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αβT cell development depends upon serial migration of thymocyte precursors through cortical and medullary microenvironments, enabling specialized stromal cells to provide important signals at specific stages of their development. Although conventional αβT cells are subject to clonal deletion in the medulla, entry into the thymus medulla also fosters αβT cell differentiation. For example, during postnatal periods, the medulla is involved in the intrathymic generation of multiple αβT cell lineages, notably the induction of Foxp3+ regulatory T cell development and the completion of invariant NKT cell development. Although migration of conventional αβT cells to the medulla is mediated by the chemokine receptor CCR7, how other T cell subsets gain access to medullary areas during their normal development is not clear. In this study, we show that combining a panel of thymocyte maturation markers with cell surface analysis of CCR7 and CCR4 identifies distinct stages in the development of multiple αβT cell lineages in the thymus. Although Aire regulates expression of the CCR4 ligands CCL17 and CCL22, we show that CCR4 is dispensable for thymocyte migration and development in the adult thymus, demonstrating defective T cell development in Aire−/− mice is not because of a loss of CCR4-mediated migration. Moreover, we reveal that CCR7 controls the development of invariant NKT cells by enabling their access to IL-15 trans-presentation in the thymic medulla and influences the balance of early and late intrathymic stages of Foxp3+ regulatory T cell development. Collectively, our data identify novel roles for CCR7 during intrathymic T cell development, highlighting its importance in enabling multiple αβT cell lineages to access the thymic medulla.
In the thymus, the development of T cells bearing the αβTCR involves the progressive migration of immature thymocyte precursors through distinct intrathymic microenvironments formed from specialized thymic stromal cells (1, 2). Although immature CD4−8− thymocytes are localized within subcapsular regions of the thymus, cortical regions house CD4+8+ thymocytes expressing low levels of αβTCR that represent precursors of conventional, natural Foxp3+ regulatory T cells (Foxp3+ nT-Reg) and CD1d-restricted invariant NKT (iNKT). In contrast, medullary areas contain more mature CD4+ and CD8+ single-positive (SP) cells that subsequently emigrate from the thymus to populate peripheral tissues (2). Importantly, the thymus medulla and medullary thymic epithelial cells (mTEC) in particular play important functional roles during the intrathymic development of multiple αβT cell subsets. For example, although several studies have highlighted the important role played by mTEC in tolerance induction via the clonal deletion of autoreactive conventional αβT cells (3, 4), the medulla also plays a key role in generating αβT cell subsets with key regulatory functions in the immune system (5). Importantly, Foxp3+ nT-Reg development has been shown to be a multistage process (6, 7), with Foxp3−CD25+ nT-Reg precursors in particular requiring interactions with mTEC for their generation (8). In addition, we recently showed that mTEC play an important role during iNKT cell development in the thymus by providing IL-15 trans-presentation to expand newly generated iNKT cells following their CD1d-dependent positive selection in the cortex (9). Collectively, such observations indicate that a key shared step in the intrathymic development of multiple αβT cell lineages is the migration of thymocytes from the cortex to the medulla.
Many studies have highlighted the importance of chemokines and their receptors during the intrathymic migration that occurs during conventional αβT cell development. In particular, CCR7 upregulation occurs during positive selection (10, 11) and plays a key role in tolerance induction, a finding compatible with its role in controlling thymocyte access to medullary microenvironments and its effects on TCR signaling (10, 12). The role of CCR7 in cortex to medulla migration of SP thymocytes fits well with expression of both CCR7 ligands CCL19 and CCL21 in the medulla (13, 14). Interestingly, a recent study also demonstrated a role for CCR7 in the medullary localization and thymic development of γδT cells (15), suggesting that it can influence multiple T cell lineages. Indeed, Foxp3+ thymocytes have been reported to accumulate in the cortex of Ccr7−/− mice (16, 17), although the impact of CCR7 deficiency on distinct nT-Reg progenitors and more mature Foxp3+ nT-Reg stages has not been fully addressed. Moreover, the chemokine receptors controlling the intrathymic migration of iNKT cells, enabling them to access the thymus medulla during their normal development, are not clear. Although CCR7 deficiency does not totally eliminate SP thymocytes from thymic medullary regions (10, 12), pertussis toxin treatment has a more profound effect (18, 19), thereby implicating other chemokines receptors in cortex to medulla migration. In line with this, positive selection is known to alter the in vitro responsiveness of thymocytes to several chemokines including CCL17 and CCL22 (20), representing ligands for CCR4 (21). Moreover, Aire expression by MHC class IIhigh mTEC is known to influence intrathymic chemokine production (22, 23), including the ligands for CCR4 (23). Indeed, impaired CCR4-mediated thymocyte migration recently has been suggested (24) to help explain defects in the development of both conventional and Foxp3+ nT-Reg that are linked to the autoimmunity seen in Aire−/− mice (22, 25). However, although CCR4 has been studied in the peripheral immune system, notably in the context of skin-homing of T cells (26), its role during the development of distinct αβT cell lineages in the adult thymus, either individually or in combination with CCR7, has not been studied.
In this study, we show that combined cell surface expression of CCR4 and CCR7 can be used to highlight multiple developmental stages of conventional Foxp3+ nT-Reg and iNKT cell lineages in the thymus. Notably, CCR7 marks early iNKT cell subsets, whereas CCR4 identifies a narrow window during the early stages of positive selection of both conventional and regulatory SP4 T cells, prior to their CCR7 expression. In addition, through analysis of single-knockout Ccr4−/− and Ccr7−/− mice and the generation of Ccr4−/−xCcr7−/− double-knockout (DKO) mice, we show that in the adult thymus, CCR4 is dispensable for thymocyte maturation, even in the context of CCR7 deficiency. Such findings argue against intrathymic redundancy of these chemokine receptors and demonstrate that Aire-mediated control of CCL17/CCL22 expression does not underlie the defective αβT cell development seen in adult Aire−/− mice (27). Moreover, we reveal previously unreported roles for CCR7 in the development of T cell lineages that arise postnatally. Thus, CCR7 is required both in the intrathymic development of iNKT cells by controlling access to mTEC-derived IL-15 and in control of the intrathymic balance of Foxp3+CD25+ nT-Reg and their Foxp3−CD25+ precursors. Such observations collectively demonstrate new roles for CCR7 during the intrathymic development of mTEC-dependent αβT cell subsets.
Materials and Methods
Wild-type (WT) CD45.2+ C57BL/6, congenic CD45.1+ C57BL/6 (BoyJ), Rag2GFP (28), C57BL/6 Foxp3GFP reporter mice (29), Ccr4−/− (30), Ccr7−/− (31), Ccr4−/−xCcr7−/−, and Zap 70−/− (32) were bred at the University of Birmingham in accordance with Home Office Regulations. Adult mice were used at 8–12 wk of age. Embryonic mice were generated by timed pregnancies and vaginal plug detection was designated day 0. All animal experiments were performed in accordance with University of Birmingham (Local Ethical Review Panel) and national United Kingdom Home Office regulations.
Abs, flow cytometry, and cell sorting
34), where square fields of 100 × 100μm were arbitrarily set within either medullary or cortical regions, at least 100 μm from the CMJ.
Bone marrow chimeras
To investigate the role of CCR4 in T cell development in a competitive setting, mixed WT/WT and Ccr4−/−/WT chimaeras were established in a manner similar to that used to analyze the role of CCR7 (10). In brief, bone marrow cells were obtained from the femurs and tibias of Ccr4−/− (CD45.2), C57BL/6 (CD45.2), or Boy J (CD45.1) mice. Bone marrow was T cell depleted using PE-labeled anti-CD3, anti-CD4, and anti-CD8 Abs and anti-PE microbeads (Miltenyi Biotec). CD45.1+ hosts were lethally irradiated (two split doses of 450 rad) and reconstituted 24 h later with 5 × 106 T cell–depleted bone marrow cells from either CD45.2+ donors (either WT C57BL/6 or Ccr4−/−) or CD45.1+ (WT C57BL/6 Boy J) donors. Mice were analyzed 5–6 wk postreconstitution.
IL-15/IL-15Rα complex administration
Soluble IL-15/IL-15Rα complexes were prepared as described previously (9, 35Ccr7−/− mice. Mice were sacrificed 4 d postinjection, and thymic samples were analyzed for iNKT cell populations by flow cytometry.
Quantitative PCR
Quantitative PCR (qPCR) analysis of freshly sorted thymocyte populations was performed exactly as described (33). Primers used were as follows: β-actin (QuantiTect Mm Actb 1SG Primer Assay Qiagen QT00095242); Foxo1, forward, 5′-TGTCAGGCTAAGAGTTAGTGAGCA-3′, and reverse, 5′-GGGTGAAGGGCATCTTTG-3′; Klf2, forward, 5′-CTCAGCGAGCCTATCTTGCC-3′, and reverse, 5′-CACGTTGTTTAGGTCCTCATCC-3′; S1pr1, forward, 5′-AAATGCCCCAACGGAGACT-3′, and reverse, 5′-CTGATTTGCTGCGGCTAAATTC-3′; Ccl17, forward, 5′-AGTGGAGTGTTCCAGGGATG-3′, and reverse, 5′-CCAATCTGATGGCCTTCTTC-3′; and Ccl22, forward, 5′-CTGATGCAGGTCCCTATGGT-3′, and reverse, 5′-GGAGTAGCTTCTTCACCCAG-3′.
CCR4 and CCR7 combine to identify serial stages in the development of both conventional and regulatory αβT cells
To identify chemokine receptors expressed at distinct stages of positive selection that may play a role in the recruitment of developing thymocytes to medullary thymic regions, we performed a flow cytometric screen of chemokine receptor expression, coupled with analysis of chemokine mRNA expression in purified TEC compartments. Specifically, to identify chemokine receptors induced by TCR-mediated positive selection, we compared CD4+8+69+ thymocytes from WT mice, representing cells undergoing selection, with CD4+8+69− thymocytes from Zap70−/− mice, which are blocked at a preselection stage of thymocyte development (32). Of the molecules analyzed and consistent with in vitro thymocyte migration data and Ccr4 mRNA analysis (20), we found that CD4+8+CD69+ thymocytes undergoing thymic selection, but not preselection CD4+8+CD69- cells, expressed readily detectable cell surface expression of CCR4 (Fig. 1A). Interestingly, analysis of CCR4 in combination with CCR7 within CD69+ SP4 thymocytes identified CCR4+CCR7−/lo and CCR4−CCR7+ subsets, whereas CD69− SP4 thymocytes were predominantly CCR4−CCR7+ (Fig. 1B). In contrast to SP4 thymocytes, αβTCR+ SP8 thymocytes lacked CCR4 expression (data not shown). Moreover, analysis with additional markers including CD24, CD62L, Qa2, and notably levels of Rag2GFP (a direct indicator of thymocyte maturational status) (36) indicated the following developmental sequence within SP4 thymocytes: CD69+CCR4+CCR7−/lo, CD69+CCR4−CCR7+, and then CD69-CCR4−CCR7+ (Fig. 1B). Such a sequence is compatible with qPCR analysis of mRNAs linked to later stage thymocyte migration and thymus egress, including Foxo1, Klf2, and S1pr1, which were notably absent from CCR4+ thymocytes (Fig. 1C). Thus, CCR4 expression identifies a narrow window of thymocyte positive selection, with SP4 thymocytes expressing CCR4 prior to their acquisition of high levels of CCR7. In line with the pattern of CCR4 expression and consistent with a potential role during cortex to medulla migration, analysis of defined TEC compartments showed that CD80+ mTEC expressed the highest levels of Ccl17 and Ccl22 mRNA (Fig. 1D).
Similar to conventional αβT cell development, Foxp3+ nT-Reg development in the thymus can be subdivided into distinct stages (6, 7). As we recently showed that the generation of CD25+Foxp3− nT-Reg precursors is dependent on contact with mTEC (8), we next examined the expression of chemokine receptors at distinct stages in Foxp3+ nT-Reg development that might play a role in their migration to medullary microenvironments. Direct gating on CD25+Foxp3+ SP4 nT-Reg and their CD25+Foxp3− SP4 precursors showed that although the former lacked CCR4 expression, CD25+Foxp3− nT-Reg precursors contained subsets of CCR4+ and CCR7+ cells (Fig. 1E, left panels). Further analysis showed that CD25+Foxp3− nT-Reg precursors were primarily contained within CD69+CCR4+CCR7−/lo and CD69+CCR4−CCR7+ SP4 thymocyte subsets (Fig. 1E, right panels). In contrast, although CD25+Foxp3+ SP4 nT-Reg cells were largely absent from the CD69+CCR4+CCR7−/lo population, they first emerged in the CD69+CCR4−CCR7+ subset and were most abundant within CD69−CCR4−CCR7+ cells. Thus, CCR4 is expressed during early stages in the development of both conventional and regulatory SP4 T cell lineages in the thymus.
CCR4 is dispensable for both conventional and Foxp3+ regulatory T cell development in the adult thymus
Unlike Ccr7−/− mice, detailed analysis of intrathymic migration and T cell development in adult Ccr4−/− mice (30) has not been performed. Although CCR4 expression mapped to a narrow developmental window in thymic positive selection, analysis of thymocyte development in adult Ccr4−/− mice revealed no major alterations in conventional SP4 and Foxp3+ regulatory αβT cell development. This included normal proportions and numbers of CD69+Qa2low,CD69−Qa2high conventional SP4 αβT cell stages (Fig. 2A–D), as well as CD25+Foxp3− nT-Reg precursors and CD25+Foxp3+ nT-Reg (Fig. 2A–D). Furthermore, confocal quantitation of the intrathymic positioning of CD4+ and CD8+ SP thymocytes in adult thymus showed a comparable density of these cells within medullary regions of WT and Ccr4−/− thymuses (Fig. 2E), arguing against a role for CCR4 in cortex-to-medulla migration of positively selected thymocytes. Moreover, in an earlier study analyzing mice at E18 of gestation, we found no overt differences between Ccr4−/− and WT embryonic thymus lobes, in terms of total thymocyte numbers, cortical and medullary architecture and proportions of immature and mature thymocytes (Ref. 37; data not shown). Thus, initial analysis suggests that absence of CCR4 during development does not have a major impact on fetal thymus populations. As the impact of CCR7 deficiency on developing thymocytes is exacerbated in the presence of WT competitors (10), we performed similar experiments to assess the impact of CCR4 deficiency in a competitive setting. T cell–depleted mixed (50:50) bone marrow chimaeras between WT (CD45.1+):Ccr4−/− (CD45.2+), or WT (CD45.1+):Ccr4−/− (CD45.2+) partners were generated as previously described (10) and harvested after 5 wk (experimental design illustrated in Fig. 3A). The capacity of Ccr4−/− thymocytes to undergo a normal program of both conventional and Foxp3+ regulatory (Fig. 3B–D) αβT cell development was not altered in the presence of competition with WT thymocytes, whereas the ability of Ccr4−/− thymocytes to gain access to medullary thymic areas was comparable to that of WT competitors (Fig. 3E). Thus, no defects in Ccr4−/− thymocyte migration and development were revealed by the presence of competition from WT counterparts.
Because the data above indicate that CCR4 and CCR7 are expressed at similar stages of intrathymic T cell development, we investigated whether compensatory mechanisms involving these two chemokine receptors could be playing a role during the intrathymic migration and development of αβT-cells. To this end, we generated Ccr4−/−xCcr7−/− DKO mice. Adult thymuses from WT, Ccr7−/− single knockout, and Ccr4−/−xCcr7−/− DKO thymuses showed comparable overall thymocyte cellularity (Fig. 4A) and proportions and numbers of conventional αβT cell subsets, including immature (CD69+Qa2low) and mature (CD69−Qa2hi) subsets of SP4 thymocytes (Fig. 4B). Although quantitation of SP thymocyte positioning by confocal analysis of frozen tissue sections confirmed a defect in medullary accumulation of Ccr7−/− SP cells (10), no additive effect was observed in adult Ccr4−/−xCcr7−/− DKO mice (Fig. 4C). Collectively, our findings show that CCR4-mediated migration does not explain the residual migration of SP thymocytes to the medullary areas seen in adult Ccr7−/− mice and that, in the context of the steady-state adult thymus, αβT cell development occurs normally in the absence of CCR4.
A role for CCR7 during the intrathymic maturation of iNKT and Foxp3+ regulatory T cells
Our previous studies have shown that the generation of both Foxp3+ nT-Reg and CD1d-restricted iNKT-cells depends on normal thymus medulla development and interactions with mTEC (8, 9). Given the role of CCR7 during thymocyte migration, we next analyzed the intrathymic generation of these lineages in mice lacking CCR7, either individually or in combination with a lack of CCR4. In contrast to that seen with normal conventional αβT-cells (Fig. 4A, 4B), we found alterations in the CD25+Foxp3+ nT-Reg lineage in Ccr7−/− mice, which were also evident to a similar degree in Ccr4−/−xCcr7−/− DKO mice. Thus, although both CCR4 and CCR7 were expressed by CD25+Foxp3− SP4 nT-Reg progenitors (Fig. 1E), their frequency in both Ccr7−/− and Ccr4−/−xCcr7−/− DKO mice remained unaltered (Fig. 4D). In contrast, the more mature CD25+Foxp3+ nT-Reg subset showed an increase in the thymuses of Ccr7−/− mice that was evident to a comparable extent in Ccr4−/−xCcr7−/− DKO mice (Fig. 4D), findings consistent with their expression of CCR7 but not CCR4 (Fig. 1E). Although the reasons for the increase are not fully understood, analysis of the proliferation marker Ki67 by CD25+Foxp3+ nT-Reg in WT and Ccr7−/− mice (Fig. 4E) argues against the idea that it is reflective of increased nT-Reg proliferation. Thus, CCR7 plays a role in controlling the size of the intrathymic CD25+Foxp3+ nT-Reg compartment but not at the level of generation of their CD25+Foxp3− nT-Reg progenitors.
Finally, given the requirements for medullary microenvironments during intrathymic iNKT cell development, we performed flow cytometric analysis of thymocytes using PBS57/CD1d tetramers (38), in combination with the iNKT cell maturation markers CD24, CD44 and NK1.1 (39). In contrast to stages in both conventional and Foxp3+ regulatory T cell development, whereas intrathymic iNKT cell development involved expression of CCR7, CCR4 was notably absent, with the former being restricted to early developmental stages (Fig. 5A). Moreover, this pattern of chemokine receptor expression correlated with a significant reduction in the proportions and frequencies of iNKT cells in the thymi of Ccr7−/− but not Ccr4−/− mice (Fig. 5B), notably stages 1–3 of development (Fig. 5C). Such defects were only slightly enhanced by combined CCR4 deficiency (Fig. 5A–C). Thus, CCR7 is expressed by early iNKT cell subsets and plays a role in their intrathymic development. We hypothesized that the defect in iNKT cell development in the absence of CCR7 may, at least in part, be a consequence of diminished CCR7 dependent thymocyte migration into the thymic medulla, a microenvironment that provides IL15 trans-presentation during iNKT cell development (9). To test this, soluble IL-15/IL-15Rα complexes were injected into WT and Ccr7−/− hosts, and iNKT cell development was examined after 4 d. Fig. 5D shows a significant increase in the frequency of total iNKT cells in Ccr7−/− hosts receiving IL-15/IL-15Rα compared with PBS injected Ccr7−/− controls, with numbers of iNKT cells equaling those observed in WT controls receiving PBS. Moreover, IL-15/IL-15Rα treatment of Ccr7−/− mice selectively increased iNKT cell numbers at stage 3 in their developmental program (Fig. 5D), correlating with the known effects of IL-15 on later stages of iNKT cell development (40). Thus, the reduction in iNKT cells seen in this study in the thymus of Ccr7−/− mice fits well with the idea that CCR7 enables developing iNKT cells to enter the thymic medulla and gain access to mTEC, known providers of IL-15/IL-15Rα (9).
The intrathymic migration of developing thymocytes through distinct cortical and medullary microenvironments plays an important role in the generation of αβT cells. With regard to the conventional αβT cell pool, CCR7 has been shown to play a prominent role in this process, with Ccr7−/− mice displaying a reduced capacity for positively selected thymocytes to enter medullary areas of the thymus (10). Interestingly, although the medullary accumulation of SP thymocytes is reduced in Ccr7−/− mice, small medullary areas are present that contain reduced numbers of SP4 and SP8 thymocytes (10), suggesting the involvement of additional chemokine receptors. With this in mind, and as positive selection is known to induce alterations in the responsiveness of developing thymocytes to multiple chemokines (20), we sought to identify additional chemokine receptors that may be involved in cortex-to-medulla migration. In addition, given that access to the thymic medulla is important during the development of other αβT cell lineages (8, 9), we compared chemokine receptor expression and requirements of Foxp3+ nT-Reg and CD1d-restricted iNKT cells with those of conventional αβT cells. We show in this study that transient upregulation of CCR4 maps to a narrow window in the positive selection of both Foxp3+ nT-Reg and conventional SP4 αβT cells but not iNKT cells. Thus, CCR4 is expressed at stages of SP4 thymocyte maturation prior to expression of high levels of CCR7 and is absent from more mature CD69− SP4 thymocytes. However, detailed analysis performed here of adult thymus showed Ccr4−/− thymocytes developed normally, either in the steady state, when in competition with WT thymocytes, or in presence of combined Ccr7 deficiency. Such findings indicate that the defective Foxp3+ nT-Reg (22) and conventional αβT cell development (27) seen in Aire−/− mice is not because of their reduced levels of the CCR4 ligands CCL17/CCL22. Whether Aire is involved in controlling the intrathymic expression of other chemokines that are required for the normal development and migration of αβT cells is not clear. In addition, the generation and analysis of adult Ccr4−/−xCcr7−/− DKO mice in this study, and the comparable medullary accumulation of SP thymocytes in Ccr7−/− and Ccr4−/−xCcr7−/− DKO mice rules out the possibility that, in the steady-state adult thymus, CCR4 partially compensates for CCR7 with regard to thymic medulla entry. Although our findings demonstrate that CCR4 is not required for adult intrathymic T cell development and migration, its expression pattern makes it a useful cell surface marker that defines a narrow and early window in the generation of both conventional SP4 thymocytes and Foxp3+ nT-Reg.
Although CCR7 has been widely studied in the context of conventional αβT cell development (10, 12, 13, 34, 41), less is known about its possible role in the intrathymic development and migration of other T cell subsets. We found that CCR7 deficiency impacted upon both Foxp3+ regulatory and iNKT αβT cell lineages. Specifically, the intrathymic development of iNKT cells was significantly reduced, a finding that correlated with CCR7 expression by early iNKT cell subsets. Given the importance of this chemokine receptor in the intrathymic migration of conventional αβT cells, the finding that iNKT cells are dependent on CCR7 is compatible with their need to enter medullary thymic microenvironments during their development to gain access to trans-presentation of IL-15 by mTEC (9). Our findings also collectively highlight CCR7 as a common mechanism for the medullary relocation that occurs during the development of conventional αβT cells (10), γδT cells (15), and now CD1d-restricted iNKT cells. Interestingly, analysis of Foxp3+ nT-Reg development showed that absence of CCR7, either individually or in combination with CCR4, did not have an impact on the generation of CD25+Foxp3− nT-Reg precursors but did result in an increase in the numbers of CD25+Foxp3+ nT-Reg in the thymus. Thus, CCR7 plays a role in controlling the balance of intrathymic nT-Reg populations but not during early stages of their development. It is currently unclear why the thymus of Ccr7−/− mice shows an increase in the CD25+Foxp3+ nT-Reg population. One possibility is that, as with αβT cells in the neonate (13), CCR7 plays a role in the emigration of Foxp3+ nT-Reg from the thymus and that increased numbers reflect their intrathymic accumulation. Such a scenario is perhaps consistent with the previously reported mislocalization of Foxp3+ cells in the thymus of Ccr7−/− mice (16, 17). However, normal numbers of CD25+Foxp3− nT-Reg precursors in Ccr7−/− mice (the generation of which is mTEC dependent) would suggest that developing nT-Reg still have sufficient access to medullary regions in the absence of CCR7. Although this may suggest roles for additional chemokine receptors in early stages of nT-Reg development, our study also rules out a role for CCR4 in this process, despite its restricted expression pattern during positive selection.
In conclusion, our study highlights the differential importance of CCR7 in the intrathymic development of multiple αβT cell lineages, and provides a mechanism by which developing iNKT cells gain access to medullary thymic microenvironments to complete their maturation. Such findings further underline the importance of cortical and medullary thymic compartmentalization during T cell development in the thymus, showing that both innate and adaptive αβT cells share a common chemokine receptor for their intrathymic migration.
The authors have no financial conflicts of interest.
We thank the National Institutes of Health Tetramer Facility for supplying CD1d/PBS57 tetramers, Prof. Antal Rot for providing Ccr7−/− mice, and the staff at the University of Birmingham Biomedical Services Unit for animal husbandry.
This work was supported by a Medical Research Council program grant (to G.A., P.J.L.L., and E.J.J.) and a Medical Research Council New Investigator award (to W.E.J.).
Abbreviations used in this article:
- double-knockout
- Foxp3+ nT-Reg
- natural Foxp3+ regulatory T cell
- iNKT
- invariant NKT
- mTEC
- medullary thymic epithelial cell
- qPCR
- quantitative PCR
- single-positive
- thymic epithelial cell
- wild-type.
- Received April 16, 2014.
- Accepted May 29, 2014.
- Copyright © 2014 The Authors
This is an open-access article distributed under the terms of the CC-BY 3.0 Unported license.


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Run Like a Girl (1999)Run Time - 60 min. | Countries - United States |
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Synopsis by Alice Day
What's it like to be an adolescent girl? American Girls - Identity and Adolescence: Run Like a Girl answers the question by interviewing a number of young girls of varied backgrounds and allowing the viewer to listen as they talk about the issues most important to their age group. These involve appearance, insecurity, friendship, relationships with boys, eating disorders, and depressive episodes. As they express their concerns, it becomes apparent that many of the girls have worked through their problems and display a resilience that is necessary to survive the adolescent years.


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Tata Memorial Hospitals to extend cashless facility to Reliance Health Insurance customers
Kerala agency's war on cancer with tech vital, say experts
Parliamentary Committee submits report to Vice President on need for expanding cancer treatment infrastructure
Kochi to get first cancer technology incubator early next year
Kerala: Indian Navy to celebrate Navy week in Kochi
In the course of the celebrations during the Navy Week-2019, the Naval Dockyard in Mumbai organised a cruise for young cancer patients from the Tata Memorial Cancer (TMC) institute.
"As part of the Navy Week 2019 celebrations, Naval Dockyard Mumbai organised a Harbour Cruise for young cancer patients of the Tata Memorial Cancer Institute. About 90 children, with attendants, participated in the cruise. This was followed by a visit onboard INS Mumbai," a press release by the PIB (Defence Wing) on Monday read.
Earlier in the day, the children along with their attendants also visited INS Shikra for a display of various helicopters used by the Indian Navy at close quarters, according to the release.
The Navy Week-2019 is being organised in the run-up to the Navy Day, which in India is celebrated on December 4 to recognize the achievements and contribution of the Indian Navy to the country.
December 4 was chosen by the Indian Navy as on that day in 1971, during Operation Trident, the Indian Navy sank four Pakistani vessels including PNS Khaibar, killing hundreds of Pakistani Navy personnel.
(This story has not been edited by Business Standard staff and is auto-generated from a syndicated feed.)


text: Translation: On every dollar there is a piece of dirt, income from the sale of weapons... On each dollar there is a trace of blood -- Lenin "(On the road signs) To China, To Iran, To Indonesia, To Greece, To Turkey " Poster Number: PP 292 Category: Communist Culture Artist: Dolgorukov, Nikolai Andreevich — Долгоруков, Николай Андреевич Poster Notes: On the road signs: To China, To Iran, To Indonesia, To Greece, To Turkey. Poster Type: Lithograph Printer: 15th Iskra Revolutsii Typography Workshop of Glavpoligrafizdat by the Council of Ministries of the USSR. Moscow Printing date: 30-Mar-49 Printer Notes: Iskra Revolutsii means "Revolutionary Spark" Publisher: Iskusstvo (Art Publishing House), Moscow-Leningrad Publishing date: 1949 Publisher Notes: Founded in 1936, Iskusstvo was formed out of Ogiz-Izogiz, the "State Art and Literature Publishing House". Editorial Information: Editor E. Povolotskaia. Technical Information on Poster: March 30, 1949. Publication No. 8675. Volume 2 sheets of paper. Order No. 424. Price 1 ruble. Print Run: 100,000 Glavlit Directory Number: A03754 Catalog Notes: PP 292 Communist Culture b


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Global Sports Composites Market size is estimated to rise at a significant CAGR in the coming years. The growth will be particularly attributed to the persistent need for continual improvement in sports equipment performance in the sports industry.
Composites have been primarily used in various industries including sports & recreation, aerospace, architecture, automotive, energy, marine, pipe & tank, transportation, etc. However, sports industry has been the primary market adopting the innovations in composite materials. Rising government investment to strengthen the sports profile of their respective countries will also help attain promising gains to the sports composites market size in the next few years.
Request for an in-depth table of contents for this report @ https://www.gminsights.com/request-toc/upcoming/2356
Composites perform fine against fatigue and are unaffected to environmental factors such as temperature fluctuations, U.V. damage, chemical or moisture exposure. These materials are used for their unique properties such as low moisture absorption, good tactile properties, perspiration outside of the body, thermal insulation.
The main applications of these materials are in sportswear with functions including temperature control, water repellence, breathability, moisture management and ease of maintenance and transportation. The most preferred methods involved in sports composites production are spread tow technology, tape weaving technology and oblique fabric technology.
The alternatives to composites have been wood, steel, metals such as iron, aluminium, etc. The anisotropic nature of wood resulted in low resistance and the variation in properties and high moisture absorption provided various deformations. In case of aluminium, even small stresses could cause great fatigue. Composites can be up to 70% lighter compared to cast steel, whose single cubic foot weighs about 490 pounds. Carbon fibres are 70% lighter, 5 times stronger than steel and are 40% lighter than aluminium.
Owing to the aforementioned factors, composites have been the most preferred materials in sports industry. The anisotropy of composites allowed the inclusion of greater resistance in applications where it is very necessary, like skis and protective gear. Carbon fibres have been contributing to the highest preference amongst the other composites materials such as boron, titanium, ceramics, glass, and other composite alloys.
Carbon fibre usage in sports equipment offer a controlled performance to the golfers, riders, skiers and other sportspersons. The boron fibre imparts good compressive properties and it is used as a raw material in the production of golf rods. Fibber glass is highly preferred in manufacture of kayaks. Yet, the sports composites market is probable to face the challenge of higher prices of advanced fibre materials as compared to metals in the forecast period. Also, any irregularity in the supply of raw materials would fluctuate the production rates.
In recent years, the developed North American and European countries have used their advantages in the field of materials and engineering technologies to the vast extents. North America with highest production capacity of carbon fiber is likely to be the most demanding region for the sports composites industry by 2024. Asia Pacific will be the fastest growing region in the product market by 2024.
The sports industry in India and China hold huge business potential, particularly in the fields of sponsorship, marketing, goods or apparel exports. Indian television sports viewership grew 30% in the year 2015 over the year 2014. The Indian sports industry was about USD 713 million in 2015 and bears huge growth aspects. The rising Indian sports industry coupled with increasing sports viewership will in turn propel the Asia Pacific sports composites market over the forecast period.
Make an inquiry for purchasing this report @ https://www.gminsights.com/inquiry-before-buying/2356
Significant market players in resistant starch manufacturing include: Aldila Inc., Head Inc., Topkey Corporation, Amer Sports, Rossignol, Zoltek Carbon Fiber, Fischer Sports GmbH, Rockwood Composites Limited, Bell Helmets Inc., Protech Composites, Entropy Resins Inc., Saint-Gobain Vetrotex, Völkl Int. GmbH, Oxeon AB, United States Ski Pole Company, UST Mamiya, Paderson Shafts, TPT Golf Inc., Perception Kayaks, BMC Group Holding AG, etc. The industry players have been gaining the competitive edge by incorporating innovative materials and refining their designs. Bell Helmets Inc. has reduced the weight of its top-of-the-line motorcycle racing helmet by 21%, using the carbon fiber reinforcement. The helmet shell was so thin, that the company could add more padding foam to its interior to enhance the rider's safety.
Read More:
Shape Memory Polymer Market By Regional Statistics & Growth Forecast 2019-2024
Metal Foam Market Statistics: 2019-2025 Global Share Report
Rahul Varpe currently writes for fractovia. A communication Engineering graduate by education, Rahul started his journey in as a freelancer writer along with regular jobs. Rahul has a prior experience in writing as well as marketing of services and products online. Apart from being an avid reader, his other hobbies include music and sketching. He can be contacted at- [email protected] | https://twitter.com/varperahul
Global Market Insights present the report on Soundproof Curtains Market evaluates the growth trends of the industry through historical study and estimates future prospects based on comprehensive research. The report extensively provides the market share, growth, trends and forecasts for the period 2... Read More
Global Market Insights present the report on Air Humidifier Market evaluates the growth trends of the industry through historical study and estimates future prospects based on comprehensive research. The report extensively provides the market share, growth, trends and forecasts for the period 2019-2... Read More
Global Market Insights present the report on Oil-Free Air Compressor Market evaluates the growth trends of the industry through historical study and estimates future prospects based on comprehensive research. The report extensively provides the market share, growth, trends and forecasts for the peri... Read More


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This business has earned a reputation as an industry leader, through quality craftsmanship and service. Licensed to conduct business throughout the northern Black Hills, this seasonal business is perfect for the individual looking for an established brand with additional growth opportunity. The firm utilizes a unique business model, that sets it apart from the competition, while easing the process for the homeowner looking for exterior coverings and construction.
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The business leases a commercial office space and yard. Plenty of space for their employees and equipment.
Assets = $30,000
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Direct competition exists, but this business has become an industry leader through aggressive and creative marketing, along with quality craftsmanship and service.
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Jeep Renegade Latitude
Here's a great deal on a 2015 Jeep Renegade Smooth gearshifts are achieved thanks to the 2.4 liter 4 cylinder engine, and for added security, dynamic Stability Control supplements the drivetrain. Turbocharger technology provides forced air induction, enhancing performance while preserving fuel economy. A wealth of standard features means that you no longer have to sacrifice. Such as remote keyless entry, 1-touch window functionality, a tachometer, an outside temperature display, front fog lights, cruise control, a roof rack, and much more. With side curtain airbags supplementing the rest of the safety network, you can be assured that you and your passengers will experience top-tier protection. We pride ourselves on providing excellent customer service. Please don't hesitate to give us a call.


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Decorate your bathroom and dry yourself off with our luxuriously soft bath towels and hand towels. Our towels are made from brushed microfiber with a 100% cotton back for extra absorption. The top of the towel has the image printed on it, and the back is white cotton. Available in three different sizes: hand towel, bath towel, and bath sheet.
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Maria L Gaglio
December 1st, 2019
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Digital Art - A seagull is vanishing. Bird with spread wings is flying around... individually version with self painted background . Lovely light colours complete the scene.
The Berlin based artist, Melanie Viola, is drawn to urban landscape and the lines she finds in architecture. Horizontal structures crossed by vertical structures increase the impression of objects and provide a rapid dynamism. Picturesquely active passages, modern colour accents and graphic nuances find a revolutionary combination in her pictures. She is keen to explore opportunities that will allow her to combine photography and art in the way she feels happy with. In addition to the decorative digital-art images you can also find a lot of classical urban images and fascinating landscape pictures in her portfolio. So many assorted art works are displayed here which cannot be more differently... HOMEPAGE:...
Our return policy is very simple:
If you're not happy with a purchase that you made on Pixels.com, for any reason, you can return it to us within 30 days of the order date. As soon as it arrives, we'll issue a full refund for the entire purchase price. Please note - Pixels does not reimburse the outgoing or return shipping charges unless the return is due to a defect in quality.
Pixels sells thousands of pieces of artwork each month - all with a 100% money-back guarantee. We take great pride in the fact that hundreds of thousands of artists have chosen Pixels to fulfill their orders, and we look forward to helping you select your next piece!
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Live Jamberoo News with Latest Updates
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Industry leaders provide a roadmap for younger women who are pursuing careers in business aviation. The past 75 years have seen an evolution in the roles of women in aviation. From the formation of the Women Airforce Service Pilots (WASPs) in 1943 to today, women have gone from being an important but underrated element of aviation in World War II, to now holding key leadership positions.
The destruction left in the wake of Hurricane Dorian – particularly in the island nation of the Bahamas – has prompted a huge, and what undoubtedly will be a long-lasting, humanitarian response from the business aviation community. Volunteers are gearing up at airports across Florida and elsewhere, ready to start flying supplies and relief missions as early as Friday to the Bahamas.
In the wake of a pair of deadly helicopter crashes in Hawaii, U.S. Rep. Ed Case (D-Hawaii) has introduced a bill designed to impose more stringent restrictions on aircrafts flying lower than 1,500 feet above ground level.
The National Weather Service, which launched the Graphical Forecasts for Aviation tool for the continental U.S. in October 2017, recently expanded the coverage area to include the Gulf of Mexico, the Bahamas, the Caribbean, and the New York Oceanic region, including Bermuda.
Flight Design, maker of the bestselling CTLS series of Light Sport aircraft, mounted a Siemens motor on the new F2 to create an electric edition for certification. The F2e made its public flight debut in Germany in June.
The Department of Homeland Security (DHS) recently issued a security alert (ICS-ALERT-19-211-01 CAN Bus Network Implementation in Avionics) warning of cyber vulnerability in general aviation aircraft that could allow nefarious entities to alter an aircraft's altitude, airspeed and angle of attack by manipulating the aircraft's Controller Area Network (CAN) bus system.
Although hurricane season runs from June 1 to Nov. 30, the vast majority of storms form from mid-August to late October. The peak is Sept. 10. Two weeks early of peak season, Gov. Ron DeSantis issued a state of emergency for 26 Florida counties on Wednesday, including all of South Florida, as Hurricane Dorian approaches.
The 5th Annual International GIAD will be held October 5, 2019, at locations around the world, hosted by both WAI chapters and corporate members.
On Aug. 9 at the Dugway Proving Ground in Utah, a Cessna 206 under robotic control flew a first solo that lasted for two hours. Alas, there were no shirttails to cut.
An 18-month-long study attempts to find out. The vision of a drone-filled future involves tiny buzzing pieces of plastic and lithium-ion filling the skies above us.
In business aviation, sales slumps are usually broken by the arrival of new products – and the latest shipment report from the General Aviation Manufacturers Association (GAMA) shows that rule still holds true.
NOAA has awarded contracts to two U.S. manufacturers for the purchase of new aircraft that will augment the agency's environmental data-gathering capabilities.
In the 100-plus years since man first slipped the surly bonds of earth, aviation has transformed our world, providing travelers with unparalleled mobility, creating millions of jobs, enabling global economic growth and bringing people together.
Each hurricane season, our members call or email AOPA for advice on how best to prepare their aircraft for the coming storm. This subject report provides tips for anchoring your aircraft to ride out the storm as safely as possible.
On September 14, 2019 from 10 am – 3 pm, Banyan Air Service based at Fort Lauderdale Executive Airport will host a customer appreciation open house.
The National Business Aviation Association (NBAA) today thanked Dr. Peggy Chabrian for her many contributions to the aviation community, most notably as the founder, president and CEO of Women in Aviation International (WAI).
The General Aviation Manufacturers Association (GAMA) today published a mid-year industry update with the release of preliminary second quarter 2019 aircraft shipment and billings data.
Today, Volocopter, the pioneer in Urban Air Mobility, presented the design for its newest air taxi model: VoloCity. This will be the fourth-generation electrical take-off and landing aircraft (eVTOL) by the German Urban Air Mobility developer.
Orlando Sanford International Airport will host its first Thunderbirds air show in 26 years in fall 2020, sending the U.S. Air Force's high-powered air squadron and other aircraft rumbling through the Central Florida sky.
On behalf of the Federal Aviation Administration (FAA), Federal Transportation Secretary Elaine Chao awarded billions of dollars to airports around the country, including ones in Florida. In total, Florida airports will receive $3.18 billion. This funding is considered to be the fourth allotment of Airport Improvement Program (AIP) funding.


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Evy Moore
A Portrait Of My Lover - Poem by Evy Moore
A thousand cuts from life's cold betrayal
In your dark eyes a sad girl's portrayal
Pain and anguish from Love's foul sickness
To crude lies your heart bears witness
Shrouded and twisted in endless despair
Soul misconstructed and black like your hair
Screams faint but loud like the cries of the dead
Emotions bled dry by the tears you once shed
Evil comatose but subsisting inside
Alone in the darkness with a wish of suicide
Comments about A Portrait Of My Lover by Evy Moore
Amanda Martiniano (11/20/2006 7:28:00 PM)
You are officially one of my favorite poets...and inspirations (Report)Reply
Jayme Peacock (10/12/2006 7:54:00 PM)
Love the imagery, very vivid. Well-put together. (Report)Reply
Vallerie Lobell (10/12/2006 6:57:00 PM)
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Evy Moore's Other Poems
Famous Poems
- Still I RiseMaya Angelou
- The Road Not TakenRobert Frost
- If You Forget MePablo Neruda
- DreamsLangston Hughes
- Annabel LeeEdgar Allan Poe
- Stopping By Woods On A Snowy EveningRobert Frost
- IfRudyard Kipling
- Do Not Stand At My Grave And WeepMary Elizabeth Frye
- I Do Not Love You Except Because I Love YouPablo Neruda
- TelevisionRoald Dahl


text: I was born and I live in Russia in the south of western Siberia in the city of Omsk. During all my life I have been praying and asking God to grant me an opportunity to work for Him. In 2004 I was granted a Messenger's Mantle of the Great White Brotherhood and received an opportunity to bring the Words of the Masters to people. During the years 2005-2011 at certain periods of time I have been receiving messages of the Ascended Masters in a special way. I am very happy that with the help of many people the messages I received have been translated into English and the English-speaking readers can become familiar with them. The only thing the Ascended Masters want is to spread their Teaching throughout the world. The Masters give their messages with the feeling of great Love. Love has no limits. There are no boundaries between the hearts of people living in different countries, there are no boundaries between the worlds. The boundaries exist only in the consciousness of people. The Masters appeal through me to every human living on planet Earth. From Siberia with Love, Tatyana Mickushina


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Disagreements: from Theory to Practice
Disagreements: from Theory to Practice.
University of Tartu, March 11th -13th, 2020
The theoretical aspects of disagreements in fields such as ontology, logic, epistemology, and ethics have already received extensive treatment in the philosophical literature. There is a plethora of views debated at an increased level of sophistication at a very high level of abstraction (e.g. conciliations vs steadfast views in the epistemology of disagreement). What is much less discussed are the practical consequences of these theoretical models of disagreement. The conference "Disagreements: from Theory to Practice" aims to bridge the gap between theory and practice and inquire into the implications of theoretical positions for real life disagreements.
We welcome abstracts on this broad theme. Possible presentation topics include but are not limited to the following:
- Value conflicts and how to resolve them
- Laypeople and disagreement among experts
- Disagreement and scientific consensus
- Metaethical disagreements
- Moral expertise and moral disagreement
- Introspective disagreements
- Practical implications of views in the epistemology of disagreement
- Practical implications of views in the metaphysics of disagreement
- Deep disagreements and practice
- Political disagreements
Abstracts should not be longer than 500 words and be prepared for blind review. The submission deadline for abstracts is January 15th. The abstracts should be submitted to the following email address: disagreements2020 [ät] ut.ee
Workshop "Postphenomenology, Technoscience and Hermeneutics"
May 4-7, 2020, workshop "Postphenomenology, Technoscience and Hermeneutics", teacher: Don Ihde, Distinguished Professorprofessor of Stony Brook University
t least since his POSTPHENOMENOLOGY, ESSAYS IN THE POSTMODERN CONTEXT (1993), Don Ihde has been known as the founder of postphenomenology, a modified phenomenology and pragmatism focused upon analysing technologies. It continues some phenomenological style, to which is added the insights of recent Pragmatism (Rorty) and classical Dewey with his emphasis upon an experimental organism/environment model of analysis. His own focal research—for now 40+ years—has been imagining technologies with a development of "material sensitivity" in relation to human experience and skill development highly influenced by Merleau-Ponty and Dreyfus. His 1998 EXPANDING HERMENEUTICS: VISUALISM IN SCIENCE looked into the human perceptual bodily relations to science practice and MATERIAL HERMENEUTICS follows these consequences. He is proposing to destroy the duality of the "Diltheyan Divide" and develop a singular hermeneutic centered style of analysis for both natural and human sciences.
Seminars take place every day at 10:00-13:30, public lectures on May 4, 5 and 6 at 16-18. One day seminars are replaced by practical work.
Schedule of the public lectures:
4 May: Why Material Hermeneutics? Otzy the iceman as a "pure" example of Material Hermeneutics
5 May: Why Civilizations Fail.
6 May: Postscript: Postphenomenology and Material Hermeneutics.
Readings for the workshop, to be read through beforehand:
Ihde, Don 2009, Postphenomenology and Technoscience
Ihde, Don 1993, Postphenomenology. Essays in the Postmodern Context, Introduction: Postphenomenology, pp 1-8
Ihde, Don 1998, Expanding Hermeneutics. Visualism in Science, pp 1-25 and part 4 Expanding hermeneutics, pp 139-198
Verbeek, Peter-Paul 2003, Material Hermeneutics. Techne 6:3, pp 91-96
Friis et al 2012, Book Symposium on Don Ihde's Expanding Hermeneutics: Visualism in Science, Philosophy and Technology, Vol 25, No. 2, pp. 249-270
The workshop is appropriate for students of all levels, beginning from advanced bachelor level, and also all other interested people are welcome. It is necessary to register beforehand. Participation is free. It is necessary to register beforehand until 3 February 2020 on the registration form (see below). Announcement of the selection process: 10 February 2020. A limited number of travel bursaries will be available for students from abroad.Students can obtain 3 ECTS for the workshop (code of course in SIS: FLFI.03.107), for which they must submit a short essay by an appointed deadline.
To prepare for the workshop we have a reading group (FLFI.03.094) where we discuss the compulsory materials. The two courses can be taken separately.
Everyone interested is welcome to participate in either/both of them.
More detailed information will become available during the spring semester.
Contact: Ave Mets (avemets [ät] ut.ee)


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Early Learning Indiana
Closed until tomorrow, 8:00 AM(317) 636-9197Get DirectionsWebsite
1776 N Meridian St Ste A Indianapolis IN 46202 US
About Us
The mission of Early Learning Indiana is to ensure the highest level of early care and education for children in Indiana.
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The pain that you feel in your joints if you're suffering from arthritis is the direct result of inflammation. There are quite a few symptoms that could arise as a result of arthritis. First of all, patients with arthritis tend to see a reduction in their red blood cell count, which could lead to anemia. Fatigue and loss of appetite are other common symptoms of arthritis. Slight fevers may also develop as a result of the condition. Osteoarthritis develops due to naturally occurring wear and tear on your bones. Genetics and injuries can also increase your odds of developing OA early or hasten its progression.
Previously, I had reviewed hemp-based beauty topicals and THC beauty products pioneering the way for cannabis in the skincare industry. This time, I tested the diverse range of CBD oil-based beauty products, which are both potent and legally available for shipping to most states. This is the new frontier in skincare—and these companies are paving the way. Go support them before Sephora hears about this.
I am Julie Ryan, a Pain Management Blogger, Surviver and Advising expert. Besides an expert blogger, I also research about CBD oil and its medical benefits. My intention with this website is to share what I have been learning about CBD oil from my researches and experiences. So the objective of my presence here is to make you well educated about CBD oil. To provide you an industry experts advice, I also seeked help from Eileen Konieczny in writing my blogs.To provide you the reliable information, I work hard with industry experts as well. With that I strongly have faith in me that I can help anyone to get the trustable information and resource about CBD oil through this site.
CBD's legality depends on a number of factors. It is legal, but that status can vary state to state and federally. One of the determining factors for legality (or not) is whether the CBD is derived from hemp or marijuana. Hemp is a legal plant; marijuana is not. Marijuana has a high THC-content (remember: THC is psychoactive and has mind-altering properties) and is federally illegal in most places.
Endoca's commitment to social responsibility is such that it even has its own non-profit, The Endoca Foundation. This organization takes in the company's Endoca Institute of Cannabis, Natural Medicine and Sustainability, which acts as a hub for education and research into medical cannabis; it also spearheads a hardship fund designated for those who wish to investigate the potential benefits of CBD oil products for themselves but lack the funds to do so. By purchasing Endoca products, then, as the producer claims on its website, "customers are not only helping to heal others but also the world." Moreover, those customers can choose from Endoca's array of skin care products and drops and even its variety of CBD oil chewing gum – all including oil from hemp plants cultivated using the company's individual seed bank.
Populum has enough confidence in what it sells to offer a 30-day trial of purchases, with the option to send products back within that period if they don't live up to expectations. That offer speaks to the company's customer-friendly ethos – and its willingness to answer any questions that people throw at it via email, phone or live chat does so, too. One customer is quoted on Populum's website as saying that the company's risk-free guarantee is "also a testament to [its] commitment to making a high-quality product." Those wanting to see what the fuss is about can sample Populum's hemp oil tincture – which comes in 250 mg, 500 mg and 1,000 mg concentrations – and cooling gel, safe in the knowledge that they are able to get their money back if need be.
Pure Hemp Botanicals has created a soothing tea made from whole-plant ingredients. The product is packaged locally and is cruelty-free and vegan. It includes the flavors of hibiscus and apple to reduce the hemp aftertaste. The hemp used to create this oil has been organically grown, and the tea contains full-spectrum plant terpenes that are entirely natural. It is also caffeine free.
This solution is water-based and also contains CBD. Liberty Lotion claims to heal a variety of illnesses. The active ingredients of this particular product are Arnica Montana and CBD oil, both known to be very effective when it comes to treating a number of issues. Distilled water, emulsifying wax, citric acid, emu oil, stearic acid, glycerin, colloidal silver, benzyl alcohol and lavender oil are also in this solution.
This is used for pain relief and its anti-inflammatory. It can also be used for neuropathic pain. It is infused with CBD, herbs and other essential oils. It has a fragrance that you will surely like. You would also love the silky feeling of this cream on your skin. The cream is easily absorbed by the skin into the bloodstream. CBD has beneficial effects without psychotic effects.
While this should by no means be taken as direct clinical evidence that CBD for weight loss is a tried-and-true remedy for obesity, the studies mentioned above should certainly warrant more investigation into the topic. And to be sure, many folks are already using CBD products to help control their appetite and lose weight. Do a little research online for customer feedback and reviews, and you'll see what we mean.
The pain that you feel in your joints if you're suffering from arthritis is the direct result of inflammation. There are quite a few symptoms that could arise as a result of arthritis. First of all, patients with arthritis tend to see a reduction in their red blood cell count, which could lead to anemia. Fatigue and loss of appetite are other common symptoms of arthritis. Slight fevers may also develop as a result of the condition. Osteoarthritis develops due to naturally occurring wear and tear on your bones. Genetics and injuries can also increase your odds of developing OA early or hasten its progression.
CBD lotion is being used by high-performance athletes and regular people alike for its' pain-reliving properties. The nearly instantaneous pain relief that you'll receive from applying this to aching joints and muscles is more effective than the vast majority of supplements and medications that are generally used for pain relief (depending on your own unique genetic makeup, of course. Some people see huge benefits, others less).
Employee-owned Lazarus Naturals touts its commitment to ethical business practices, including pricing that aims to enable anyone to sample its range. Indeed, for veterans and those with long-term disabilities, there's even a commendable 40-percent discount on the company's wares. Customers wanting more bang for their buck could, then, try Lazarus Naturals' 3,000 mg flavorless CBD tincture, a 60 ml bottle of which costs $125, though an even more potent 6,000 mg tincture is also available. Plus, the supplier's handy and inexpensive taster packs may be a boon for more indecisive or novice CBD oil users. These packs come in regular and high-potency varieties, both of which contain samples of the company's tinctures, capsules and CBD-infused coconut oil.
CBD oil has been scientifically proven to reduce stress. It combats stress both psychologically and physically. Psychologically, it levels out your brains cortisol levels. Cortisol is the stress hormone, and high levels are associated with chronic depression, anxiety, and low immune function. Whenever you get stressed out, your body produces excessive amounts of the chemical.
Generally, CBD oil is made by combining an extract with a carrier fluid or oil. This question is best answered by looking at how the CBD oil was extracted. CBD oil can be extracted using CO2 systems or by using chemical solvents. Both methods produce a CBD oil byproduct that is then combined with a fluid like MCT oil, coconut oil, or olive oil so that it can be delivered to the body. Always check to make sure you know the CBD content of the products you purchase.


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The match between Australia and Lithuania ended with the score 90:64 (17:26, 13:22, 13:22, 21:20).
"We lost and we have to swallow the bitter pill. We know how much basketball means in Lithuania. We gave all our strength, but sometimes things are what they are, all we can do is thank those who supported and believed in us," defender Renaldas Seibutis said after the match.
Team Lithuania advanced to the quarter-finals with three victories and two defeats.
Since 1992, when Lithuania débuted in Summer Olympics, its basketball team would always make it to the top 4, except in 2012 in London. It has three Olympic bronze medals.
The game quarter-by-quarter
Since the beginning of the match, Lithuania failed to stop the Australian attacks and, at the end of the first quarter, the Australians had a nine-point lead, 26:17.
During the second quarter, Lithuania managed to score only 13 points while their rivals scored 22.
At the halftime, the faces of the Lithuanian players said everything when the scoreboard showed 48-30. It was seen that the Lithuanian players lost their way on the floor and didn't know what to do in order to return to the game.
The third quarter was a re-run of the second one when Australia won with the same score, 22-13, and increasing the gap to 27 points. During the last quarter, Lithuania finally woke up, but it was too little and too late. Team Lithuania managed to win the quarter by a single point, 21-20, but the final score was 90-64.
Despite the loss, team Lithuania got the gratefulness from the nation that hugged the disappointed players who promised to up their game in the next big tournament. After the match, the captain of the national team, Robertas Javtokas, announced his retirement from the national squad. He did not play in Wednesday's match.
Mantas Kalnietis and Antanas Kavaliauskas led the Lithuanian team with 11 points each with Jonas Valančiūnas grabbing eight rebounds and scoring six points.
First official planespotting site in Lithuania opened
A planespotting site having no analogues in the country has been officially opened in the territory...
Christmas fairs in Kaunas: from sustainable living to traditional crafts
There's more than one festive fair or bazaar going on every weekend in December, and we've...
Kaunas turns into mysterious planet for Christmas (1)
The Kaunas Christmas tree and Cosmic village will be truly special this year in another aspect –...
Unique amber with prehistoric insects will be auctioned
50 rare amber stones will be put up for bidding in Kaliningrad . The auction can be participated...
Chess queen-like Christmas tree lit up in Vilnius
The Lithuanian capital has officially started the holiday season by lighting up the Christmas tree...


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Kaspersky for Android bundles on-demand malware scans and active web protection. Additionally, it implements a host of anti-theft features including remote locate, lock and wipe, as well as the ability to sound an alarm. You can even lock apps behind a PIN as an extra preventative measure in case your phone gets stolen. Although you need a paid Kaspersky Internet Security subscription to use all its features, some basic functionality is available for free. It's a top choice for Android antivirus apps. Droid App
There are a lot of apps out there that pay lip service to security and privacy, but Signal was built from the ground up with the goal of letting people easily communicate without having to worry about being overheard. The Signal app is a complete phone and SMS client replacement (though it works just fine as a standalone app, too) for sending and receiving encrypted calls and messages. A recent update has greatly improved the app's look and feel, proving that security and usability don't have to be at odds. New Droid Apps
Your Gmail inbox is not secure. Use ProtonMail instead. This encrypted email platform offers a range of pricing options, including a free tier, so you have no excuse not to join. Security and privacy features aside, ProtonMail works like any other email app. Make sure to also check out ProtonVPN for securing your network traffic; it's an Editors' Choice for free VPNs. Droid App
Let's be honest: Tasker is very intimidating. But we recognize that this is perhaps the most powerful app available in Google Play. With it, you can script basic actions for your Android to perform when specific conditions are met—like flash the LED when you receive a text message. Learning to use something this powerful can be tricky, but the rewards seem worth it. New Droid Apps
Tasker is a glorious application if you have the patience to learn how to use the app. What it does is allow users to create custom made commands and then use them in various places. There are many apps out there that have Tasker support and you can even use Tasker to create very complex commands for NFC tags. It's difficult to truly explain what this app can do because it can do so many things. Between the apps supported, plugins you an add, and the sheer volume of stuff that you can do, there aren't many apps out there as useful as this one. IFTTT is another excellent automation app. In some cases, it may even be better than Tasker thanks to its simplicity and wide range of uses.
The library is an often-overlooked public resource (and some are modernizing quickly), but OverDrive brings it back into the fold with the newly redesigned, Libby. Supported by over 30,000 libraries worldwide, the app lets you access your local library's array of available ebooks and audiobooks. Use it to download titles, place holds on titles not yet available, and consume your borrowed content. The only requirement is a library card (or its digital equivalent). Droid Apps
You now know how to download Google Play Store to ensure you have the latest version. This method will work on almost any Android device, but there may be slight variances depending on your Android version and OEM. Do keep in mind that this will not work on Amazon Kindle Fire devices. That's an entirely different process which may also require you to root your device. Droid Apps
With its huge inventory of music, Spotify has become a mainstay for free, legal music streaming. You can shuffle through smartly selected songs, find your favorite albums, or explore one of the service's excellent playlists. Close ties with artists means that new tracks are sometimes available the same day they hit the shelves in record shops. Those still exist, right? Spotify also offers a limited free version. Droid App
Streaming services are all the rage, but if you still keep music locally, BlackPlayer is worth your attention. We particularly like the Discover option, which shuffles through 10-second previews of seldom-played tracks in your library continuously, until you decide on what to listen to next. There's also a built-in tag editor and a ton of customization options for button styles, fonts, themes, and now playing notifications. New Droid Apps
With its slick, streamlined interface, Flipboard is one of the best apps for reading the news. With it, you browse the articles, videos, podcasts, and other media that matter most to you. The app's signature magazine-style interface lets you explore the day's headlines in a gorgeous environment. The Daily Edition feature gives you the most important news along with themed stories for each day of the week. Flipboard has been one of our top picks for years, and it's easy to see why. New Droid Apps
Sophos consistently ranks high on watchdogs such as AV-Test. In addition to perfect protection, it offers a varied feature set. The highlight: the app is totally free, even from ads (Sophos makes its money from business customers). Malware protection is based on an online database that also checks the reputation of individual apps and recommends alternatives.
There is probably a greater enticement for developers because Slide ME offers the chance to target a wider international audience. Apps are also more likely to be discovered, thanks to the search filter options. Developers can set prices and choose to offer their apps for free. There is also a Slide ME ad network, and it is even possible to get a greater percentage of the revenue generated by your app with Slide ME — potentially more than the standard 70 percent you'll get through most other app stores. Droid Apps
Initially, we weren't going to put any launchers on this list. Nova Launcher seems to be extend beyond what normal launchers are. It's been around for years, it's been consistently updated, and thus it's never not been a great option for a launcher replacement. It comes with a host of features, including the ability to backup and restore your home screen set ups, icon theming for all of your Android apps, tons of customization elements for the home screen and app drawer, and more. You can even make it look like the Pixel Launcher if you want to. If you go premium, you can tack on gesture controls, unread count badges for apps, and icon swipe actions. Those looking for something simpler may want to try Lawnchair Launcher, Hyperion Launcher, and Rootless Launcher as well.
Ben Moore is an Analyst for PCMag's software team. He has previously written for Laptop Mag, Neowin.net, and Tom's Guide. Ben holds a degree in New Media and Digital Design from Fordham University at Lincoln Center, where he served as the Editor-in-Chief of The Observer, the student-run newspaper. He spends his free time taking photos and reading b... See Full Bio Android App
Untappd helps you record each brew you try along with a rating and tasting notes. Think of it as Swarm for beer! No more staring at a tap list, trying to remember which ones you've had before. The app also has a vibrant community of beer drinkers that can point you toward new discoveries and an extensive list of beers. It's also a handy way to find your favorite brew near your current location. While it's not perfect, it can change the way you think of beer.
Asana is the 800-pound gorilla of task management for teams, dwarfing other popular services like the capable Trello. Asana is all about workflows and checkbox tasks that can be assigned to individuals. The Android app lets you take your tasks on the go and offline, syncing your progress when you're back on the network. It's a powerful tool with an excellent interface, and new features are added regularly.
If an event isn't being organized on Facebook, it's probably being done through Eventbrite. This service makes it easy to manage invites, RSVPs, and even ticket purchasing. Attendees get handy reminders about the events they planned to attend, organizers can see guest lists. We particularly like that Eventbrite can generate QR codes, making event check-in a breeze. Droid App
With its slick, streamlined interface, Flipboard is one of the best apps for reading the news. With it, you browse the articles, videos, podcasts, and other media that matter most to you. The app's signature magazine-style interface lets you explore the day's headlines in a gorgeous environment. The Daily Edition feature gives you the most important news along with themed stories for each day of the week. Flipboard has been one of our top picks for years, and it's easy to see why. New Droid Apps
Facebook Messenger Kids might seem like it's built for kids, but it's really built for parents. With the app, parents have control over their kids' contact list, and messages can't be deleted — meaning that parents can check in to take a look whenever they want. Of course, there are some pretty sweet features for kids, like kid-appropriate stickers, GIFs, emojis, and so on.
With millions of apps on the Play Store, searching for the best among them is daunting. Whether you need a great weather app or a new tool to speed up your productivity, there are always dozens available. We've spent hours curating the top Android apps for every purpose, and compiled only the best of them into one list. We've just added a mobility section for the best apps from getting you from A to B in 2019.
Though its interface is a bit tricky, Stitcher connects you with just about every podcast out there. Podcasts are organized by subject, or you can just search for a familiar title. Shows can, in turn, be organized into playlists for a continuous stream. Once you've exhausted all your favorites, let Stitcher recommend something new. With a special emphasis on sources like NPR, CNN, and ESPN, Stitcher has a little bit of everything. Droid App
You'll find a huge repository of free apps here for various platforms including Android. GetJar is fairly basic, but apps are divided into categories and sub-categories within the store to make it easier to find what you want. You'll also find likes or dislikes and comments on each app from users. GetJar can also connect with Facebook. On the downside, you will encounter sponsored recommendations, but they are clearly marked. Droid App


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Re: Typing texts at the wheel is incredibly dangerous
Holding your phone IS illegal. Eating or drinking while driving IS illegal. Tapping on your laptop, reading the newspaper or a book, or even watching a movie on your laptop or cellphone while driving IS illegal. ( Yes, i've seen all of these happen, unfortunately ).
What I find interesting is that you can be fined for eating an apple, but smoking a cigarette (which involves picking one out of a pack, and fiddling with lighter and ashtray) is perfectly OK. That, to me, is really BS.


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Top recommendations from localsFrom sightseeing to hidden gems, find out what makes the city unique with the help of the locals who know it best.
"Great place to eat and then go across the road and enjoy the River Murray views. If you go at night you will be sure to see the possums in the trees in the park across the road from the Hotel."
- 2 locals recommend
"Great meals and right on the foreshore of the River Murray. Eat on the balcony and enjoy the views. Great nightlife on weekends for the young at heart."
- 5 locals recommend
Point of Interest
"Great Indian Food, the Renmark Club is also good with a deck overlooking the river. The Renmark Hotel also serves good meals."
- 2 locals recommend
"It is next to the river. There is a library, playground and a large shady, grassed area for picnics or sitting and watching the river.It has a path which can be used for walks or jogging."
- 1 local recommends
Natural Feature
"There are many different types of native birds to be seen. It is along a creek and is pretty and quiet."
- 1 local recommends
Shopping Mall
"Woolworths, Big W, Rockmans, Subway, Reject shop, Chemist, Flightcentre, Optometrist, Telstra Shop, Optus shop, Art Gallery with work of local artists, "
- 1 local recommends
Clothing Store
"Lucca provides a broad range of clothes for women from 15 to 50. Their clothes are reasonably priced and very unique. There is something for every one."
- 1 local recommends
"information about the local area, booking for river cruise and bus terminal"
- 2 locals recommend
Natural Feature
"There are a huge variety of birds on this walk along the creek. It is very close to the town. For a walk or jog there is a path along the river that goes to the Paringa Bridge and past The Renmark Club."
- 1 local recommends


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