, macrophage populations exhibit "plasticity" of phenotype (52, 58), regardless of whether they are found in vivo or derived in vitro. As macrophages adapt or change their functions, they can simultaneously express markers of M1 and M2 activation, including NOS2 and arginase-1 (12, 59, 60). For example, tissue macrophages (and MDM) from Mycobacterium tuberculosis-infected cynomolgus macaques have been observed to co-express functional NOS and arginase enzymes (12). We suggest that macrophages display a spectrum of activation phenotypes, and it is the relative (and not absolute) proportion of M1 or M2 markers that we can use as a 'handle' to determine the type of activation state.
Effect of Disease and Trauma on Macrophage Activation
Blood monocytes from healthy volunteers do not usually need to produce NOS or arginase, so it is not surprising that many studies have not detected NOS or arginase in these cells (10, 14, 21, 22, 29, 30, 37). However, studies performed on tissue or cells from people undergoing stress, trauma [e.g., burns (29)], pregnancy (16), or disease {such as infection [e.g., tuberculosis (12, 24, 26) or filarial infection (28)], atherosclerosis (7, 10, 17, 25), autoimmune diseases (27, 36) and cancer (42, 61)} demonstrate that human macrophages (and sometimes monocytes) can produce active forms of the arginine-metabolizing enzymes (Table 1).
Trauma results in a pattern of gene expression in macrophages that is consistent with a wound-healing response, with an initial increase in NOS followed by decreased NOS production and activity, elevated IL-4, IL-10, and TGFβ levels, and increased arginase expression and activity, resulting in decreased plasma arginine levels (28, 29, 62).
Disease, however, causes different patterns of gene expression. For example, monocytes from multiple sclerosis sufferers not only have higher levels of arginase-1 and increased urea production, but also have increased NOS2 mRNA and nitrite production (particularly when stimulated by M1 cytokines or LPS) (36). Macrophages from patients with inflammatory diseases, such as tuberculosis, malaria, or rheumatoid arthritis, have increased levels of NOS2 mRNA and active protein (4, 8, 24, 26, 63), which may contribute to elevated plasma NO levels (64). Atherosclerosis is another inflammatory disease with a considerable macrophage contribution, with oxidized low-density lipoproteins taken up by macrophages during their transformation into foam cells. Plaque macrophages express NOS2 RNA and protein, as well as markers of NOS activity (including the presence of nitrotyrosine or ceroid) (4, 7, 17, 25). Plaque macrophages and foam cells express arginase-1 (10), and macrophages laser-dissected from plaque have upregulated levels of arginase-2 and NOS2 (65). Macrophages present in some neoplastic diseases also produce active NOS2 (4, 42, 66). Reducing the local levels of arginine has been proposed as a treatment for these diseases, by reducing inflammation-triggered immune dysfunction, tumor escape, fibrosis, and immunosuppression (61). Possible pharmacological interventions include treatment with arginine degrading enzymes, NOS competitors and inhibitors, asymmetric dimethylarginine, NO-releasing aspirins, cyclooxygenase, and phosphodiesterase or arginase inhibitors (8, 61). These studies suggest that an inflammatory environment is necessary in order to observe NOS or arginase in human monocytes and macrophages. The in vitro experiments that do not demonstrate arginase or NOS expression may simply be lacking the additional cues needed for expression rather than demonstrating an inability to actually express these factors.
The modulation of macrophages to express NOS or arginase has clear benefits for treating disease in humans (and other species). To do this, one needs to either determine suitable signals to stimulate these pathways or obtain a sufficient number of human macrophages (e.g., by tissue culture) that function like tissue macrophages.
Because macrophages from different inbred strains of mice vary greatly in their macrophage NOS and arginase balance, one would predict similar variability to be found in humans as well. In addition, the source of the macrophages being studied has been found to be important. Several groups have reported that human monocytes from healthy volunteers that have been differentiated or manipulated in vitro using current protocols tend not to have detectable levels of arginase and NOS enzymes, whereas MDM from diseased or stressed individuals or tissue macrophages obtained from normal, diseased, or stressed individuals do express NOS and/or arginase. Together these observations suggest that the current system of differentiating macrophages from human peripheral monocytes in vitro needs further refinement before it can be considered to be an accurate model of human macrophage behavior in vivo (63). In turn, we need to understand the differences and similarities between the different species and the cells being studied to develop experimental models that will answer some of the outstanding questions regarding macrophage M1/M2 or other activation states: What regulates macrophage activation in tissues? What mechanisms regulate macrophage plasticity and stability? How does plasticity of phenotype affect tissue macrophages? What are the full in vivo ramifications of the M1/M2 paradigm?
Further work is important to be sure that our observations of the human system in vitro are real, and not due to our cell source, measurements, or manipulations. We suggest that macrophages obtained from mice remain useful for investigating aspects of these questions in humans/human macrophages. So, although mice are not men (as Robert Koch observed), we agree with Rudolf Virchow that "Between animal and human medicine there is no dividing line – nor should there be. The object is different but the experience obtained constitutes the basis of all medicine" [Rudolph Virchow, 1821–1902].
Conflict of Interest Statement
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Anita C. Thomas is supported by funding from the British Heart Foundation. Joshua T. Mattila is supported in part by NIH RO1A103785 and Bill and Melinda Gates Foundation grants to JoAnne Flynn (University of Pittsburgh).
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21. Cameron ML, Granger DL, Weinberg JB, Kozumbo WJ, Koren HS. Human alveolar and peritoneal macrophages mediate fungistasis independently of l-arginine oxidation to nitrite or nitrate. Am Rev Respir Dis (1990) 142:1313–9. doi:10.1164/ajrccm/142.6_Pt_1.1313
22. Schneemann M, Schoedon G, Hofer S, Blau N, Guerrero L, Schaffner A. Nitric oxide synthase is not a constituent of the antimicrobial armature of human mononuclear phagocytes. J Infect Dis (1993) 167:1358–63. doi:10.1093/infdis/167.6.1358
23. Kapetanovic R, Fairbairn L, Beraldi D, Sester DP, Archibald AL, Tuggle CK, et al. Pig bone marrow-derived macrophages resemble human macrophages in their response to bacterial lipopolysaccharide. J Immunol (2012) 188:3382–94. doi:10.4049/jimmunol.1102649
24. Nicholson S, Bonecini-Almeida MG, Lapa e Silva JL, Nathan C, Xie QW, Mumford R, et al. Inducible nitric oxide synthase in pulmonary alveolar macrophages from patients with tuberculosis. J Exp Med (1996) 183:2293–302. doi:10.1084/jem.183.5.2293
25. Cromheeke KM, Kockx MM, De Meyer GRY, Bosmans JM, Bult H, Beelaerts WJF, et al. Inducible nitric oxide synthase colocalizes with signs of lipid oxidation/peroxidation in human atherosclerotic plaques. Cardiovasc Res (1999) 43:744–54. doi:10.1016/S0008-6363(99)00148-0
26. Choi HS, Rai PR, Chu HW, Cool C, Chan ED. Analysis of nitric oxide synthase and nitrotyrosine expression in human pulmonary tuberculosis. Am J Respir Crit Care Med (2002) 166:178–86. doi:10.1164/rccm.2201023
27. Rouzaut A, Subira ML, de Miguel C, Domingo-de-Miguel E, Gonzalez A, Santiago E, et al. Co-expression of inducible nitric oxide synthase and arginases in different human monocyte subsets. Apoptosis regulated by endogenous NO. Biochim Biophys Acta (1999) 1451:319–33. doi:10.1016/S0167-4889(99)00106-8
28. Babu S, Kumaraswami V, Nutman TB. Alternatively activated and immunoregulatory monocytes in human filarial infections. J Infect Dis (2009) 199:1827–37. doi:10.1086/599090
29. Kobayashi M, Jeschke MG, Shigematsu K, Asai A, Yoshida S, Herndon DN, et al. M2b monocytes predominated in peripheral blood of severely burned patients. J Immunol (2010) 185:7174–9. doi:10.4049/jimmunol.0903935
30. Martinez FO, Gordon S, Locati M, Mantovani A. Transcriptional profiling of the human monocyte-to-macrophage differentiation and polarization: new molecules and patterns of gene expression. J Immunol (2006) 177:7303–11. doi:10.4049/jimmunol.177.10.7303
31. Semnani RT, Mahapatra L, Moore V, Sanprasert V, Nutman TB. Functional and phenotypic characteristics of alternative activation induced in human monocytes by interleukin-4 or the parasitic nematode Brugia malayi. Infect Immun (2011) 79:3957–65. doi:10.1128/IAI.05191-11
32. Raes G, Brys L, Dahal BK, Brandt J, Grooten J, Brombacher F, et al. Macrophage galactose-type C-type lectins as novel markers for alternatively activated macrophages elicited by parasitic infections and allergic airway inflammation. J Leukoc Biol (2005) 77:321–7. doi:10.1189/jlb.0304212
33. Zhang X, Laubach VE, Alley EW, Edwards KA, Sherman PA, Russell SW, et al. Transcriptional basis for hyporesponsiveness of the human inducible nitric oxide synthase gene to lipopolysaccharide/interferon-gamma. J Leukoc Biol (1996) 59:575–85.
34. Shen Y, Yang S, Shi Z, Lin T, Zhu H, Bi F, et al. SeMet mediates anti-inflammation in LPS-induced U937 cells targeting NF-kB signaling pathway. Inflammation (2014). doi:10.1007/s10753-014-9984-0
35. Weinberg JB, Misukonis MA, Shami PJ, Mason SN, Sauls DL, Dittman WA, et al. Human mononuclear phagocyte inducible nitric oxide synthase (iNOS): analysis of iNOS mRNA, iNOS protein, biopterin, and nitric oxide production by blood monocytes and peritoneal macrophages. Blood (1995) 86:1184–95.
36. Christophi GP, Panos M, Hudson CA, Christophi RL, Gruber RC, Mersich AT, et al. Macrophages of multiple sclerosis patients display deficient SHP-1 expression and enhanced inflammatory phenotype. Lab Invest (2009) 89:742–59. doi:10.1038/labinvest.2009.32
37. Padgett EL, Pruett SB. Evaluation of nitrite production by human monocyte-derived macrophages. Biochem Biophys Res Commun (1992) 186:775–81. doi:10.1016/0006-291X(92)90813-Z
38. Erdely A, Kepka-Lenhart D, Clark M, Zeidler-Erdely P, Poljakovic M, Calhoun WJ, et al. Inhibition of phosphodiesterase 4 amplifies cytokine-dependent induction of arginase in macrophages. Am J Physiol Lung Cell Mol Physiol (2006) 290:L534–9. doi:10.1152/ajplung.00326.2005
39. Liu CY, Wang YM, Wang CL, Feng PH, Ko HW, Liu YH, et al. Population alterations of l-arginase- and inducible nitric oxide synthase-expressed CD11b+/CD14(-)/CD15+/CD33+ myeloid-derived suppressor cells and CD8+ T lymphocytes in patients with advanced-stage non-small cell lung cancer. J Cancer Res Clin Oncol (2010) 136:35–45. doi:10.1007/s00432-009-0634-0
40. Ozer L, Elgun S, Ozdemir B, Pervane B, Ozmeric N. Arginine-nitric oxide-polyamine metabolism in periodontal disease. J Periodontol (2011) 82:320–8. doi:10.1902/jop.2010.100199
41. Debats IB, Wolfs TG, Gotoh T, Cleutjens JP, Peutz-Kootstra CJ, van der Hulst RR. Role of arginine in superficial wound healing in man. Nitric Oxide (2009) 21:175–83. doi:10.1016/j.niox.2009.07.006
42. Suriano F, Santini D, Perrone G, Amato M, Vincenzi B, Tonini G, et al. Tumor associated macrophages polarization dictates the efficacy of BCG instillation in non-muscle invasive urothelial bladder cancer. J Exp Clin Cancer Res (2013) 32:87. doi:10.1186/1756-9966-32-87
43. Bogdan C. Response to 'Species differences in macrophage NO production are important'. Nat Immunol (2002) 3:102. doi:10.1038/ni0202-102a
44. Galkina E, Kadl A, Sanders J, Varughese D, Sarembock IJ, Ley K. Lymphocyte recruitment into the aortic wall before and during development of atherosclerosis is partially l-selectin dependent. J Exp Med (2006) 203:1273–82. doi:10.1084/jem.20052205
45. Yeramian A, Martin L, Serrat N, Arpa L, Soler C, Bertran J, et al. Arginine transport via cationic amino acid transporter 2 plays a critical regulatory role in classical or alternative activation of macrophages. J Immunol (2006) 176:5918–24. doi:10.4049/jimmunol.176.10.5918
46. Shalhoub J, Falck-Hansen MA, Davies AH, Monaco C. Innate immunity and monocyte-macrophage activation in atherosclerosis. J Inflamm (2011) 8:9. doi:10.1186/1476-9255-8-9
47. Hart PH, Bonder CS, Balogh J, Dickensheets HL, Donnelly RP, Finlay-Jones JJ. Differential responses of human monocytes and macrophages to IL-4 and IL-13. J Leukoc Biol (1999) 66:575–8.
48. Geelhaar-Karsch A, Schinnerling K, Conrad K, Friebel J, Allers K, Schneider T, et al. Evaluation of arginine metabolism for the analysis of M1/M2 macrophage activation in human clinical specimens. Inflamm Res (2013) 62:865–9. doi:10.1007/s00011-013-0642-z
49. Mills CD. Macrophage arginine metabolism to ornithine/urea or nitric oxide/citrulline: a life or death issue. Crit Rev Immunol (2001) 21:399–425. doi:10.1615/CritRevImmunol.v21.i5.10
50. Epelman S, Lavine KJ, Randolph GJ. Origin and functions of tissue macrophages. Immunity (2014) 41:21–35. doi:10.1016/j.immuni.2014.06.013
51. Hussell T, Bell TJ. Alveolar macrophages: plasticity in a tissue-specific context. Nat Rev Immunol (2014) 14:81–93. doi:10.1038/nri3600
52. Ravasi T, Wells CA, Hume DA. Systems biology of transcription control in macrophages. Bioessays (2007) 29:1215–26. doi:10.1002/bies.20683
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55. Albina JE, Mills CD, Barbul A, Thirkill CE, Henry WL, Mastrofrancesco B, et al. Arginine metabolism in wounds. Am J Physiol (1988) 254: E459–67.
56. Daley JM, Brancato SK, Thomay AA, Reichner JS, Albina JE. The phenotype of murine wound macrophages. J Leukoc Biol (2010) 87:59–67. doi:10.1189/jlb.0409236
57. Johnson JL, Newby AC. Macrophage heterogeneity in atherosclerotic plaques. Curr Opin Lipidol (2009) 20(5):370–8. doi:10.1097/MOL.0b013e3283309848
58. Hume DA. Plenary perspective: the complexity of constitutive and inducible gene expression in mononuclear phagocytes. J Leukoc Biol (2012) 92:433–44. doi:10.1189/jlb.0312166
59. Poljakovic M, Porter DW, Millecchia L, Kepka-Lenhart D, Beighley C, Wolfarth MG, et al. Cell- and isoform-specific increases in arginase expression in acute silica-induced pulmonary inflammation. J Toxicol Environ Health A (2007) 70:118–27. doi:10.1080/15287390600755075
60. Arora S, Olszewski MA, Tsang TM, McDonald RA, Toews GB, Huffnagle GB. Effect of cytokine interplay on macrophage polarization during chronic pulmonary infection with Cryptococcus neoformans. Infect Immun (2011) 79:1915–26. doi:10.1128/IAI.01270-10
61. Munder M. Arginase: an emerging key player in the mammalian immune system. Br J Pharmacol (2009) 158:638–51. doi:10.1111/j.1476-5381.2009.00291.x
62. Ochoa JB, Bernard AC, O'Brien WE, Griffen MM, Maley ME, Rockich AK, et al. Arginase I expression and activity in human mononuclear cells after injury. Ann Surg (2001) 233:393–9. doi:10.1097/00000658-200103000-00014
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64. Boutlis CS, Tjitra E, Maniboey H, Misukonis MA, Saunders JR, Suprianto S, et al. Nitric oxide production and mononuclear cell nitric oxide synthase activity in malaria-tolerant papuan adults. Infect Immun (2003) 71:3682–9. doi:10.1128/IAI.71.7.3682-3689.2003
65. Tuomisto TT, Korkeela A, Rutanen J, Viita H, Brasen JH, Riekkinen MS, et al. Gene expression in macrophage-rich inflammatory cell infiltrates in human atherosclerotic lesions as studied by laser microdissection and DNA array: overexpression of HMG-CoA reductase, colony stimulating factor receptors, CD11A/CD18 integrins, and interleukin receptors. Arterioscler Thromb Vasc Biol (2003) 23:2235–40. doi:10.1161/01.ATV.0000102551.91154.96
66. Pantano F, Berti P, Guida FM, Perrone G, Vincenzi B, Amato MM, et al. The role of macrophages polarization in predicting prognosis of radically resected gastric cancer patients. J Cell Mol Med (2013) 17:1415–21. doi:10.1111/jcmm.12109
Keywords: arginase, human macrophage, macrophage polarization, nitric oxide synthase, macrophage activation
Citation: Thomas AC and Mattila JT (2014) "Of mice and men": arginine metabolism in macrophages. Front. Immunol. 5:479. doi: 10.3389/fimmu.2014.00479
Received: 01 August 2014; Accepted: 19 September 2014;
Published online: 07 October 2014.
Edited by:Charles Dudley Mills, BioMedical Consultants, USA
Reviewed by:Charles Dudley Mills, BioMedical Consultants, USA
Sidney Morris, University of Pittsburgh, USA
Copyright: © 2014 Thomas and Mattila. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: email@example.com


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March 4, 2020 | 8:00am- 1:30pm
ServiceMaster Global Service Center
150 Peabody Place
Memphis, TN 38103
The InventHER Conference is designed to create a hands-on experience for high school girls to engage and explore opportunities in STEM focused careers. This one-day event focuses on connecting women excelling in STEM careers to the next generation of female leaders in our community
Audrey P. Willis
Innovator | Entrepreneur | Change Agent
Audrey P. Willis is a social innovator who breathes life into big ideas. Tapping into her over 15 years of innovation and entrepreneurship experience, Mayor Lee Harris appointed Audrey as the first Manager of Innovation and Performance Analysis for Shelby County Government in Tennessee. There, she is responsible for transforming government, talent, transportation, and IT to better serve the citizens of Shelby County. As a founder of businesses, Audrey has led, raised capital, and scaled businesses including most recently CodeCrew, a code school exposing underrepresented children in Memphis to computer science skills where she presently serves on the board. With a passion for talent development Audrey serves on the boards of Veritas College Prep, STS Enterprise, and LITE Memphis. She is a New Memphis Institute and Leadership Memphis Fellow. She has been recognized globally by the BBC during their "100 Women" series documenting her in "Young, geeky and black in Memphis". Audrey was named one of the "185 Black Women in Tech to Follow" for her work as a tech inclusion advocate and evangelist. More recently she was featured in a national ad campaign for Prudential Insurance, "The State of Us", highlighting her entrepreneurial contributions to making Memphis one of the cities with the highest rate of female business ownership.


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The Pillar is a recessed outdoor light constructed from stainless steel with a glass diffuser. A robust item, built to last and suitable for walkover applications. Available in square and round options.
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All information and content is subject to copyright Landscapes Direct 2020. All rights reserved.


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EU will 'catch up on two decades of technological evolution' with the new rules
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Body of Lies
Netflix 2008 62% rating
The CIA's hunt is on for the mastermind of a wave of terrorist attacks. Roger Ferris is the agency's man on the ground, moving from place to place, scrambling to stay ahead of ever-shifting events.
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YONEX Women Royal EZONE Iron Set RH 6-PW, AW, AS, SW Graph Reg-Lady – Irons are the most common type of club; a standard set of 14 golf clubs will usually contain between 7 and 11 irons, including wedges.
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Pipe leaks spur water woes in Jamshedpur
Over 5,000 households on the eastern fringe of the steel city are being deprived of drinking water for weeks because the main pipeline of a Rs 108 crore water project, commissioned barely a month ago to serve those areas, have developed leaks due indiscriminate road digging by various government and private entities.
Water released from four water towers of the Chhotagovindpur drinking water project is meant for 17,000 households of the area. But as many as 5,000 homes spread over Govindpur, Chhotagovindpur, Sarjamda and Sopdera, aren't getting water after the pipes were damaged by a panchayat samiti of Chotagovindpur, a private school and a private telecom company.
Drinking water and sanitation department executive engineer of Jamshedpur Shishir Kumar Soren said the private telecom company was laying cables at the very spot where the department had laid pipes.
"The agency assigned the work by the department had dug the ground to a depth of 3 metre to lay pipes. The private telecom company is finding it easy to lay cables there as the soil is still soft," Soren told The Telegraph, adding that they had complained to the district administration and BSNL, the mother feeder for the private company, to take necessary action.
But so far, no action has been taken.
Sources said a private school damaged the main pipeline while erecting a pillar for a boundary wall. The pipeline was damaged at another point while workers were digging the ground to build a panchayat bhavan at Sarjamda.
The Chhotagovindpur drinking water project aims to providing tap water to 24,900 households in the eastern part of the steel city, including densely-populated areas of Govindpur, Chhotagovindpur, Sarjamda, Sopodera and Parsudih. Water is being drawn from the Subernarekha river and is processed at a filtration plant near Hudco Lake.
The water department has already constructed four water towers in Sarjamda (2.5 million litres), Gadra (3 million litres), Haludvani (2.3 million litres) and Chhotagovindpur (2.2 million litres). A fifth water tower (2 million litres) is under construction at Parsudih.
As of now, 17,000 households have been given water connections. The remaining 7,900 houses will be given connections once the Parsudih water tower is ready.
Soren said the department was trying to plug the leaks, but added that it was a time consuming effort. "We have to release water through the day to detect the leaks," he said.
Local residents are naturally upset. "We are having to fetch water from far off places. The water department must take stern action against those responsible for damaging the pipeline," said Judhistir Sardar, a resident of Sarjamda.


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Salwar Size Guide
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The educational process of handicapped children is based on a good individual diagnosis and an appropiate individual educational plan. At the Don-Bosco-Schule teachers, parents and therapists - if necessary - are involved in the teaching process.
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These Are Now Buffett's 3 Favorite Banks
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After significantly cutting his stake again in Wells Fargo, there is a new pecking order among Buffett's favorite banks.
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Coronavirus infections are dropping across the UK as the latest R rate fell to between 0.8 and 1. Last week, the r value, which indicates levels of Covid-19 transmission, was between 0.9 and 1. The Government's Sage advisory group, which publishes the data, suggested the second wave of the pandemic is starting to subside across the country, as it found that the number of new infections is shrinking by between one and three per cent every day.
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As well as your day-to-day nutrition playing a crucial role in the strength of your immune system, your nutrition during and immediately after heavy exercise also plays a critical role in preventing infection, which we discuss in the next section of this resource.
At TORQ we've always been advocates of a healthy balanced diet. In a previous resource when we discussed this subject we said:
"…the constituents of a universally healthy balanced diet must never be ignored. For intestinal health and the delivery of vital micronutrients such as vitamins and minerals, you should always look to consume plenty of fibrous fresh fruit and vegetables. These foods ease the passage of all of your nutrients through the intestine, helping to keep it clean, healthy and functional. We won't focus on any specific vitamins or minerals here, because you need them all and the beauty of ensuring that you consume fresh fruit and vegetables is that you will indeed get them all.
At TORQ, we love organic food, which is why a number of our products are Soil Association Certified Organic and we advise, where possible, to choose organic options. From a pure performance perspective, the difference between 'natural' and 'organic' foods is likely to be negligible, but from a long term health perspective there are probably going to be benefits to going down the organic route with more concentrated vitamin/mineral levels and less pollution from undesirable herbicides and pesticides. It's a personal choice of course and there's a valid argument to say that 'going organic' makes no appreciable difference to health either – this is our opinion and perhaps time will tell. It would certainly be a valid approach to consume organic produce where practical – perhaps a halfway house?"
We stand by these sentiments and just as it applied to Performance Nutrition then, it applies to immunity now. The problem with popping supplements right left and centre is that it becomes easy to ignore the bigger picture of holistic health if you're just focussing on one variable – and if you focus on every possible health variable, the paranoia may lead you to popping an awful lot of pills! We would also suggest that a movement towards consuming less meat and incorporating more high quality plant-based proteins into your diet would be a healthy thing to do.
We mention some specific supplements (Vitamin C, L-Glutamine and Echinacea) in the Fighting an Infection section of these resources, because there's strong scientific evidence backing the notion of supporting your immune system with supplementation once you have become infected with a virus or bacteria, but this is entirely different to the concept of building an immune defence to bat away infection before it has a chance to take hold.
There are a couple of supplements you could consider in building strong immune defences and they are Vitamin D and Zinc. We suggest Vitamin D on the basis that colds and flu tend to do the rounds in the winter months when our skin gets to see very little sunlight (sunlight is an essential source of Vitamin D), so it makes sense from a holistic health perspective to consider supplementing with Vitamin D over the winter months. The easiest way to supplement Vitamin D is with a one-a-day Cod Liver Oil tablet and as soon as the sun comes out, you can stop taking them, assuming of course you can expose your skin to a sensible amount of sunshine. Vitamin D is inked to immune system health and if you're interested you can read more about it HERE.
A review paper by Shankar and Prasad (1998) looked at how Zinc functions within the immune system and it seems that zinc deficiency negatively affects a number of the immune system cells available to do battle. According to the authors, ensuring sufficient Zinc intake is vital in supporting these immune cells and increasing resistance to infections. With Zinc we're talking about just 15mg per day, so it's a tiny dose. If you're going to pop a pill daily throughout the year, this is perhaps the one to take if you're not confident your healthy balanced diet will deliver. Achieving 15mg of Zinc per day will be more difficult through a vegetarian or vegan diet.


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Research Repository
Effects of dietary antioxidants on training and performance in female runners
Braakhuis, AJ, Hopkins, William ORCID: 0000-0002-7066-4000 and Lowe, TE (2014) Effects of dietary antioxidants on training and performance in female runners. European Journal of Sport Science, 14 (2). 160 - 168. ISSN 1536-7290Full text for this resource is not available from the Research Repository.
Official URL: https://www.tandfonline.com/doi/abs/10.1080/174613...
|Item Type:
|Uncontrolled Keywords:
|antioxidant drinks; vitamin C; blackcurrant juice; oxidative stress; athletic performance; running
|Current > FOR Classification > 0104 Statistics
Current > FOR Classification > 1106 Human Movement and Sports Science
Current > Division/Research > College of Sports and Exercise Science
|Depositing User:
|Symplectic Elements
|Date Deposited:
|19 Sep 2018 07:46
|Last Modified:
|19 Sep 2018 07:46
|ePrint Statistics:
|View download statistics for this item
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|42 - View on Scopus
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|View Item


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Turn back the clock with a retro point-and-click adventure.
The Epic Games Store has been rife with a selection of free games since its inception, and now it's home to 2017's retro-inspired point and click game Thimbleweed Park.
It's currently free via the Epic Games Store, bringing that classic Day of the Tentacle-like flavor back to a PC near you. Based on the classic SCUMM engine adventure games, it finds you exploring the town of Thimbleweed Park with five different characters: FBI agents, a clown, a dead guy, etc. It's pretty silly, but it's also pretty fun.
The official synopsis goes something like this:
"A haunted hotel, an abandoned circus, a burnt-out pillow factory, a dead body pixelating under the bridge, toilets that run on vacuum tubes… you've never visited a place like this before.
Five people with nothing in common have been drawn to this rundown, forgotten town. They don't know it yet, but they are all deeply connected. And they're being watched."
You can download it for free right now if you have an Epic Games Store account, and it'll be available until March 7. You'll need to make sure you register for an Epic Games account, though, and make sure you don't delete it at any point. Because if you do, you'll lose out on those cool freebies up for grabs each month. Once Thimbleweed Park is gone, it'll be time for Slime Rancher to take over, so don't let this indie adventure pass you by.
Brittany has been covering games and tech for over a decade for the likes of G4, Popular Science, IGN, Empire, Kotaku, Rolling Stone and GameSpot – as well as appearing as a speaker at video game conventions such as PAX East. When she's not writing or gaming, she's looking for the next great visual novel.


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Earlier this week, President Donald Trump and his opponent, ex-vice president and Democratic hopeful Joe Biden held their second and last election debate, disputing the pandemic response, immigration, the economy, and foreign policy.
US President Donald Trump casts his ballot in early voting for the 2020 presidential election in Palm Beach, Florida. A day before, his running mate, Vice President Mike Pence also voted early, casting his ballot in Indianapolis, Indiana.
While millions of Americans have already voted by mail this year due to the coronavirus pandemic, the president and his party have stressed it may lead to election fraud and urged voters to come to the polls on 3 November.
Sourse: sputniknews.com


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How to date on dating websites
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Saga, Japan
Joe M
I travel the world and capture it's beauty on my a7rIII. I aspire to be a travel photographer. Food is an amazing way to see a new culture, and eating the weird stuff always peaks my interest. I take photos of my friends if they fall asleep. www.joeswanderlust.com
Giving Advice Photography, Japanese language, books, conversation.
Meeting Up Eat, take pictures, explore, contemplate life.Hosting
Interests adventure travel, adventures, animals, backpacking, beer, camping, culture, diving, eating, globetrotting, hiking, movies, photography, reading, road trips
Spot Map
Joe M's travel map
Save your travel map
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Celebrate Earth Day- Ongoing beach survey of plastic pollution, bioblitz and clean up, volunteer program to care for our National Park – and lets make it butt and smoke free!Read More
Bay Day 2018- Beach Survey and Clean Up
Help Celebrate the Bay with Shark Stewards and our partners Save the Bay.
Join our beach survey and clean up at Aquatic Park San Francisco in our National Maritime Heritage Area at Aquatic Park.Read More
Aquatic Park Stewards: Marine Debris & Bay Ecosystem Conservation
The Aquatic Park Stewards program with Shark Stewards is helping lead the Bay marine debris prevention effort through clean-ups and quantifying marine debris at Aquatic Park, San Francisco. Cleanups are the […]Read More
Aquatic Park Beach Clean Ups
Meet outside the Dolphin Club, west end. Join us cleaning and supporting Aquatic Park National Marine Heritage Area.Read More
Earth Day of Action With Shark Stewards
Join Shark Stewards celebrating the ocean and earth and taking action in Pacifica with the Pacifica Beach Coalition.Read More
What Color is Your Easter Egg?
"No water, no life. No blue, no green." Dr. Sylvia Earle Nearly 10 % of the planet has some level of protection, yet the oceans have only around 3%. Shark […]Read More
Beach Cleanups Aquatic Park
Clean ups are every other Saturday at Aquatic Park. Next clean up is January 28, at 0900 AM. Meet outside the Dolphin Club, west end. Join us cleaning and supporting Aquatic Park […]Read More


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User Recommended
Average Rating
Total Reviews
Products Reviewed
I wanted to try this because of all the ingredients it contains and also what it does not contain was also important to me. Dogs Love it and so do I
We have been purchasing this beyond superfood for about a year now ... I have 5 dogs from 5 lbs to about 80lbs and they all love it every kibble gone every meal and I love it cause it's good for them
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My dog had been getting tired of his food while we were living abroad, and I could only get him to eat it with hidden treats, so as soon as I came back to the U.S. I was in the market for an affordable natural dog food that he would actually like. He LOVES this one, and not just the salmon, but all the Purina Beyond flavors. He gobbles up his breakfast and dinner in a second. This is honestly the first food he's come running for when he hears it being poured into his bowl. I'm so happy he likes it!
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With two absolutely particular animals out of three enthusiastically embracing the fish dog food we feel there is no more high praise that can be given
I bought this to replace a dehydrated food that my pet didn't like...she is not much of an eater. She has gobbled up every piece morning and evening!
I just bought this for my dog who has food sensitive issues. I have gone through so many brands trying to find one that agrees with him.He went crazy when I introduced him to it and seems to being doing very well on it so far. Hoping this will be the last I have to try. (This review was collected as part of a promotion.)
it took a while to find this in my area but so happy that we did as our dog loves it and his sensitive stomach also loves it!


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- Commercial malls
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- First 10 line road


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A compact, native shrub with a rounded growth habit. Has silvery-grey foliage and produces clusters of yellow, semi-double flowers in spring. Plant in a sunny position where soils are well-drained, suitable for dry sites. Will tolerate frost and naturally grows in coastal locations. Evergreen.
Grows to about 1m x 1m.