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ISBN : 978-1-63315-205-2
Short Views on Insect Biochemistry and Molecular Biology Vol.(1), October 2014
2014
Section II
Insect Growth
IO
NAL B OO
SION
MIS
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Invited Review
Chapter 8
The Ben May Department for Cancer research, The University of Chicago, Chicago, IL, 60637,USA.
2
Department of Biochemistry & Molecular Biophysics, Kansas State University,
Manhattan, KS 66506, USA
Abstract
In this short review, it will be focused on the regulatory biosynthetic pathway of juvenile
hormone (JH). JH is involved in the regulation of nearly every aspect of insect life, such as
metamorphosis, reproduction, diapause, lifespan and polyphenisms. To perform these
biological functions, the biosynthesis of JH is under a rigorous control by insulin/TOR
signaling pathway, neuropeptides, neurotransmitters and other bioactive molecules, modulating
enzymes in the JH biosynthetic pathway. JH acid methyl transferase (JHAMT) converts JH acid
to JH or farnesoic acid (FA) to methyl farnesoate (MF) at final steps in JH biosynthesis.
JHAMT is the key regulatory enzyme for JH biosynthesis, which is activated by DPP-mediated
TGF signaling and also targeted by other regulators for JH biosynthesis. Targeting genes in JH
synthetic pathway, especially the late insect specific steps, was already broadly applied in studies
on JH function and is a promising strategy for pest control.
Key words: juvenile hormone, JH acid methyl transferase, Neuropeptides, Insulin/TOR signaling
pathway
*For Correspondence (email:
zsheng@uchicago.edu or zhtsheng911@gmail.com)
1. Introduction
Juvenile hormones (JH) are a family of insect sesquiterpenoids mainly
synthesized by the corpora allata (CA) (1) and also produced in the gonads, especially
the male accessory gland in Hyalophora cecropia, Apriona germari and Aedes
aegypti (reviewed in 2,3). To what extent do tissues other than CA work as sources
for JH biosynthesis should be easily determined by measuring the expression level
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Overview
1. Introduction
2. The regulatory biosynthetic pathway of JH
2a. Early steps
2b. Late steps
3. Regulators of JH biosynthesis
3a. Insulin / TOR nutrient signaling
3b. Neurotransmitters and neuropeptides
4. Conclusion and future direction
5. Acknowledgement
6. References
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Despite its multiple functions, the molecular basis of JH action was unknown
until recently. New studies had established that the binding of JH to the bHLH-PAS
family transcription factor, methoprene-tolerant (Met) help the heterodimerization of
Met with other members of this family including steroid receptor coactivator (SRC),
Cycle and Clock, which in turn binds to the JH response elements (JHRE) present in
the promoters of JH-response genes. Even these studies defined Met as the JH
receptor, it is still poorly understood about the way JH produces lots of functional
outputs, possibly through heterodimerization with different bHLH-PAS partners,
recruitment of other alternative transcription factors, crosstalk with insulin or other
canonical signaling pathways or affecting on epigenetic programming (1,4,6,12-19).
There are already several excellent reviews about the molecular basis for JH action (1,
4, 14). Therefore, I will focus on the regulatory biosynthetic pathway of JH, another
important aspect for controlling JH function, which is applied by insects and may be
used in pest control (20, 21).
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are broadly expressed in many tissues, because farnesol and farnesal homeostasis are
vital for cells in majority of insects tissues. FPPP was first isolated from
D. melanogaster (40) and its homologue was recently characterized in A. aegypti (41).
All three identified A. aegypti FPPases (AaFPPase) are abundantly expressed in the
CA, but only AaFPPase-1 and -2 can functionally convert FPP to farnesol and only
the reduction of AaFPPase-1 by RNAi caused a significant decrease on JH
biosynthesis, implying AaFPPase-1 is the real FPPase involved in JH biosynthesis
(41). FOLD, oxidizing farnesol into farnesal, is a NADP+ dependent short-chain
dehydrodenase, first identified from A.aegypti. FOLD is expressed not only in the
CA but showing an expression pattern correlated with the JH synthesis level after
adult emergence (39). This study suggested that the oxidation of farnesol might be a
regulatory step in JH synthesis in adult mosquitoes. NAD+ dependent long-chain
aldehyde dehydrogenase 3 (ALDH3), working as a FALD, converts farnesal into
farnesoic acid in the mosquito CA (42). The knockdown of the expression of ALDH3
abolished the FALD enzyme activity and impaired JH biosynthesis (42). This study
functionally characterized ALDH3 as FALD in A. aegypti. The CYP15, belonging to
P450 family, catalyzes the expoxidization of FA in Lepidoptera or MF in other insect
species based on their substrate specificity, resulting in the variation of the last two
steps of JH (Fig.2) biosynthesis in insects (25,26,43,44). CYP15A1, from D. punctata,
specifically converts MF to JH III, not using various other MF-related compounds
such as farnesol, FA and farnesyl methyl ether as substrates. CYP15A1 is specifically
expressed in the CA and only during peak JH production.
Interestingly, selected 1,5-disubstituted imidazoles, known to inhibit JH
biosynthesis and cause accumulation of MF in cockroach CA, displayed parallel
effectiveness in inhibiting the activity of recombinant CYP15A1 and in vitro JH
biosynthesis by isolated CA (25). CYP15C1, cloned from B. mori, is also CA-specific
but converting FA to JHA in a highly stereospecific manner. CYP15C1 was
constitutively expressed from larval to adult stages, suggesting that the transcriptional
regulation of CYP15C1 is not important for the temporal regulation of JH production
in B. mori. JHAMT, responsible for the adding of methyl group to JHA or FA, was
first cloned from B. mori and appears critical for metamorphosis. At the beginning of
the fifth instar, the expression of JHAMT in CA reaches the highest level then
declines rapidly to undetectable by day 4 and remains so until pupation (28). During
the period of adult emergence, a high expression level of JHAMT was also observed
in female CA (24, 30).The expression pattern of JHAMT during the postembryonic
development exhibits a good correlation with JH synthesis by CA (45). In M. sexta,
B. mori and Samia cynthica ricini, only CA of female adults synthesizes JH, while
male CA produces JH acid (JHA). The authors believed it was because female, but
not male, CA, has JHAMT activity (24,30,46). Cause MF but not JHA or FA showed
functional activity, JHAMT mediated methylation should be more important than the
CYP15 catalyzed epoxidation for the regulation of JH biosynthesis and insect life
cycle. The JH-deficient CYP15C1 null mutant silkworm undergoes a precocious
metamorphosis with fewer larvallarval molts, which is because the possible presence
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3. Regulators of JH biosynthesis
3a. Insulin/TOR Nutrient signaling
During the control of life span, reproductive maturation, reproductive diapause
and caste fate, nutrient status and nutritional signaling pathway, insulin/TOR pathway,
affects the JH biosynthesis by regulating the expression of genes in the JH
biosynthesis pathway. In D. melanogaster, the mutant of insulin receptor (InR)
yielded dwarf females with extended lifespan. These insects exhibited decreased JH
levels and undeveloped ovarioles that resembled the ovarioles of wild-type flies that
are in diapause, a delay in development in response to adverse environmental
conditions (47). An exogenous application of JH analog methoprene to InR mutant
dwarf females initiated vitellogenesis and restored lifespan similar to that of
wild-type flies (47). The A. aegypti larvae reared in inadequate nutrient conditions
developed into smaller adults and showed an inactivated TOR pathway, lower JH
levels, and needed additional blood meals to complete gonadotropic cycle (48).
Application of JH III to these small mosquitoes could initiate vitellogenesis with only
one blood meal (48). RNA interference (RNAi)-mediated gene knockdown of IRS
(InR substrate) and TOR abolished a JH peak and resulted in a worker phenotype in
larvae reared on queen diet which is destined to queen development (49). Recently, in
B. germanica, FOXO was shown to inhibit JH biosynthesis without modifying the
expression of HMGS1, HMGS2, HMGR or MFO (CYP15A1) in the CA (50). But it
is shown that TOR mediated nutrition status affects JH levels in this insect by
modulating the expression of HMGS1, HMGS2 and HMGR (36). The expression of
genes encoding JH biosynthetic enzymes (HMGS, HMGR, IPPI, FPPS, JHAMT and
MFO) is decreased and the JH biosynthesis is reduced when rapamycin, RNAi
against TOR, and PI3K inhibitor were applied in an in vitro culture of CC-CA
complex in A. aegypti (51). The transcriptional level of HMGR in CA of
D. melanogaster is also modulated by the insulin signal transduction pathway and a
transcriptional factor sterol regulatory element binding protein (SREBP) (52).
3b. Neurotransmitters and Neuropeptides
Neurotransmitters, such as dopamine and glutamate, were shown to regulate the
JH biosynthesis by the isolated CA in vitro (53,54). Studies in cockroach and
Drosophila revealed that glutamatergic nerves innervate CA cells and that the
N-methyl-D-aspartate subtype of glutamate receptors (NMDAR) are expressed in
both brain and CA (54). A recent study well established the molecular mechanisms of
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receptor (NGPR), BNGR-A16, was first identified as the AT receptor in silkworm (75),
by comprehensive cloning and systematic analyses of all neuropeptide G protein
coupled receptors (GPCRs). Interestingly, BNGR-A16 was not highly expressed in the
CA, but instead was predominantly transcribed in the corpora cardiaca (CC), an organ
adjacent to the CA. Furthermore, Yamanaka and coauthors (75) purified and
characterized novel allatoregulatory peptides produced in AT receptor-expressing CC
cells, which may mediate the indirect effect of AT on the CA. These novel
allatoregulatory peptides belong to the short neuropeptide F (sNPF) family, showing
clearly inhibition of JH synthesis in an in vitro CA culture (75). Receptors of sNPF
are also GPCRs: BNGR-A10 and A11 in silkworm, and so do the receptor for
Manse-AST: BNGR-A1 in silkworm and Drostar1 and 2 in D. melanogaster (75, 76).
sNPF suppressed the expression of MVAK and HMGR, and Manse-AST suppressed
the expression of MVAK, HMGR and IPPI, which were initially believed to inhibit
JH synthesis by blocking the transfer or cleavage of citrate to yield of acetyl-CoA
(53). So neuropeptide GPCRs and their peptide ligands, including AT, AST and sNPF,
play a critical role in the control of JH biosynthesis by modulating the expression of
JH biosynthesis enzymes. But it is still not clear how neuropeptide GPCRs regulate,
stimulate or inhibit JH biosynthesis enzymes at transcriptional level.
AT receptors were also isolated from A. aegypti, T. castaneum and M. sexta.
Based on their expression patterns, the working model of AT receptors maybe not the
same in different insect species and the function of AT and its receptors is not only to
stimulate the biosynthesis of JH (77-79). Unlike the CC-specific expression pattern of
AT receptor in B. mori, the A. aegypti AT receptor was expressed in the CA but not
the CC, showing a expression pattern resembles the changes in JH biosynthesis (77).
Besides the presumable expression in the CA (showed with the expression in the
head), the T. castaneum AT receptor was surprisingly highly expressed in
reproductive system with a higher level in the male (78). Whereas in M.sexta AT
receptor was expressed in multiple tissues in feeding larvae, the highest level is in the
Malphighian tubules (79).
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it is necessary to investigate their effects on the enzyme activity and measure their
influence on JH biosynthetic capability by analyzing the changes of JH precursor
pools (42, 80).
5. Acknowledgements
The authors thanks to Drs. Sheng Li, Chinese Academy of Sciences, China, and
Prof. Noriega, F.G., Florida National University, USA and Dr. Hua Bai, Brown
University, USA for their critical readings and valuable comments.
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tribute to Berta Scharrer. Insect Biochem. Mole. Biology 30: 653-662.
64. Barbara, S. and Tobe, S.S. (2007) The role of allatostatins in juvenile hormone synthesis in insects and
crustaceans. Annu. Rev. Entomol. 52: 277-299.
65. Gde, G., and Hoffmann, K.H. (2005) Neuropeptides regulating development and reproduction in insects.
Physiol. Entomol. 30(2): 103-121.
66. Kataoka, H., Toschi, A., Li, J.P., Carney, R.L., Schooley, D.A., and Kramer, S.J. (1989). Identification of
an allatotropin from adult Manduca sexta. Science, 243(4897): 1481-1483.
67. Kramer, S.J., Toschi, A., Miller, C.A., Kataoka, H., Quistad, G.B., Li, J.P., Carney, R.L., and
Schooley, D.A. (1991) Identification of an allatostatin from the tobacco hornworm Manduca sexta. Proc.
Natl. Acad. Sci. 88(21): 9458-9462.
68. Taylor, P.A., Bhatt, T.R., and Horodyski, F.M. (1996) Molecular characterization and expression analysis
of Manduca sexta allatotropin. Euro. J. Biochem. 239(3): 588-596.
69. Jansons, I.S., Cusson, M., Mcneil, J.N., Tobe, S.S., Bendena, W.G. (1996) Molecular characterization of a
cDNA from Pseudaletia unipuncta encoding the Manduca sexta allatostatin peptide (Mas-AST). Insect
Printed in the Unitated States of America, 2014
ISBN: 978-1-63315-205-2
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Invited Review
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Invited Review
Praying mantis
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Invited Review
2014
Section III
Insect Immunity
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Table Contents
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Page No.
Preface
Forward message
Contributors
Reviewers
Acknolwedgement
i
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iv
v
Volume1
Section I: Insect Biochemical approaches
Raman Chandrasekar, P.G., Brintha, Enoch Y.Park, Paolo Pelsoi, Fei Liu,
Marian Goldsmith, Anthony Ejiofor, B.R., Pittendrigh, Y.S., Han,
Fernando G. Noriega, Manickam Sugumaran, B.K., Tyagi, Zhong Zheng Gui,
Fang Zhu, Bharath Bhusan Patnaik, and P. Michailova
2.
57
Sahayaraj, K.
3.
75
4.
99
5.
127
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149
Manickam Sugumaran
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217
Insect Immunity
233
253
271
291
317
331
Paraskeva V. Michailova
355
Dhanenjeyan, K. J., Paramasivam, R., Thanmozhi, V., Chandrasekar,R., and Tyagi, B.K.
Index
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Volume2
Section V:
373
385
in Lepidoptera.
409
Section VI:
429
449
473
497
509
Ronald J. Nachman
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Section VII:
533
549
Usha Rani, P.
575
595
Section VIII:
Insect Bioinformatics
621
633
685
Jitrayut Jitonnom
Index
709
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Book Mission Project # 2: Initiated on June 2010; Completed on March 2014 and Published on Oct. 2014.
PREFACE
Entomology as a science of inter-depended branches like biochemistry, molecular entomology, insect
biotechnology; has made rapid progress in its attributes in the light of modern discoveries. This also
implies that there is an urgent need to manage the available resources scientifically for the good of man.
In the past five decades, entomology in the world/country has taken giant steps ahead. Continued
research has evolved better pest management through molecular approaches. The aim of the Short
Views on Insect Biochemistry and Molecular Biology book is to integrate perspectives across
biochemistry and molecular biology, physiology, immunology, molecular evolution, genetics,
developmental biology and reproduction of insects. This century is proclaimed as the Era of
Biotechnology and its consists of all types of Mol-Bio applications, which is an essential component for
a through understanding of the Insect Biology. This volume 1 & 2 (8 section with 30 chapters)
establishes a thorough understanding of physiological and biochemical functions of proteins, genes in
insects life processes; the topics dealt with in the individual chapters include chemistry of the insect
cuticle, hormone and growth regulators; biochemical defenses of insects; the biochemistry of the toxic
and detoxification action; modern molecular genetics and evolution; inter- and intra-specific chemical
communication and behavior; insect pheromone and molecular architecture, phylogeny and chemical
control of insect by using insect pheromones biotechnology; insect modern biology and novel plant
chemical and microbial insecticides for insect control, followed by a discussion of the various
mechanisms of resistance (both behavioral and physiological) and resistance management; modern insect
pest management through biochemical and molecular approaches; Mimetic analogs of insect
neuropeptide for pest management; entomo-informatics and computer-aided pesticide designing. In short
this book provides comprehensive reviews of recent research from various geographic areas around the
world and contributing authors area recognized experts (leading entomologist/scientist) in their
respective filed of molecular entomology. We will miss this collaboration now it has ended, but will feel
rewarded if this book is appreciated by our team/colleagues and remarkable mile stone in entomology
field.
This book emphasizes upon the need for and relevance of studying molecular aspects of entomology in
Universities, Agricultural Universities and other centers of molecular research. To encompass this
knowledge and, particularly disseminate it to the scientific community free of cost, was the major
inspiring force behind the launch of Short Views on Insect Biochemistry and Molecular Biology.
Editors
Raman Chandrasekar
Brij Kishore Tyagi
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iv
vi
ShortViewson
InsectBiochemistryand
MolecularBiology
Editedby
Raman Chandrasekar, Ph.D.,
Kansas State University, USA.
B.K.Tyagi, Ph.D.,
Centre for Research in Medical Entomology (ICMR), India.
Zhong Zheng Gui, Ph.D.,
Jiangsu University of Science and Technology,
Sericultural Research Institute, Chinese Academy of
Agricultural Sciences, China.
Gerald R. Reeck, Ph.D.,
Kansas State University, USA.
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Contributing Authors
Dr. B.K.Tyagi
Prof.Fernando G. Noriega
Prof. K. Sahayaraj
Prof.Yanyuan Bao
Institute of Insect Science,
Zhejiang University, China.
Prof. PatriciaY.Scaraffia
Department of Tropical Medicine,
Tulane University, New Orleans,
LA 70112, USA.
Dr. P. Somasundaram
Central Sericultural Germplasm Resources Centre,
P.B.No.44, Thally Road,
Hosur-635109,
Tamilnadu, India.
College of Forestry,
Northwest A & F University
Yangling, Shaanxi 712100, China
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Dr. R. Srinivasan
School of Biotechnology,
Trident Academy of Creative Technology
(TACT), Bhubaneswar 751013 Odisha, India.
School of Science
University of Phayao, Thailand.
Department of Entomology,
University of Illinois, Urbana-Champaign, IL,
61801, USA
.
Prof. K. Murugan
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Acknowledgements
Writing and publishing a book requires the assistance of individuals who are
creative, talented, and hard-working. All of these qualities were present in the
individuals assembled to produce this book volume. I would like to express my
heartfelt gratitude to my former teacher Prof. Seo Sook Jae, (GSNU, South Korea),
Prof. Subba Reddy Palli (University of Kentucky, USA), and other external mentors
Prof. Marian R. Goldsmith (University of Rhode Island, USA), Prof. Enoch Y. Park
(Shizuoka University, Japan), Prof. M. Kobayashi (Nagoya University, Japan), Prof.
CHU Jang Hann (National University of Singapore, Singapore), Prof. Thomas W.
Sappington (USDA-ARS, USA), Prof. Fernando G. Noriega (Florida International
University, USA), Dr. Srinivasan Ramasamy, AVRDC, The World Vegetable
Center, Taiwan), Dr. H.C. Sharam (ICRISAT, India), who inspiration and
supported me at many ways for the commencement of this International Book
Mission Program. The book mission program was initiated on May 2010,
completed on March 2014 and published on October 2014. I have no words to
express my feeling for all those who provided valuable contributions from USA,
South Korea, Japan, China, India, Thailand, Taiwan, Bulgaria, France, Iseral, and
Portugal (Contributors name list, see page no. v) and made the completion of this
book possible. We express our appreciation to the following people (Reviewer
name list, see page no. vii) who reviewed various part of the manuscript as it was
being developed and improved quality of each chapter. I thank the ICMR, New
Delhi, and Chinese Academy of Agricultural, China, and Kansas State University for
support from several aspects. Many others (scientists and publishers) have also
allowed us to use their materials in the various chapters, their color image have then
been converted to gray color/BW. Iam especially indebted to International Book
Mission Organization, Academic Publishing Services for the production of book. I
thank my Co-Editors for their continuous vigilance over the book project and for
always giving advance notice of the editing and proofreading schedules. I thank also
my Brintha, P.G., (my wife), who in all possible way, encouragement helped
transform our original efforts into an acceptable final form. I apologize to those
whose work could not be cited owing to space considerations limitation. Further, I
wish to recognize the moral support extended by colleagues and friends. I hope that
this volume will inspire interest on the diverse aspects of insect biochemistry and
molecular biology in aspiring and established scientists.
Raman Chandrasekar
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Book Series
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