[논문]곤충 면역반응을 중개하는 프로스타글란딘의 분자적 기작과 해충방제 응용

김용균

doi:10.5656/ksae.2022.01.1.067

곤충 면역반응을 중개하는 프로스타글란딘의 분자적 기작과 해충방제 응용
Molecular Action of Prostaglandin to Mediate Insect Immunity and Its Application to Develop Novel Insect Control Techniques 원문보기

한국응용곤충학회지 = Korean journal of applied entomology, v.61 no.1, 2022년, pp.173 - 195

초록
AI-Helper

척추동물과 유사하게 곤충도 인지질분해효소(phospholipase A₂)의 촉매 작용으로 다양한 아이코사노이드를 합성한다. 그러나 일련의 아이코사노이드 생합성과정은 척추동물과 차이를 보이는데, 이는 곤충의 인지질에는 전구물질인 아라키도닉산의 함량이 낮기 때문이다. 대신에 비교적 풍부하게 존재하는 다가불포화지방산인 리놀레익산을 기반으로 사슬 연장 및 불포화반응으로 아라키도닉산을 합성하여 척추동물과 같이 아이코사노이드 전구물질로 이용하는 것 같다. 이렇게 해서 형성된 아라키도닉산은 다시 척추동물의 cyclooxygenase와 유사한 peroxynectin이 PGH₂ 형태의 프로스타글란딘(prostaglandin: PG) 전구물질을 형성하게 된다. 이후 여러 이성체 효소들의 특이적 반응에 의해 PGA₂, PGD₂, PGE₂, PGI₂, TXB₂의 다양한 PG가 생성된다. 반면에 또 다른 형태의 아이코사노이드인 에폭시아이코사트리에노익산(epoxyeicosatrienoic acid: EET)은 척추동물과 유사한 단일산화효소의 산화반응으로 아라키도닉산을 전구물질로 5,6-EET, 8,9-EET, 11,12-EET, 14,15-EET를 형성하게 된다. 그러나 세 번째 아이코사노이드 부류인 류코트리엔(leukotriene)의 경우 곤충 체내 존재는 확인되었지만 생합성 과정은 아직 밝혀지지 않았다. 이들 아이코사노이드가 곤충의 대사, 배설, 면역 및 생식에 관여하는 생리작용을 중개한다. 따라서 아이코사노이드 생합성 과정을 교란하는 물질 탐색은 새로운 살충제 개발 전략이 된다. 본 종설은 이 가운데 PG의 곤충 면역 중개 기작을 소개한다.

Abstract ▼ AI-Helper

Like vertebrates, insects synthesize various eicosanoids after the committed catalytic step of phospholipase A2 (PLA2). However, the subsequent biosynthetic steps exhibit some deviation from those of vertebrates. Due to little composition of arachidonic acid in insect phospholipids, PLA2 releases linoleic acid, which is another polyunsaturated fatty acid and relatively rich in insect phospholipids, to synthesize arachidonic acid via chain extension and desaturation. Resulting arachidonic acid is then oxygenated into a prostaglandin (PG), PGH2, by a specific peroxidase called peroxynectin, but not by cyclooxygenase. PGH2 is then isomerized to various PGs such as PGA2, PGD2, PGE2, PGI2, and a thromboxane (TXB2). All four epoxyeicosatrienoic acids such as 5,6-EET, 8,9-EET, 11,12-EET, and 14,15-EET are also synthesized from arachidonic acid by oxygenation of vertebrate types of monooxygenases. However, the other type of eicosanoids called leukotrienes are found in insect tissues but their synthetic pathway is unclear. Eicosanoids mediate various insect physiological processes such as metabolism, excretion, immunity, and reproduction. Thus, identification of novel compounds interrupting eicosanoid biosynthesis would be a novel approach to develop insecticides. This review focuses on PGs and their immune mediation.

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표/그림 (9)

그림 Fig. 1. Cross-talks of cellular and humoral immune signaling pathways. Upon microbial infection, nonself signal of pattern recognition receptor ('PRR') is propagated by immune mediators such as plasmatocyte-spreading peptide ('PSP'), octopamine ('OA'), 5-hydroxytryptamine ('5-HT'), and nitric oxide ('NO') via eicosanoids. Eicosanoids activate cellular immune responses by activating integrin to stimulate cell-to-cell interactions via tetraspanin. Eicosanoids also influence on Toll/IMD signaling to modulate expression of antimicrobial peptide ('AMP') genes. PRR also activates the upstream signaling components of Toll pathway such as modular serine protease ('MSP'), spätzle activating enzyme ('SAE'), and spätzle processing enzyme ('SAE'). SAE catalyzes inactive components such as serine protease homolog ('SPH') and phenoloxidase ('PO')-activating protease ('PAP'). Toll signaling also activates PLA₂ to produce PGE₂, which binds to its receptor ('PGR') and turns on Ca2+ signaling. The secondary messenger, Ca2+, activates sodium-potassium-chloride cotransporter ('NKCC') to generate ion concentration gradient between inside and outside of the oenocyte, which is followed by water incurrent via aquaporin ('AQP'), resulting in oenocytoid cell lysis ('OCL') to release proPO.
표 Table 1. Different prostaglandins (PGs) detected in insects
그림 Fig. 2. Domain structure of three PLA₂ genes of Spodoptera exigua. In sPLA₂, 'SP' and 'CAT' represent signal peptide and catalytic site, respectively. In iPLA₂, 'ANK' represents ankyrin motif.
그림 Fig. 3. Biosynthetic pathway of PGs in insects. PLA₂ activated by Ca²⁺ and MAP kinase ('MAPK') is translocated to membrane and catalyzes phospholipid to release linoleic acid ('LA'). LA is then elongated by long chain elongase ('ELO') to form C20 fatty acid and desaturated by desaturase ('DES') to form arachidonic acid ('AA'). AA is oxygenated by cyclooxygenase-like peroxinectin ('Pxt') to form PGH₂, which is the substrate to be isomerized to various PGs including PGE₂ by PGE₂ synthase ('PGES'), PGD₂ by PGD₂ synthase ('PGDS'), PGI₂ by PGI₂ synthase ('PGIS'), and TXB₂ by TXA₂ synthase ('TXAS').
그림 Fig. 4. Hemocyte-spreading behavior driven by PGE₂ in hemocytes of Spodoptera exigua . Diagram demonstrates actin cytoskeleton rearrangement by F-actin growth and volume change by aquaporin (AQP) activation. Photo shows spread hemocytes in which F-actin is labeled by FITC (green) and nucleus is stained with DAPI (blue).
그림 Fig. 5. Symbiotic life cycle of nematode and bacteria. Nematodes include Steinernema and Heterorhabditis. Bacteria include Xenorhabdus and Photorhabdus.
그림 Fig. 6. Immunosuppression of Spodoptera exigua by infection of a pathogenic bacterium, Photorhabdus temperata temperata. Hemocyte-spreading behavior post-infection (PI) is active at infection of a non-pathogenic bacterium, Escherichia coli. Photos show F-actin detected by FITC (green) and nucleus stained with DAPI (blue). Scale bar indicates 10 μm. Figure is modified from Kim et al. (2018b).
표 Table 2. Specific association of entomopathogenic nematodes and symbiotic bacteria
그림 Fig. 7. Secondary metabolites produced by Xenorhabdus nematophia, X. hominickii, and Photorhabdus temperata temperata.

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