Insect Neuropeptides. Chemistry, Biology, and Action

262 Pages·1991·4.68 MB·English

by  Julius J. Menn

A CS S Y M P O S I UM SERIES 453
Insect Neuropeptides
Chemistry, Biology, and Action
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3.f Julius J. Menn, EDITOR
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1-04 Thomas J. Kelly, EDITOR
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9 Edward P. Masler,
1 EDITOR
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1/ U.S. Department of Agriculture
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5,
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ar Developed from a symposium sponsored
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American Chemical Society, Washington, DC 1991
Library of Congress Cataloging-in-Publication Data
Insect neuropeptides: chemistry, biology, and action / Julius J. Menn,
editor, Thomas J. Kelly, editor, Edward P. Masler, editor,
p. cm.—(ACS symposium series; 453)
1 "Developed from a symposium sponsored by the 1989 International
00 Chemical Congress of the Pacific Basin Societies, December 17-22,
w 1989."
3.f Includes bibliographical references and index.
5
4 ISBN 0-8412-1919-2
0
1- 1. Neuropeptides—Congresses. 2. Insect hormones—Congresses.
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9 I. Menn, Julius J. II. Kelly, Thomas J. III. Masler, Edward P., 1948- .
k-1 IV. International Chemical Congress of Pacific Basin Societies (1989:
b Honolulu, Hawaii) V. Series.
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10 QL495.I4965 1991
0. 595.7'01'88—dc20 90-24674
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5, The paper used in this publication meets the minimum requirements of American National
y 2 Standard for Information Sciences—Permanence of Paper for Printed Library Materials, ANSI
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Ja Copyright © 1991
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Foreword
iHE ACS SYMPOSIUM SERIES was founded in 1974 to provide
a medium for publishing symposia quickly in book form. The
format of the Series parallels that of the continuing ADVANCES
IN CHEMISTRY SERIES except that, in order to save time, the
papers are not typeset, but are reproduced as they are submit
01 ted by the authors in camera-ready form. Papers are reviewed
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w under the supervision of the editors with the assistance of the
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5 Advisory Board and are selected to maintain the integrity of the
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1- symposia. Both reviews and reports of research are acceptable,
9
9 because symposia may embrace both types of presentation.
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bk- However, verbatim reproductions of previously published
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2 papers are not accepted.
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Preface
FIFTEEN YEARS HAVE ELAPSED since Alvin N. Starratt and Brian E.
1
00 Brown released their pioneering publication announcing the initial deter
3.pr mination of a primary structure for an insect neuropeptide—the penta-
5
4 peptide proctolin. In the early 1980s, only a few American, Japanese, and
0
91- European universities, as well as the Zoecon Corporation in the United
9
k-1 States, were engaged in insect neuropeptide research. Neuropeptide
b
1/ discoveries in vertebrates and molluscs—and the advent of exquisitely sen
2
10 sitive techniques for isolation, sequence determination, and synthesis of
0.
1 neuropeptides—rapidly accelerated discoveries in both the mammalian
oi: and insect arenas. More than 50 insect neuropeptides have been
d
1 | sequenced and reported in the literature, and that number is increasing
9
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1 rapidly. Concomitant with these developments have been several sympo
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2 sia, conferences, and workshops that address various aspects of insect
y
ar neuroscience.
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a Three years ago, with the exponential growth of insect neuropeptide
J
e: identification, we felt that the time had arrived to hold a comprehensive
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n D symposium to assess the state of the science in bioactive insect neuropep
o
ati tides and to identify the most promising directions for future research.
c
bli The appropriate occasion arose in conjunction with the International
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P Chemical Congress of Pacific Basin Societies. Twenty-four leading

researchers in insect neurobiology, biochemistry, chemistry, and molecular
biology participated in this symposium. Recent results of research on
developmental, homeostatic, behavioral, reproductive, and metabolic neu
ropeptides were presented and discussed. Important papers were also
presented on neuropeptide synthesis, tertiary structure conformation,
gene identification, and expression of foreign peptides in baculoviruses—
the latter having potential as a means of introducing neuropeptide regu
lating genes into living insects.
This volume reports the current state of our knowledge. We hope that
it will be read and perused by both the initiated and those just entering
this challenging field of research.
ix
As the senior editor (Julius J. Menn), I extend my deepest appreciation
to Herbert Roller, whose foresight, knowledge, and inspiration were
instrumental in introducing me to the exciting field of insect neuropep­
tides.
JULIUS J. MENN
THOMAS J. KELLY
EDWARD P. MASLER
Plant Science Institute
U.S. Department of Agriculture
Beltsville, MD 20705
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The cover illustration is based on a model of two related insect neuropeptide active
core regions in a β-turn conformation generated from molecular dynamics on
a Cray supercomputer.
x
Abbreviations and Nomenclature
In addition to abbreviations and acronyms which are individually identified in the
01 following manuscripts, two systems are used throughout this volume:
0
pr
53. I. Insect peptide nomenclature follows the rules prescribed in Raina and GMe,
4
0 Insect Biochem. 1988,18,785-787.
1-
9
9
1 II. Single-letter amino acid codes:
k-
b
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2
0
1 Ala A Leu L
0.
1
oi: Arg R Lys K
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9 Asn N Met M
9
1
25, Asp D Phe F
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nu Cys C Pro P
a
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ate: Gin Q Ser S
D
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o Glu E Thr T
ati
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bli Gly G Trp W
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His H Tyr Y
lie I Val V
xi
Chapter 1
Neuropeptide Research in Historical Perspective
Berta Scharrer
Department of Anatomy and Structural Biology, Albert Einstein College
of Medicine, 1300 Morris Park Avenue, New York, NY 10461
1
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0
h
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53. The impressive advances made over the years in the identification
04 and functional interpretation of bioactive neuropeptides are the
91- result of a broadly based comparative and multidisciplinary
9
1 approach. The insights gained from studies in insects and verte
bk- brates show remarkable parallelisms between the two groups. What
21/ we have learned is that neuropeptides are engaged in multiple
0
1 forms of intercellular communication in the control of a variety of
10. biologically important integrative functions.
oi: At the outset, it appeared that proteinaceous products dis
d
1 | patched by a special class of neurosecretory neurons, such as those
9 of the pars intercerebralis of insects, function exclusively in a
9
25, 1 nKeuorpoehćo (r1m92o2na) l ocfa paa bcritayin. hForomlloonwein cgo nthtreo lclliansgs iicnasle cdti smcoevtaemryo rbphyo sis
ary much information has been gained on a number of essential neuro
nu hormonal functions carried out by neuropeptides in the control of
a
J postembryonic development as well as reproductive, metabolic,
ate: muscular, and additional activities.
n D These blood-borne messengers may reach terminal effector sites
o directly in a one-step operation, or they may accomplish their task
cati indirectly by signals to non-neuronal glands of internal secretion
bli (corpus allatum, prothoracic gland). These two-step operations are
u
P analogous to those in vertebrates where the dispatch of adenohypo
physial hormones is controlled by hypothalamic signals. The
effective mode of operation of this neuroendocrine connection, i.e.,
the delivery of neural directives to endocrine glands proper by
means of hormones provided by "nonconventional" neurons is a major
concern of the discipline of neuroendocrinology.
An interesting feature which the neuroendocrine axis of
vertebrates and insects has in common is the interaction of stimu
latory and inhibitory neurohormonal directives to the respective
glands of the endocrine system. Their existence has been demon
strated by a combination of structural and experimental studies.
The sites of production (neuronal perikarya) and release (axon
terminals) of the respective, selectively stained messenger sub
stances could be ascertained by tracing their disposition throughout
0097-6156/91/0453-0002$06.00/0
© 1991 American Chemical Society
1. SCHARRER Neuropeptide Research in Historical Perspective 3
the neurosecretory neuron. The inhibitory (allatostatic) role
performed by a group of these neurons became apparent from the
effect of the surgical interruption of this pathway which prevents
the proximo-distal transport of the messenger substance and thus
abolishes its action.
A major step forward, which extended our understanding of the
range of neuropeptide activities, occurred after the advent of
electron microscopy. The observation of synapse-like ("synaptoid")
contacts between peptidergic neuron terminals and endocrine cells
(in the corpus allatum as well as the adenohypophysis) clearly
indicated that close-range signals from neural to endocrine cells
contribute to the process of neuroendocrine communication. Sub
sequently, such peptidergic contacts turned out not to be restricted
to endocrine cells; they also occur on various somatic cells.
More importantly, they establish connections between conventional
1
00 as well as peptidergic neurons.
h
c At this juncture, a new type of interneuronal communication,
3.
5 i.e., the capacity of neuropeptides to act as neurotransmitters,
4
1-0 had become established. Moreover, the structural parameters of
9 peptidergic innervation were seen to include junctional complexes
9
k-1 between two peptidergic neurons that seem to enable a reciprocal
b exchange of signals.
1/
2 Some peptidergic terminals are separated from neuronal elements
0
0.1 by a narrow gap of intercellular space, an arrangement suitable for
oi: 1 ap opssairbalcer inpee pftoidrme- mofe drieatgeudl antieounr.o moTdhuisl atsipoant, iail. ep.,r otxhiem iftyin em-atkuens ing of
d
1 | signals between two synaptically linked neurons by the neuro
99 chemical intervention of a third neuron. Less can be said about
1
5, the possible functional significance of neuropeptides that are
y 2 colocalized with conventional neurotransmitters in a great number
ar of neurons, or of those produced by glial elements.
u
n
a
e: J The presence of neuropeptides at these and many other sites has
at been ascertained by use of immunocytochemical techniques, made
D
n possible by the increasing availability of appropriate peptide
o
ati antibodies. This development took place in conjunction with rapid
ublic badiovaanccteivse inp etphteid ecsh eamnidc atl heid eprnotdifuicctaiotnio no f ofs yan tghreotwici nga nnaulmogbse.r of
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Numerous tests carried out in insects with antibodies raised against
mammalian neuropeptides revealed reaction products within and out
side of the nervous system. Conversely, certain neuropeptides
first identified in invertebrates were shown to occur also in
mammals and other vertebrates. These commonalities are indicative
of a long evolutionary history, as well as a wide distribution of
active neuropeptides in neural and non-neural tissues. One of the
new insights gained from the use of multiple antisera and carefully
conducted specificity tests was the immunocytochemical detection in
the insect brain of molecules closely resembling mammalian ACTH,
prolactin, and insulin,. The localization of these substances sug
gests a neurotransmitter!ike or neuromodulatory rather than a
hormonal role.
4
INSECT NEUROPEPTIDES: CHEMISTRY, BIOLOGY, AND ACTION
In recent years, the improvement of appropriate techniques has
facilitated the chemical identification of a number of insect neuro
peptides as well as comparison with their respective counterparts
in vertebrates. Among these are metabolic, myotropic, allatotropic,
and allatostatic factors.
In insects as well as vertebrates, the effectiveness of regulatory
neuropeptides is known to depend on stereospecific high-affinity
binding sites located in the plasma membrane of the respective
cells to be addressed. Such receptors have been demonstrated for
several opioid peptides by means of radio-ligand techniques in the
brain and digestive tract of insects. Differences in binding-site
density in the brain were shown to be correlated with the cyclicity
of the female's reproductive activity, those in the midgut with its
autonomic nerve supply. The existence of complex decoding processes
for such receptor-mediated stimuli has been carefully documented in
1
0 vertebrates. Recent evidence indicated that a comparable second-
0
ch messenger system, involving inositol lipids and mobilization of
53. intracellular calcium, operates also in insects, i.e., in the
04 peptidergic modulation of the intrinsic rhythmicity of the heart.
1-
9
9
1 Among recent developments in the elucidation of neuropeptide
bk- activities in non-neuronal tissues and cells, those in the immune
21/ system are receiving the greatest attention. Fascinating results
0
1 obtained in invertebrates are in line with those in mammals.
0.
1 Various neuroactive principles have been shown to participate in
oi: cell-mediated immune responses in insects as well as molluscs and
d
1 | mammals. These messenger substances, in addition to lymphokines,
9 are instrumental in the bidirectional exchange of signals betweer
9
5, 1 the neuroendocrine apparatus and the immune system. Furthermore,
2 neuropeptides, among them particularly endogenous opioids, in
y
ar fluence autoregulatory activities of immunocompetent cells. These
nu include adherence and migratory behavior of these cells. The
a
ate: J sctoinmfourlmataiotino noafl lcohcaonmgeost or(yf laatctteinviintyg iasn dp rfeocremdaetdi obny odf isptsienucdtoipveo dia-
n D like cellular processes). There is evidence for the presence, in
o immunocytes as well as cell-free hemolymph, of Met-enkephalin which
cati appears to play a distinctive role in these activities, perhaps by
bli interaction with a special subtype of delta receptor. Moreover,
u
P the inhibitory effect of the opioid antagonist naloxone on these
processes indicates that they are receptor mediated.
There is ample documentation, obtained primarily in mammals, for
the concept that the biosynthesis of the great majority of regula
tory neuropeptides occurs by way of enzymatically cleaved large
precursor molecules produced under the direction of mRNA templates.
Future work will have to show to which extent these and the sub
sequent steps (posttranslational processing, release, and degrada
tion of these products) also occur in insects.
The wide distribution of regulatory neuropeptides throughout the
animal kingdom, and their close chemical similarity or even identity