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62 Hemolymph Proteins and Functional Peptides: Recent Advances in Insects and Other Arthropods Vol. 1, 2012, 62-77<br />

Immune Response of Insects and Crustaceans<br />

Akira Goto 1,2 and Shoichiro Kurata 1,*<br />

Muhammad Tufail and Makio Takeda (Eds)<br />

All rights reserved-© 2012 <strong>Bentham</strong> <strong>Science</strong> Publishers<br />

CHAPTER 4<br />

1 2<br />

Graduate School of Pharmaceutical <strong>Science</strong>, Tohoku University and Graduate School of Life <strong>Science</strong>s,<br />

Tohoku University, Sendai, 980-8578, Japan<br />

Abstract: Innate immune systems, which are essential for all metazoans, provide the first-line of defense<br />

against pathogens and comprise mainly humoral and cellular immune responses. Over the last two decades,<br />

genetic studies from a model insect, the fruit fly Drosophila melanogaster, have significantly contributed to<br />

elucidate the complex molecular mechanisms underlying the activation of humoral innate immune<br />

responses from pathogen recognition by pattern recognition receptors (PRRs) such as peptidoglycan<br />

recognition proteins (PGRPs), infection-induced nuclear factor (NF)-B intracellular activation, and to the<br />

subsequent induction of hundreds of effector genes, including antimicrobial peptides (AMPs). In addition,<br />

many biochemical studies in Drosophila as well as in other arthropods such as the horseshoe crab recently<br />

uncovered the detailed molecular mechanisms of hemocyte-mediated immune responses, such as<br />

hemolymph coagulation or melanization, which have led us to reconsider the importance of cellular immune<br />

responses. This review describes recent progress towards understanding the molecular mechanisms of these<br />

innate immune responses mainly from studies of Drosophila, horseshoe crab, or other arthropods, and<br />

discusses the fundamental similarities and differences from mammalian innate immune systems.<br />

Keywords: Drosophila melanogaster, fruit fly, horseshoe crab, innate immunity, peptidoglycan<br />

recognition, PGRP, Toll, imd, antimicrobial peptides.<br />

INTRODUCTION<br />

Arthropods are the most diverse animal species on earth. More than 75% of all animal species are<br />

arthropods, and they therefore represent the largest group of invertebrates. Insects, crustaceans, arachnids,<br />

and horseshoe crabs are all classified as arthropods. Among the more than 1.3 million different kinds of<br />

invertebrates with various evolution histories, body pattern developments, and ecological niches,<br />

arthropods are a highly successful species. One of the main reasons for their success is undoubtedly their<br />

efficient innate immune systems.<br />

The innate immune system involves germline-encoded factors that recognize and eliminate invading<br />

pathogens, and comprises three major steps; 1) Recognition of pathogens or non-self, 2) Signal<br />

amplification and activation of the signaling pathways, 3) Induction of effector gene expression. Pathogen<br />

recognition is mediated by a number of pattern recognition receptors (PRRs) that recognize the conserved<br />

molecular signatures of pathogens, termed pathogen-associated molecular patterns (PAMPs) [1-3].<br />

Representative PAMPs include lipopolysaccharides (LPS) from Gram-negative bacteria, lipoteichoic acid<br />

(LTA) from Gram-positive bacteria, peptidoglycans and lipoproteins from many bacterial cell walls, glucan<br />

from fungal cell walls, double-stranded RNA from many viruses, etc. PRRs are broadly categorized<br />

into three functional receptors: signaling PRRs transduce the infection signal and activate evolutionarily<br />

conserved innate immune signaling pathways. Phagocytic PRRs are expressed on the cell surface and are<br />

involved in pathogen uptake. Secreted PRRs are surveillance receptors that detect pathogens and activate<br />

complement systems, such as opsonization [4]. Upon detection of PAMPs by PRRs, signals are transduced<br />

into host cells where they trigger the expression of numerous effector genes.<br />

Due to well-established powerful genetic and RNA interference (RNAi) technologies and the high<br />

evolutionarily conserved mammalian innate immune systems, the fruit fly Drosophila melanogaster is an<br />

*Address correspondence to Shoichiro Kurata: Graduate School of Pharmaceutical <strong>Science</strong>, Tohoku University, Sendai, 980-8578,<br />

Japan; Tel: 81-22-795-5916; Fax: +81-22-795-6802; Email: kurata@mail.pharm.tohoku.ac.jp

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