luc Maroteaux

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The Control of Vascular No Production by the 5-HT2B Receptor is Involved in Blood Pressure Regulation

Journal of Hypertension, Jun 1, 2004

Serotonin and Cardiovascular Diseases

Elsevier eBooks, 2019

Serotonergic dysfunction is mainly associated with neuropsychiatric and cardiovascular disorders ... more Serotonergic dysfunction is mainly associated with neuropsychiatric and cardiovascular disorders but has also been linked with many other pathological conditions. This review provides an overview of the recent pharmacological developments involving 5hydroxytryptamine (5-HT; serotonin), released from blood platelets, in the human cardiovascular system. The acute cardiovascular response to serotonin, named the Bezold-Jarish reflex, leads to intense bradycardia associated with atrioventricular block, and involves 5-HT 3 , 5-HT 1B , 5-HT 7 and 5-HT 2A/2B receptors. The contribution of serotonin and its receptors (5-HT 4 and 5-HT 2A/2B ) in cardiac development, in cardiovascular tissue remodeling, with a particular emphasis on cardiac hypertrophy, fibrosis and failure, in valve degeneration, is explored in this review. Some new aspects of 5-HT 1B , 5-HT 7 and 5-HT 2A/2B receptors in vasomotor tone regulation and the interaction between endothelial and smooth muscle cells are also be discussed. As well, serotonin, its transporter, 5-HT 1B , and 5-HT 2B receptors participation in pulmonary hypertension and pulmonary vascular remodeling are presented. Finally, the contribution of bone marrow derived endothelial progenitors in the pathogenesis of pulmonary hypertension and interactions with bone morphogenic protein type 2 receptor signaling are highlighted. The aim of this review is thus to emphasize the vascular, cardiac and pulmonary effects caused by serotonin receptor activation, and to highlight their possible prevention by the development of new drugs targeting this system. cardiovascular regulation. In periphery, this mediator contributes to control many cardiovascular functions since, in addition to effects on platelet aggregation and hematopoietic stem cells differentiation, serotonin can modulate blood pressure, heart rate and respiration. Finally, it has a major pathophysiological role in metastatic neuroendocrine carcinoid tumors. Carcinoid tumors are neuroendocrine tumors originating from enterochromaffin cells that develop within the digestive tract and secrete large amounts of serotonin . The cardiovascular functions of serotonin, often unrecognized, have recently been re-evaluated in particular in cerebral and pulmonary circulatory beds with development of "serotonergic" drugs for treatment of migraines and likely in a near future in primary pulmonary arterial hypertension (PAH) and valvular heart diseases (VHD). It may be surprising that a unique mediator possesses so many different functions. This diversity can be explained by the fact that serotonin acts through numerous receptors. , working on guinea pig ileum, proposed that serotonin acts, on two pharmacologically different receptors, M (muscarin-sensitive) and D (dibenzylinsensitive) receptors located on smooth muscles and neurons respectively. During early 90s, identification of serotonin receptor genes has progressed considerably with gene cloning and sequencing techniques. It now appears that there are 15 different receptor genes, divided into four subtypes: 5-HT 1/5 , 5-HT 2 , 5-HT 3 , and 5-HT 4/6/7 . These receptors are grouped according to their intracellular coupling mode. 5-HT 3 receptors are ion channels whereas all others are G-protein coupled receptors: Gi proteins for 5-HT 1/5 , Gs for 5-HT 4/6/7 and finally Gq for 5-HT 2 . Subfamilies have been distinguished within these categories. Thus, for example, there are three members in 5-HT 2 receptors (5-HT 2A , 2B , 2C ). In this chapter, we will present current state of knowledge about roles of serotonin, its transporter and its receptors in physiology and pathology of cardiovascular system. Serotonin is present at early stages of embryogenesis and is synthesized during zygotic cleavages, gastrulation and neurulation . Serotonin appears necessary for the formation of heart: immunolabeling studies with antibody directed against serotonin revealed that the myocardium of mice embryos (E8-11) grown in the presence of serotonin can take up this mediator avidly. Thereafter, uptake decreases to be restricted to endocardial cushions. Thymidine incorporation experiments show that serotonin, reuptake inhibitors, fluoxetine or sertraline, inhibit proliferation of cardiac mesenchyme, endocardium and myocardium especially at E9 (before cushion formation). Serotonin therefore participates in cardiac morphogenesis before and during formation of endocardial cushions (see below). In mice, expression of 5-HT 2A and 5-HT 2B receptors is observed during active phases of morphogenesis in neuroepithelium, notochord, somites, neural crest cells, craniofacial mesenchyme, myocardium and endocardial cushions, dental germs, hair follicles, cartilage and striated muscles . In mouse embryo culture experiments, antagonists of high affinity for 5-HT 2B receptor such as ritanserin (but not ketanserin, a 5-HT 2A preferential antagonist) cause morphological deficits in cephalic, cardiac and neural tube regions. These treatments interfere with migration of neural crest cells and cause their apoptosis . With regard to heart formation, these treatments lead to an abnormal organization of sarcomeres of subepicardial layer and to the absence of myocardial trabeculation.

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Hypertension artérielle pulmonaire, moelle osseuse, précurseurs endothéliaux et sérotonine

Biologie Aujourd'hui, 2016

Titre courant : Sérotonine et hypertension artérielle pulmonaire Résumé -La sérotonine et les cel... more Titre courant : Sérotonine et hypertension artérielle pulmonaire Résumé -La sérotonine et les cellules souches dérivées de la moelle osseuse participent de concert à l'hypertension artérielle pulmonaire. Notre travail a montré que l'absence du récepteur 5-HT 2B génère des altérations permanentes de la composition du sang et de la moelle osseuse dans les lignées myéloïdes et, en particulier, dans les cellules progénitrices endothéliales. Dans l'hypertension artérielle pulmonaire (HTAP), les récepteurs 5-HT 2B portés par les cellules de la moelle osseuse jouent un rôle fondamental. Ils contribuent à la différenciation / prolifération / mobilisation des précurseurs endothéliaux de la moelle. Ces cellules dérivées de la moelle osseuse ont une fonction critique dans le développement de l'HTAP et du remodelage vasculaire pulmonaire. Ces données indiquent que les cellules de la moelle osseuse jouent un rôle clé dans la pathogenèse de l'HTAP et suggèrent que les interactions entre la sérotonine et BMPR2 (bone morphogenic protein type 2 receptor) pourraient se situer dans la moelle osseuse. Mots-clés : Plaquettes sanguines / récepteurs de la sérotonine / précurseurs endothéliaux / cellules souches hématopoïétiques / hypertension artérielle pulmonaire.

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5-HT2B Receptors

Springer eBooks, 2021

The Receptors book Series, founded in the 1980's, is a broad-based and wellrespected series on al... more The Receptors book Series, founded in the 1980's, is a broad-based and wellrespected series on all aspects of receptor neurophysiology. The series presents published volumes that comprehensively review neural receptors for a specific hormone or neurotransmitter by invited leading specialists. Particular attention is paid to in-depth studies of receptors' role in health and neuropathological processes. Recent volumes in the series cover chemical, physical, modeling, biological, pharmacological, anatomical aspects and drug discovery regarding different receptors. All books in this series have, with a rigorous editing, a strong reference value and provide essential up-to-date resources for neuroscience researchers, lecturers, students and pharmaceutical research.

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Pharmacological mechanisms involving serotonergic 5-HT2 receptors in drug-induced valvulopathy

European Heart Journal, Aug 2, 2013

ABSTRACT Recent safety data emphasized the close link between chronic therapeutic use of serotone... more ABSTRACT Recent safety data emphasized the close link between chronic therapeutic use of serotonergic 5-HT2B receptors agonists and cardiac valve degeneration but the pharmacological mechanisms have not been clearly understood. Histopathological lesions show cell proliferation with extracellular matrix deposition that can drive to valvular leaflets stenosis, severe cardiac insufficiency and surgical valve replacement. In this work, we evaluated the effects of chronic administration of nordexfenfluramine (NDF) a highly selective 5-HT2B receptors agonist, on mice mitral valve leaflets. Adult male 129sv mice (aged 12 weeks) were randomly assigned in 10 different groups and underwent a subcutaneous micro-osmotic pump implantation delivering either NDF (1mg/kg/day) or vehicle (sterile water) for 4 weeks. Four groups were designed with wild-type mice: vehicle; NDF; NDF + ritanserine (2mg/kg/day); NDF + SB 206553 (1mg/kg/day) and two groups of each type of these knockout mice: 5-HT2A-/-; 5-HT2B-/-; 5-HT2A/2B-/- receiving either NDF or vehicle. Mice were monitored by regular body weight, food and water intake, blood pressure measurement and echocardiography. At the end of the experiment, urinary 5-HIAA was also monitored to appreciate serotonin synthesis. Hearts were harvested for histological analysis to quantify mitral valve leaflets thickness and cellularity. The comparison between wild-type vehicle treated and NDF mice didn&#39;t reveal any significant differences in terms of food or water intake, body weight, echocardiography parameters, beside of a slight blood pressure increase from 115±5 mmHg vs 124±6 mmHg for NDF treated mice. We also pointed out a significant increase of urinary amounts of 5-HIAA (52.65±7.38 mcg/L vs 74.18±4.53 mcg/L. Histological analysis of the heart revealed significant increased mitral valve leaflets thickness (35±6 micrometer vs 60±6 micrometer) and cellularity in NDF group (+20% vs control). These histological lesions are prevented in part by a pharmacological co-treatment with ritanserine and SB 206553 but not in NDF transgenic 5-HT2A-/- treated mice. However NDF induced mitral lesions were fully prevented in 5-HT2B -/- and 5-HT2A/2B -/- mice. In conclusion, we demonstrate the crucial role of 5-HT2B receptors in NDF induced valvulopathy. Experiments are currently evaluating cellular and molecular mechanisms.

The role of 5-HT<sub>2B</sub>receptors in mitral valvulopathy: bone marrow mobilization of endothelial progenitors

British Journal of Pharmacology, Oct 15, 2017

Valvular heart disease (VHD) is highly prevalent in industrialized countries. Chronic use of anor... more Valvular heart disease (VHD) is highly prevalent in industrialized countries. Chronic use of anorexigens, amphetamine or ergot derivatives targeting the 5-HT system is associated with VHD. Here, we investigated the contribution of 5-HT receptors in a model of valve degeneration induced by nordexfenfluramine, the main metabolite of the anorexigens, dexfenfluramine and benfluorex. Nordexfenfluramine was infused chronically (28 days) in mice ((WT and transgenic Htr 2B -/-, Htr 2A -/-, and Htr 2B/2A -/-) to induce mitral valve lesions. Bone marrow transplantation was also carried out. Haemodynamics were measured with echocardiography; tissues and cells were analysed by histology, immunocytochemistry, flow cytometry and RT -qPCR. Samples of human prolapsed mitral valves were also analysed. Chronic treatment of mice with nordexfenfluramine activated 5-HT 2B receptors and increased valve thickness and cell density in a thick extracellular matrix, mimicking early steps of mitral valve remodelling. Lesions were prevented by 5-HT 2A or 5-HT 2B receptor antagonists and in transgenic Htr 2B À/À or Htr 2A/2B À/À mice. Surprisingly, valve lesions were mainly formed by numerous nonproliferative CD34 + endothelial progenitors. These progenitors originated from bone marrow (BM) as revealed by BM transplantation. The initial steps of mitral valve remodelling involved mobilization of BM-derived CD34 + CD31 + cells by 5-HT 2B receptor stimulation. Analysis of human prolapsed mitral valves showing spontaneous degenerative lesions, demonstrated the presence of non-proliferating CD34 + /CD309 + /NOS3 + endothelial progenitors expressing 5-HT 2B receptors. BM-derived endothelial progenitor cells make a crucial contribution to the remodelling of mitral valve tissue. Our data describe a new and important mechanism underlying human VHD.

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5-HT2B Receptor, the Heart and Blood Vessels

5-HT2B Receptors, 2021

NKD, a developmentally regulated tachykinin receptor in Drosophila

Journal of Biological Chemistry, 1992

-Restriction endonucleases, avian myeloblastosis virus reverse transcriptase, DNA polymerase I, T... more -Restriction endonucleases, avian myeloblastosis virus reverse transcriptase, DNA polymerase I, T4 polynucleotide kinase, T4 DNA ligase, and T3 or T7 RNA polymerase were purchased from Bethesda Research Laboratories, New England Biolabs, Boehringer purchased from U. S. Biochemical Corp. Taq polymerase for PCR' Mannheim, and Stratagene. Sequenase and sequencing reagents were was purchased from Perkin-Elmer Cetus, and reactions were performed in a Perkin-Elmer Cetus cycler. Peptides were purchased from Bachem, when available. Other chemicals were of the highest purity available. 32P-Labeled nucleotides were from Du Pont-New England Nuclear, and my~-[~H]inositol was from Amersham Corp.

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Serotonin 2B Receptors in Mesoaccumbens Dopamine Pathway Regulate Cocaine Responses

The Journal of Neuroscience, 2017

Addiction is a maladaptive pattern of behavior following repeated use of reinforcing drugs in pre... more Addiction is a maladaptive pattern of behavior following repeated use of reinforcing drugs in predisposed individuals, leading to lifelong changes. Common among these changes are alterations of neurons releasing dopamine in the ventral and dorsal territories of the striatum. The serotonin 5-HT2Breceptor has been involved in various behaviors, including impulsivity, response to antidepressants, and response to psychostimulants, pointing toward putative interactions with the dopamine system. Despite these findings, it remains unknown whether 5-HT2Breceptors directly modulate dopaminergic activity and the possible mechanisms involved. To answer these questions, we investigated the contribution of 5-HT2Breceptors to cocaine-dependent behavioral responses. Male mice permanently lacking 5-HT2Breceptors, even restricted to dopamine neurons, developed heightened cocaine-induced locomotor responses. Retrograde tracing combined with single-cell mRNA amplification indicated that 5-HT2Breceptor...

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Cardiovascular remodeling and the peripheral serotonergic system

Archives of Cardiovascular Diseases, 2017

Plasma 5-hydroxytryptamine (5-HT; serotonin), released from blood platelets, plays a major role i... more Plasma 5-hydroxytryptamine (5-HT; serotonin), released from blood platelets, plays a major role in the human cardiovascular system. Besides the effect of endogenous serotonin, many drugs targeting serotonergic receptors are widely used in the general population (antiobesity agents, antidepressants, antipsychotics, antimigraine agents), and may enhance the cardiovascular risk. Depending on the type of serotonin receptor activated and its location, the use of these compounds triggers acute and chronic effects. The acute cardiovascular response to 5-HT, named the Bezold-Jarish reflex, leads to intense bradycardia associated with atrioventricular block, and involves 5-HT3, 5-HT1B/1D, 5-HT7 and 5-HT2A/2B receptors. The chronic contribution of 5-HT and its receptors (5-HT4 and 5-HT2A/2B) in cardiovascular tissue remodeling, with a particular emphasis on cardiac hypertrophy, fibrosis and valve degeneration, will be explored in this review. Finally, through the analysis of the effects of sarpogrelate, some new aspects of 5-HT2A receptor pharmacology in vasomotor tone regulation and the interaction between endothelial and smooth muscle cells will also be discussed. The aim of this review is to emphasize the cardiac side effects caused by serotonin receptor activation, and to highlight their possible prevention by the development of new drugs targeting this system.

New therapeutic opportunities for 5-HT2 receptor ligands

Pharmacology & Therapeutics, 2017

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Hypertension artérielle pulmonaire, moelle osseuse, précurseurs endothéliaux et sérotonine

Biologie Aujourd'hui, 2016

Reçu le 18 mars 2016 Résumé -La sérotonine et les cellules souches dérivées de la moelle osseuse ... more Reçu le 18 mars 2016 Résumé -La sérotonine et les cellules souches dérivées de la moelle osseuse participent de concert à l'hypertension artérielle pulmonaire. Notre travail a montré que l'absence du récepteur 5-HT2B génère des altérations permanentes de la composition du sang et de la moelle osseuse dans les lignées myéloïdes et, en particulier, dans les cellules progénitrices endothéliales. Dans l'hypertension artérielle pulmonaire (HTAP), les récepteurs 5-HT2B portés par les cellules de la moelle osseuse jouent un rôle fondamental. Ils contribuent à la différenciation/prolifération/mobilisation des précurseurs endothéliaux de la moelle. Ces cellules dérivées de la moelle osseuse ont une fonction critique dans le développement de l'HTAP et du remodelage vasculaire pulmonaire. Ces données indiquent que les cellules de la moelle osseuse jouent un rôle clé dans la pathogenèse de l'HTAP et suggèrent que les interactions entre la sérotonine et BMPR2 (bone morphogenic protein type 2 receptor) pourraient se situer dans la moelle osseuse.

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Serotonin 2B receptor slows disease progression and prevents degeneration of spinal cord mononuclear phagocytes in amyotrophic lateral sclerosis

Acta neuropathologica, Jan 7, 2016

Microglia are the resident mononuclear phagocytes of the central nervous system and have been imp... more Microglia are the resident mononuclear phagocytes of the central nervous system and have been implicated in the pathogenesis of neurodegenerative diseases such as amyotrophic lateral sclerosis (ALS). During neurodegeneration, microglial activation is accompanied by infiltration of circulating monocytes, leading to production of multiple inflammatory mediators in the spinal cord. Degenerative alterations in mononuclear phagocytes are commonly observed during neurodegenerative diseases, yet little is known concerning the mechanisms leading to their degeneration, or the consequences on disease progression. Here we observed that the serotonin 2B receptor (5-HT2B), a serotonin receptor expressed in microglia, is upregulated in the spinal cord of three different transgenic mouse models of ALS. In mutant SOD1 mice, this upregulation was restricted to cells positive for CD11b, a marker of mononuclear phagocytes. Ablation of 5-HT2B receptor in transgenic ALS mice expressing mutant SOD1 resul...

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Contribution of serotonin to cardiac remodeling associated with hypertensive diastolic ventricular dysfunction in rats

Journal of Hypertension, 2015

Objective: Left-ventricular hypertrophy and interstitial fibrosis are the main pathophysiological... more Objective: Left-ventricular hypertrophy and interstitial fibrosis are the main pathophysiological factors of heart failure with preserved ejection fraction. Blockade of the serotonin 5-HT 2B receptor (5-HT 2B R) has been shown to reduce cardiac hypertrophy, oxidative stress, and extracellular cell matrix activation. In this study, we evaluated the effects of the 5-HT 2B R blockade, on hemodynamic and cardiac remodeling, in spontaneously hypertensive rats (SHRs) that display a diastolic dysfunction with preserved ejection fraction. Method: Thirty-seven-week-old SHRs were randomized in four groups receiving either saline, the selective 5-HT 2B R antagonist RS-127445 (1 mg/kg per day), a calcium channel blocker nicardipine (6 mg/kg per day), or RS-127445 R nicardipine. During the 14 weeks of treatment period, cardiac function and blood pressure were monitored by echocardiography and tail-cuff. Finally, electrocardiograms and invasive hemodynamics were obtained before blood collection. Heart was analyzed for morphology and mRNA expression. A complementary study evaluated the cardiac and vascular effects of serotonin on wild-type and mice knockout for the 5-HT 2B R (Htr 2B À/À ) and/or the 5-HT 2A R (Htr 2A À/À ). Results: Despite the left ventricular 5-HT 2B R overexpression, 5-HT 2B R blockade by RS-127445 did not affect left ventricular hypertrophy and fibrosis in SHRs. This antagonist did not improve diastolic dysfunction, neither alone nor in combination with nicardipine, although it induced plasma brain natriuretic peptide decrease. Moreover, RS-127445 amplified subendocardial fibrosis and favored left ventricular dilatation. Finally, a subendocardial left ventricular fibrosis was induced by chronic serotonin in wild-type mice, which was increased in Htr 2B À/À animals, but prevented in Htr 2A À/À and Htr 2A/2B À/À mice, and could be explained by a contribution of the endothelial 5-HT 2B Rs to coronary vasodilatation. This work is the first to identify a cardioprotective function of the 5-HT 2B R in an integrated model of diastolic dysfunction with preserved ejection fraction.

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[The 5-HT2B receptor: a main cardio-pulmonary target of serotonin]

Journal de la Société de biologie, 2004

In agreement with previous data in the literature, our results indicate that serotonin, a monoami... more In agreement with previous data in the literature, our results indicate that serotonin, a monoamine neurotransmitter, can also regulate cell proliferation, cell movements and cell differentiation. We have recently shown that serotonin is required for embryonic heart development. Genetic ablation of the 5-HT2B receptor leads to partial embryonic and postnatal lethality with abnormal heart development. Similar molecular mechanisms seem to be involved in adult cardiomyocytes since mutant mice surviving to adulthood display a dilated cardiomyopathy. Furthermore this receptor appears to be involved in survival of cardiomyocytes. The 5-HT2B receptor is also implicated in systemic hypertension. Furthermore, mice with pharmacological or genetic ablation of 5-HT2B receptor are totally resistant to hypoxia-induced pulmonary hypertension, indicating that this receptor is regulating the pathologic vascular proliferation leading to this disease. Underlying mechanisms are still to be discovered.

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The mouse 5-HT2B receptor: possible involvement in trophic functions of serotonin

Cellular and molecular biology (Noisy-le-Grand, France), 1994

The novel serotonin receptor 5-HT2B shows the highest homology to the 5-HT2 family of receptors. ... more The novel serotonin receptor 5-HT2B shows the highest homology to the 5-HT2 family of receptors. The pharmacological profile of membranes from 5-HT2B cDNA stably transfected LMTK- cell line, corresponds to a new 5-HT2-like receptor named 5-HT2B, although some difference exists between the mouse and rat pharmacology. A similar pharmacological profile is detected on the immortalized teratocarcinoma-derived cell line 1C11 upon 2 days of serotoninergic differenciation by cAMP. In both cell lines, the analysis 125I-DOI binding reveals the presence of a single class of sites, the affinity of which is one order of magnitude lower than the one reported for the 5-HT2A receptor. This demonstrates that the 5-HT2B receptor is functionally expressed before the complete serotoninergic differentiation of 1C11 cells. These observations are in good agreement with the presence of 5-HT2B mRNA in early mouse embryonic development. Furthermore, the major sites of 5-HT2B mRNA embryonic expression are in ...

Serotonin receptors and therapeutics

ABSTRACT

Drosophila 5-HT2 serotonin receptor: coexpression with fushi-tarazu during segmentation

Proceedings of the National Academy of Sciences, 1995

Serotonin, first described as a neurotransmitter in invertebrates, has been investigated mostly f... more Serotonin, first described as a neurotransmitter in invertebrates, has been investigated mostly for its functions in the mature central nervous system of higher vertebrates. Serotonin receptor diversity has been described in the mammalian brain and in insects. We report the isolation of a cDNA coding for a Drosophila melanogaster serotonin receptor that displays a sequence, a gene organization, and pharmacological properties typical of the mammalian 5-HT2 serotonin receptor subtype. Its mRNA can be detected in the adult fly; moreover, a high level of expression occurs at 3 hr of Drosophila embryogenesis. This early embryonic expression is surprisingly organized in a seven-stripe pattern that appears at the cellular blastoderm stage. In addition, this pattern is in phase with that of the even-parasegment-expressed pair-rule gene fushi-tarazu and is similarly modified by mutations affecting segmentation genes. Simultaneously with this pair-rule expression, the complete machinery of se...

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The Natural Mutation Encoding a C Terminus-Truncated 5-Hydroxytryptamine2B Receptor Is a Gain of Proliferative Functions

Molecular Pharmacology, 2005

List of non-standard abbreviations (6)-[ 125 I]1-(2,5-dimethoxy-4-iodophenyl)-2-aminopropane HCl ... more List of non-standard abbreviations (6)-[ 125 I]1-(2,5-dimethoxy-4-iodophenyl)-2-aminopropane HCl ([ 125 I]DOI; G-protein coupled receptor (GPCR); G-protein coupled receptor kinase (GRK); human herpes virus 8 (HHV8); mitogen-activated protein kinase (MAPK); Multi-PDZ domain containing protein 1 (MUPP1); nuclear factor kappa B (NFkappa B); nitric oxide synthase (NOS); PSD-95, Dlg, ZO-1 (PDZ); serum response element (SRE) MOL#8268

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Evidence for a Control of Plasma Serotonin Levels by 5-Hydroxytryptamine2B Receptors in Mice

Journal of Pharmacology and Experimental Therapeutics, 2006

HAL is a multi-disciplinary open access archive for the deposit and dissemination of scientific r... more HAL is a multi-disciplinary open access archive for the deposit and dissemination of scientific research documents, whether they are published or not. The documents may come from teaching and research institutions in France or abroad, or from public or private research centers. L'archive ouverte pluridisciplinaire HAL, est destinée au dépôt et à la diffusion de documents scientifiques de niveau recherche, publiés ou non, émanant des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

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