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Ecological and Environmental
Physiology of Insects
This page intentionally left blank
Ecological and Environmental
Physiology of Insects

Jon F. Harrison
School of Life Sciences, Arizona State University
H. Arthur Woods
Division of Biological Sciences, University of Montana
Stephen P. Roberts
Department of Biology, Central Michigan University

1
1
Great Clarendon Street, Oxford ox2 6dp
Oxford University Press is a department of the University of Oxford.
It furthers the University’s objective of excellence in research, scholarship,
and education by publishing worldwide in
Oxford New York
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With offices in
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South Korea Switzerland Thailand Turkey Ukraine Vietnam
Oxford is a registered trade mark of Oxford University Press
in the UK and in certain other countries
Published in the United States
by Oxford University Press Inc., New York
© Jon F. Harrison, H. Arthur Woods, and Stephen P. Roberts 2012
The moral rights of the authors have been asserted
Database right Oxford University Press (maker)
First published 2012
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
without the prior permission in writing of Oxford University Press,
or as expressly permitted by law, or under terms agreed with the appropriate
reprographics rights organization. Enquiries concerning reproduction
outside the scope of the above should be sent to the Rights Department,
Oxford University Press, at the address above
You must not circulate this book in any other binding or cover
and you must impose the same condition on any acquirer
British Library Cataloguing in Publication Data
Data available
Library of Congress Cataloging in Publication Data
Data available
Typeset in Adobe Garamond Pro by Cenveo Publisher Services
Printed and bound by
CPI Group (UK) Ltd, Croydon, CR0 4YY
ISBN 978–0–19–922594–1 (Hbk)
ISBN 978–0–19–922595–8 (Pbk)
10 9 8 7 6 5 4 3 2 1
 We dedicate this book first to our wives, Julie, Creagh, and Jennifer,
who nurtured our families and paid the bills while we got to bury our
heads in the sands of insect environmental physiology.
Second, we dedicate this to our scientific “mothers” and “fathers”
who collected the data and developed the conceptual models we’ve
stolen and amended: V.B. Wigglesworth, Torkel Weis-Fogh,
Liz Bernays, Mary Chamberlin, George Bartholomew, Peter Miller,
Reg Chapman, Neil Hadley, John Phillips, Todd Gleeson, Joel Kingsolver,
Ray Huey, John Lighton, Tim Bradley and Stefan Hetz come to mind,
but there are many others.
 Ecological and Environmental Physiology Series (EEPS)
Series Editor: Warren Burggren, University of North Texas

This authoritative series of concise, affordable volumes provides an integrated

overview of the ecological and environmental physiology of key taxa including
birds, mammals, reptiles, amphibians, insects, crustaceans, mollusks, and fish.
Each volume provides a state-of-the-art review and synthesis of topics that are
relevant to how that specific group of organisms have evolved and coped with the
environmental characteristics of their habitats. The series is intended for stu-
dents, researchers, consultants, and other professionals in the fields of physiology,
physiological ecology, ecology, and evolutionary biology.
A Series Advisory Board assists in the commissioning of titles and authors,
development of volumes, and promotion of the published works. This Board
comprises more than 50 internationally recognized experts in ecological and
environmental physiology, providing a combination of both depth and breadth
to proposal evaluation and series oversight.
The reader is encouraged to visit the EEPS website for additional information
and the latest volumes (http://www.eeps-oxford.com/). If you have ideas for new
titles in this series or just wish to comment on EEPS, please do not hesitate
to contact the Series Editor, Warren Burggren (University of North Texas;
[email protected]).

Volume 1: Ecological and Environmental Physiology of Amphibians

Stanley S. Hillman, Philip C. Withers, Robert C. Drewes, Stanley D. Hillyard

Volume 2: Ecological and Environmental Physiology of Birds

J. Eduardo P.W. Bicudo, William A. Buttemer, Mark A. Chappell,
James T. Pearson, Claus Bech

Volume 3: Ecological and Environmental Physiology of Insects

Jon F. Harrison, H. Arthur Woods, Stephen P. Roberts
Contents

Acknowledgments ix

1 Introduction 1
1.1 Overview of this Book 1
1.2 The Ecological Importance of Insects 2
1.3 Insects as Models for the Study of Ecological Responses 3
1.4 Insect Evolutionary History and Phylogenetic Diversity 5
1.5 Themes of this Book 8

2 Basic Insect Functional Anatomy and Physiological Principles 16

2.1 Insect Lifecycles 16
2.2 The Cuticle 17
2.3 Insect Muscles 23
2.4 Insect Respiratory Systems 31
2.5 Insect Circulatory Systems 39
2.6 Metabolic Systems in Active and Resting Insects, and in Diapause 47
2.7 Insect Feeding and Digestive Systems 50
2.8 Insect Renal Systems 56
2.9 Insect Nervous System 57
2.10 Energizing Nutrient and Water Transport across Insect Epithelia 62

3 Temperature 64
3.1 Defining the Problem 64
3.2 Sensing Temperature and Control 72
3.3 Heat Balance and Mechanisms of Thermoregulation 76
3.4 Thermotolerance Mechanisms 88
3.5 Evolution of Thermoregulation and Thermotolerance 97

4 Water 102
4.1 Defining the Problem 102
4.2 Whole-Insect Water Balance 103
4.3 Evolution of Desiccation Resistance in Drosophila 113
4.4 Hormonal Control of Water Balance 117
4.5 Controlling Where Water Goes: Aquaporins 122
viii | Contents

4.6 Insects Living in Water 128

4.7 Why are There No Marine Insects? 137

5 Nutrition, Growth, and Size 140

5.1 Defining the Problem 140
5.2 Behavioral Responses to Food 161
5.3 Post-ingestive Responses of Individual Insects to Food 169
5.4 Thermal Effects on the Rate and Efficiency of Feeding and Growth 180
5.5 Adaptations of Feeding and Digestive Systems in
Populations and Species 183
5.6 Regulation of Growth and Development 198
5.7 Body Size 210

6 Oxygen 216
6.1 Defining the Problem 216
6.2 Responses to Hypoxia 233
6.3 Sensing and Responding to Hypoxia 247
6.4 Living at High Altitudes 251
6.5 Living in Water 256

7 Techniques and Applications 263

7.1 Insect Molecular Biology 263
7.2 Working with Insects–Behavior and Physiology 264
7.3 Metabolism, Gas Exchange, Internal Gases, Air Flow 268
7.4 Water, Ions, Transport 274
7.5 Temperature 278
7.6 Cardiac Physiology 280
7.7 Issues with Using the Comparative Method in Insects 282

8 Conclusions and Future Directions 283

8.1 Insects and Anthropogenic Perturbation of the Environment 283
8.2 Lifecycle Approach 284
8.3 Interactions of Behavior, Plasticity, and Evolution 284
8.4 Environmental Physiology of Insects from the Field 285
8.5 Paleophysiology 287
8.6 Fundamental Principles of Scaling 288
8.7 Remote monitoring of Insect Physiology, Behavior, and Environment 288
8.8 High-end Imaging 289
8.9 Internationalization of Environmental and Ecological Insect Physiology 290

Bibliography 291
Index 367
Acknowledgments

We thank and acknowledge the current community of working insect physiolo-

gists and comparative and evolutionary physiologists, whose hard work is pushing
the boundaries of our field so fast that any text can only be a blurry snapshot.
Warren Burggren deserves a special thank you for envisioning and directing this
series, which acknowledges that evolutionary history makes the environmental
physiology of each taxon in some way unique. We thank our students and col-
leagues who made comments on portions of this manuscript, including Arianne
Cease, Thomas Förster, Melanie Frazier, Sue Nicolson, Brent Sinclair, Peter
Piermarini, James Waters, John VandenBrooks, Jaco Klok, Sydella Blatch, and
Keaton Wilson. Melanie Frazier and Lee McCoy graciously hosted and fed us
through a week of furious writing at their lovely home on Beaver Creek, and pro-
vided many helpful scientific comments and discussions. Their dog Pierce gra-
ciously allowed us into the living room and permitted us to pet him. The Whiteley
Center at the Friday Harbor Laboratories was a wonderful and peaceful location
for two weeks of intensive writing. Milcho Penchev, Elana Niren, Adam House,
Stephanie Heinrich, and Brian Foster did the yeoman’s work of helping with the
reference library and preparing figures. The IOS Division of the National Science
Foundation has funded much of the research described here, including most of
the work from our own labs. We thank our institutions for being supportive of
research and scholarly activity, especially a sabbatical from ASU to JFH.
This page intentionally left blank
 1
Introduction

1.1 Overview of this Book

Ecological and environmental physiology is the study of how physiological sys-

tems respond, over different timescales, to variation in physical and biological
environments. This introduction describes the ecological importance of insects,
some of the advantages of insects as experimental models, a brief introduction to
insect phylogeny and evolution, and the themes of the book. Chapter 2 is
designed for the physiological ecologist without a background in entomology. It
introduces some basic anatomy and physiology of insects by system, and provides
references for detailed information on these topics. The heart of the book
(Chapters 3–6) examines responses of insects to the classic environmental param-
eters that vary across the planet and that determine insect distributions. Chapter
3 examines the pervasive effects of temperature on insect physiology, ecology, and
evolution, covering heat balance, thermoregulation, and mechanisms of thermo-
tolerance. Chapter 4 examines the role of water, covering organismal balancing
and mechanisms for coping with deficiency or excess. Chapter 5 examines the
role of nutrient availability and its effects on growth, body size, and reproduc-
tion. Chapter 6 examines the role of oxygen, focusing on the structure and func-
tion of the respiratory system and its plasticity in response to changing supply
and demand. In choosing these variables we excluded others of interest, includ-
ing light, pollutants, and the biotic environment (social interactions, parasites,
and disease), but we could not include a careful coverage of these in a space-
constrained book. The concluding chapters of the book consider insect-specific
methodological approaches in environmental physiology (Chapter 7) and pro-
pose a set of conclusions and future directions (Chapter 8).
There are multiple ways to write such a book. One approach is the comprehen-
sive summary. However, the field is simply too large, and the coverage too uneven,
to make such an approach feasible. For a few species (e.g. Drosophila
melanogaster) there is a wealth of genetic and cellular studies, and some on envi-
ronmental physiology. Many other fundamental questions have been addressed
only in a few scattered species. We have therefore taken a different approach that,
we hope, will both summarize important ideas and highlight gaps.
2 | Introduction

In each chapter, topics are organized around a set of fundamental questions.

Each major chapter begins by ‘defining the problem’. In this section, we cover
what is known about pathologies that ensue when responses to environmental
change are inadequate, and provide theoretical background. We hope to use this
section to introduce the functional and theoretical relevance of the physiological
responses described later in each chapter.
Three themes run through each chapter. The first is that different physiological
responses occur over different timescales. These responses include passive buffer-
ing, short- and medium-term active responses, developmental plasticity, cross-
generational plasticity (i.e. epigenetic inheritance), multigenerational evolution
(within population selection), and deep evolutionary responses. The second
theme is that active responses require control systems. Here we address what is
known about sensors, integrators, and effectors for each type of response. The
third theme is trade-offs, because all organisms have limited resources (e.g. mate-
rials, energy, space, time) the enhanced capacity for one function may reduce the
capacity for others. Indeed, trade-offs have also constrained our writing of this
book: we have sought to identify provocative patterns and interesting directions
for future research at the expense of providing exhaustive summaries.
Such an approach will inevitably run into the problem of data scarcity in some
areas and overload in others. Our solution is to point out holes in the literature
and to give short shrift to many other areas. We ask our un-cited colleagues for
their forgiveness.

‘It is a capital mistake to theorize before one has data. Insensibly one begins
to twist facts to suit theories, instead of theories to suit facts.’
Sherlock Holmes in A Scandal in Bohemia by Arthur Conan Doyle.

We attempt to follow Sherlock Holmes’s advice, but—reader beware!

1.2 The Ecological Importance of Insects

Insects often are the most important heterotrophs in terrestrial ecosystems.

During outbreaks of the gypsy moth, Lymantria dispar (L.) (Lepidoptera:
Lymantriidae), nearly 100% of available host leaves can be consumed (Kosola
et al., 2001). A recently observed locust swarm in Morocco was 230 km long, at
least 150 m wide, and contained an estimated 69 billion locusts (Ullman, 2006).
While insect outbreaks occur infrequently, they are common in many ecosystems
and can have long-lasting effects on communities. During such outbreaks, insects
are thought to function as keystone species that reduce abundance of other dom-
inant species, which can subsequently increase biodiversity (Carson et al., 2004).
Insects also have a major role in secondary production due to their high
 Insects as Models for the Study of Ecological Responses | 3

abundance and assimilation efficiencies relative to vertebrates (Price, 1997). In a

study of energy flow through old temperate fields, insects (primarily Orthoptera
and Hymenoptera) accounted for 80% of total energy assimilation by heterotro-
phs (Wiegert and Evans, 1967). In agriculture, insecticide use is reported to
increase plant production by 30% or more, suggesting that insect herbivory exerts
tremendous loads (Price, 1997). These pesticides can have major effects on
humans and non-target wildlife (Matson et al., 1997), indicating that we need to
better understand and control insects.
In other ecological situations, insects typically have more modest effects on
nutrient transfer through ecosystems. A variety of studies have suggested that the
standing crop of insects is relatively low compared to plant and vertebrate bio-
mass, and that, in most ecosystems (particularly forests), insects consume less
than 10% of net primary productivity (Weisser and Siemann, 2004). However,
these studies have emphasized that even when abundances are low, insects do
have strong effects on plant investment in secondary metabolites and structures,
and on rates of mineralization and availability of phosphorus and nitrogen
(Weisser and Siemann, 2004). In grasslands, insect herbivory has been shown to
increase nutrient cycling and plant productivity (Belovsky and Slade, 2000).
These studies together suggest that the most important effects of insects on eco-
systems are indirect; for example, selective herbivory and pollination can strongly
affect plant species’ composition (Weisser and Siemann, 2004). Pollination is a
particularly critical contribution of insects. In addition to being essential for most
wild angiosperms, insects pollinate 84% of the world’s food crops, representing
one third of global food production, with an estimated annual replacement value
of $68–433M for the USA alone (Allsopp et al., 2008).

1.3 Insects as Models for the Study of Ecological Responses

The jagged exoskeletons, protruding horns, large compound eyes, lacey wings,
feathery antennae, sharp stings, and painful bites of insects have always fascinated
humans. Their extremes in size, shape, and lifestyle have even served as models
for our concept of alien. In one sense, these divergent life forms provide an
opportunity to understand how the physical and chemical aspects of the environ-
ment shape all life; for example similar biomechanical principles can explain
wing-to-body ratios in birds and insects (Dudley, 1992). Despite the obvious
differences, many profound similarities exist between humans and insects, such
as the virtually identical organization of major biochemical pathways (glycolysis,
Krebs cycle, transcription, and translation). We (humans and insects) shared a
common ancestor around 600 million years ago, more than half of insect and
vertebrate genes are orthologous, and these have a mean sequence identity of
about 46% (Consortium, 2006). This fundamental similarity is one reason why
4 | Introduction

it is so difficult to find insect-specific pesticides. Some of the most significant

questions about how life works have first been asked of insects, with the answers
later found to apply to all eukaryotes.
All organisms respond to environmental variation, and the goal of environ-
mental physiology is to determine the proximate mechanisms and functional
significance of such responses, as well as the mechanisms that allow different spe-
cies to occupy different niches. Proximate questions ask about mechanisms: how
do organisms sense change? What regulatory or compensatory responses occur?
What role does the neuroendocrine system play in the response? Are specific
genes activated? How effective is compensation? Ultimate questions ask why
organisms show such responses, and the answers often represent amalgams of
current selection and historical constraint. Do responses enhance fitness, or do
they indicate stress-induced pathologies? Do organisms evolve different types of
physiologies in response to spatial scale (local vs. global climate change)?
What historical pathways have made some insects sensitive to change and
others hardy? What fitness trade-offs are associated with different physiological
phenotypes?
Insects are uniquely suited for the study of physiological responses to environ-
mental variation because one often can ask both proximate and ultimate ques-
tions. Although the small size of insects can make physiological measurements
challenging (e.g. sampling blood repeatedly from individuals), sensor miniatur-
ization is overcoming these limitations, and most standard physiological tools
can be applied to at least some insects. The genomes of more than 20 insects have
been completely sequenced and many more (more than100) are in the pipeline.
The genes of no other animal are currently manipulated as easily as those of
Drosophila melanogaster. Ease and cost-effectiveness of rearing many individuals
in small spaces makes insects uniquely suited for powerful experimental designs.
Insects can often be easily observed in large numbers in the field and then col-
lected and reared in the laboratory. The ethics of using insects for physiological
research are less complicated than for vertebrates, and therefore many animal care
and use committees actively promote insects as model organisms.
Many insect species are also exceptionally suited to answering ultimate ques-
tions. The short lifespan of insects allows students to examine development or
evolution during a graduate thesis, an option not available for many vertebrate
studies. The wide diversity and number of species allows comparative studies
with much greater scope than is possible in most vertebrate groups (though the
lack of phylogenies for many insect groups remains a challenge). Many insects
can be reared and mated in the lab, allowing long-term natural and artificial
selection experiments. Such laboratory selection studies can only be conducted in
a handful of small vertebrate species. The ability to address ultimate questions
with repeatable laboratory experiments provides researchers with exceptionally
powerful tools for understanding evolutionary responses of insects.
 Insect Evolutionary History and Phylogenetic Diversity | 5

1.4 Insect Evolutionary History and Phylogenetic Diversity

Insects evolved at least 400 million years ago, probably in the early Devonian or
late Silurian, and therefore are among the earliest land animals (Grimaldi and
Engel, 2005; Glenner et al., 2006). Recent molecular evidence suggests that
hexapods are most closely related to branchiopods, a freshwater crustacean group
that includes water fleas and fairy shrimp (Glenner et al., 2006; Mallatt and
Giribet, 2006). This finding suggests that the common ancestor of branchiopods
and hexapods evolved in fresh water (Fig. 1.1), and that crustaceans, rather than
myriapods, (centipedes, millipedes) are the most closely related major taxa to
insects (Glenner et al., 2006; Bradley et al., 2009). Thus insects can be considered
terrestrial crustaceans.
Most modern insect orders and some modern families had evolved by the
Cretaceous (ca. 100 million years ago, Fig. 1.2). This long evolutionary history
has important implications for insect environmental physiology. In particular,
major groups of insects are only distantly related. For example, polyneuropterans
such as grasshoppers and cockroaches have not shared a common ancestor with
Panorpida (including flies and moths) since the Carboniferous, more than 300
million years ago. This deep evolutionary history means that many physiological
characteristics of insects diverge strongly along taxonomic lines.
Consideration of insect evolution can help us understand which features are
derived in insects. The most primitive insects are the Archeognatha (bristletails)
and the Zygentoma (silverfish) (Fig. 1.3). These groups possess basic insect char-
acteristics (segmented, six-legged adults, external mouthparts, antennae without
muscles beyond the initial segment, a chordotonal organ in the base of the anten-
nae, compound eyes, ocelli, tracheal system, Malpighian tubules). However, they

Decapods Branchiopods Hexapods

Crustaceans

Land
Million years ago

Ocean Freshwater
360
Devonian
410
Silurian
440
Ordovician 500
Cambrian 570

Fig. 1.1 Evolution of hexapods. From Glenner et al. (2006), used with permission from
AAAS.
6 | Introduction

400 300 200 100 million years ago (mya) 0

P A L E O Z O I C M E S O Z O I C Cenozoic
Silurian Devonian Carboniferous Permian Triassic Jurassic Cretaceous
Entognatha
Archaeognatha
Zygentoma
Insecta

Ephemeroptera
Protodonata
Odonata
Pterygota: wings

Palaeodictyopterida

Plecoptera
Embiodea
Zoraptera

Polyneoptera
Dermaptera
Grylloblattodea
Mantophasmatodea
Orthoptera
Phasmatodea

Dictyoptera
Blattaria
Isoptera
Mantodea
Neoptera: wing folding

Paraneoptera
Psocoptera
Phthiraptera
Thysanoptera
Hemiptera

Neuropterida Antliophora
Coleoptera
Raphidioptera
Megaloptera
Neuroptera
Hymenoptera
larvae
Holometabola

Mecoptera

Panorpida
Siphonaptera
Diptera
Strepsiptera
Trichoptera
Lepidoptera

Fig. 1.2. Insect phylogeny in relation to geologic time. From Grimalidi and Engels (2005),
with permission from Cambridge University Press.

also exhibit several key primitive characteristics, including lack of wings, a single
mandibular articulation, and projections on the abdomen that may be remnants
of ancestral legs (Grimaldi and Engel, 2005). Like other relatively primitive
insects, they are hemimetabolous, without a pupal stage marking a profound
change between juvenile and adult stages.
Phylogenetic and fossil data provide the best support for the hypothesis that
insects evolved first on land, with aquatic forms evolving later, and indepen-
dently, in many groups. One class of evidence for this hypothesis is that all of the
known non-insect hexapods (Entognatha, which have internal mouthparts and
include springtails, proturans, and diplurans) are terrestrial. This suggests that
terrestriality evolved in ancestors to the Entognatha and insects (the Ectognatha),
after fresh-water derived hexapods colonized land in the late Devonian (Glenner
et al., 2006). In addition, the earliest true insects (Archaeognatha: bristletails and
Zygentoma: silverfish) are completely terrestrial. The oldest insect fossils, from
the Devonian, Carboniferous, and Permian, are all terrestrial, with no clear
aquatic insect fossils known before the late Triassic (Grimaldi and Engel, 2005).
More support for this hypothesis is that the air-filled tracheal system of insects
would have been unlikely to evolve in aquatic organisms. Nearly all insects have
 Insect Evolutionary History and Phylogenetic Diversity | 7

Fig. 1.3 A bristletail, Nesomachilis sp. [Archaeognatha: Meinertellidae], illustration by

Geoff Thompson, reprinted with permission from the Queensland Museum, 1985.

a terrestrial adult stage, and those that don’t—certain beetles and hemipterans—
are clearly derived from terrestrial ancestors. Finally, insects are virtually excluded
from marine environments, suggesting that the insect bauplan and physiology
are poorly suited for deep waters. Most likely, this is because possession of a tra-
cheal respiratory system precludes deep diving due to excessive buoyancy
(Maddrell, 1998) or hydrostatic pressure effects on tracheal system function.
While the major orders of insects are at least 100 million years old, extant species
are much younger, with most estimates ranging from 2 to 10 million years (Grimaldi
and Engel, 2005). Extensive studies from the Pleistocene have demonstrated
8 | Introduction

that beetles dramatically changed their distributions as glaciers advanced and

receded, allowing them to maintain similar climate and plant niches, but at dif-
ferent places and times (Coope, 1979). These data strongly support the hypoth-
esis that characteristics of species, such as preferred temperature, can be relatively
invariant over millions of years, as long as dispersal allows habitat-tracking.
The most remarkable aspect of insects is their diversity. About one million
species of insects have been described, which represents at least half of docu-
mented global species diversity (Gullan and Cranston, 2005). High insect
diversity likely relates to their environmental physiology. First, small size allows
insects to occupy small ecological niches compared to vertebrates, and to better
mitigate macroenvironmental conditions by choosing microhabitats. Flight
provides tremendous advantages relative to terrestrial animals in foraging and
dispersal. High reproductive output provides the grist for natural selection, and
the capacity to rapidly produce new, successful biotypes. The tremendous flexi-
bility of insect morphology and physiology, especially regarding capacities to
feed and reproduce on different plants, certainly plays a key role in the great
biodiversity of this group.

1.5 Themes of this Book

1.5.1 Timescale of Environmental Responses

During environmental challenges, organismal responses will depend strongly
on the duration of exposure. For the sake of simplicity, physiological responses
can be divided into six functionally distinct stages, based on temporal duration
(Fig. 1.4).
These six temporal classes clearly overlap (e.g. the distinction between macro-
molecular remodeling and developmental plasticity relies on sufficient time for
altering a developmental pathway, a continuous rather than binary response), but
these definitions provide a useful paradigm for understanding the duration of
exposure on organismal response.

1.5.1.1 Passive Buffering

Changes in environment produce changes in internal state. When sunlight falls
on a shaded insect, temperature rises to a new steady-state. Consumption of a
base-generating food quickly raises the blood and tissue pH. The result is a
passive, physical and chemical change in the state of the organism from A0 to A1
that stem from no active response by the animal.
However, state changes for given environmental changes are not the same for
all organisms, as they depend on many different organismal characteristics. We
divide them arbitrarily into structural properties and biochemical equilibria.
 Themes of this Book | 9

Time Sequence of responses

Environmental
A0 impact

Instantaneous Passive buffering

to seconds
A1
Subseconds Behavioral or biochemical
to hours switching

A2

Seconds Macromolecular
to days remodeling

A3

Throughout Developmental plasticity

ontogeny
A4

Generations
Within species evolution
within species
A5

Eras Macroevolution

B1 B2 B3

Fig. 1.4 Effect of duration of exposure on type of response to the environment.

Structural characteristics are important components of passive buffering. For

example, the size of an insect determines how quickly it will heat or cool and how
quickly it will dehydrate. For an insect suddenly exposed to sunlight, its exposed
surface area, color, and reflectance determines its new steady-state temperature.
For a phytophagous insect consuming a base-generating leaf, a smaller pH change
will occur when hemolymph and tissue space are larger.
Biochemical equilibria also contribute to passive physical and chemical buffer-
ing. According to Le Chatelier’s principle, in an equilibrium reaction:

A+B↔C D (1)

Increasing A leads to increased formation of C and D. A classic example of

such an effect is buffering of acid.

H+ + A − ↔ HA (2)
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