Invertebrate Immunity

338 Pages·2010·10.112 MB·English

by  René Augustin, Thomas C. G. Bosch (auth.), Kenneth Söderhäll (eds.)

Invertebrate Immunity
ADVVVANCESINEXPERIMENTTTAL MEDICINEAND BIOLOGY
Editorial Board:
NATHAAANBACK,State University of New York at Buffalo
IRUUUNR. COHEN, The Weizmann Institute of Science
ABEL LAJTHA, N.S. Kline Institute for Psychiatric Research
JOHN D. LAMBRIS,University of Pennsylvania
RODOLFO PAOLETTI,University of Milan
Recent Volumes in this Series
Volume 700
REGULATION OF MICRORNAS
Helge Großhans
Volume 701
OXYGEN TRANSPORT TO TISSUE XXXII
Duane F. Bruley and J.C. LaManna
Volume 702
RNA EXOSOME
Torben Heick Jensen
Volume 703
INFLAMMATION AND RETINAL DISEASE
John D. Lambris and Anthony P. Adamis
Volume 704
TRANSIENT RECEPTOR POTENTIAL CHANNELS
Md. Shahidul Islam
Volume 705
THE MOLECULAR IMMUNOLOGY OF COMPLEX CARBOHYDRATES-3
Albert M. Wu
Volume 706
ADHESION-GPCRS: STRUCTURE TO FUNCTION
Simon Yona and Martin Stacey
Volume 707
HORMONAL AND GENETIC BASIS OF SEXUAL DIFFERENTIATION DISORDERS
AND HOT TOPICS IN ENDOCRINOLOGY
Maria I. New and Joe Leigh Simpson
Volume 708
INVERTEBRATE IMMUNITY
Kenneth Söderhäll
AContinuation Order Plan is available for this series. Acontinuation order will bring delivery of each new volume
immediately upon publication. Volumes are billed only upon actual shipment.For further information please contact
the publisher.
Invertebrate Immunity
Edited by
Kenneth Söderhäll
Department of Comparative Physiology, Uppsala University, Uppsala, Sweden
Springer Science+Business Media, LLC
Landes Bioscience
Springer Science+Business Media,LLC
Landes Bioscience
Copyright ©2010Landes Bioscience and Springer Science+Business Media, LLC
All rights reserved.
No part of this book may be reproduced or transmitted in any form or by any means, electronic or mechanical,
including photocopy, recording, or any information storage and retrieval system, without permission in writing
(cid:2)(cid:3)(cid:4)(cid:5)(cid:6)(cid:7)(cid:8)(cid:9)(cid:6)(cid:10)(cid:11)(cid:12)(cid:13)(cid:14)(cid:15)(cid:8)(cid:9)(cid:3)(cid:16)(cid:6)(cid:17)(cid:14)(cid:7)(cid:8)(cid:6)(cid:7)(cid:8)(cid:9)(cid:6)(cid:9)(cid:18)(cid:19)(cid:9)(cid:10)(cid:7)(cid:14)(cid:4)(cid:20)(cid:6)(cid:4)(cid:2)(cid:6)(cid:21)(cid:20)(cid:22)(cid:6)(cid:5)(cid:21)(cid:7)(cid:9)(cid:3)(cid:14)(cid:21)(cid:13)(cid:6)(cid:15)(cid:11)(cid:10)(cid:10)(cid:13)(cid:14)(cid:9)(cid:23)(cid:6)(cid:15)(cid:10)(cid:9)(cid:19)(cid:14)(cid:24)(cid:19)(cid:21)(cid:13)(cid:13)(cid:22)(cid:6)(cid:2)(cid:4)(cid:3)(cid:6)(cid:7)(cid:8)(cid:9)(cid:6)(cid:10)(cid:11)(cid:3)(cid:10)(cid:4)(cid:15)(cid:9)(cid:6)(cid:4)(cid:2)(cid:6)(cid:12)(cid:9)(cid:14)(cid:20)(cid:25)(cid:6)(cid:9)(cid:20)(cid:7)(cid:9)(cid:3)(cid:9)(cid:23)(cid:6)
and executed on a computer system; for exclusive use by the Purchaser of the work.
Printed in theUSA.
Springer Science+Business Media, LLC, 233 Spring Street, New York, New York 10013,USA
http://www.springer.com
Please address all inquiries to the publishers:
Landes Bioscience, 1806Rio Grande,Austin, TTTexas 78701,USA
Phone: [phone]; FAX: [phone]
http://www.landesbioscience.com
The chapters in this book are available in the MadameCurie Bioscience Database.
http://www.landesbioscience.com/curie
Invertebrate Immunity, edited byKenneth Söderhäll.Landes Bioscience / Springer Science+Business Media,
LLC dual imprint / Springer series:Advances in Experimental Medicine and Biology.
ISBN: 978-1-4419-8058-8
(cid:26)(cid:8)(cid:14)(cid:13)(cid:9)(cid:6)(cid:7)(cid:8)(cid:9)(cid:6)(cid:21)(cid:11)(cid:7)(cid:8)(cid:4)(cid:3)(cid:15)(cid:16)(cid:6)(cid:9)(cid:23)(cid:14)(cid:7)(cid:4)(cid:3)(cid:15)(cid:6)(cid:21)(cid:20)(cid:23)(cid:6)(cid:10)(cid:11)(cid:12)(cid:13)(cid:14)(cid:15)(cid:8)(cid:9)(cid:3)(cid:6)(cid:12)(cid:9)(cid:13)(cid:14)(cid:9)(cid:27)(cid:9)(cid:6)(cid:7)(cid:8)(cid:21)(cid:7)(cid:6)(cid:23)(cid:3)(cid:11)(cid:25)(cid:6)(cid:15)(cid:9)(cid:13)(cid:9)(cid:19)(cid:7)(cid:14)(cid:4)(cid:20)(cid:6)(cid:21)(cid:20)(cid:23)(cid:6)(cid:23)(cid:4)(cid:15)(cid:21)(cid:25)(cid:9)(cid:6)(cid:21)(cid:20)(cid:23)(cid:6)(cid:7)(cid:8)(cid:9)(cid:6)(cid:15)(cid:10)(cid:9)(cid:19)(cid:14)(cid:24)(cid:19)(cid:21)(cid:7)(cid:14)(cid:4)(cid:20)(cid:15)(cid:6)(cid:21)(cid:20)(cid:23)(cid:6)(cid:11)(cid:15)(cid:21)(cid:25)(cid:9)(cid:6)
of equipment and devices, as set forth in this book, are in accord with current recommendations and practice
at the time of publication, they make no warranty, expressed or implied, with respect to material described
in this book. In view of the ongoing research, equipment development, changes in governmental regulations
and the rapid accumulation of information relating to the biomedical sciences, the reader is urged to carefully
review and evaluate the information provided herein.
Library of Congress Cataloging-in-Publication Data
Invertebrate immunity / edited byKenneth Söderhäll.
p. ; cm. -- (Advances in experimental medicine and biology ; v. 708)
Includes bibliographical references and index.
ISBN 978-1-4419-8058-8
1. Invertebrates--Immunology. I. Söderhäll, Kenneth. II. Series: Advances in experimental medicine and
biology ; v. 708. 0065-2598
[DNLM: 1. Invertebrates--immunology. W1AD559 v.708 2010 / QLL362.85]
QL362.85.I58 2010
571.9’612--dc22
2010042470
DEDICAAATION
This volume is dedicated to all my present and former PhD students, post-docs
and fellow scientists who worked in my laboratory and who have contributed to
our research on invertebrate immunity.
v
PREFFFACE
The total number of animal species range up to around 1,000,000 species, of which
invertebrates constitute more than 95%, including a vast diversity of organisms from
unicellular protozoans to the much more complex echinoderms and protochordates.
Insects are by far the largest group of animals within the invertebrates and have received
a large amount of research interest due to their importance as vectors for human and
animal diseases, such as malaria, and the serious harm done by insects to crops and food.
Thus a long-standing research interest in diseases, pathogens and immune responses
of insects there has been. The finding by the late Professor Hans G. Boman and colleagues
that Drosophila responded to a challenge with dead or live bacteria by the synthesis of
antibacterial peptides initiated an intense research interest on the mechanism of this
induction. In 1996Jules Hoffmann, Bruno Lemaitre and colleagues published a paper
in which they for the first time showed that TTToll was involved in the production of the
antifungal peptide drosomycin. Subsequently, this led to the discovery of TTToll-like
receptors (TLRs) in vertebrates and their importance in immune responses, particularly
in mammals.Research on the role of TLRs in immune responses is very intense and the
importance of these receptors is probably greater in mammals than in most invertebrates.
The finding that TTToll was involved in induction of an antimicrobial peptide opened the
possibility of performing detailed genetic studies of the signaling pathways involved in the
production of antimicrobial peptides. The completion of the Drosophila genome in 2000
made it possible to carry out powerful molecular genetic analysis of the immune system of
this insect.For the past 10 years several genomes of insects and other invertebrates have
been sequenced, making comparisons between invertebrates possible, not only between
Drosophila and mammals. Several chapters in this book deal with immune responses
in different groups of insects. It is evident that the immune responses are very similar
between insects, but that there are also differences (Chapters 8-12). Interesting research is
now being performed on mosquitoes because they are vectors for many human diseases.
One important aspect, for example, is how malaria avoids any immune responses while
entering into and being inside the vector mosquitoes (Chapter 12). Interesting studies are
now carried out on developmental biology and innate immunity inHydra (Chapter1),
and there is now interest in studying immune reactions in other invertebrate groups,
vii
viii PREFFFACE
especially in the model organism Caenorhabiditis elegans(Chapter 6), but also in leeches
and earthworms (Chapters 4 and 5).
For the past 10 years research has increasingly been focused on crustaceans (shrimp,
crabs and crayfish) (see Chapter 13) and mollusks (scallop, oysters and mussels) (see
Chapters 2-3), mainly because of their importance as farmed species for consumption.
In these aquatic animals antibacterial peptides have a greater variation and each peptide
appears to be produced in several isoforms induced by different bacterial species,
indicating that there seems to be some sort of specific response to different bacterial
species. The ways in which hemocytes are synthetized have also been studied in detail
in crayfish, and the finding of a family of astakines similar to prokineticins, which are
involved in hematopoiesis, show that these animals may be well suited for studies on
hemocytes and their synthesis (Chapter 13).
Recent research has shown that there are great differences in immune responses
between different invertebrate groups, but of course also that there are many similarities.
TTToll receptors are present in some invertebrate groups but so far the importance of their
function has been mainly studied in insects, while sea urchins, which have more than
200 TTToll receptors still await further studies in toll receptor function (Chapter 14). The
significance of another group of molecules, the so-called Dscams, which seem to be
restricted to insects and crustaceans and are also present in vertebrates, is difficult to
define. Their exact role in immunity is unkown at this moment.
TTTo further emphasize the diversity between different invertebrate animal groups,
the clotting reaction can serve as a good example and is described in two chapters
(Chapters 7 and 13). In crustaceans it comprises a transglutaminase and a clotting
protein, whereas in horse-shoe crabs clotting it is induced by microbial polysaccharides
(LPS and beta-1,3-glucans) so that a proteolytic cascade is activated and terminates with
coagulogen being cleaved by the proclotting enzyme to form the clot (Chapter 7).There
is no similarity between the clotting proteins of crustaceans and horse-shoe crabs, the
process is totally different. The only similarity is that transglutaminase is involved in
both groups of animals.
The melanization reaction is an important innate immune response and which is
present in most invertebrates. It was first noted by Söderhäll and Unestam in 1977
that this process, i.e., activation of prophenolocxidase, was induced in the presence of
beta-1,3-glucans. Subsequently, this was shown in insects and several other invertebrates.
Recently Lee and colleagues managed to show that the prophenoloxidase activating
system and the induction of the TTToll pathway in an insect share the same proteolytic
cascade (Chapter 9). Interestingly, melanization is a highly conserved immune response
which is present in nearly all vertebrates where it provides protection against UVVV-light
and other stressors. Melanization in vertebrates is catalysed by tyrosinase, which has
no homology with prophenoloxidase except for the copper binding sites. However, the
reactions tyrosinase and phenoloxidase catalyze are exactly the same, converting phenols
to quinones and subsequently melanin. Melanization is responsible for skin color in
humans, and in other animals, and it is also an important component in the brain and eyes.
Another innate immune system which is present in both vertebrates and invertebrates
is the complement system, and to date a complement-like system seems to be operable
in echinoderms (Chapter 14), horseshoe crabs (Chapter 7) and tunicates (Chapter 15). In
insects the so-called thiolester-containing proteins are proposed to be complement-like
PREFFFACE ix
and have been shown to function as opsonic proteins in mosquitoes (Chapter 12). It
remains to be shown whether true complement proteins are present in other protostome
groups and not only in horse-shoe crabs.
AA growing interest in invertebrate immunity is the study of the link between
physiology and immunity. Surely diet, ageing, reproduction, reproductive behavior, time
of day, and use of pathogens are all likely to have an effect on immunity and immune
studies. The time of day for immune and challenge studies are important, and both in
Drosophila and crayfish it has been shown that the immune system varies in efficiency
during a day.This means that scientists should be aware of these facts when planning
or making experiments.
In this book I have gathered scientists who are working with different invertebrates,
and it can be seen that the insects are the still attracting most research and researchers.
However, an increasing interest is emerging to study new invertebrate groups, especially
those where the genome is known, as seen in Chapters 1, 5, 14 and 15. Even though
Drosophila has been and still is an excellent model for immune studies, it is now clear
that there are great differences between immune responses in Drosophila and that of
several other invertebrates, which indeed calls for more research on other invertebrates.
Kenneth Söderhäll
Department of Comparative Physiology, Uppsala University
Uppsala, Sweden
ABOUT THE EDITOR...
KENNETH SÖDERHÄLLL studied at Uppsala University and obtained his MSc
degree in 1972 and PhD in 1978.He was promoted to associate professor (Docent) in
1980.After receiveing his PhD he worked in many different laboratories as a post-doc
or as a guest scientist at the University of Montpellier/St.Christol, the Medical Cell
Biology department at TTTromsö University, the Millport Marine Biological Station in
Scotland, UniversityCollege of Wales in Swansea, the Marine BiologicalLaboratory at
Woods Hole and JohnHopkinsUniversity. In 1986 he held aRoyal Society Fellowship
at Swansea. In collaboration with Valerie J. Smith, he developed a new method to isolate
and separate blood cells from invertebrates,which was based on using an anticoagulant
with a low pHand EDTA, a method now used to isolate most invertebrate blood cells.
When he returned to Uppsala he took a position as a researcher at the Swedish Science
ResearchCouncil in 1987, obtaining a chair and appointment to professor and head of
department at Uppsala University in 1989. He continued with research mainly on the
proPO-system in arthropods. His group was first to clone proPOfrom an invertebrate and
he has worked with this so-called melanization reaction for several years.His research
team has also deciphered the clotting reaction in crustaceans and shown that it consists of
a clotting protein present in plasma and a clotting enzyme, a transglutaminase present in
the blood cells. This clotting system is distinct from that of a horseshoe crabs’ in which a
xi