Abstract
The adhesion and migration of leukocyte play an important role in immune responses and pathogenesis of inflammatory diseases including asthma and atherosclerosis. Very-late-antigen-4 (VLA-4)-mediated adhesion of eosinophils to vascular-cell adhesion molecules-1 (VCAM-1) may contribute to their selective recruitment to tissues and thus is a key event in the pathogenesis of asthma. The cross-talk between lymphocyte function-associated antigen-1 (LFA-1) and VLA-4 on monocytes may be crucial for a coordinated regulation of the cellular adhesion at the complex process of transendothelial migration during the process of atherogenesis. Reactive oxygen species (ROS) are ubiquitous reactive derivatives of oxygen metabolism found in the environment and in all biological system. ROS including superoxide (O2-), hydrogen peroxide (H2O2), and nitric oxide (NO), are abundant in the airway of allergic patients. How ROS participate in the pathogenesis of inflammatory diseases, however, remained to be investigated. We used an eosinophil cell line, HL-60 clone 15, in an adhesion assay to VCAM-1 to investigate the effect of ROS on integrin-mediated leukocyte adhesion. Phorbol 12-myristate 13-acetate (PMA)-stimulated neutrophils, superoxide, NO, and H2O2 were added to analyze their effects on cell binding. Stimulated neutrophils significantly increased eosinophil binding to VCAM-1 but NO and H2O2 decreased the adhesion. Neutrophils from a chronic granulomatous disease (CGD) patient lacked this activity in enhancing eosinophil adhesion. The activity of ROS to enhance cellular adhesion was through activation of integrins because we showed that superoxide increased the expression of activated conformation of VLA-4 on HL-60 clone 15 cells as well as freshly isolated eosinophils. Not only do ROS play a role in intercellular communication, we also found that ROS are important in intracellular cross-talk between different integrins. By using monoclonal antibodies and recombinant intercellular adhesion molecule-1 (ICAM-1) to engage LFA-1 on monocytic cells, we found that the cellular adhesion to VCAM-1 mediated by VLA-4 was suppressed and the suppression depended on the presence of ROS. Inhibition of production of ROS through the use of inhibitor of the nicotinamide adenine dinucleotide phosphate (NADPH) oxidase, but not inhibitors of mitochondrial electron transport chain or xanthine oxidase, revealed that this suppression on VLA-4-mediated cellular binding was mediated by ROS produced by phagocyte NADPH oxidase. Activation of phosphoinositol-3 kinase and Akt appears to mediate this NADPH oxidase activation through p47phox phosphorylation and Rac 1 activation. Our results suggest that the balance between ROS molecules in different tissue microenvironments may change the integrin-mediated leukocyte adhesion and is likely to be a key factor in leukocyte recruitment in inflammation diseases. Our results also provide a novel pathway in which ROS play a critical role in regulating leukocytes adhesion.

中文摘要
了解白血球的黏著和移動，在治療疾病上扮演著很重要的角色。尤其是在氣喘 (asthma) 和粥狀動脈硬化 (atherosclerosis) 這類發炎性疾病上。血球從血管移動到組織，必須靠黏著分子 (adhesion molecules) 幫忙。在氣喘的患者中有大量的嗜酸性球 (eosinophil)，藉由very late antigen 4 (VLA-4) 黏著分子，累積在呼吸道。 另一方面lymphocyte function-associated antigen 1 (LFA-1) 和VLA-4 黏著分子互相聯絡而影響粥狀動脈硬化中單核球細胞 (monocyte)浸潤到組織中也有著舉足輕重的角色。在身體中因為呼吸及代謝作用產生的活性氧分子包含超氧化物 (O2-)、過氧化氫 (H2O2) 和一氧化氮 (NO)。其中在氣喘病人所呼出的氣體，和肺泡沖洗液中，含有大量的一氧化氮及超氧化物。但是對活性氧分子在發炎疾病中所扮演的角色仍待更進一步的了解。我們使用了HL-60 clone 15嗜酸性球細胞株，和vascular-cell adhesion molecules-1 (VCAM-1)蛋白質進行黏著試驗並觀察活性氧分子對血球黏著的影響。Phorbol 12-myristate 13-acetate (PMA)刺激後的嗜中性球 (neutrophil)，超氧化物，一氧化氮和過氧化氫都用來測試對細胞黏著的影響。刺激後的嗜中性球可以增加嗜酸性球黏到VCAM-1上但是相反的過氧化氫，與一氧化氮可以降低黏著。另外慢性肉芽腫病人((chronic granulomatous disease (CGD)) 的嗜中性球，缺乏增加嗜酸性球黏著的能力。其中了解到：超氧化物是經由使VLA-4黏著分子由非活化態構型轉變成活化態的方式，來達到增加黏著能力，而且這現象在真正的嗜酸性球中被觀察到。活性氧分子不只會去影響細胞內的溝通而且我們也發現活性氧中間產物對不同的黏著分子的互相聯絡也很重要。藉著使用intercellular adhesion molecule-1 (ICAM-1) 蛋白質或單株抗體刺激單核球細胞上的LFA-1，發現在單核球細胞上LFA-1和VLA-4黏著分子，會透過活性氧分子互相聯絡。當單核球細胞上的LFA-1受到刺激後，會透過活性氧分子降低VLA-4黏到VCAM-1的能力。只有抑制nicotinamide adenine dinucleotide phosphate (NADPH) oxidase酵素，而非粒線體電子傳遞鏈，或Xanthine oxidase酵素所產生的氧活性分子時會抑制VLA-4所媒介的黏著。當LFA-1受刺激時，先活化phosphoinositol-3 kinas (PI3K) 和Akt訊息傳遞路徑，接著活化對NADPH oxidase酵素活化過程中很重要的Rac 1 蛋白質。透過一連串訊息傳遞路徑，使得黏著分子之間可以互相影響。經由這些實驗，我們了解到：這些活性氧分子，在發炎組織的微環境中如何取得一個平衡，對這整個調控機制是重要的。透過這些研究，發現在血球細胞的黏著上重要且獨特的調控方式。

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