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When exposed to hypoxia, Sema7A expression is significantly upregulated in vascular endothelial cells and promotes neutrophil adhesion and transmigration to the subendothelial layer Morote-Garcia et al. Besides, Sema7A was reported to stimulate monocyte chemotaxis and cytokine production. Suzuki et al. Given the wide expression of Sema7A and its receptors in blood cells Garcia-Areas et al. Interestingly, the reduction in the plaque size was primarily in the aortic arch exposed to d-flow.

Further studies showed that Sema7A is upregulated in the lesser curvature of mouse aortic arch endothelium, carotid artery exposed to d-flow in mice subjected to partial carotid artery ligation PCL , and human umbilical venous endothelial cells HUVECs under oscillatory shear stress, potentially associated with the inhibition of cAMP response element-binding protein CREB signaling. Mechanistic diagram of Sema7A in atherosclerosis.

Endothelial Sema7A is upregulated in response to d-flow in the lesser curvature of mouse aortic arch, which is potentially regulated by the inhibition of cAMP response element-binding protein CREB signaling. In addition to Sema3A, Sema3E, Sema4D, and Sema7A that have been extensively described for their role in the development of atherosclerosis, other members of the semaphorin family are potentially involved in atherogenesis due to their role in endothelial activation and dysfunction, neovascularization, and immune response.

Semaphorin 6A Sema6A , a transmembrane semaphorin protein expressed in endothelial cells, sustains the homeostasis of endothelial cells by regulating the vascular endothelial growth factor receptor 2 VEGFR2 expression and signaling Segarra et al. Segarra et al.

Atherosclerosis is the consequence of a chronic inflammatory response in the arterial wall and the activation of the innate immune system has been recognized to be fundamentally involved in atherogenesis Chavez-Sanchez et al. More recently, evidence has been provided that adaptive immunity is also involved in the initiation and progression of atherosclerosis Luchtefeld et al. In particular, Sema4A has been investigated extensively.

Expressed on dendritic cells and B-lymphocytes, Sema4A increases the activity of T lymphocytes Suzuki and Kumanogoh, , including cell activation and differentiation, and regulates the generation of T lymphocytes both in vitro and in vivo Kumanogoh et al.

However, there is no report on whether Sema4A participates in atherosclerosis. Sema4D has been shown to be critical for B-lymphocyte Kuklina et al. Sema3A is highly expressed on many immune cells and responsible for T-cell and dendritic cell activation, inhibiting T-cell proliferation and proinflammatory cytokines production Lepelletier et al.

Therefore, semaphorins in the immune system may provide a link between immune response and atherosclerosis. How such mechanisms of semaphorins would be started up in response to inflammatory injury in the context of atherosclerosis remains to be investigated. Semaphorins, netrins, slits, and ephrins make up the four families of neuronal guidance cues that regulate neuronal growth and migration via multiple signaling pathways.

Growing evidence suggests that members from other families participate in the regulation of atherosclerosis. Similar to Sema3A, netrin-1 was found to be involved in the initiation of atherosclerosis. Netrin-1 was detected by immunostaining in endothelial cells of the greater curvature but was significantly downregulated in endothelial cells at the lesser curvature of the aortic arch.

In vitro , netrin-1 suppresses leukocyte recruitment by inhibiting the production of proatherogenic chemokines monocyte chemotactic protein-1 MCP-1 and fractalkine. A consistent result was observed in the adhesion of monocytes to endothelial cells, suggesting that netrin-1 functions as a negative regulator of atherosclerosis van Gils et al. Besides, recent studies showed that Slit2 inhibits monocyte adhesion on activated human endothelial cells and chemotaxis of monocytes to chemokines stromal cell-derived factor-1 SDF-1 and MCP In addition, the inhibition of monocyte recruitment by supplementing Slit2 was found to delay atheroprogression in mice Mukovozov et al.

On the other hand, ephrinB2 was demonstrated to be upregulated in endothelial cells by proatherogenic stimuli and remarkably promote monocyte recruitment Poitz et al. Growing evidence indicates that semaphorins have distinct biological activities, which are not only limited to the nervous system but also branching into other pathophysiological processes, including inflammation, immune response, angiogenesis, and especially atherogenesis, of which several mechanisms work in concert to modulate the initiation, progression, and regression of atherosclerosis.

The upregulation of Sema7A and downregulation of Sema3A and Sema3E function as proatherogenic factors to induce cell adhesion and transmigration via several mechanotransduction pathways. Meanwhile, inflammatory stimulation may induce platelet Sema4D activation, enhancing leukocyte adhesion and transmigration into the subendothelium to accelerate the development of atherosclerosis. In the plaques, Sema4D may shed from activated T lymphocytes and interact with PlexinB1 on endothelial cells, causing endothelial cell activation, leukocyte migration, and intraplaque angiogenesis.

Subsequently, Sema3E interacts with PlexinD1 on macrophages to impair their emigration from advanced plaques, thereby promoting macrophage retention and chronic inflammation. Upon plaque rupture, platelet Sema4D binds to leukocytes and injured endothelial cells to exacerbate thrombosis and inflammation Figure 6. Future investigation using the state-of-the-art technologies on the role of additional semaphorin family members in atherogenesis, their potential network regulation in atherosclerosis and their association with other proatherosclerotic or antiatherosclerotic factors will not only shed light on the full picture of the functional role of semaphorins in atherogenesis but also identify potential therapeutic targets and pharmacological interventions for atherosclerosis.

Network regulation of semaphorins in atherosclerosis. When exposed to oxLDL or injured by disturbed blood flow, endothelial cells undergo major phenotypic changes, e. Accumulating immune cells including macrophages and T cells fuel the immune response, leading to the transition of a fatty streak to a complex atherosclerotic plaque.

Plaque rupture and thrombus formation may lead to fatal cardio-cerebrovascular events. Following cell infiltration and plaque formation, intraplaque Sema4D, shedding from activated T lymphocytes, interacts with PlexinB1 on endothelial cells, resulting in endothelial cell migration and intraplaque angiogenesis. In the advanced plaque, Sema3E interacts with PlexinD1 on macrophages to regulate macrophage emigration.

Upon plaque rupture, platelet Sema4D binds leukocytes as well as injured endothelial cells to support thrombus formation, exacerbating the pathological process. SH selected topics and wrote the review. LZ reviewed the manuscript and modified the content.

All authors critically read and commented on the manuscript. 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. Aghajanian, H. Semaphorin 3d and semaphorin 3e direct endothelial motility through distinct molecular signaling pathways. Alto, L. Semaphorins and their signaling mechanisms. Methods Mol. Barger, A. III, Lainey, L. Hypothesis: vasa vasorum and neovascularization of human coronary arteries, a possible role in the pathophysiology of atherosclerosis.

Basile, J. Class IV semaphorins promote angiogenesis by stimulating Rho-initiated pathways through plexin-B. Cancer Res. Bellingan, G. In vivo fate of the inflammatory macrophage during the resolution of inflammation: inflammatory macrophages do not die locally, but emigrate to the draining lymph nodes. Google Scholar. Bougeret, C. Increased surface expression of a newly identified kDa dimer early after human T lymphocyte activation.

PubMed Abstract Google Scholar. Bribian, A. Capparuccia, L. Semaphorin signaling in cancer cells and in cells of the tumor microenvironment—two sides of a coin. Cell Sci. Chapoval, S. Semaphorins 4A and 4D in chronic inflammatory diseases. Chavez-Sanchez, L. Innate immune system cells in atherosclerosis. Christensen, C. Transcription of a novel mouse semaphorin gene, M-semaH, correlates with the metastatic ability of mouse tumor cell lines.

Conrotto, P. Sema4D induces angiogenesis through met recruitment by Plexin B1. Blood , — Czopik, A. Semaphorin 7A is a negative regulator of T cell responses. Immunity 24, — Dent, E. Netrin-1 and semaphorin 3A promote or inhibit cortical axon branching, respectively, by reorganization of the cytoskeleton. Dzau, V. Vascular proliferation and atherosclerosis: new perspectives and therapeutic strategies.

Esnault, S. Endogenous semaphorin-7A impedes human lung fibroblast differentiation. PLoS One e Garcia-Areas, R. Semaphorin7A promotes tumor growth and exerts a pro-angiogenic effect in macrophages of mammary tumor-bearing mice. Semaphorin 7A: branching beyond axonal guidance and into immunity. Suppression of tumor-derived semaphorin 7A and genetic ablation of host-derived semaphorin 7A impairs tumor progression in a murine model of advanced breast carcinoma.

Geovanini, G. Atherosclerosis and inflammation: overview and updates. Gibson, M. Lipid and non-lipid factors affecting macrophage dysfunction and inflammation in atherosclerosis. Giordano, S. The semaphorin 4D receptor controls invasive growth by coupling with Met. Cell Biol. Gomez, D. Smooth muscle cell phenotypic switching in atherosclerosis. Gu, C. Semaphorin 3E and plexin-D1 control vascular pattern independently of neuropilins. Science , — Guttmann-Raviv, N. Semaphorin-3A and semaphorin-3F work together to repel endothelial cells and to inhibit their survival by induction of apoptosis.

Hall, K. Human CD, a novel leukocyte semaphorin that promotes B-cell aggregation and differentiation. Hashimoto, M. Regulation of semaphorin 3A expression in neurons of the rat spinal cord and cerebral cortex after transection injury.

Acta Neuropathol. Herman, J. Increased class 3 semaphorin expression modulates the invasive and adhesive properties of prostate cancer cells. Holmes, S. Sema7A is a potent monocyte stimulator. Hou, S. Sustained up-regulation of semaphorin 3A, Neuropilin1, and doublecortin expression in ischemic mouse brain during long-term recovery. Hu, S. Vascular semaphorin 7A upregulation by disturbed flow promotes atherosclerosis through endothelial beta1 integrin.

Jaimes, Y. Janssen, B. Structural basis of semaphorin-plexin signalling. Nature , — Ji, J. Expression and function of semaphorin 3A and its receptors in human monocyte-derived macrophages. Jonasson, L.

Regional accumulations of T cells, macrophages, and smooth muscle cells in the human atherosclerotic plaque. Arteriosclerosis 6, — Jonsson-Rylander, A. Karayan-Tapon, L. Cancer 99, — Kashiwagi, H. Negative regulation of platelet function by a secreted cell repulsive protein, semaphorin 3A.

Kikutani, H. Semaphorins in interactions between T cells and antigen-presenting cells. Kim, J. Genes Dev. Kolodkin, A. Neuropilin is a semaphorin III receptor. Cell 90, — The semaphorin genes encode a family of transmembrane and secreted growth cone guidance molecules. Cell 75, — Komatsu, M. R-Ras is a global regulator of vascular regeneration that suppresses intimal hyperplasia and tumor angiogenesis. Kuklina, E. Involvement of semaphorin Sema4D in T-dependent activation of B cells.

Kumamoto, M. Intimal neovascularization in human coronary atherosclerosis: its origin and pathophysiological significance. Kumanogoh, A. Immune semaphorins: a new area of semaphorin research. Lange, C. New eukaryotic semaphorins with close homology to semaphorins of DNA viruses. Genomics 51, — Lepelletier, Y. Immunosuppressive role of semaphorin-3A on T cell proliferation is mediated by inhibition of actin cytoskeleton reorganization. Liu, H. Cell , — Llodra, J. Emigration of monocyte-derived cells from atherosclerotic lesions characterizes regressive, but not progressive, plaques.

Luchtefeld, M. Chemokine receptor 7 knockout attenuates atherosclerotic plaque development. Circulation , — Luo, Y. Collapsin: a protein in brain that induces the collapse and paralysis of neuronal growth cones. Luque, M.

CD and plexins B2 and B1 mediate monocyte-endothelial cell adhesion and might take part in atherogenesis. Phage display identification of CD in human atherosclerotic plaque macrophages and foam cells. PLoS One 8:e Maestrini, E. A family of transmembrane proteins with homology to the MET-hepatocyte growth factor receptor. Martin-Satue, M. Identification of semaphorin E gene expression in metastatic human lung adenocarcinoma cells by mRNA differential display.

Moreno, P. Neovascularization in human atherosclerosis. Moretti, S. Semaphorin3A signaling controls Fas CD95 -mediated apoptosis by promoting Fas translocation into lipid rafts. Moriya, J. Inhibition of semaphorin as a novel strategy for therapeutic angiogenesis.

Morote-Garcia, J. Endothelial semaphorin 7A promotes neutrophil migration during hypoxia. Moulton, K. Plaque angiogenesis and atherosclerosis. Inhibition of plaque neovascularization reduces macrophage accumulation and progression of advanced atherosclerosis.

Mukovozov, I. The neurorepellent Slit2 inhibits postadhesion stabilization of monocytes tethered to vascular endothelial cells. Neufeld, G. The semaphorins: versatile regulators of tumour progression and tumour angiogenesis. Cancer 8, — Nishide, M. The role of semaphorins in immune responses and autoimmune rheumatic diseases. Nogi, T. Structural basis for semaphorin signalling through the plexin receptor. Oh, W. The respondent alleges that it sold the device to distributors who in turn, as principals, sold it to consumers for their own account.

Consequently, SARS auditors sought various information and documents from the respondent and a chain of correspondence flowed between the parties in this regard. It set out what SARS believed formed evidence in support of the findings and its legal conclusions.

However, SARS was not swayed. No revised assessments were issued regarding income tax. As a result, it invoked its rights under rule 3 of the rules of the tax court 2 and requested the commissioner to furnish reasons for the assessment. The request was contained in two letters dated 25 and 26 July , which bore 97 detailed questions targeted at the three items of assessment. SARS was not inclined to indulge the respondent. It sought an order that its requests for reasons be remitted to the commissioner for reconsideration with directions issued by the tax court to ensure that the commissioner provided adequate reasons therefor.

Thus, he argued, the response fell within the purview of s 5 2 of the Promotion of Administrative Justice Act 3 of PAJA which requires the administrator to whom the request for reasons is made to give adequate reasons in writing for the administrative action. I am now in a position to decide whether that decision has involved an unwarranted finding of fact, or an error of law, which is worth challenging. He should do so in clear and unambiguous language, not in vague generalities or the formal language of legislation.

The appropriate length of the statement covering such matters will depend upon considerations such as the nature and importance of the decision, its complexity and the time available to formulate the statement. Often those factors may suggest a brief statement of one or two pages only. But I do not think that they have the meaning that the respondent ascribes to them for present purposes.

This was so, it was argued, because the commissioner, whose perception of the facts differed dramatically to that of the respondent, had failed to disclose the reasoning process which led to his conclusion. Thus, the cogency or rationality of the reasons is not yet in the balance. It is only when the objection itself is adjudicated under judicial review that the PAJA test which the respondent wants imposed comes into play.

What it did instead, as Fyfe properly acknowledged, was reply in fine detail as to why it disagreed with the reasoning and findings and clearly had no difficulty responding to them. It explained further why no revised assessments would be issued in respect of income tax.

There is absolutely no reason why the respondent would be unable to formulate its objection, if it has any, in the circumstances. And I am inclined to agree with the commissioner that this litigation is merely a delaying tactic. Appealability and related issues. The relevant parts of the section read:. There is as yet no dispute between the parties over the merits of the assessment. It remains uncertain if the matter will proceed to the objection stage. And if further litigation should eventuate, it would have no bearing whatsoever on the order of the tax court.

Another relevant fact is that the tax court cannot alter its order; it is, therefore, final in that regard.

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It is mandatory to procure user consent prior to running these cookies on your website. View all cookies used on this site. Focus on essential Goal Setting practices in with Twigg. Twigg makes is easy to set goals, track progress, and give or receive real-time feedback. Learn More. We believe that performance management is a conversation. Request a Demo. See what our clients have to say about Sprigg! Read Reviews. Tab 1 Tab 2 Tab 3. It has a wide range of functionality, especially with regards to Performance Management including the ability to: set-up competencies, create and customize performance reviews based on company, team and individual goals, as well as, create one-on-one or team meetings in real time and to store all meeting minutes.

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Year after year as our clients grow their businesses, they renew with us to keep goals and growth tracking. What we believe We believe that Performance Management is a series of conversations that include real-time feedback and regular touch-point meetings. Everything we do at SpriggHR is about enabling those conversations by making the simplest and best of online technologies available. Helping our clients stay focused on their business priorities. It describes itself as an exclusive importer of branded electric home products which it imports from Taiwan and sells through distributors in South Africa and Namibia.

At the material time it sold an electricity-saving device called Electro Smart the device. The respondent alleges that it sold the device to distributors who in turn, as principals, sold it to consumers for their own account. Consequently, SARS auditors sought various information and documents from the respondent and a chain of correspondence flowed between the parties in this regard. It set out what SARS believed formed evidence in support of the findings and its legal conclusions.

However, SARS was not swayed. No revised assessments were issued regarding income tax. As a result, it invoked its rights under rule 3 of the rules of the tax court 2 and requested the commissioner to furnish reasons for the assessment. The request was contained in two letters dated 25 and 26 July , which bore 97 detailed questions targeted at the three items of assessment. SARS was not inclined to indulge the respondent. It sought an order that its requests for reasons be remitted to the commissioner for reconsideration with directions issued by the tax court to ensure that the commissioner provided adequate reasons therefor.

Thus, he argued, the response fell within the purview of s 5 2 of the Promotion of Administrative Justice Act 3 of PAJA which requires the administrator to whom the request for reasons is made to give adequate reasons in writing for the administrative action. I am now in a position to decide whether that decision has involved an unwarranted finding of fact, or an error of law, which is worth challenging. He should do so in clear and unambiguous language, not in vague generalities or the formal language of legislation.

The appropriate length of the statement covering such matters will depend upon considerations such as the nature and importance of the decision, its complexity and the time available to formulate the statement. Often those factors may suggest a brief statement of one or two pages only. But I do not think that they have the meaning that the respondent ascribes to them for present purposes. This was so, it was argued, because the commissioner, whose perception of the facts differed dramatically to that of the respondent, had failed to disclose the reasoning process which led to his conclusion.

Thus, the cogency or rationality of the reasons is not yet in the balance. It is only when the objection itself is adjudicated under judicial review that the PAJA test which the respondent wants imposed comes into play. What it did instead, as Fyfe properly acknowledged, was reply in fine detail as to why it disagreed with the reasoning and findings and clearly had no difficulty responding to them.

It explained further why no revised assessments would be issued in respect of income tax. There is absolutely no reason why the respondent would be unable to formulate its objection, if it has any, in the circumstances. And I am inclined to agree with the commissioner that this litigation is merely a delaying tactic. Appealability and related issues. The relevant parts of the section read:. There is as yet no dispute between the parties over the merits of the assessment.

It remains uncertain if the matter will proceed to the objection stage.