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Modulation of the Inflammatory Response in Murine Macrophages by Mastic Essential Oil

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Modulation of the Inflammatory Response in Murine Macrophages by Mastic Essential Oil MODULATION OF THE INFLAMMATORY RESPONSE IN MURINE MACROPHAGES BY MASTIC ESSENTIAL OIL Georgia A. Kolezaki Supervisor: Dr. K. Chlichlia Alexandroupolis, 2016 2 CONTENTS Acknowledgements …………………………………………………...................................... 6 Abstract ……………………………………………………………………………………………………….. 7 1.Introduction ……………………………………………………………………………………………… 8 1.1 Inflammation …………………………………………………………………………………………….. 8 1.1.1Definition ………………………………………………………………………………………………… 8 1.1.2 Inflammatory Mediators …………………………………………………………………………. 9 1.1.2.1 Nitrite Oxide (NO) ……………………………………………………………………………….. 9 1.1.2.2 Tumor Necrosis Factor-α (ΤΝF-α)………………………………………………………….10 1.1.2.3 PGE2………………………………………………………………………………………………………11 1.1.3 LPS-induced inflammatory response………………………………………………………..13 1.1.4 Phagocytosis and the inflammatory response………………………………………….14 1.2 Pistacia lentiscus var. Chia…………………………………………………………………………..17 1.2.1 General Characteristics…………………………………………………………………………….17 1.2.2 Taxonomy………………………………………………………………………………………………..18 1.2.3 Mastic Oil…………………………………………………………………………………………………19 1.2.3.1 Essential Oils …………………………………………………………………………………………19 1.2.3.2 Chemical Composition…………………………………………………………………………..20 1.2.4 Biological Activities of mastic tree oil and resin………………………………………..20 1.2.4.1 Anti-cancer Activity……………………………………………………………………………….20 1.2.4.2 Anti-oxidant Activity……………………………………………………………………………..21 1.2.4.3 Antibacterial Activity…………………………………………………………………………….21 3 1.3 Anti-inflammatory properties of mastic oil’s major components………………..22 1.4 Aim……………………………………………………………………………………………………………..23 2. Materials and Methods………………………………………………………………………………24 2.1 Mastic oil’s major components…………………………………………………………………..24 2.2 Cell Cultures………………………………………………………………………………………………..27 2.3 SRB Assay…………………………………………………………………………………………………….29 2.4 Griess Test…………………………………………………………………………………………………..30 2.5 Enzyme Linked Immunosorbent Assay (ELISA)……………………………………………..31 2.6 Prostaglandin E2 (PGE2) Assay………………………………………………………………………33 2.7 Comet Assay………………………………………………………………………………………………..35 2.8 Phagocytosis Assay………………………………………………………………………………………38 2.9 Fluorescence Microscopy……………………………………………………………………………40 2.10 Optical Microscopy…………………………………………………………………………………….41 2.11 Statistical Analysis……………………………………………………………………………………..41 3. Results………………………………………………………………………………………………………..42 3.1 Determinaton of the maximum non toxic concentrations of mastic oil and its major components…………………………………………………………………………………………….42 3.2 Mastic and b-pinene reduce significantly the production of NO, PGE2 and ΤΝF-α in supernatants of LPS-induced RAW264.7 cells………………………………………………..43 3.3 Mastic, limonene and linalol reduce the %DNA-damage caused by LPS in RAW264.7 cells………………………………………………………………………………………………….46 3.4 Inhibition of Morphological Changes in LPS-stimulated RAW264.7 Macrophages by mastic oil………………………………………………………………………………………………………45 4 3.5 Limonene, linalol and mastic oil reduce the rate of phagocytosis of L. casei by RAW 264.7 cells………………………………………………………………………………………………..49 4. Discussion…………………………………………………………………………………………………..52 5. References………………………………………………………………………………………………….55 5 Acknowledgements This study was conducted for my thesis as a requirement for the completion of my undergraduate studies in Molecular Biology and Genetics. First of all, I would like to thank my supervisor Dr. K. Chlichlia for her continued support, interest and time that she dedicated in order this study to be carried out. I would also like to express my gratitude to the PhD student K. Spyridopoulou for her help and useful instructions either during the laboratory work or the writing of this dissertation. Also, I would like to thank all the laboratory team for their help and support as well as Dr. A. Pappa and the PhD student E. Fitsiou for their help in carrying out the Comet assay experiments. 6 Abstract Pistacia lentiscus var. Chia (mastic tree) is a member of the species Pistacia lentiscus that is grown in Chios island in Greece. The mastic resin is produced from the bark of the trees and, although it is considered to have significant biological activities such as antimicrobial, antioxidant and anticancer and has been used in pharmaceutical products and nutritional supplements, little is known about its role in modulating the inflammatory response. In this study the modulation of the inflammatory response by the essential oil of Chios mastic resin and its major constituents was comparatively evaluated using an in vitro inflammation model based on LPS-induced murine macrophages RAW264.7. After the determination of the non-toxic concentrations of mastic oil or its major components (b-pinene, α-pinene, myrcene, linalol and limonene) to RAW264.7 cells, their anti-inflammatory potency was investigated by assaying their effects on inflammation mediators production/secretion and inflammatory responses induced in RAW264.7 cells by LPS stimulation. Mastic oil and some of its bioactive constituents suppressed a) NO production, b) prostaglandin E2 secretion and c) TNF-alpha production in LPS- stimulated RAW264.7 cells. Moreover, the percentage of RAW264.7 cells that underwent LPS-induced morphological changes was smaller after pre-tretment of the the cells with either mastic or some of its constituents. Additionally, mastic oil, linalol and limonene were found to possess a protective role against DNA-damage caused by LPS. Finally, mastic oil and its major constituents reduced the rate of phagocytosis on murine macrophage. Thus, mastic oil and its bioactive components modulate the inflammatory response, suggesting that mastic oil promotes an anti- inflammatory transition in LPS-stimulated murine macrophages. 7 1. INTRODUCTION 1.1 Inflammation 1.1.1 Definition Inflammation is part of the complex biological response of our body tissues to harmful stimuli such as pathogens or damaged cells. It is a protective response involving immune cells, blood vessels and molecular mediators. The role of inflammation is to eliminate the initial cause of injury, clear out necrotic cells and repair tissues [1]. The classical signs of acute inflammation are heat, pain redness, swelling and loss of function. Inflammation is a generic response and therefore it is considered as a mechanism of innate immunity. It can be classified as either acute or chronic. Acute inflammation is achieved by the increased movement of plasma and leukocytes from blood into tissues, whether chronic inflammation is characterized by simultaneous destruction and healing of the tissue [2]. Figure 1: Acute vs Chronic Inflammation. (Baniyash, Michal. "TCR ζ-chain downregulation: curtailing an excessive inflammatory immune response." Nature Reviews Immunology 4.9 (2004): 675-687.) 8 1.1.2 Inflammatory Mediators 1.1.2.1 Nitrite Oxide (NO) An important mediator in acute and chronic inflammation is nitric oxide (NO). NO is generated via the oxidation of the terminal guanidino nitrogen atom of L-arginine by the enzyme, nitric oxide synthase (NOS). Three major isoforms of NOS have been identified. Two expressed constitutively, are calcium/calmodulin-dependent and are classified together as constitutive NOS isoforms (cNOS). The third is a cytokine- inducible, calcium/calmodulin-independent isoform of NOS (iNOS). NO has been shown to increase the production of pro-inflammatory prostaglandins in vitro, ex vivo and in vivo studies, potentially by S-nitrosation of cysteine residues in the catalytic domain of cyclo-oxygenase (COX) enzymes. In addition, NO can also react with superoxide anion to form peroxynitrite (ONOO-), a potent oxidizing molecule capable of eliciting lipid peroxidation and cellular damage. These findings suggest that NO has the ability to exert multiple cytotoxic effects during inflammatory responses including an increase PG production as well as the formation of ONOO- [3]. Figure 2: Nitrite Oxide (NO) synthesis reaction (Lauer T., Kleinbogard P., Kelm M. Indexes of NO Bioavailability in Human Blood. American Physiological Society, 2002, 17: 251-255). 9 . 1.1.2.2 Tumor Necrosis Factor-α (TNF-α) TNF-α was discovered in 1975 as an endotoxin-inducible molecule that caused necrosis of tumors in vitro. It is a transmembrane 26kDa protein expressed by activated NK and T-cells, but also by a diverse array of non-immune cells such as endothelial cells and fibroblasts. The production of TNF-α mRNA is transcriptionally regulated by NF-κΒ, c-Jun, AP-1 and NFAT, consistent with the presence of these transcription factor binding sites within the promoter region of the TNF gene [4]. A plethora of in vitro studies have revealed complex and divergent TNF-R signaling pathways which account for all aspects of TNF’s ability to induce both cell death and/or co stimulation and cell activation [5]. TNFs’ stimulation of globally activating transcription factors such as NF-kB, and its signaling via bio-active lipids that induce arachidonic acid, 5-HETE and ultimately leukotrienes and prostaglandins, explain its effects on diverse cells within almost every human physiological system. They also explain TNFs powerful proinflammatory capacity, especially within immune cells capable of producing a cascade of downstream cytokines and chemokines [6]. For example TNF promotes monocyte/macrophage differentiation and enhances activated B cell proliferation concomitant with an autocrine increase in TNFR expression. It promotes the proliferation of fibroblasts and is a powerful inducer of inflammation, often acting together with IL-1b [7]. 10 Figure 3: The TNF-α pathway. (Victor F.C., Gottlieb A.B., TNF-α
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