mimmunМедицинская иммунологияMedical Immunology (Russia)1563-06252313-741XSPb RAACI10.15789/1563-0625-2019-1-9-20mimmun-1695Research ArticleОБЗОРЫREVIEWSТРОМБОЦИТЫ КАК АКТИВАТОРЫ И РЕГУЛЯТОРЫ ВОСПАЛИТЕЛЬНЫХ И ИММУННЫХ РЕАКЦИЙ. ЧАСТЬ 2. ТРОМБОЦИТЫ КАК УЧАСТНИКИ ИММУННЫХ РЕАКЦИЙBLOOD PLATELETS AS ACTIVATORS AND REGULATORS OF INFLAMMATORY AND IMMUNE REACTIONS. PART 2. THROMBOCYTES AS PARTICIPANTS OF IMMUNE REACTIONSСеребрянаяН. Б.SerebryanayaN. B.

д.м.н., профессор, ведущий научный сотрудник отдела общей патологии и патологической физиологии ФГБНУ «Институт экспериментальной медицины»

197376, Санкт-Петербург, ул. Академика Павлова, 9а.Тел.: 8 (812) 234-15-83; Факс: 8 (812) 234-94-93.

PhD, MD (Medicine), Professor, Leading Research Associate, Department of General Pathology and Pathophysiology, Institute of Experimental Medicine

197376, St. Petersburg, Acad. Pavlov str., 9а. Phone: 7 (812) 234-15-83; Fax: 7 (812) 234-94-93.

nbvma@mail.ruШанинС. Н.ShaninS. N.

к.м.н., cтарший научный сотрудник отдела общей патологии и патологической физиологии

Санкт-Петербург

РhD (Medicine), Senior Research Associate, Department of General Pathology and Pathophysiology

St. Petersburg

ФомичеваЕ. Е.FomichevaE. E.

к.б.н., старший научный сотрудник отдела общей патологии и патологической физиологии

Санкт-Петербург

PhD (Biology), Senior Research Associate, Department of General Pathology and Pathophysiology

St. Petersburg

ЯкуцениП. П.YakutseniP. P.

д.б.н., главный научный сотрудник Центра перспективных исследований

Санкт-Петербург

PhD, MD (Biology), Chief Research Associate, Center for Advanced Studies

St. Petersburg

ФГБНУ «Институт экспериментальной медицины»; ФГБОУ ВО «Санкт-Петербургский государственный университет»; ФГБОУ ВО «Северо-Западный государственный медицинский университет имени И.И. Мечникова»РоссияInstitute of Experimental Medicine; St. Petersburg State University; I. Mechnikov North-Western State Medical UniversityRussian FederationФГБНУ «Институт экспериментальной медицины»РоссияInstitute of Experimental MedicineRussian FederationФГАОУ ВО «Санкт-Петербургский политехнический университет Петра Великого»РоссияPeter the Great St. Petersburg Polytechnic UniversityRussian Federation201922012019211920Copyright © Серебряная Н.Б., Шанин С.Н., Фомичева Е.Е., Якуцени П.П., 20192019Серебряная Н.Б., Шанин С.Н., Фомичева Е.Е., Якуцени П.П.Serebryanaya N.B., Shanin S.N., Fomicheva E.E., Yakutseni P.P.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://www.mimmun.ru/mimmun/article/view/1695

Тромбоцитам принадлежит ведущая роль в сопряжении тромбоза, воспаления и врожденных иммунных реакций. Тромбоциты устанавливают стабильное адгезионное взаимодействие с иммунными клетками. Активированные тромбоциты экспрессируют CD40L (CD154), мембранный гликопротеин семейства фактора некроза опухоли (TNF), причем тромбоциты представляют собой главный источник sCD40L в плазме крови. Тромбоцитарный CD154 может взаимодействовать с рецептором CD40 на эндотелии, вызывая воспалительный ответ, и В-лимфоцитах, усиливая продукцию ими иммуноглобулинов. CD154 тромбоцитов в комплексе с другими сигналами может вызвать созревание и активацию дендритных клеток (ДК). Тромбоциты обладают функциональными рецепторами TLR2, TLR4, TLR7 и TLR9. На тромбоцитах присутствуют Fc-рецепторы, в том числе FcγRIIA, FcεRI и FcαRIA. FcγRIIA на тромбоцитах может вовлекать их в защиту от бактерий. Перекрестная связь FcαRI на тромбоцитах приводит к продукции ими протромботических и провоспалительных медиаторов, таких как тканевой фактор и IL-1β. Активация тромбоцитов через FcεR1 вызывает высвобождение хемокина RANTES и серотонина, которые способствуют провоспалительному ответу других иммунных клеток. Тромбоциты обладают рецепторами для активированных компонентов комплемента (CR2, CR3, CR4, C1qR) и C1-ингибитора, факторов D и H. Активированные тромбоциты способствуют активации системы комплемента через высвобождение протеинкиназ и ATФ, а также путем фосфорилирования C3 и C3b. В α-гранулах тромбоцитов хранятся хемокины, которые представляют и самую многочисленную группу антимикробных белков тромбоцитов (киноцидины), а в цитоплазме тромбоцитов имеется антимикробный белок семейства дефензинов – hBD-1. Продуктами тромбоцитов, относящимися к семейству цитокинов и их рецепторов, признаны лиганд и рецептор суперсемейства TNF (TRAIL и LIGHT), хемокин SDF-1 (CXCL12), интерлейкины IL-1β, IL-8 и растворимый рецептор IL-6 (sRIL-6). Белок HMGB-1, классифицированный как воспалительный цитокин, экспрессируется в активированных тромбоцитах и вызывает формирование нейтрофилами внеклеточных ловушек. Тромбоциты являются источником многочисленных факторов роста, включая EGF-α и EGF-β1, EGF-β2, TGF-α и TGF-β1, TGF-β2, PDGF, HGF, FGF-β, IGF, про и антиангиогенные факторы, например VEGF-F и ангиопоэтины Ang-1 и Ang-2. Реализация иммунных функций тромбоцитов осуществляется при их взаимодействии с лейкоцитами, которые привлекаются к месту инфекции и воспаления и удерживаются при формировании «иммунного тромба» в условиях высокого напряжения сдвига. Тромбоциты могут не только поддерживать и направлять иммунный ответ, но и инициировать его. Они способны презентировать антиген в контексте молекул MHC I класса и активировать наивные CD8+Т-лимфоциты. В обзоре рассмотрены некоторые последствия взаимодействия тромбоцитов с нейтрофилами, моноцитами, дендритными клетками и лимфоцитами.

Thrombocytes keep a leading role in conjugating thrombosis, inflammation and congenital immune responses. The platelets provide stable adhesion and interaction with immune cells. Activated platelets express CD40L (CD154), a membrane glycoprotein of tumor necrosis factor (TNF) family. Hence, the platelets are the main source of sCD40L in blood plasma. Platelet CD154 may interact with CD40 receptor on endothelial cells, causing an inflammatory response, and enhancing production of immunoglobulins by B-lymphocytes. Membrane and soluble CD154 of platelets combined with other signals can induce maturation and activation of dendritic cells (DC). The platelets possess functional receptors, e.g., TLR2, TLR4, TLR7 and TLR9 they also bear Fc-receptors, including FcγRIIA, FcεRI and FcαRIA. FcγRIIA on platelets mediate protection against bacteria. Cross-linking of FcαRI on platelets results in production of prothrombotic and pro-inflammatory mediators such as tissue factor and IL-1β. Activation of platelets via FcεR1 causes release of chemokine RANTES and serotonin, which contribute to the pro-inflammatory response of other immune cells. Platelets possess receptors for activated complement components and its fragments (CR2, CR3, CR4, C1q, C1 inhibitor and factors D and H). Activated platelets trigger the complement system through the release of protein kinases and ATP, and also by phosphorylation of C3 and C3b. α-granules of platelets contain chemokines which represent the most numerous group of antimicrobial proteins of platelets (kinocidins), and there is an antimicrobial protein of the defensin family – hBD-1 in the cytoplasm of platelets. Ligand and receptor of the TNF superfamily (TRAIL and LIGHT), the SDF-1 chemokine (CXCL12), the IL-1βinterleukins, IL-8 and the soluble IL-6 receptor (sRIL-6) are recognized as platelet products belonging to the family of cytokines and their receptors. The HMGB-1 protein classified as an inflammatory cytokine, is expressed by activated platelets and causes formation of the extracellular traps by neutrophils. Platelets produce numerous growth factors, including EGF-α and EGF-β1, EGF-β2, TGF-α and TGF-β1, TGF-β2, PDGF, HGF, FGF-β, IGF, pro- and antiangiogenic factors, e.g., VEGF-F and angiopoietins Ang-1 and Ang-2. Fulfillment of immune functions by the platelets is carried out by their interaction with leukocytes, which are attracted to the site of infection and inflammation and retained during the development of an “immune thrombus” under conditions of high shear stress. Platelets can not only maintain and guide the immune response, but also initiate these events. They are able to present the antigen in the context of MHC class I molecules, and activate naїve CD8+ T lymphocytes. Potential consequences of platelet interaction with neutrophils, monocytes, dendritic cells and lymphocytes are discussed in the review article.

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The authors declare that there are no conflicts of interest present.