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Animal Athletes
Animal Athletes
An Ecological and Evolutionary
Approach

Duncan J. Irschick
Professor, Department of Biology, University of Massachusetts Amherst, USA

Timothy E. Higham
Associate Professor, Department of Biology, University of California, Riverside, USA

1
1
Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
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. Oxford is a registered trade mark of
Oxford University Press in the UK and in certain other countries
© Duncan J. Irschick & Timothy E. Higham 2016
The moral rights of the authors have been asserted
First Edition published in 2016
Impression: 1
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, by licence 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 work in any other form
and you must impose this same condition on any acquirer
Published in the United States of America by Oxford University Press
198 Madison Avenue, New York, NY 10016, United States of America
British Library Cataloguing in Publication Data
Data available
Library of Congress Control Number: 2015944343
ISBN 978–0–19–929654–5 (hbk.)
ISBN 978–0–19–929655–2 (pbk.)
Printed and bound by
CPI Group (UK) Ltd, Croydon, CR0 4YY
Links to third party websites are provided by Oxford in good faith and
for information only. Oxford disclaims any responsibility for the materials
contained in any third party website referenced in this work.
 Foreword

This book examines how animal athletes have evolved. While many of us
might normally think of the activities associated with “athletes” as consisting
of only running and jumping, we set out to show that animals perform many
tasks to an amazing degree of accomplishment. Some of the examples might
seem obscure at first—for example, how can the rattling tail of a rattlesnake be
considered an “athletic feat”? Once one realizes that the snakes vibrate their
tail at 90 Hz for over an hour, then one’s view changes. If you don’t believe us,
try shaking your hand as quickly as you can for an hour! In other words, the
mundane world of animals is filled with extraordinary accomplishments, and
our book sets out to celebrate this wonderful diversity by focusing not just on
spectacular feats of running and jumping but also on feeding, vocalization,
diving, flying, and many other feats. Each of these facets operates in a larger
evolutionary and ecological context, and therefore we wanted to examine not
only the “how” of amazing animal athletic feats but also the “why.” Why do
seals dive so deep? Why do some lizards live on twigs? Why do frogs vocalize
for long periods of time when, in so doing, they expend a tremendous amount
of energy? Our belief is that only in the broader context of ecology and evolu-
tion can these questions be answered and the broader pattern of animal ath-
letic traits be understood.
Therefore we delve into many areas—such as, for example, how natural
selection and sexual selection operate on performance traits, the musculoskel-
etal basis of these traits, and how such traits vary in different geographical or
evolutionary settings. Our examples are wide-ranging and are drawn from
a range of invertebrates and vertebrates, including frogs, lizards, sharks, ro-
dents, bats, birds, insects, spiders, and more! Our goal was not to be compre-
hensive in our analysis of different systems, and therefore our approach is ex-
emplary rather than exhaustive. We took this approach to enhance readability,
and readers are recommended to read many of the cited papers and books to

v
 vi Foreword

learn more on a certain topic. Our language was intended to be simple, and
therefore not every study is covered in the exhaustive detail one might expect
to find in a typical academic paper. Our book was hopefully written so that
many could access it, but we include enough detail that (we hope) will please
specialists. We sincerely hope that this book inspires a new way of thinking
about animal athletics and spurs new research into the integrative biology of
complex functional traits.
This book could not have been written without the assistance of many in-
dividuals. A number of individuals read key chapters and provided invalua-
ble assistance. These individuals include Anthony Herrel, Scott Kelly, Rodger
Kram, Henry Astley, Kiisa Nishikawa, William Hopkins, Jerry Husak, Jona-
than Losos, Daniel Moen, Jennifer Grindstaff, Simon Lailvaux, Sheila Patek,
Raoul Van Damme, Roberto Nespolo, and others. Several discussions with
members of the Irschick and Higham lab groups solidified some ideas regard-
ing performance as well as helped identify idiosyncrasies that frequently arise
in science. Our editors Lucy Nash and Ian Sherman were especially helpful in
guiding us throughout the book-writing process by keeping us on schedule
and providing invaluable assistance in the editing process. Their patience dur-
ing the ten (!) years of writing and during our many twists and turns is greatly
appreciated. Finally, we could not have written this book without the support
of our families. Specifically, Duncan Irschick thanks Jitnapa Suthikant, as well
as Darwin and Calder Irschick for their support and patience. Tim Higham
would like to thank Melissa, Daphne, Iris, and Violet for their encouragement,
enthusiasm, and endless support.
Contents

1 Animal performance: An overview 1

1.1 Why study performance? 1
1.2 Definitions of performance 6
1.3 The hierarchical nature of biological systems 10
1.4 Variability, repeatability, and heritability 11
1.5 The ecological context of performance 13
1.6 The evolution of performance 15
1.7 Behavior and performance 17
1.8 Mechanistic and energetic constraints on
performance 21

2 The ecology of performance I: Studies

of fitness 24
2.1 Natural selection, sexual selection, and performance 24
2.2 Path diagrams of selection 25
2.3 Definitions of fitness 27
2.4 Quantifying the impact of natural and sexual
selection 28
2.5 Does high performance result in higher fitness? 29
2.6 Is selection on morphology stronger than for
performance capacity? 35
2.7 Manipulative studies of animal performance
and selection 38
2.8 Complex forms of selection on performance 43

3 The ecology of performance II: Performance

in nature 47
3.1 Why study animal performance in nature? 47
3.2 What is ecological performance? 49

vii
viii Contents

3.3 Impact of behavior on ecological performance 50

3.4 Environmental impacts on ecological performance 53
3.5 What percentage of maximum capacity do animals
use in nature? 59
3.6 Emergent behaviors in the field: Defying
physiological constraints 61
3.7 Slackers and overachievers: Behavioral
compensation in nature 62

4 The ecology of performance III: Physiological

ecology 67
4.1 Environmental influences on performance 67
4.2 Energetics and movement 68
4.3 Aerobic versus anaerobic metabolism 71
4.4 Born to run: Variation among animal species
in the energetics of movement 73
4.5 Feel the burn: Harmful byproducts of movement 76
4.6 Temperature and animal performance 78
4.7 Acclimation and temperature 82
4.8 The ecological context of temperature 85

5 The evolution of performance I: Mechanism

and anatomy 91
5.1 Mechanistic limits on performance 91
5.2 The mechanism and anatomy of trade-offs 92
5.3 Variation in anatomical structure as a foundation
of animal performance 96
5.4 Creative arrangement of anatomical structure to
enable high performance in suboptimal settings 99
5.5 Anatomical structure as a way of overcoming
physical limits 103
5.6 Highly specialized performance: How do they
do that? 108
5.7 Elastic mechanisms for powering rapid movements 110

6 The evolution of performance II: Convergence,

key innovations, and adaptation 117
6.1 How does performance evolve? 117
6.2 Quantitative analysis of evolutionary data 119
6.3 A model for understanding the evolution of
performance 120
6.4 Convergent evolution of morphology and
performance traits 120
 Contents ix

6.5 Tinkering and animal performance: How variation

among species has resulted in evolution of novel
performance capacities 130
6.6 Key innovations, adaptive radiation, and the invasion
of novel habitats 135

7 Trade-offs and constraints on performance 143

7.1 Trade-offs, performance, and optimization 143
7.2 Why expect a trade-off? 144
7.3 Trade-offs within and among species 146
7.4 Mechanical constraints on performance 147
7.5 Innovations, constraints, and trade-offs 155
7.6 Ecological and reproductive constraints on
performance 158
7.7 Overcoming trade-offs: The role of behavior 160

8 Sexual selection and performance 163

8.1 What is sexual selection? 163
8.2 A functional approach to sexual selection 164
8.3 Male competition and performance 165
8.4 Are sexual signals honest signals of male
performance? 167
8.5 Sexual selection imposing costs on performance 171
8.6 Female choice and performance 174
8.7 Asymmetry and performance 177

9 Extreme performance: The good, the bad,

and the extremely rapid 181
9.1 Extreme performance: If the Olympics were open
to oribatid mites 181
9.2 Extreme performance: Overcoming limits 182
9.3 Need for speed: Extremely rapid movements 184
9.4 The evolution of morphological novelties 190
9.5 Morphological and physiological mechanisms of
extreme performance 192
9.6 Behavioral means for greatly enhancing
performance 195

10 Genetics, geographic variation, and community

ecology 200
10.1 Animal athletics from a population perspective 200
10.2 The genetic basis of animal athletics 202
10.3 Geographic variation in performance capacity 205
x Contents

10.4 The role of community structure in molding animal

athletics 208
10.5 Many-to-one mapping, and communities 214

11 Human athletics: A link to nonhuman

animals? 220
11.1 Humans versus other animals 220
11.2 Performance specialists versus generalists 221
11.3 Training impacts 222
11.4 Steroids and performance 226
11.5 Are athletes anatomically different? 229
11.6 Extreme sports and human mortality 231

12 Conclusion 236

Artwork credit lines 239

Index 243
 1 Animal performance
An overview

1.1 Why study performance?

What is performance ability? If one were to ask this question in any class-
room or lecture hall (and we have!), one would immediately be greeted by a
showing of hands, followed by a variety of different opinions. In the common
vernacular, good “performance” can mean almost anything, such as how well
one dances, sings, or even earns money. Simply put, “performance” can mean
many things to many people, but a concept is only as useful as it is specific and
explanatory. An overly broad use of the term “performance” is meaningless
unless it allows us to examine a common body of knowledge. Fortunately, in
the world of functional morphology and evolution, this term has enjoyed a
more specific meaning that we will focus on throughout this book: perform-
ance capacity is any quantitative measure of how well an organism performs
an ecologically relevant task that is vital for survival. As explained below,
there are some more stringent assumptions behind this definition, but let’s
put that aside for now. Some classic examples include how fast an animal can
run, jump, bite, fly, or perform nearly any athletic feat. You could even think
of animal performance as being analogous to the human Olympics, although
in the nonhuman animal world, the consequences of a bad day at the track
are far more dire than any for even the most heartbroken athlete who has lost
his or her event. Indeed, one of the hallmarks of animal performance is that
nonhuman animals can place themselves at tremendous risk when perform-
ing “animal Olympics”; a cheetah or lion running after its prey can have its

Animal Athletes. Duncan J. Irschick and Timothy E. Higham. © Duncan J. Irschick and Timothy E.
Higham 2016. Published in 2016 by Oxford University Press.
 2 Animal Athletes

jaw dislocated if it miscalculates during the closing phase, and a miscalculated

bite on a hard object can irreversibly damage a dog’s jaws. One could specu-
late about the level of performance we might see at the human Olympics if
lions were chasing those sprinters!
A study we had each heard of when we were graduate students crystal-
lizes the difference between nonhuman animal and human performance. In
a study in the 1980s, the physiologist Todd Gleeson performed a simple ex-
periment with two groups of fence lizards (Sceloporus; a lizard commonly
seen basking on trees, fences, and rocks in the western United States). For the
“control” group, he allowed the lizards to stay in their small cages for about
6 weeks (Gleeson 1979). For the “experimental” group, he induced them to
perform aerobic exercise on a treadmill in a training regimen for the same
length of time. After 6 weeks had passed, he compared the aerobic capaci-
ties of both groups (the control and experimental) and found, much to his
surprise, that the two groups did not differ! Obviously, if one performed the
same experiments with humans, one would find that the group that had been
trained would not only have superior aerobic capacities compared to the con-
trol group but also possess several physiological differences, such as increased
capillary networks adjacent to large skeletal muscles, increased maximum aer-
obic capacity, improved blood pressure, and so forth. While not every animal
shows this lack of a response to exercise (see chapter 11), for those species that
do, it signifies the profound divide between human and nonhuman animal
physiology. Sceloporus lizards rely on both burst speed and aerobic capacity to
elude predators, capture prey, and defend territories, and the relentless pres-
sure of natural selection over millions of years has not only removed many of
the “weakest” individuals, it has done so to such an extent that natural selec-
tion has a far more limited menu to choose from. In other words, performance
capacity is life and death for animals and has shaped their morphology, be-
havior, and physiology over vast stretches of evolutionary time.
The concept of performance also holds a key place in the human conscious-
ness, both in terms of our own lives and for the natural world around us.
Much of our obsession with performance dates to Charles Darwin and his
synthetic theory of natural selection. A central thrust of his theory of natural
selection is that some organisms will be more fit than others, and one of the
main reasons for this variation in fitness is the ability of animals to perform
such feats as running, jumping, biting, and so forth. Consequently, one impli-
cation of his theory is that some organisms are better performers than others
and that this variation is vitally important to survival. His theory has also been
extrapolated to include social aspects of human behavior—“Darwinism” or
the “survival of the fittest,” the idea that only the strong survive—an appeal-
ing, though somewhat misguided, view of human culture. Nonetheless, the
idea of performance capacity as an arbiter of who lives or dies, or reproduces
 Animal performance: An overview 3

or not, has percolated in our collective consciousness and has been a spur for
biological research into the process of adaptation.
We are fascinated with the traces that, in our everyday lives, reflect our ob-
session with performance. Each year, people lose millions of dollars betting
on which dog or horse will be the winner at the racetrack, and entire countries
have been known to explosively celebrate or fall into despair at the success or
defeat of their favorite athletes at the Olympics. It is performance that drives
almost 200,000 people to attend a single football (termed “soccer” in North
America) match in Brazil. At such events, one can clearly see the range of
performance abilities in nonhuman animals and humans; while some horses
or humans are superior runners, others lag behind. But the artificial world of
nonhuman animal and human athletics is not the only window into the mys-
teries of animal performance, as the natural world offers many strange and
wonderful examples of the extremes to which animals have evolved various
kinds of performance traits. Even the most jaded observer is fascinated when
watching a small frog, only as small as a thumbnail, emit a piercing call that
echoes over great distances. Most of us marvel at the rapid speeds of chee-
tahs when watching them chase their prey on the African plains. We are often
caught pointing up in the air as hummingbirds zip by on a nice spring day,
saying, “Wow, look at that!” Even in the mundane setting of our homes, the
powerful crunch of a dog’s jaw on a bone is a reminder of the extreme biting
capacities of man’s best friend. Evolution has seemingly endowed animals
with amazing abilities that can exceed the limitations of animal morphology.
This book examines the broad scope of animal performance and integrates in-
formation on morphology, behavior, ecology, and evolution to understand the sig-
nificance of performance in the lives of animals. We have two primary goals: first,
to provide a perspective on the range of performance capacities present in na-
ture; and, second, to understand the ecological and evolutionary context in which
these performance capacities have evolved. To address these broad goals, we will
explore how five factors, namely, behavioral, ecological, evolutionary, morpho-
logical, and physiological, act as innovators and constraints for generating diver-
sity in animal performance. These factors allow us to identify how different parts
of an animal work in concert to execute behaviors. It is during this behavior that
we can define the kind of performance we are interested in. One theme we will
explore is the underlying complexity of animal performance. One cannot fully
comprehend the diversity of animal performance either by reducing an animal
to its parts (reductionism) or by focusing solely on each factor (behavioral, eco-
logical, evolutionary, morphological, and physiological) without considering
interactions among these parts and how the whole organism behaves in nature.
This integrative approach is important because performance capacities emerge
from the whole organism, not from individual parts, and it is the whole animal
that lives and dies and that passes on its genes (or not) to the next generation.
 4 Animal Athletes

On the other hand, there is much to be learned by studying a few components in

detail; but the important point is that reductionist studies are only pertinent when
we also understand how the whole organism functions.
Beyond the aesthetic pleasure one derives from watching animals run,
swim, jump, or fly, why should biologists study animal performance and how
will such a focus provide insights into the broader fields of evolution and ecol-
ogy? To address this, let’s consider how natural selection works. Within any
animal population over a span of time, individuals will either live or die, and
some of those individuals will reproduce, while others will not. For the most
part, factors that cause death or that influence the probability of reproduction,
such as temperature, food, predators, and so forth, act on the whole organism.
Let’s consider an example that explains this point. Imagine a male frog calling
on a summer night. The main reason that male frogs call is to attract mates.
For most frogs, males that call the loudest and the longest are most likely to
attract female frogs (Wells and Taigen 1989) and hence are more likely to re-
produce and increase their overall fitness. Based on this line of logic, it seems
reasonable that every male should have evolved the ability to call both loudly
and for long periods; however, an inspection of both variables (call loudness
and call duration) shows this not to be the case. As with many biological sys-
tems, there is tremendous variation in both traits, with some males being able
to call both louder and longer than other males, often by large margins. The
reason for this inequality is that calling in frogs is energetically expensive
(fig. 1.1) and requires substantial aerobic investment, which is driven by mus-
cles that are specialized for long-duration aerobic movements (Wells and
Taigen 1989). Further, frog vocalization is a complex process, involving both
a variety of muscle groups and complex behaviors, such as extension of the
throat sac. However, because females choose males on the basis of call loud-
ness and call duration, natural selection will operate primarily on these aspects
of male performance (in this case, a specific form of natural selection called
“sexual selection”) and only secondarily on other aspects of this complex sys-
tem. In other words, one can state with certainty that “natural selection favors
frogs with large aerobic muscles” for long bouts of calling, but in fact selec-
tion favors individual frogs that can call for long durations. Female frogs care
nothing for the muscles involved in calling. That feature is favored by natural
selection only because it is necessary for frogs to call for long periods of time.
In short, performance abilities are the “face” that animals put forth in the nat-
ural world, with many other supporting features “hidden” in the organism
(although in some cases, natural selection does act directly on morphology or
behavior alone). This example underscores how one cannot understand the
evolution of unique morphological traits (i.e., the throat sac and the special-
ized muscle groups) or behavior (i.e., vocalization) without understanding the
resultant performance capacity (call intensity and duration) or vice versa.
 Animal performance: An overview 5

A 3.0
Y = 0.18 + 0.0004X
r2 = 0.95
VO2 WHILE CALLING, ml/(g·h)

2.5

2.0

1.5

1.0

0.5
·
0
0 1000 2000 3000 4000 5000 6000 7000
NOTE RATE, notes/h

B 50
No stimulus Playbacks
n = 11 n = 50
40 Median = 1.73 ml/(g·h) Median = 1.55 ml/(g·h)
FREQUENCY, %

30

20

10

0.8 1.2 1.6 2.0 2.4 2.8

·
VO2, ml/(g·h)

Fig. 1.1 A, A plot showing the relationship between note rate and the rate of oxygen consumption
for the frog Hyla microcephala during vocalization. B, A frequency distribution of different oxygen
consumption rates (VO2) for the frog Hyla microcephala during vocalization. The open and filled
bars indicate vocalizations in response to playback or without a stimulus, respectively. Note that, in
A, the higher the note rate, the higher the rate of oxygen consumption. Redrawn from Wells and
Taigen (1989) with permission from Springer publishing. Image is from Wikimedia Commons

Now let’s examine a common predator-prey scenario that reveals both the
importance and subtlety of animal performance for survival. Small fish often
elude predators (e.g., a larger fish) by using rapid bursts of acceleration (Webb
1976), often in the stereotyped form of a C-start, a circular escape motion in
which the fish swims off in the opposite direction from which it was first stimu-
lated (by, for example, a touch on the tail, or the sensation of water pushing up
against its tail). The salient point is that predators only have a very brief period
of time to capture their prey, given this mode of rapid escape, and this time is
usually on the order of milliseconds (a millisecond is one thousandth of a sec-
ond). Whether the predator successfully captures the prey or, from the prey’s
 6 Animal Athletes

point of view, whether the prey lives to see another day depends on many
factors, including the initial attack distance and accuracy of the predator, the
relative sizes of the predator and prey, and the relative performance capacities
of each (i.e., how fast each can accelerate). Clearly, both predator and prey in
such circumstances would evolve many specializations for maximizing their
chances of success (capturing prey or eluding capture, respectively), but one
of the most prominent specializations is the ability to accelerate quickly either
to capture an evasive prey or to evade a quick predator. In other words, we
can think of this simple yet ubiquitous dynamic as a coevolutionary arms race
between contestants for performance supremacy. Performance often comes at
a cost, so it isn’t surprising to see this high level of performance relaxed when
competition is absent. Indeed, if you find a lake where fish live without loom-
ing predators, their ability to accelerate could be reduced, as should the size of
the anatomical parts that contribute to acceleration.

1.2 Definitions of performance

Before proceeding further, let’s revisit the definition of performance capacity
and flesh out some of the underlying assumptions. The ability to quantify a
task is essential and distinguishes performance from other characteristics: one
can measure how hard a dog can bite, count the number of seconds a frog will
croak, measure the adhesive ability of geckos to cling to a surface, and quan-
tify how long it takes Usain Bolt to run 100 m. This ability to quantify perform-
ance is essential for comparing different individuals, or comparing the same
individuals at different times. However, based on this simple definition, many
other kinds of performance can be defined, including how quickly an animal
can digest food, how many offspring a female mammal can raise in a year,
and so forth. In other words, “performance,” if not carefully defined, can be
extrapolated to a myriad of seemingly disconnected actions. From an evolu-
tionary perspective, whole-organism performance holds little meaning if we
attempt to compare “apples and oranges,” such as comparing the maximum
sprinting speed of a cheetah to the maximum book-reading performance of a
teenage boy. In the former case, the task is essential for survival and pushes
the entire physiological and muscular system to its limits whereas, in the latter
case, extreme book-reading performance may be simply a function of a few
heightened senses (e.g., eye coordination) and can hardly be considered as es-
sential to survival (although we heartily advocate reading).
Because the goal of this book is to examine comparable kinds of perform-
ance that have significance in the lives of animals, we adopt a dynamic, func-
tional, and whole-organism view of animal performance; such a view has been
the center of conceptually related studies over the past several decades. The
utility of this seemingly narrow definition can be understood by considering
 Animal performance: An overview 7

our own human Olympics, which measures human performance at a myriad

of tasks, including sprinting, weightlifting, rowing, and so forth. However,
the human Olympics do not measure many other potentially valid measures
of performance, such as how many books an individual can read in a year or
how much money a person can earn in a year. Similarly, this above definition
of performance precludes suborganismal measures of performance, such as
biochemical and physiological functions within organisms; these measures
might include aspects such as the ability of enzymes to catalyze reactions, for
example. A dynamic view of performance necessarily emphasizes movement,
such as vocalization, locomotion, feeding. By contrast, aspects of performance
such as digestion, metabolism, and other such measures are not considered
here. In addition, we as humans intuitively consider performance to be a fac-
tor that can be increased with training, which would typically preclude things
like digestion.
The logic behind this emphasis on the whole organism relates to the hier-
archical nature of biological systems. Animals exhibit behaviors and functional
capacities that are “emergent” properties at the organism level and cannot be
fully understood by only examining individual components (fig. 1.2). Moreo-
ver, as noted above, it is the whole organism that is visible to the environmen-
tal forces that dictate life or death. Therefore, although understanding how an
individual enzyme’s function may provide some insight into a larger physio-
logical process, it is the functioning of the larger system (e.g., the organism) that
is of paramount importance for evolution. A final criterion is best illustrated
by the Castanza family from the TV comedy “Seinfeld.” Instead of celebrating
Christmas, they celebrate “Festivus” in which “feats of strength” take place
(alongside the presence of a bare iron rod). In other words, an important aspect

Morphological Physiology
structure

Performance

Behavior

Fig. 1.2 A heuristic diagram showing the interrelationships between morphological structures,
physiology, and behavior for influencing performance in animals. The image of the jumping lizard
Anolis valencienni was taken by Esteban Toro with permission.
 8 Animal Athletes

of this adopted definition is that high performance is both physically diffi-

cult and potentially costly. Sprinting at peak speeds is difficult, because of the
extreme coordination of numerous body parts (e.g., heart, lungs), and costly,
because of the energetic demands on the physiological system. The study of
performance would become meaningless if we were to compare performance
traits that are physically easy to those that are physically challenging.
The distinction between performance and nonperformance traits as defined
here can often be subtle based on the criterion of physical difficulty or cost. For
example, at first glance, there would seem to be little distinction between bird
song and frog vocalizations, but a closer look reveals some important differ-
ences. For many frogs, vocalization is energetically expensive, and long peri-
ods of vocalization are highly exhausting (Wells and Taigen 1989; Chappell
et al. 1995). Therefore, many frogs have evolved calling structures that allow
them to vocalize for long periods, with large amounts of slow-twitch muscle.
Because frog vocalization is challenging, it is also a basis by which female
frogs choose mates, with females selecting males that vocalize the longest.
In other words, frog vocalization is a “feat of strength” and, like many other
performance traits, varies among individuals. By contrast, bird song, while
often remarkably complex, is apparently not energetically expensive, at least
in some species (Chappell et al. 1995; Horn et al. 1995), although there is pos-
sibly variation among species in this regard. While there are anatomical and
structural costs to the evolution of bird song, many kinds of bird song can be
more analogized to piano playing than to an Olympian performance feat such
as running or jumping, although the concept of bird song is still useful for
demonstrating trade-offs, which we explore in chapter 7.
However, even if one accepts this definition, within the field of functional
morphology, one is confronted with an array of seemingly overlapping and
confusing terms. One distinction is between “performance” and “function.” In
its broadest sense, “function” refers to the biological role of a morphological
trait (see Lauder 1996 for a discussion of this and other terms below). By bio-
logical “role,” we mean the action that natural selection originally favored. Ac-
cordingly, “function” tends to be broadly defined, such as “grasping” or “ma-
nipulation” in the case of the human hand. This broad definition is essential
because morphological traits are often “multifunctional” and appear suited
for many different biological tasks. Consequently, a result of this definition
is that a single morphological trait can have multiple “functions,” depending
on how specifically that function is defined. By contrast, performance is any
quantitative measure of how well a function is accomplished and thus provides
a necessary quantitative face on the rather amorphous concept of function.
For example, a key function of the lower limbs in bipeds is locomotion (e.g.,
jumping or running). We can quantify locomotion in any number of ways, such
as by measuring speed (how fast an animal runs), endurance (how long an
 Animal performance: An overview 9

animal runs), and so forth, and therefore no single measure of performance

will completely encompass all aspects of a broader function. Since dynamic
movements arise from morphological structures, it is also useful to understand
the relationship among function, performance, and two other terms describing
morphological shape, namely, “structure” and “design.” “Structure” refers to
the underlying arrangement of parts in a particular morphological trait. By ar-
rangement, we mean the configuration of parts, such as the attachment of ten-
dons to bones, the arrangement of muscle fibers, and so forth. Throughout this
book, we will refer to structure as the primary arrangement of morphological
parts within organisms. By comparison, “design” is more difficult to define.
Design overlaps with structure in referring to the arrangement of morpho-
logical parts, but the former refers also to an underlying (or predetermined)
process, either conscious or unconscious. Because of this unfortunate historical
connotation, we will no longer refer to design in this book. A final distinction
is between performance and behavior. Behavior refers to what animals do, not
how well they do it. An example from the human Olympics is instructive. In
the high jump, Olympic jumpers used to jump over the bar belly first, whereas
Dick Fosbury pioneered the “back-first” “Fosbury flop” that is widely used
today (fig. 1.3). Hence, whereas the “behavior” of jumping has changed, the
underlying metric of performance (jump height) has not. Note also the diffe-
rence here between behavior and function, as jumping is a function, whereas
how that jumping is done (belly first or back first) represents behavior, and
how well that jumping is done (jump height) is the relevant performance trait.

Fig. 1.3 An image of the Fosbury flop. Image from Wikimedia Commons.
 10 Animal Athletes

1.3 The hierarchical nature of biological systems

A pervading theme throughout this book is the concept of the hierarchy of bio-
logical systems; this hierarchy is manifested at two levels. First, performance
capacity represents the final output from the underlying working components
of complex functional systems. For example, many toads exhibit remarkable
tongue-projection performance capacities, with accelerations approaching
300 m/s2. Examination of the ecology of these toads provides insight into
why these remarkable performance capacities have evolved. Many toads are
largely defenseless because of their poor locomotor abilities and hence largely
rely on crypticity (as well as some excreted toxins) to protect themselves from
predators. However, this impaired locomotor ability also places the toads at
a distinct disadvantage for capturing active prey, such as insects. Most toads
have resolved this conflict by modifying evolutionarily a complex set of mus-
cles and nerves that enable their tongues to project rapidly and accurately
(Nishikawa 1999). However, this ability to project the tongue at extreme accel-
erations is only the final output derived from a complex set of interacting mor-
phological structures. For example, key nerves innervate muscle groups that
play a key role in tongue projection. The visual system in toads is also highly
attuned to moving prey and thus is synchronized to track moving prey and
enable accurate tongue projection. Thus, one cannot easily separate the under-
lying structures that produce performance from the evolution of performance
capacity. Hence, while not dwelling on the precise mechanism of how per-
formance is produced, we will examine the morphological and physiological
structures associated with performance while also considering broader evolu-
tionary and ecological implications.
The complex nature of morphological structures, physiological processes,
and their effects on performance means that predicting how different mor-
phological structures should influence performance is tricky. Because they
emerge from a complex set of lower-level traits, performance traits do not
always show proportional changes in relation to changes in traits such as
morphology, behavior, or physiology. Small changes in morphology can have
large effects on whole-organism performance, or vice versa. Such unpredict-
able effects are termed “nonlinear” because they don’t occur predictably from
one another in the way that one might expect in a classic linear relationship
(Emerson et al. 1990). Two key factors that drive these nonlinear relationships
are behavior and the environmental context of animal performance. For ex-
ample, simple alterations in the way that animals use their body can have pro-
found consequences on performance, even when the underlying morphology
remains the same. This idea is familiar to anyone who has participated in or-
ganized sports—after all, how much time and effort is spent training athletes
to move their bodies in certain ways to maximize success at a certain event? In
 Animal performance: An overview 11

the animal kingdom, where even slight differences in performance can make a
difference between life and death, animals may make adaptive and conscious
behavioral decisions that alter the basic relationship between morphology and
performance. A classic example is flying frogs, which glide through the forest
canopies of tropical forests in South America and Southeast Asia by the use of
flaps of skin on their feet and flanks. Flying frogs cannot engage in powered
flight like birds, but they can successfully glide for reasonably long distances
by manipulating the position of their limbs, a discovery by Sharon Emerson
and colleagues, who used plasticine models in wind tunnels to study the re-
lationship between gliding ability and flap area (Emerson et al. 1990). They
found that even slight alterations in the physical configuration of the mod-
els resulted in dramatic differences in gliding ability; but the effects of these
changes were nonlinear, as other larger changes seemed to have less impact.
Second, measures of performance exhibit a hierarchical structure. For ex-
ample, consider the case of crabs, as some males will fight, sometimes fiercely,
to acquire and defend territories. Crabs exhibit several stages in fighting, be-
ginning with an assessment stage, in which the crabs first evaluate the size
and strength of their opponent, and then followed by intense grappling and
wrestling with their two large front claws, which can result in severe injuries.
In this system, one can define many potentially overlapping and hierarchical
measures of performance. One could define performance relatively narrowly
in measuring the pinching strength of claws by using force transducers, for
example. Alternatively, one could take a broader view and measure fighting
ability, which can be quantified as fighting success. However, note that, in this
case, the broadest measure of performance (fighting success) is itself dictated
by many other variables, including body size, the relative (i.e., size-adjusted)
pinching strength of the claws, and so forth.
The choice of performance variables is as varied as the study subjects examined.
Some researchers are interested in sexual selection and factors that determine male
fighting success, or access to females, whereas other researchers are interested in
understanding the mechanistic basis of how frogs jump. We will be inclusive in
examining multiple, potentially disparate measures of performance capacity, for
the reason that each measure can provide information on the underlying ecology
and evolution of species. In some cases, we will even extract performance from
studies that may not even have been considered their measures as performance.
Below we describe several key examples that highlight this approach.

1.4 Variability, repeatability, and heritability

Three characteristics are vital for performance traits to play an important role
during evolution: variability, repeatability, and heritability. Variability is the
concept that a performance measure should vary among individuals within
 12 Animal Athletes

an interbreeding population. Performance capacities are distinctive from be-

havioral or morphological traits in sometimes (Le Galliard et al. 2004) display-
ing highly skewed distributions (fig. 1.4A), although many performance traits
also show more normal-shaped patterns. This skewed pattern is generated
by the presence of a large number of individuals who exhibit low or inter-
mediate performance, and a few individuals who are outstanding perform-
ers. Contrast this pattern to the typical pattern for a morphological trait such

A 100

80
Number of individuals

60

40

20

0
0 400 800 1200 1600
Initial endurance (s)
B 100

80
Proportion (counts)

60

40

20

N = 403
0
0 5 10 15 20
Femur length (mm)

Fig. 1.4 A, A histogram showing the distribution of values of endurance for the lizard Lacerta
vivipara. Taken from Le Galliard et al. (2004) with permission from Nature Publishing group. B, A
histogram of femur lengths of green anole lizards, Anolis carolinensis, from Southern Louisiana
(Irschick, unpublished data).
 Animal performance: An overview 13

as limb length, in which the distribution is nearly bell shaped (i.e., normal;
see fig. 1.4B). A second important feature of performance traits is repeatabil-
ity, or the tendency for the same individual to consistently display the same
level of performance capacity. Consider an example of two track stars running
the 100 m dash, for which a world-class time would be 10 s. Further imagine
that one of these two is an exceptional runner (track star A), with a personal
best time of 9.90 s, and the other one is less exceptional (track star B), with a
personal best of 10.20 s. If these fictitious track stars ran side-by-side 100 m
dashes once per day for 5 straight days, and the times (in seconds) for runners
A and B were (in order of days) 9.90, 9.89, 9.91, 9.90, and 9.88 (track star A),
and 10.20, 10.22, 10.21, 10.20, and 10.19 (track star B), then it’s easy to see (and
statistical tests would verify this, but that’s not necessary for this example)
that maximum speed is repeatable in these individuals, with A being consist-
ently better than B, no matter how often they race. Repeatability is an essential
ingredient for performance traits to evolve because it means that a particular
performance value for an individual is not a chance result but rather reflects
basic physiological, morphological, behavioral, or genetic qualities.
A third factor is heritability, or the degree to which traits within a population
have a genetic basis. A trait with a high heritability, typically measured on a
scale of 0 (no genetic basis) to 1 (completely genetically controlled), is a trait that
is likely to be inherited by an offspring from its parents. For example, consider
a male alligator that has a particularly strong bite and happens to mate with
a female who also has a very strong bite. If bite force shows high heritability
(i.e., values closer to 1 than to 0), then the offspring of these two alligators will
also likely be a hard biter, although other factors, such as the environment in
which the offspring was raised, are relevant. All three of these traits (variability,
repeatability, and heritability) are crucial to the evolutionary process, as traits
that show high values for all three factors are also more likely to evolve via nat-
ural selection. By contrast, the efficacy of natural selection on performance traits
that show low values for any of these three factors will be severely dampened.

1.5 The ecological context of performance

At the most basic level, performance capacities have evolved to enable ani-
mals to survive in their chosen habitats, whether in water, land, air, or the
canopy of trees. Because of this basic requirement, body shape, physiology,
and performance in animals are often matched to enhance chances of survival
in different habitats. This phenomenon is glaringly obvious when one exam-
ines animals that occur in “extreme” environments, such as regions of extreme
cold, heat, or salinity. For example, cave-dwelling organisms frequently pos-
sess enhanced organs and structures for touch or smell but reduced visual
organs, whereas diurnal active hunters frequently rely on their visual abilities.
 14 Animal Athletes

Large predators in the open landscapes of Africa or Australia often have excel-
lent locomotor abilities (aerobic or anaerobic) that enable them to either am-
bush or chase down prey. Consequently, one can study the general nature of
adaptation, or the process by which species evolve to match their habitats, by
examining the performance capacities of species that occur in different habi-
tats. Further, we can look within species and examine how individual vari-
ation in performance allows some animals to occupy some preferred habitats,
while poor performers are more limited.
Let’s ponder an example with butterflies, as it demonstrates nicely the eco-
logical role of performance. A long-standing question in the evolution of flight
concerns the effect of wing size on flying performance. Among others, two
simple aspects of wing shape contribute to flight: the aspect ratio (wing length
divided by wing width) and the relative loading of a system (body mass relative
to wing area). Butterflies offer the opportunity to manipulate these parameters
because one can surgically and nontraumatically remove certain amounts of
the wing, as well as add loads, each of which can impair (though reversibly so)
flight. Based on biomechanical predictions, one would expect that butterflies
that have their wing area reduced should have reduced survival relative to
butterflies whose wing areas were unmanipulated (Kingsolver 1999). In fact,
laboratory studies seem to confirm this prediction; butterflies whose wing
areas were reduced suffer dramatically reduced flight performance. But what
are the consequences of this reduced flight performance for animals in nature?
By releasing both manipulated (i.e., wings cut) and unmanipulated butterflies,
Joel Kingsolver from the University of North Carolina made a surprising dis-
covery; butterflies whose wings were surgically reduced did not suffer higher
mortality rates compared to unmanipulated butterflies, despite the obvious
performance disadvantage (fig. 1.5). Even butterflies whose wings were re-
duced by as much as 30% did not suffer reduced fitness relative to unma-
nipulated butterflies. This example highlights an important feature of the evo-
lutionary process, namely, that evolution does not necessarily concern itself
with decrements in performance if the animal can perform “well enough” in
its environment to survive. In this case, it’s clear that the butterflies, even with
a large chunk of their wings missing, were able to move around effectively.
However, field observational studies of butterflies showed another com-
peting role of butterfly wings, namely, thermoregulation. Many butterflies use
their wings to thermoregulate; during cool mornings, butterflies spread their
wings to gain maximum exposure to the rising sun and, during the hotter
parts of the day, butterflies close their wings to avoid overheating. The reason
for this careful thermoregulation is that flight is largely dictated by tempera-
ture. Flight performance is maximized across a relatively narrow range of tem-
peratures, ranging from about 28°C to 32°C. Larger wings enable butterflies to
effectively thermoregulate by increasing the surface area that is exposed to the
 Animal performance: An overview 15

May-June 1993

1.0

Reduced-wing
0.8
Survival Probability

0.6
Unman+Control
0.4

0.2

0.0
0 1 2 3 4 5
Time Period

Fig. 1.5 Survival probabilities for the butterfly Pontia occidentalis (inset), for animals whose wings
had been reduced in size (dashed line) and for those butterflies whose wings had not been altered
(solid line). Note that despite the small difference in survival probability, there was no significant
difference between the reduced-wing and control animals. Redrawn from Kingsolver (1999) with
permission from Wiley-Blackwell Press. Image of the butterfly is from Wikimedia Commons.

sun. Therefore, an ecological perspective on performance (in this case, flight)

can reveal the complex nature of morphological traits, and the performance
traits they have evolved for. A priori, one would expect that the most obvious
function of butterfly wings would be flight, but the presence of other roles
(e.g., thermoregulation) suggests that not all aspects of wing morphology
have evolved as adaptations for flight. This plurality of morphological struc-
tures is a recurring theme in the evolution of performance.

1.6 The evolution of performance

One of the most notable features of the animal kingdom is the tremendous
variability within and among species in body shape and performance. For ex-
ample, within anurans (frogs), there is variation in the length of the hindlimb
(which directly powers jumping in frogs) and in jumping ability (i.e., how
far a frog can jump). Many frogs can only jump a few inches, but the record
frog jump at the Calaveras Frog Jumping Contest was over 2 m long (Ast-
ley et al. 2013)! Whereas we can learn a great deal about the muscular and
neural factors involved in jumping by studying single species, we can learn
far more about how jumping has evolved by taking a comparative approach.
The comparative approach in evolutionary biology is a heuristic way of test-
ing ideas about adaptation by comparing species in divergent environments
and thereby testing how different selective regimes have resulted in different
 16 Animal Athletes

morphological and performance traits. For instance, when one compares spe-
cies of fish that occur in polar regions with those from more temperate areas,
one of the reasons why the polar fish can survive in freezing waters (with-
out freezing) becomes obvious: some arctic fish possess ice-nucleating agents
that prevent their internal tissues from freezing (and therefore killing the fish),
whereas fish in warmer regions typically don’t require such an adaptation.
Only by comparing different species can one clearly understand the connec-
tion between the environment and adaptive features for avoiding freezing.
Just as we can use the comparative approach to understand why some fish
can survive freezing and others cannot, we can profitably apply this idea to-
ward explaining variation in performance capacity among and within animal
species.
One of the most useful ways to study how traits evolve is to understand
the selective pressures that have driven the change in the first place. One of
the driving forces for variation in locomotor performance is the threat of pre-
dation. Because many animals flee predators using locomotion, much of the
variation in peak running speed has evolved as a direct response to pressure
from predators of varying locomotor abilities and strategies. One instructive
example comes from studies of escape strategies in damselflies, genus Enal-
lagma, that were examined in the Northern Lakes region in the USA by Mark
McPeek and his colleagues (McPeek et al. 1996; McPeek 1999). There are two
important predators for these invertebrates: dragonfly larvae and fish, each of
which requires different strategies for effective escape. The ancestral and pre-
dominant ecological context for damselflies is the occupation of lakes with fish
predators but, in some cases, damselflies have successfully colonized lakes
that contain a novel predator (dragonfly larvae). Whereas damselflies avoid
active locomotion as an escape strategy in the presence of much faster and
larger fish (doing so would only provide a quick snack!), rapid bursts of loco-
motion are more effective as a strategy for eluding the ambushing dragonfly
larvae. This novel and intense form of predation pressure from dragonfly lar-
vae has had several physiological, morphological, and functional evolution-
ary consequences for damselflies, including the evolution of a greater number
of tail fins for propelling locomotion (called lamellae) as well as the evolution
of enhanced burst speeds, and enhanced biochemical activity for several key
enzymes that enhance burst speed.
Many aspects of morphology and physiology influence burst speed; how-
ever, in this case, three enzymes seem to play an especially important role:
pyruvate kinase, lactate dehydrogenase (both used in glycolysis), and argin-
ine kinase (involved in reestablishing the pool of available ATP). Fitting a pri-
ori predictions, damselflies that have colonized lakes with dragonfly larvae
predators show significantly greater activity for arginine kinase, but not for
the two other enzymes. Because these damselflies move only in brief bursts
 Animal performance: An overview 17

when escaping, the increased activity of arginine kinase may extend the pe-
riod of maximal exertion for several seconds, providing a crucial window of
opportunity. Although indirect, this example suggests that variation among
species in arginine kinase activity (physiology) is adaptive because of its role
in influencing burst speed (performance) that has arisen as a consequence of
variation of different habitats (lakes with fish predators vs. lakes with dragon-
fly predators). Further, this damselfly example demonstrates the coevolution-
ary interaction between prey and predators and shows how the presence of
different predators can alter escape behavior (rapid escape vs. lying still) and
consequently both performance levels (high vs. low burst speed) and physio-
logical characteristics.

1.7 Behavior and performance

As exemplified by the Fosbury flop, described in section 1.2, behavior and
performance are often inextricably linked. Our thesis is that behavior plays
a key role in determining both whether animals undertake a particular per-
formance task and how effective an animal is at a particular performance task.
Particularly germane to this idea is the notion that animals make conscious
choices about how to move their body to maximize certain aspects of per-
formance (and perhaps minimize other, more deleterious aspects). The idea
of behavioral choices, or cognition, in animal performance is a relatively new
one. The field of biomechanics, the study of the physics of animal movement,
stresses the stereotyped nature of neuromuscular movements and examines
the optimal ways in which animals can achieve peak performance by moving
in a specific manner. However, functional morphologists are now aware that
individuals employ different strategies for maximizing performance and that
there may not always be “one” way to perform a particular task. This phe-
nomenon becomes obvious when comparing different individuals or species
with different phenotypes. A popular example comes from human athletics:
the relatively slight Janet Evans found success during swimming events by
rapidly cycling her arms like a windmill in order to keep up with (and defeat
in many cases) much larger swimmers who took fewer, longer arm and leg
strokes to swim. In her case, the decision to adopt a different swimming strat-
egy was a conscious and successful modification of behavior. The same is true
for professional basketball, which has undergone an evolution with respect to
the way in which free-throw shots are performed. Rick Barry, a member of the
NBA Hall of Fame, is famous for employing an underhand shooting method
and achieving a 90% success rate throughout his career. The fact that his ap-
proach has now been completely replaced by the overhand shot, with great
success, again highlights that there can be multiple ways through which to
achieve the same level of performance.
 18 Animal Athletes

One performance trait thought to be stereotyped is jumping; but even for

this task, there is evidence for the overarching role of behavior. Jumping has
been particularly well studied in frogs and lizards, both of which commonly
use jumping to escape predators or simply move around their habitats. One
group of lizards in which jumping has been well studied is the diverse genus
Anolis (Irschick et al. 1997; Losos 2011). The arboreal insectivorous lizards
from this genus present a wide diversity of morphological forms. Some spe-
cies have long hindlimbs and are fast, active runners, whereas other species
have short hindlimbs and are slow, cryptic twig specialists (fig. 1.6). Such
morphological and behavioral adaptations have occurred largely because the
different species occupy arboreal perches of different diameters and heights.
Whereas some species occur on broad perches (e.g., tree trunks) close to the
ground, other species occur high in the forest canopy on narrow twigs. These
different surfaces impose divergent functional demands for movements such
as running and jumping. Jumping is an important activity for these arbo-
real animals, as up to 50% of their movements will be jumps, often between
branches. However, anole species vary in how far they can jump, with some
species being outstanding, and others mediocre. Why do anole species vary in
jumping capacity, and which factors enable some species to be better jumpers
than others?
A basic understanding of the biomechanics of jumping is necessary to an-
swer this question. Jumping in limbed vertebrates can be deconstructed into
several stages (fig. 1.7). Four primary stages exist: the preparatory phase, the
takeoff phase, the suspended phase, and the landing phase. How far a lizard
can jump is largely a consequence of three factors: the takeoff velocity, the

Trunk-crown Trunk-ground Trunk

Grass-bush Trunk Twig

Fig. 1.6 Images of anole lizards, Anolis spp., from the Caribbean. Note the diversity in body form
and habitat use, including variation among species in head, body, limb, and tail dimensions. The
titles above each lizard indicate their preferred habitat. Images by Duncan J lrschick.
Exploring the Variety of Random
Documents with Different Content
As to Porphyry it was doubtless his more practical temperament that
led him to modify the doctrine of Plotinus concerning ecstasy. With
Porphyry the mind does not lose, in that state of exaltation, its
consciousness of personality. He calls it a dream in which the soul,
dead to the world, rises to an activity that partakes of the divine. It is
an elevation above reason, above action, above liberty, and yet no
annihilation, but an ennobling restoration or transformation of the
individual nature.[27]
Gower. One of Porphyry’s notions about the spirits of the air, of
which you told me in our walk yesterday, quite haunted me
afterwards. It contains a germ of poetry.
Kate. By all means let us have it.
Gower. Our philosopher believed in a certain order of evil genii who
took pleasure in hunting wild beasts,—dæmons, whom men
worshipped by the title of Artemis and other names, falsely
attributing their cruelty to the calm and guiltless gods, who can never
delight in blood. Some of these natures hunted another prey. They
were said to chase souls that had escaped from the fetters of a body,
and to force them to re-enter some fleshly prison once more. How I
wish we could see a design of this by David Scott! Imagine the soul
that has just leaped out of the door of that dungeon of ignorance and
pain, the body, as Porphyry would term it, fluttering in its new
freedom in the sunshine among the tree-tops, over wild and town—
all the fields of air its pleasure-ground for an exulting career on its
upward way to join the journeying intelligences in their cars above.
But it sees afar off, high in mid-air, a troop of dark shapes; they seem
to approach, to grow out of the airy recesses of the distance—they
come down the white precipices of the piled clouds, over the long
slant of some vapour promontory—forms invisible to man, and, with
them, spectre-hounds, whose baying spirits alone can hear. As they
approach, the soul recognises its enemies. In a moment it is flying
away, away, and after it they sweep—pursuers and pursued, shapes
so ethereal that the galleries of the ant are not shaken as hunters
and quarry glide into the earth, and not a foam-bell is broken or
brushed from the wave when they emerge upon the sea, and with
many a winding and double mount the air. At last hemmed in, the
soul is forced—spite of that desperate sidelong dart which had all but
eluded them—down into a body, the frame of a beggar’s babe or of a
slave’s; and, like some struggling bird, drawn with beating wings
beneath the water, it sinks into the clay it must animate through
many a miserable year to come.
Willoughby. I wish you would paint it for us yourself. You might
represent, close by that battle of the spirits, a bird singing on a
bough, a labourer looking down, with his foot upon his spade, and
peasants dancing in their ‘sunburnt mirth’ and jollity—wholly
unconscious, interrupted neither in toil nor pleasure by the conflict
close at hand. It might read as a satire on the too common
indifference of men to the spiritual realities which are about them
every hour.
Mrs. Atherton. The picture would be as mysterious as an Emblem
by Albert Durer.
Gower. It is that suggestiveness I so admire in the Germans. For
the sake of it I can often pardon their fantastic extravagances, their
incongruous combinations, their frequent want of grace and
symmetry.
Atherton. So can I, when an author occupies a province in which
such indirectness or irony, such irregularity, confusion, or paradox,
are admissible. Take, as a comprehensive example, Jean Paul. But
in philosophy it is abominable. There, where transparent order
should preside, to find that under the thick and spreading verbiage
meaning is often lacking, and, with all the boastful and fire-new
nomenclature, if found, is old and common,—that the language is
commonly but an array of what one calls

Rich windows that exclude the light,

And passages that lead to nothing;—

This puts me out of all patience.

Gower. The fault you object to reminds me of some Flemish
landscape-pieces I have seen; there are trees, so full of grand life,
they seem with their outstretched arms to menace the clouds, and as
though, if they smote with their many hundred hands, they could
beat away the storm instead of being bowed by it; and underneath
these great ones of the forest, which should shadow nothing less
than a woodland council of Titans or a group of recumbent gods, the
painter places only a rustic with a cow or two, an old horse, a
beggar, or some other most every-day of figures.
Mrs. Atherton. And you mean that the German words are large-
looking as the trees, and the ideas worn and ordinary as the figures?
What will Mr. Willoughby say to that?
Atherton. I think Willoughby will agree with me that it is high time
that we should go back to our theurgic mysticism and Iamblichus.
Here is a letter of his:—

Iamblichus to Agathocles.
I assure you, my friend, that the efforts of Porphyry, of whom you
appear disposed to think so highly, will be altogether in vain. He
is not the true philosopher you imagine. He grows cold and
sceptical with years. He shrinks with a timid incredulity from
reaping in that field of supernatural attainment which theurgy has
first opened, and now continually enlarges and enriches.
Theurgy, be sure of it, is the grand, I may say, the sole path to
the exaltation we covet. It is the heaven-given organum, in the
hands of the wise and holy, for obtaining happiness, knowledge,
power.
The pomp of emperors becomes as nothing in comparison with
the glory that surrounds the hierophant. The priest is a prophet
full of deity. The subordinate powers of the upper world are at his
bidding, for it is not a man, but a god who speaks the words of
power. Such a man lives no longer the life common to other men.
He has exchanged the human life for the divine. His nature is the
instrument and vehicle of Deity, who fills and impels him
(ὄργανον τοῖς ἐπιπνέουσι θεοῖς.) Men of this order do not
employ, in the elevation they experience, the waking senses as
do others (οὔτε κατ᾽ αἴσθησιν ἐνεργοῦσιν οὔτε ἐγρηγόρασι).
They have no purpose of their own, no mastery over themselves.
They speak wisdom they do not understand, and their faculties,
absorbed in a divine power, become the utterance of a superior
will.
Often, at the moment of inspiration, or when the afflatus has
subsided, a fiery Appearance is seen,—the entering or departing
Power. Those who are skilled in this wisdom can tell by the
character of this glory the rank of the divinity who has seized for
the time the reins of the mystic’s soul, and guides it as he will.
Sometimes the body of the man subject to this influence is
violently agitated, sometimes it is rigid and motionless. In some
instances sweet music is heard, in others, discordant and fearful
sounds. The person of the subject has been known to dilate and
tower to a superhuman height; in other cases, it has been lifted
up into the air. Frequently, not merely the ordinary exercise of
reason, but sensation and animal life would appear to have been
suspended; and the subject of the afflatus has not felt the
application of fire, has been pierced with spits, cut with knives,
and been sensible of no pain. Yea, often, the more the body and
the mind have been alike enfeebled by vigil and by fasts, the
more ignorant or mentally imbecile a youth may be who is
brought under this influence, the more freely and unmixedly will
the divine power be made manifest. So clearly are these
wonders the work, not of human skill or wisdom, but of
supernatural agency! Characteristics such as these I have
mentioned, are the marks of the true inspiration.
Now, there are, O Agathocles, four great orders of spiritual
existence,—Gods, Dæmons, Heroes or Demi-gods, and Souls.
You will naturally be desirous to learn how the apparition of a
God or a Dæmon is distinguished from those of Angels,
Principalities, or Souls. Know, then, that their appearance to man
corresponds to their nature, and that they always manifest
themselves to those who invoke them in a manner consonant
with their rank in the hierarchy of spiritual natures. The
appearances of Gods are uniform (μονοειδῆ), those of Dæmons
various (ποικίλα). The Gods shine with a benign aspect. When a
God manifests himself, he frequently appears to hide sun or
moon, and seems as he descends too vast for earth to contain.
Archangels are at once awful and mild; Angels yet more
gracious; Dæmons terrible. Below the four leading classes I have
mentioned are placed the malignant Dæmons, the Anti-gods
(ἀντιθέους).
Each spiritual order has gifts of its own to bestow on the initiated
who evoke them. The Gods confer health of body, power and
purity of mind, and, in short, elevate and restore our natures to
their proper principles. Angels and Archangels have at their
command only subordinate bestowments. Dæmons, however,
are hostile to the aspirant,—afflict both body and mind, and
hinder our escape from the sensuous. Principalities, who govern
the sublunary elements, confer temporal advantages. Those of a
lower rank, who preside over matter (ὑλικά), display their bounty
in material gifts. Souls that are pure are, like Angels, salutary in
their influence. Their appearance encourages the soul in its
upward efforts. Heroes stimulate to great actions. All these
powers depend, in a descending chain, each species on that
immediately above it. Good Dæmons are seen surrounded by
the emblems of blessing, Dæmons who execute judgment
appear with the instruments of punishment.
There is nothing unworthy of belief in what you have been told
concerning the sacred sleep, and divination by dreams. I explain
it thus:—
The soul has a twofold life, a lower and a higher. In sleep that
soul is freed from the constraint of the body, and enters, as one
emancipated, on its divine life of intelligence. Then, as the noble
faculty which beholds the objects that truly are—the objects in
the world of intelligence—stirs within, and awakens to its power,
who can be surprised that the mind, which contains in itself the
principles of all that happens, should, in this its state of liberation,
discern the future in those antecedent principles which will make
that future what it is to be? The nobler part of the soul is thus
 united by abstraction to higher natures, and becomes a
participant in the wisdom and foreknowledge of the Gods.
Recorded examples of this are numerous and well authenticated;
instances occur, too, every day. Numbers of sick, by sleeping in
the temple of Æsculapius, have had their cure revealed to them
in dreams vouchsafed by the god. Would not Alexander’s army
have perished but for a dream in which Dionysus pointed out the
means of safety? Was not the siege of Aphutis raised through a
dream sent by Jupiter Ammon to Lysander? The night-time of the
body is the day-time of the soul.
What I have now said—with little method, I confess—sets before
you but a portion of the prerogatives in which the initiated glory.
There is much behind for which words are too poor. I have
written enough, I am sure, to kindle your ambition, to bid you
banish doubt, and persevere in the aspirations which so
possessed you when I saw you last.[28] Farewell.

Gower. That explanation of prophetic dreams and the temple sleep

is very curious and characteristic. No doubt the common phenomena
of mesmerism may have been among the sacred secrets preserved
by the priests of Egypt and of Greece.
Kate. The preference for young and weakly persons, who would
possess an organization more susceptible of such influences, makes
it look very likely.
Atherton. Observe how completely the theurgic element, with
Iamblichus, supersedes the theosophic. In the process of time the
philosophical principles on which the system of Plotinus rested are
virtually surrendered, little by little, while divination and evocations
are practised with increasing credulity, and made the foundation of
the most arrogant pretensions. Plotinus declared the possibility of an
absolute identification of the divine with the human nature. Here was
the broadest basis for mysticism possible. Porphyry retired from this
position, took up narrower ground, and qualified the great mystical
principle of his master. He contended that in the union which takes
place in ecstasy, we still retain the consciousness of personality.
Iamblichus, the most superstitious of all in practice, diminished the
real principle of mysticism still farther in theory. He denied that man
has a faculty inaccessible to passion, and eternally active.[29]
Willoughby. And so the metaphysics and the marvels of mysticism
stand in an inverse ratio to each other. But it is not unnatural that as
the mystic, from one cause or another, gives up those exaggerated
notions of the powers of man and those mistaken views of the
relationship between man and God, which went together to make up
a mystical system of philosophy, he should endeavour to indemnify
himself by the evocations of theurgy, so as to secure, if possible,
through a supernatural channel, what speculation had
unsuccessfully attempted.
Atherton. True; but in this case I should invert the order, and say
that as the promise of theurgy exercised an attraction of growing
strength on an order of mind less fitted for speculation, such
temperaments would readily drop the speculative principle of
mysticism in their eagerness to grasp the illusive prize—apparently
so practical—which a commerce with superior natures held out.
Willoughby. And so the intellectual ambition and the poetical spirit,
so lofty in Plotinus, subside, among the followers of Iamblichus, into
the doggrel of the necromancer’s charm.
Gower. Much such a descent as the glory of Virgil has suffered,
whose tomb at Pausilipo is now regarded by the populace of
degenerate Naples less with the reverence due to the poet than with
the awe which arises from the legendary repute of the mediæval
magician.
Atherton. So the idealism of strong minds becomes superstition in
the weak. In the very shrine where culture paid its homage to art or
science, feebleness and ignorance, in an age of decline, set up the
image-worship of the merely marvellous.
Mrs. Atherton. I think you mentioned only one other of these
worthies.
Atherton. Proclus. He is the last great name among the Neo-
Platonists. He was the most eclectic of them all, perhaps because
the most learned and the most systematic. He elaborated the trinity
of Plotinus into a succession of impalpable Triads, and surpassed
Iamblichus in his devotion to the practice of theurgy. Proclus was
content to develop the school in that direction which Iamblichus—
(successful from his very faults)—had already given it. With Proclus,
theurgy was the art which gives man the magical passwords that
carry him through barrier after barrier, dividing species from species
of the upper existences, till, at the summit of the hierarchy, he arrives
at the highest. According to him, God is the Non-Being who is above
all being. He is apprehended only by negation. When we are raised
out of our weakness, and on a level with God, it seems as though
reason were silenced, for then we are above reason. We become
intoxicated with God, we are inspired as by the nectar of Olympus.
He teaches philosophy as the best preparation for Quietism. For the
scientific enquirer, toiling in his research, Proclus has a God to tell of,
supreme, almighty, the world-maker and governor of Plato. For him
who has passed through this labour, a God known only by ecstasy—
a God who is the repose he gives—a God of whom the more you
deny the more do you affirm.
Willoughby. And this is all! After years of austerity and toil, Proclus
—the scholar, stored with the opinions of the past, surrounded by the
admiration of the present—the astronomer, the geometrician, the
philosopher,—learned in the lore of symbols and of oracles, in the
rapt utterances of Orpheus and of Zoroaster—an adept in the ritual
of invocations among every people in the world—he, at the close,
pronounces Quietism the consummation of the whole, and an
unreasoning contemplation, an ecstasy which casts off as an
incumbrance all the knowledge so painfully acquired, the bourne of
all the journey.
Mrs. Atherton. As though it were the highest glory of man,
forgetting all that his enquiry has achieved, hidden away from the
world,—to gaze at vacancy, inactive and infantine;—to be like some
peasant’s child left in its cradle for a while in the furrow of a field,
shut in by the little mound of earth on either side, and having but the
blue æther above, dazzling and void, at which to look up with smiles
of witless wonder.
 Note to page 103.

Iamblichus, de Mysteriis, sect. x. cc. 1, 4, 6; iii. 4, 8, 6, 24; i. 5, 6; ii.

3; iii. 31; ii. 4, 6, 7; iii. 1, 3. These passages, in the order given, will
be found to correspond with the opinions expressed in the letter as
those of Iamblichus.
The genuineness of the treatise De Mysteriis has been called in
question, but its antiquity is undoubted. It differs only in one or two
very trivial statements from the doctrines of Iamblichus as
ascertained from other sources, and is admitted by all to be the
production, if not of Iamblichus himself, of one of his disciples,
probably writing under his direction. Jules Simon, ii. 219.
For the opinions ascribed to Porphyry in this letter, see his Epistola
ad Anebonem, passim. He there proposes a series of difficult
questions, and displays that sceptical disposition, especially
concerning the pretensions of Theurgy, which so much scandalized
Iamblichus. The De Mysteriis is an elaborate reply to that epistle,
under the name of Abammon.
In several passages of the De Mysteriis (ii. 11; v. 1, 2, 3, 7; vi. 6)
Iamblichus displays much anxiety lest his zeal for Theurgy should
lead him to maintain any position inconsistent with the reverence due
to the gods. He was closely pressed on this weak point by the
objections of Porphyry. (Ep. ad Anebon. 5, 6.) His explanation in
reply is, that the deities are not in reality drawn down by the mere
human will of the Theurgist, but that man is raised to a participation
in the power of the gods. The approximation is real, but the apparent
descent of divinity is in fact the ascent of humanity. By his long
course of preparation, by his knowledge of rites and symbols, of
potent hymns, and of the mysterious virtues of certain herbs and
minerals, the Theurgist is supposed to rise at last to the rank of an
associate with celestial powers; their knowledge and their will
become his, and he controls inferior natures with the authority of the
gods themselves.
Iamblichus supposes, moreover, that there is an order of powers in
the world, irrational and undiscerning, who are altogether at the
bidding of man when by threats or conjurations he chooses to
compel them. De Myst. vi. 5.
 BOOK THE FOURTH
MYSTICISM IN THE GREEK CHURCH
 CHAPTER I.

Questi ordini di su tutti s’ammirano

E di giù vincon si che verso Iddio
Tutti tirati sono e tutti tirano.
E Dionisio con tanto disio
A contemplar questi ordini si mise,
Che li nomò e distinse com’ io.[30]

Dante.

Kate. I have been looking at the pictures in Mrs. Jameson’s Sacred

and Legendary Art, of those strange creatures, the hermit saints—
the Fathers of the desert. Only see this one, what a mane and claws!
The two lions digging the grave there are own brothers to the holy
men themselves.
Atherton. Yet they claimed powers as much above humanity as, to
look at them, you would think them beneath it.
Gower. Religious Nebuchadnezzars.
Willoughby. No shavelings, at any rate, like the smooth-faced
sanctities of the later calendar.
Atherton. You will find among these anchorites almost all the
wonder-working pretensions of mediæval mysticism in full
development, thus early;—the discernment of spirits, gift of
prophecy, miraculous powers of various kinds, ecstasy, exorcism,
&c. &c. I should take St. Antony as a fair specimen of the whole
class.[31]
Mrs. Atherton. Look, here is his picture; there he stands, with
crutch and bell and pig.
Atherton. The bell denotes his power over evil spirits, and the pig
the vanquished dæmon of sensuality. In his life, by Athanasius, there
is a full account of his battle with many dæmons in the shape of
lions, bulls, and bears. He passed twenty years in an old castle
which he found full of serpents. The power of the saint expelled
those unpleasant aborigines. That nose, you see there, was
supposed to possess the faculty of detecting by its miraculous
keenness of scent the proximity of an evil spirit. There is an odour of
iniquity, you must know, as well as an odour of sanctity. This
disposition to literalize metaphors gave currency to the monkish
stories of after times concerning the refreshing fragrance found to
arise from the remains of disinterred saints. In fact, the
materialization of the spiritual, or what passes for such, is the
characteristic principle of the theurgic mysticism within the Roman
Catholic Church. St. Antony, on one occasion, sees his own soul,
separated from the body, carried through the air.
Gower. A striking instance, I should say, of the objectivity of the
subject.
Atherton. One of his visions is not without grandeur. The brethren
had been questioning him one day concerning the state of departed
spirits. The following night he heard a voice saying, ‘Antony, get up;
go out and look!’ He obeyed, and saw a gigantic figure, whose head
was in the clouds, and whose outstretched arms extended far across
the sky. Many souls were fluttering in the air, and endeavouring, as
they found opportunity, to fly upward past this dreadful being.
Numbers of them he seized in the attempt, and dashed back upon
the earth. Some escaped him and exulted above, while he raged at
their success. Thus sorrowing and rejoicing were mingled together,
as some were defeated and others triumphant. This, he was given to
understand, was the rise and fall of souls.
Willoughby. That picture would be really Dantesque, if only a little
more definite. Macarius is another great name, too, among these
Christian ascetics and theurgists—the one who retired to the deserts
of Nitria in the fourth century.
Atherton. He is not only famous for his measure of the
supernatural powers ascribed to his brethren, but his homilies have
been appealed to by modern theopathetic mystics as an authority for
Quietism. He teaches perfectionist doctrine, certainly, but I do not
think his words will bear the construction Poiret and others would
give them. He was at least innocent of the sainte indifférence.[32]
Mrs. Atherton. You said we were to discuss Dionysius the
Areopagite this evening.
Kate. Pray introduce me first. I know nothing about him.
Atherton. No one does know who really wrote the books which
passed under that name. It is generally admitted that the forgery
could not have been committed earlier than the middle of the fifth
century, probably somewhat later. So all I can tell you is, that
somewhere or other (it is not unlikely at Constantinople, but there is
no certainty), about the time when Theodoric was master of Italy—
when the Vandal swarms had not yet been expelled from northern
Africa—while Constantinople was in uproar between the greens and
the blues, and rival ecclesiastics headed city riots with a rabble of
monks, artizans, and bandit soldiery at their heels—while orthodoxy
was grappling with the Monophysite and Eutychian heresies on
either hand, and the religious world was rocking still with the
groundswell that followed those stormy synods in which Palestine
and Alexandria, Asia and Constantinople, from opposite quarters,
gathered their strength against each other—a monk or priest was
busy, in his quiet solitude, with the fabrication of sundry treatises and
letters which were to find their way into the Church under the all-but
apostolic auspices of that convert made by the Apostle of the
Gentiles when he spoke on Mars Hill. The writings would seem to
have been first appealed to as genuine in the year 533. As heretics
cited them, their authority was disputed at the outset; but being
found favourable to the growing claims of the hierarchy, and likely to
be useful, they were soon recognised and employed accordingly.[33]
Willoughby. Proclus could not have been long dead, and his
reputation must have been still at its height, when this anonymous—
let us call him Dionysius at once—was writing his Platonized
theology.
Atherton. With the divines of Byzantium Proclus represented the
grand old world of Greek thought. Even those who wrote against him
as a heathen betray the influence he exercised on their doctrines.
The object of Dionysius evidently was to accommodate the
theosophy of Proclus to Christianity. Another aim, not less
conspicuous, was to strengthen all the pretensions of the priesthood,
and to invest with a new traditionary sanction the ascetic virtues of
the cloister.
 CHAPTER II.

They that pretend to these heights call them the secrets of the
kingdom; but they are such which no man can describe; such
which God hath not revealed in the publication of the Gospel;
such for the acquiring of which there are no means prescribed,
and to which no man is obliged, and which are not in any man’s
power to obtain; nor such which it is lawful to pray for or desire;
nor concerning which we shall ever be called to account.—
Jeremy Taylor.

‘I have here,’ said Atherton on the next evening, ‘some notes on the
doctrine of this pretended Areopagite—a short summary; shall I read
it?’
‘By all means.’
So the following abstract was listened to—and with creditable
patience.[34]
(1.) All things have emanated from God, and the end of all is return
to God. Such return—deification, he calls it—is the consummation of
the creature, that God may finally be all in all. A process of evolution,
a centrifugal movement in the Divine Nature, is substituted in reality
for creation. The antithesis of this is the centripetal process, or
movement of involution, which draws all existence towards the point
of the Divine centre. The degree of real existence possessed by any
being is the amount of God in that being—for God is the existence in
all things. Yet He himself cannot be said to exist, for he is above
existence. The more or less of God which the various creatures
possess is determined by the proximity of their order to the centre.
(2.) The chain of being in the upper and invisible world, through
which the Divine Power diffuses itself in successive gradations, he
calls the Celestial Hierarchy. The Ecclesiastical Hierarchy is a
corresponding series in the visible world. The orders of Angelic
natures and of priestly functionaries correspond to each other. The
highest rank of the former receive illumination immediately from God.
The lowest of the heavenly imparts divine light to the highest of the
earthly hierarchy. Each order strives perpetually to approximate to
that immediately above itself, from which it receives the transmitted
influence; so that all, as Dante describes it, draw and are drawn, and
tend in common towards the centre—God.
The three triads of angelic existences, to whom answer the ranks of
the terrestrial hierarchy, betrays the influence of Proclus, whose
hierarchy of ideas corresponds, in a similar manner, to his hierarchy
of hypostases.
Gower. The system reminds one of those old pictures which are
divided into two compartments, the upper occupied by angels and
cherubs on the clouds, and the lower by human beings on the earth,
gazing devoutly upward at their celestial benefactors.
Atherton. The work of Christ is thrown into the background to make
room for the Church. The Saviour answers, with Dionysius, rather to
the Logos of the Platonist than to the Son of God revealed in
Scripture. He is allowed to be, as incarnate, the founder of the
Ecclesiastical Hierarchy; but, as such, he is removed from men by
the long chain of priestly orders, and is less the Redeemer, than
remotely the Illuminator, of the species.
Purification, illumination, perfection,—the three great stages of
ascent to God (which plays so important a part in almost every
succeeding attempt to systematise mysticism) are mystically
represented by the three sacraments,—Baptism, the Eucharist, and
Unction. The Church is the great Mystagogue: its liturgy and offices
a profound and elaborate system of symbolism.
(3.) The Greek theory, with its inadequate conception of the nature of
sin, compels Dionysius virtually to deny the existence of evil.
Everything that exists is good, the more existence the more
goodness, so that evil is a coming short of existence. He hunts sin
boldly from place to place throughout the universe, and drives it at
last into the obscurity of the limbo he contrives for it, where it lies
among things unreal.
All that exists he regards as a symbolical manifestation of the super-
existent. What we call creation is the divine allegory. In nature, in
Scripture, in tradition, God is revealed only in figure. This sacred
imagery should be studied, but in such study we are still far from any
adequate cognizance of the Divine Nature. God is above all negation
and affirmation: in Him such contraries are at once identified and
transcended. But by negation we approach most nearly to a true
apprehension of what He is.
Negation and affirmation, accordingly, constitute the two opposed
and yet simultaneous methods he lays down for the knowledge of
the Infinite. These two paths, the Via Negativa (or Apophatica) and
the Via Affirmativa (or Cataphatica) constitute the foundation of his
mysticism. They are distinguished and elaborated in every part of his
writings. The positive is the descending process. In the path
downward from God, through inferior existences, the Divine Being
may be said to have many names;—the negative method is one of
ascent; in that, God is regarded as nameless, the inscrutable
Anonymous. The symbolical or visible is thus opposed, in the
Platonist style, to the mystical or ideal. To assert anything concerning
a God who is above all affirmation is to speak in figure, to veil him.
The more you deny concerning Him, the more of such veils do you
remove. He compares the negative method of speaking concerning
the Supreme to the operation of the sculptor, who strikes off
fragment after fragment of the marble, and progresses by diminution.
(4.) Our highest knowledge of God, therefore, is said to consist in
mystic ignorance. In omni-nescience we approach Omniscience.
This Path of Negation is the highway of mysticism. It is by refraining
from any exercise of the intellect or of the imagination—by self-
simplification, by withdrawal into the inmost, the divine essence of
our nature—that we surpass the ordinary condition of humanity, and
are united in ecstasy with God. Dionysius does not insist so much on
Union as the later mystics, but he believes, at all events, that the
eminent saint may attain on earth an indescribable condition of soul
—an elevation far transcending the reach of our natural faculties—an
approach towards the beatific vision of those who are supposed to
gaze directly on the Divine Essence in heaven. His disciple is
perpetually exhorted to aspire to this climax of abstraction—above
sight, and thought, and feeling, as to the highest aim of man.
Willoughby. What contradictions are here! With one breath he
extols ineffable ignorance as the only wisdom; with the next he
pretends to elucidate the Trinity, and reads you off a muster-roll of
the heavenly hierarchies.
Gower. And are not these, supplemented by the hierarchy of
ecclesiastics, his real objects of worship? No man could make an
actual God of that super-essential ultimatum, that blank Next-to-
Nothingness which the last Neo-Platonists imagined as their
Supreme. Proclus could not; Dionysius could not. What then? A
reaction comes, which, after refining polytheism to an impalpable
unity, restores men to polytheism once more. Up mounts
speculation, rocket-like: men watch it, a single soaring star with its
train of fire, and, at the height, it breaks into a scattering shower of
many-coloured sparks. From that Abstraction of which nothing can
be predicated, nothing can be expected. The figment above being is
above benignity. So the objects of invocation are gods, demi-gods,
dæmons, heroes; or, when baptized, cherubim, seraphim, thrones,
dominions, powers, archangels, angels, saints; in either case,
whether at Athens or at Constantinople, the excessive subtilisation of
the One contributes toward the worship of the Manifold.
Atherton. The theology of the Neo-Platonists was always in the first
instance a mere matter of logic. It so happened that they confounded
Universals with causes. The miserable consequence is clear. The
Highest becomes with them, as he is with Dionysius, merely the
most comprehensive, the universal idea, which includes the world,
as genus includes species.[35]
Mrs. Atherton. The divinity of this old Father must be a bleak affair
indeed—Christianity frozen out.
Gower. I picture him to myself as entering with his philosophy into
the theological structure of that day, like Winter into the cathedral of
the woods (which an autumn of decline has begun to harm already);
—what life yet lingers, he takes away,—he untwines the garlands
from the pillars of the trees, extinguishes the many twinkling lights
the sunshine hung wavering in the foliage, silences all sounds of
singing, and fills the darkened aisles and dome with a coldly-
descending mist, whose silence is extolled as above the power of
utterance,—its blinding, chill obscureness lauded as clearer than the
intelligence and warmer than the fervour of a simple and scriptural
devotion.
Atherton. You have described my experience in reading him,
though I must say he suggested nothing to me about your cathedral
of the woods, &c. His verbose and turgid style, too, is destitute of all
genuine feeling.[36] He piles epithet on epithet, throws superlative on
superlative, hyperbole on hyperbole, and it is but log upon log,—he
puts no fire under, neither does any come from elsewhere. He
quotes Scripture—as might be expected—in the worst style, both of
the schoolman and the mystic. Fragments are torn from their
connexion, and carried away to suffer the most arbitrary
interpretation, and strew his pages that they may appear to illustrate
or justify his theory.
Gower. How forlorn do those texts of Scripture look that you discern
scattered over the works of such writers, so manifestly transported
from a region of vitality and warmth to an expanse of barrenness.
They make the context look still more sterile, and while they say
there must be life somewhere, seem to affirm, no less emphatically,
that it is not in the neighbourhood about them. They remind me of
those leaves from the chestnut and the birch I once observed upon a
glacier. There they lay, foreign manifestly to the treeless world in
which they were found; the ice appeared to have shrunk from them,
and they from the ice; each isolated leaf had made itself a cup-like
cavity, a tiny open sarcophagus of crystal, in which it had lain,
perhaps for several winters. Doubtless, a tempest, which had been
vexing some pleasant valley far down beneath, and tearing at its
trees, must have whirled them up thither. Yet the very presence of
the captives reproached the poverty of the Snow-King who detained
them, testifying as they did to a genial clime elsewhere, whose
products that ice-world could no more put forth, than can such frozen
speculations as this of Dionysius, the ripening ‘fruits of the Spirit.’
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