Physiological Systems in Insects

686 Pages·2013·22.143 MB·English

by  Marc J. Klowden (Auth.)

PHYSIOLOGICAL
SYSTEMS IN INSECTS
PHYSIOLOGICAL
SYSTEMS IN
INSECTS
THIRD EDITION
Marc J. Klowden
Division of Entomology, University of Idaho, Moscow, Idaho
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Dedicated to the memory of Arden O. Lea, who made me think,
and to Alex, Avi, Keira, and Shea, who make me smile.
Preface to the Third Edition
Sir Vincent Wigglesworth, the presci- the cold war with the policy of mutually
ent founder of the field of insect physiol- assured destruction, we knew that insects
ogy, introduced his 1934 treatise, Insect were different enough to survive the inevi-
Physiology, with the comment, ‘The fun- table nuclear holocaust and repopulate
damental processes of vital activity, the the planet without us. The major reason to
ordered series of physical and chemical study insects was to find new ways to kill
changes which liberate energy and main- them that did not kill us.
tain the “immanent movement” of life, In 1948, Wigglesworth presented a well-
are probably the same wherever “living documented justification for using insects as
matter” exists.’ In the 134 pages that fol- models for studying general animal physi-
lowed in this first insect physiology text- ology. However, insects were still largely
book, Wigglesworth described what was relegated to the category of pests we aim
known about the systems of insects based to tolerate or eradicate. Through five edi-
on what he estimated to be 2000 publica- tions of the classic Destructive and Useful
tions. Despite his prophetic introduction, Insects, the value of insects to humans was
few related comparisons of insect systems summarized in a short chapter, while their
followed in the little book, most likely destructive side dominated the remainder.
because there were no comparisons to make, Ten examples of how insects were beneficial
as few others considered there to be much or useful to humans were discussed, with
in common between arthropods and ver- the last being the limited use of insects and
tebrates. The experiments by Kopec´ (1922) insect products in medicine. This included
demonstrating that the insect brain was a maggot therapy for the treatment of
source of hormones were largely ignored wounds, the use of insect venoms for treat-
until Wigglesworth rediscovered them. Who ing rheumatism and arthritis, and the use
besides another insect physiologist would of insect extracts and products such as royal
believe that these simple creatures had hor- jelly as medicines. However, over the last
mones, let  alone hormones produced by 15 years there has been an unprecedented
an insect brain the size of a poppy seed? explosion of information as molecular tech-
The pioneering work by Berta and Ernst niques have yielded information unobtain-
Scharrer (1944) made a strong case for neu- able by conventional biochemistry and
rosecretion in both insects and vertebrates, physiology alone. Now that the genomes
but they too had difficulty with the scien- of humans and many species of insects
tific community accepting the concept that can be compared, the similarities are truly
nerve cells in any animal could produce remarkable and emphasize the wisdom of
hormones. Insects were evolutionarily dis- Wigglesworth’s remarks in 1934. Given that
tant from humans, classified in a primitive the arthropod lineage diverged from that
phylum, with a strange basic body ground of vertebrates more than 600 million years
plan and physiological makeup. During ago, parallels between the physiological
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systems of insects and humans are enough insect nervous system shares numerous
to make your respiratory system inspire, similarities with that of humans. Insects
give your integument chills, and cause your may not dream, but they certainly sleep.
tarsi to twitch. It appears that as many as They meet the criteria established for sleep:
75% of the genes that are associated with a period of quiescence that is associated
human genetic diseases have homologies in with a species-specific posture; a reduced
Drosophila. These similarities have altered responsiveness to external stimuli; a rapid
the focus of insect science, with insects seen reversibility to wakefulness; a homeostasis
more as model systems for studying human based on a longer recovery period following
physiology, and a better understanding periods of sleep deprivation; and an appro-
of them able to be applied to ourselves to priate expression of clock genes. Drosophila
advance the pace of human disease research. engage in periods of quiescence that are
A reason often given for using insects in characterized by changes in brain activ-
research is that they are simple. With few ity and the specific expression of numer-
parts, moving or otherwise, an insect system ous genes. Dozing flies undergo sustained
can be studied without the complications periods of quiescence during the night,
of ancillary and redundant components, but when prevented from sleeping are less
and is therefore much easier to dissect and adept at performing their usual tasks. Just
manipulate. The Drosophila genome, on its as older humans sometimes have problems
four pairs of chromosomes, has been fully sleeping, old flies show disturbances in their
identified and encoded by fewer than 14 000 sleep patterns. Although in older Drosophila
genes – about half that of humans. The other total amounts of sleep do not decrease,
obvious aspects of short lifecycles, little their sleep–wake cycles tend to be more
space required for rearing, considerably less fragmented with age. The observation that
food needed than if one was working with the roundworm Caenorhabditis elegans also
elephants, and the advantage of being com- sleeps indicates that sleep is a basic biologi-
pletely off the radar screens of university cal phenomenon common to many living
animal care and use committees, are addi- things, and that Drosophila can be an effec-
tional points in their favor. Unfortunately, tive model for studying aging and sleep
this view of simplicity often tends toward in humans, and even the phenomenon of
a pejorative label: insects are small, unso- sleep-related restless leg syndrome.
phisticated, and have little in common with Several human neurodegenerative dis-
us more complicated vertebrates. However, eases, including Alzheimer’s disease,
their size belies their complexity, because Parkinson’s disease, muscular dystro-
their incredible success has not been in phy, and Huntington’s disease, lack effec-
spite of their simplicity but because of it. To tive treatments and have undetermined
downplay this simplicity is also to conclude causes. Given the identification of several
that an IBM system 360 computer that once homologous regulatory genes involved in
filled an entire air-conditioned room accom- brain development in both humans and
panied by a colossal 8 Mb of storage was Drosophila, the use of insects to examine the
more complex than a present-day iPhone genetic dissection of the developing brain
with 64 Gb of memory that fits in your may expand our knowledge of how gene
pocket. mis-expression or loss of function might be
Ignoring the possibilities of conscious- countered. The presence of the Drosophila
ness and personality, the far less complex homologue of the microtubule-associated
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protein that is related to Alzheimer’s dis- controlling cell and organ size, and deter-
ease could establish insects as suitable mining overall longevity make insects ideal
models for Alzheimer’s. The kynurenine model systems for understanding growth-
pathway for the degradation of the amino related processes in vertebrates. The steroid
acid tryptophan has been studied in trans- insect hormone 20-hydroxyecdysone (20E)
genic flies as a model for the treatment of that initiates molting of the exoskeleton may
Huntington’s disease. also benefit the vertebrate skeleton as an
Because insects do not have vertebrate- anti-osteoporosis drug. Rats fed 20E over 3
type breathing organs and, except for the months showed increases in bone mineral
largest insects, appear to be free from hav- density over controls. Rats also benefited
ing to take the deep breaths that we do, from increases in the cross-sectional area of
Aristotle, without the benefit of today’s sci- muscles after ingesting 20E isolated from
entific instrumentation in the ancient Greece plant material. In neither case were any side
of 350 BC, can be forgiven for characterizing effects noted.
them as animals that did not breathe. It is An immunity to intestinal disease is
thus satisfying that the branching morpho- essential, given the widespread exposure
genesis of the developing insect tracheal of insects to microbial organisms acquired
system is now recognized as a paradigm for while feeding on fermenting substrates.
the development of branching in mamma- There are many similarities to the human
lian lung and vascular systems. Drosophila intestinal mucosa which involve both
has also been proposed as a model system physical and molecular mechanisms that
for the study of asthma-susceptibility genes maintain a resident flora and discourage
and the innate immune responses of airway pathogenic bacteria. Modeling human intes-
epithelial cells – perhaps even more useful tinal disease in Drosophila has been pro-
than the traditional mouse model. posed, as many signaling pathways that
Given the health concerns about our regulate disease, as well as gut development
increasing waistlines, Drosophila metabolism and regeneration, have been conserved in
and energy homeostasis may yield insights human and insect systems.
into human obesity and the related patholo- It is odd that at the same time as insects
gies, such as diabetes. In mammals, insulin are being utilized more and more as model
and leptin signaling to centers in the brain systems for what we consider to be higher
regulates our metabolism and food intake. organisms, departments of entomology
Insulin has been recognized in insects for and the insect-specific courses associ-
many years, but it is only recently that the ated with them are being eliminated. As
role of insulin signaling has been shown to insects become increasingly used as mod-
be phylogenetically conserved. Although els for understanding human ailments, it
a natural epidemic of obesity in the insect is essential that we understand more about
population has yet to be identified, trans- them. As Barbara McClintock, the winner
genic flies with certain blocked neurons of the 1983 Nobel Prize in Physiology or
store more fat, and suggest that the insect Medicine, so passionately stated, above all
brain apparently measures the level of fat one must have ‘a feeling for the organism’
stores by similar mechanisms as does our with which one is working. There is and
own. The physiological roles of insulin for will be an increasing large cadre of scientists
both humans and insects in the sensing of who work with insects but do not have the
nutritional state and triglyceride storage, opportunity to feel very much about them.
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I hope this text will provide a feeling for the DiAngelo JR, Bland ML, Bambina S, Cherry S,
beauty and complexity of physiological sys- Birnbaum MJ: The immune response attenu-
ates growth and nutrient storage in Drosophila by
tems in insects for those scientists, as well as
reducing insulin signaling, Proc Natl Acad Sci USA
for the traditional entomologists who began
106:20853–20858, 2009.
their careers already so inspired. Dow JA, Romero MF: Drosophila provides rapid mod-
Rather than this third edition evolving into eling of renal development, function, and disease,
a multivolume reference, I chose to main- Am J Physiol Renal Physiol 299:F1237–F1244, 2010.
Freeman A, Pranski E, Miller RD, et al: Sleep fragmen-
tain it as a textbook that is an introduction to
tation and motor restlessness in a Drosophila model
physiological systems in insects. To meet the
of Restless Legs Syndrome, Curr Biol 22:1142–1148,
needs of readers who require more detail, 2012.
there is an extensive bibliography at the end Gilbert LI: Drosophila is an inclusive model for human
of each chapter to serve as a guide to further diseases, growth and development, Mol Cell
Endocrinol 293:25–31, 2008.
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C H A P T E R
1
Signaling Systems
It is essential for cells to know where they are and what the environment around them is
like. Single-celled organisms may have to detect changes in nutrients, temperature, mechan-
ical pressure, electromagnetic fields, light, and the metabolic products of other cells of the
same or different species, and make appropriate responses. This process becomes more criti-
cal for the cells of multicellular organisms, because beginning a few moments after the fer-
tilized egg cell first divides, the individual cells must begin communicating with each other
to coordinate their activities. As development proceeds, cells not only need to respond to
changing conditions but also to exchange information that will determine their specialized
identity and the positions they will assume in the mature organism. The information from
the outside of the cell must be translated and converted to specific cellular responses in a
series of steps referred to as signal transduction pathways.
In multicellular organisms these signals are most frequently chemical in nature, emanat-
ing from a signaling cell and detected by specific receptor proteins on a target cell. There
are hundreds of different signal molecules, but each cell responds selectively depending on
its particular function within the organism. Its response, or failure to respond, depends on
whether or not it possesses a receptor for that signal. The receptor molecules are virtually
always proteins that span the cell membrane. If the correct receptor receives this extracel-
lular signal, the target cell converts it to an intracellular signal that affects cell physiology,
such as changes in cell shape, metabolism, or gene expression. The external signal is thus
internalized, amplified, and distributed to several internal targets (Figure 1.1). The chemi-
cal signals may consist of many types of molecules, categorized by their range and speed of
activity. After the signal is transduced and arrives inside the cell, its target might be either
already-synthesized proteins that become activated by the message, or changes in gene
expression, either of which can alter cell physiology and behavior (Figure 1.2).
Autocrine signaling is the most private of the signaling modes, with a cell signaling itself
by producing a chemical that activates receptors within its own cytoplasm or on its sur-
face (Figure 1.3A). An example is the prothoracic gland, which during some developmental
stages activates its own production of ecdysteroids.
Contact-dependent signaling relies on direct contact between neighboring cells, with
signaling molecules embedded in the cell membranes or passed directly through pores
(Figure 1.3B). The signaling cell may produce a molecule that binds to a receptor in the
Physiological Systems in Insects.
1
DOI: http://dx.doi.org/10.1016/B978-0-12-415819-1.00001-5 Copyright © 2013, 2007, 2002 Elsevier Inc. All rights reserved.