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 Rare Earth:
Why Complex Life is
Uncommon in the Universe

Peter D. Ward
Donald Brownlee

COPERNICUS BOOKS
ward_FM_i-xxviii 3/10/03 14:01 Page i

Praise for Rare Earth . . .

“ . . . brilliant and courageous . . . likely to cause a revolution in thinking.”

—William J. Broad
The New York Times “Science Times”

“A pleasure for the rational reader . . . what good books are all about . . . ”
—Associated Press

“If Ward and Brownlee are right it could be time to reverse a process that has been
going on since Copernicus.”
—The Times (London)

“Although simple life is probably abundant in the universe, Ward & Brownlee say,
‘complex life—animals and higher plants—is likely to be far more rare than is
commonly assumed’.”
—Scientific American, Editor’s Choice

“ . . . a compelling argument [and] a wet blanket for E.T. enthusiasts . . . ”

—Discover

“Peter Ward and Donald Brownlee offer a powerful argument . . . .”

—The Economist

“[Rare Earth] has hit the world of astrobiologists like a killer asteroid . . . .”
—Newsday

“A very good book.”

—Astronomy

The notion that life existed anywhere in the universe besides Earth was once
laughable in the scientific community. Over the past thirty years or so, the laugh-
ter has died away . . . . [Ward and Brownlee] argue that the recent trend in scien-
tific thought has gone too far . . . . As radio telescopes sweep the skies and earth-
bound researchers strain to pick up anything that might be a signal from
extraterrestrial beings, Rare Earth may offer an explanation for why we haven’t
heard anything yet.”
—CNN.com

“ . . . a sobering and valuable perspective . . . ”

—Science
ward_FM_i-xxviii 3/10/03 14:01 Page ii

“Movies and television give the (optimistic) impression that the cosmos is teeming
with civilizations. But what if it isn’t? . . . Life elsewhere in the universe may never
reach beyond microbes, which, the authors note, could be much more widespread
than originally believed.”
—Sky & Telescope

“It’s brilliant . . . courageous. . . . It’s rare in literature and science that a stance goes
so far against the grain.”
—Dr. Geoffrey W. Marcy
Extra-solar planet discoverer
University of California at Berkeley

“It’s a thought that grips most everyone who stares into the unfathomable depths of
a star-speckled night: Is there anybody out there? The odds, say Peter Ward and
Don Brownlee, are probably more remote than you think.”
—The Seattle Times

“Alien life is more likely to resemble the stuff you scrub off the tiles in your shower
than Klingons, Wookies or Romulans, say Ward and Brownlee.”
—Popular Mechanics

“Ward and Brownlee have taken an issue that is much in the public domain and
treated it thoughtfully and thoroughly, but with a lightness of touch that draws
the reader on . . . . Rare Earth is an excellent book for both specialists and non-
specialists.”
—The Times Higher Education Supplement (UK)

“A provocative, significant, and sweeping new book . . . Rare Earth is a fast-paced,

thought-provoking read that I gobbled like popcorn. It’s one of those rare books that
is at once delightful, informative, and important: an end-of-the-millennium synthesis
of science that tackles the central question of our past, place, and destiny.”
—Northwest Science & Technology

“ . . . well thought out and intriguing . . . ”

—Icarus

“ . . . a startling new hypothesis . . . Highly recommended.”

—Library Journal

“Rare Earth will surely appeal to those who would dare to disagree with icons Carl
Sagan and George Lucas.”
—San Gabriel Valley Tribune
ward_FM_i-xxviii 3/10/03 14:01 Page iii

“ . . . a timely, entirely readable account.”

—Toronto Globe & Mail

“ . . . a stellar example of clear writing . . . ”

—American Scientist

“ . . . thought-provoking and authoritative . . . ”

—Physics Today

“In this encouraging and superbly written book, the authors present a carefully rea-
soned and scientifically statute examination of the age-old question—‘Are we alone
in the universe?’ Their astonishing conclusion that even simple animal life is most
likely extremely rare in the universe has many profound implications. To the aver-
age person, staring up at a dark night sky, full of distant galaxies, it is simply incon-
ceivable that we are alone. Yet, in spite of our wishful thinking, there just may not
be other Mozarts or Monets.”
—Don Johanson
Director, Institute of Human Origins
Arizona State University

“A fabulous book! If we’re to believe what we see in the movies, extraterrestrials

thrive on every world. But this unique book, written by two of the top scientists in
the field, tells a different story. As we know it on Earth, complex life might be very
rare, and very precious. For those of us interested in our cosmic heritage, this book
is a must-read.”
—David Levy
Co-discoverer of Comet Shoemaker-Levy

“Ward and Brownlee take us on a fascinating journey through the deep history of
our habitable planet and out into space; in the process they weave a compelling ar-
gument that life at the level of an animal should be vastly rarer in the universe than
life at the level of a lowly bacterium.”
—Steven M. Stanley,
Author of Children of the Ice Age and Earth and Life Through Time
The Johns Hopkins University

“Microbial life is common in the universe, but multicellular animal life is rare. A
controversial thesis, but one that is well-researched and well-defended. A must-read
for anyone who is interested in whether life exists beyond Earth.”
—James Kasting
Pennsylvania State University
ward_FM_i-xxviii 3/11/03 10:34 Page iv

Rafted ice covering the subterranean ocean of Europa (moon of planet Jupiter), a possible life habitat in
the outer solar system. NASA image from the Galileo spacecraft. Courtesy of NASA.
ward_FM_i-xxviii 3/10/03 14:01 Page v

RARE
EARTH
Why
Complex Life
Is Uncommon in
the Universe

Peter D. Ward
Donald Brownlee

COPERNICUS BOOKS
AN IMPRINT OF SPRINGER-VERLAG
ward_FM_i-xxviii 3/10/03 14:01 Page vi

First softcover printing, 2003.

© 2000 Peter D. Ward and Donald Brownlee.

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, electronic, mechanical, photocopying, recording,
or otherwise, without the prior written permission of the publisher.

Published in the United States by Copernicus Books,

an imprint of Springer-Verlag New York, Inc.
A member of BertelsmannSpringer Science+Business Media GmbH

Copernicus Books
37 East 7th Street
New York, NY 10003
www.copernicusbooks.com

Library of Congress Cataloging-in-Publication Data

Ward, Peter D.
Rare earth: why complex life is uncommon in the universe / Peter D.
Ward, Donald Brownlee.
p. cm.
Includes bibliographical references and index.
ISBN 0-387-95289-6 (pbk.: alk. paper)
1. Life on other planets. 2. Exobiology. I. Brownlee, Donald.
II. Title.
QB54.W336 2003
576.839—dc21 2003043437

Manufactured in the United States of America.

Printed on acid-free paper.

9 8 7 6 5 4 3 2 1

ISBN 0-387-95289-6 SPIN 10838285

ward_FM_i-xxviii 3/10/03 14:01 Page vii

To the memory of
Gene Shoemaker
and
Carl Sagan
This page intentionally left blank
ward_FM_i-xxviii 3/10/03 14:01 Page ix

Contents
Preface to the Paperback Edition ........................................................................ x
Preface to the First Edition.............................................................................. xiii
Introduction: The Astrobiology Revolution and the Rare Earth Hypothesis ... xvii
Dead Zones of the Universe ........................................................................ xxix
Rare Earth Factors .................................................................................... xxxi

1 Why Life Might Be Widespread in the Universe .... 1

2 Habitable Zones of the Universe ........................... 15
3 Building a Habitable Earth ..................................... 35
4 Life’s First Appearance on Earth ............................. 55
5 How to Build Animals ............................................ 83
6 Snowball Earth ..................................................... 113
7 The Enigma of the Cambrian Explosion .............. 125
8 Mass Extinctions and the Rare Earth Hypothesis .. 157
9 The Surprising Importance of Plate Tectonics .... 191
10 The Moon, Jupiter, and Life on Earth .................. 221
11 Testing the Rare Earth Hypotheses ..................... 243
12 Assessing the Odds .............................................. 257
13 Messengers from the Stars ................................... 277
References ................................................................................................ 289
Index ...................................................................................................... 319

ix
ward_FM_i-xxviii 3/10/03 14:01 Page x

Preface to the
Paperback Edition

O
n November 12, 2002, Dr. John Chambers of the NASA Ames Re-
search Center gave a seminar to the Astrobiology Group at the
University of Washington. The audience of about 100 listened
with rapt attention as Chambers described results from a computer study of
how planetary systems form. The goal of his research was to answer a decep-
tively simple question: How often would newly forming planetary systems
produce Earth-like planets, given a star the size of our own sun? By “Earth-
like” Chambers meant a rocky planet with water on its surface, orbiting
within a star’s “habitable zone.” This not-too-hot and not-too-cold inner re-
gion, relatively close to the star, supports the presence of liquid water on a
planet surface for hundreds of million of years—the time-span probably nec-
essary for the evolution of life. To answer the question of just how many
Earth-like planets might be spawned in such a planetary system, Chambers
had spent thousands of hours running highly sophisticated modeling pro-
grams through arrays of powerful computers.
x
ward_FM_i-xxviii 3/10/03 14:01 Page xi

Preface to the Paperback Edition

The results presented at the meeting were startling. The simulations

showed that rocky planets orbiting at the “right” distances from the central
star are easily formed, but they can end up with a wide range of water con-
tent. The planet-building materials in a habitable zone include dry materials
that form locally, as well as water-bearing materials that originate further from
the star and have to be scattered inward, mostly in the form of comets. With-
out water-bearing comet impacts, Earth-wannabes would just stay wannabes—
they would never contain any water.
The model showed that the inbound delivery of water worked best in
planetary systems where the intermediate planets, in the position of our giants
Jupiter and Saturn, were far smaller. In solar systems such as our own, the effi-
ciency of water being conveyed to the surface of an inner, Earth-like planet is
relatively small. Yet in systems where the intermediate planets were much
smaller—perhaps Uranus- or Neptune-sized—water delivery was relatively
frequent. But then another problem arises: in such a system, the rate of water-
bearing comet impacts is great; the rate of asteroid impacts, however, is also so
great that any evolving life might soon be obliterated. And oddly, it is not only
the asteroid impacts, with their fireballs, dust storms, meteor showers, and “nu-
clear winters,” that cause a problem. An excess of water-bearing impacts can
amount, in effect, to too much of a good thing: too much water produces plan-
ets entirely covered with water, and such an environment is not conducive to
the rich evolution seen on our planet. Earth seems to be quite a gem—a rocky
planet where not only can liquid water exist for long periods of time (thanks to
Earth’s distance from the sun as well as its possession of a tectonic “thermostat”
that regulates its temperature), but where water can be found as a heathy
oceanful—not too little and not too much. Our planet seems to reside in a benign
region of the Galaxy, where comet and asteroid bombardment is tolerable and
habitable-zone planets can commonly grow to Earth size. Such real estate in
our galaxy—perhaps in any galaxy—is prime for life. And rare as well.
We, the authors of Rare Earth, were in the audience that November day.
One of us raised his hand and asked the question: What does this finding
mean for the number of Earth-like planets there might be—planets with not
only water and bacterial life, but with complex multi-cellular life? Chambers
scratched his head. Well, he allowed, it would certainly make them rare.
xi
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R A R E E A R T H

There was one other aspect of the lecture that struck us. Chambers matter-of-
factly spoke of the necessity of planets having plate tectonics to be habitable,
and of the effect of mass extinctions. We know that plate tectonics provides a
method of maintaining some sort of planetary thermostat that keeps planets at a
constant temperature for billions of years. We know, too, that mass extinctions
can end life on a planet abruptly, at any time, and that the number of mass ex-
tinctions might be linked to astronomical factors, such as the position of a planet
in its galaxy. Prior to the publication of the first edition of Rare Earth in January
2000, neither of these concepts had publicly appeared in discussions of planetary
habitability. Now they do, as a matter of course, and this has been a great satisfac-
tion to us. Our hypothesis that bacteria-like life might be quite common in the
Universe, but complex life quite rare, may or may not be correct. But the fact that
we’ve been able to bring new lines of evidence into the debate, evidence that was
once controversial but is now quite mainstream, has been extremely gratifying.
With its initial publication, Rare Earth struck chords among a wide commu-
nity. Because it took a rather novel position about the frequency of complex life,
the discussion spurred by the book often left the realm of scientific discourse,
where we’d intended it to take place, and entered the arenas of religion, ethics,
and science fiction. Science has progressed since the publication, yet nothing we
have read or discovered in the years since has caused us to change our minds.
One of the most remarkable developments has been the continual discovery of
new planets orbiting other stars (the count is now over 100). While this shows
that planets are common, it also shows how complex and varied planetary sys-
tems are, and how difficult it is to make a stable Earth-like planet. Most of the
extra-solar planets that have been discovered are giant planets in orbits that pre-
clude the possibility of water-covered Earths with long-term stability.
This edition, then, is changed only in the removal of several egregious
and sometimes hilarious typos and errors. We stand by our initial assessment
and are proud to see that Rare Earth continues to spawn heated debate even as
it makes its way into textbooks as accepted dogma.

Peter D. Ward, Donald Brownlee

Seattle, February 2003

xii
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Preface to the
First Edition

his book was born during a lunchtime conversation at the University of

T Washington faculty club, and then it simply took off. It was stimulated
by a host of discoveries suggesting to us that complex life is less perva-
sive in the Universe than is now commonly assumed. In our discussions, it be-
came clear that both of us believed such life is not widespread, and we de-
cided to write a book explaining why.
Of course, we cannot prove that the equivalent of our planet’s animal
life is rare elsewhere in the Universe. Proof is a rarity in science. Our argu-
ments are post hoc in the sense that we have examined Earth history and then
tried to arrive at generalizations from what we have seen here. We are clearly
bound by what has been called the Weak Anthropic Principle—that we, as
observers in the solar system, have a strong bias in identifying habitats or fac-
tors leading to our own existence. To put it another way, it is very difficult to
do statistics with an N of 1. But in our defense, we have staked out a position
rarely articulated but increasingly accepted by many astrobiologists. We

xiii
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R A R E E A R T H

have formulated a null hypothesis, as it were, to the clamorous contention

of many scientists and media alike that life—barroom-brawling, moral-
philosophizing, human-eating, lesson-giving, purple-blooded bug-eyed mon-
sters of high and low intelligence—is out there, or that even simple worm-like
animals are commonly out there. Perhaps in spite of all the unnumbered stars,
we are the only animals, or at least we number among a select few. What has
been called the Principle of Mediocrity—the idea that Earth is but one of a
myriad of like worlds harboring advanced life—deserves a counterpoint.
Hence our book.
Writing this book has been akin to running a marathon, and we want to
acknowledge and thank all those who offered sustaining draughts of infor-
mation as we followed our winding path. Our greatest debts of gratitude we
owe to Jerry Lyons of Copernicus, who invested so much interest in the proj-
ect, and to our editor, Jonathan Cobb, who fine-tuned the project on scales
ranging from basic organization of the book to its numerous split infinitives.
Many scientific colleagues gave much of themselves. Joseph Kirschvink
of Cal Tech read the entire manuscript and spent endless hours thrashing
through various concepts with us; his knowledge and genius illuminated our
murky ideas. Guillermo Gonzalez changed many of our views about planets
and habitable zones. Thor Hansen of Western Washington University de-
scribed to us the concept of “stopping plate tectonics.” Colleagues in the De-
partment of Geological Sciences, including Dave Montgomery, Steve Porter,
Bruce Nelson, and Eric Cheney, discussed many subjects with us. Many
thanks to Victor Kress of the University of Washington for reading and cri-
tiquing the plate tectonics chapter. Dr. Robert Paine of the Department of
Zoology saved us from making egregious errors about diversity. Numerous
astrobiologists took time to discuss aspects of the science with us, including
Kevin Zahnlee of NASA Ames, who patiently explained his position—one
contrary to almost everything we believed—and in so doing markedly ex-
panded our understanding and horizons. We are grateful to Jim Kasting of
Penn State University for long discussions about planets and their forma-
tions. Thanks as well to Gustav Ahrrenius from UC Scripps, Woody Sullivan
(astronomy) of the University of Washington, and John Baross of the School

xiv
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Preface to the First Edition

of Oceanography at the University of Washington. Jack Sepkoski of the

University of Chicago generously sent new extinction data, Andy Knoll of
Harvard contributed critiques by E-mail; Sam Bowring spent an afternoon
sharing his data and his thoughts on the timing of major events in Earth his-
tory; Dolf Seilacher talked with us about ediacarans and the first evolution of
life; Doug Erwin lent insight into the Permo/Triassic extinction; Jim Valen-
tine and Jere Lipps of Berkeley gave us their insights into the late Precam-
brian and animal evolution; David Jablonski described his views on body plan
evolution. We are enormously grateful to David Raup, for discussions and
archival material about extinction, and to Steve Gould for listening to and
critiquing our ideas over a long Italian dinner on a rainy night in Seattle.
Thanks to Tom Quinn of UW astronomy for illuminating the rates of obliq-
uity change and to Dave Evans of Cal Tech, with whom we discussed the Pre-
cambrian glaciations. Conway Leovy talked to us about atmospheric matters.
With Bob Berner of Yale, we discussed matters pertaining to the evolution of
the atmosphere through time. Steve Stanley of Johns Hopkins gave us insight
into the Permo/Triassic extinction. Walter Alvarez and Allesandro Montanari
talked with us about the K/T extinction. Bob Pepin gave us insight into at-
mospheric effects.
Ross Taylor of the Australian University provided useful information to
us, and Geoff Marcy and Chris McKay discussed elements of the text. Doug
Lin of U.C. Santa Cruz discussed the fate of planetary systems with “Bad”
Jupiters. We are grateful to Al Cameron for use of his lunar formation results.

Peter D. Ward, Donald Brownlee

Seattle, August 1999

xv
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ward_FM_i-xxviii 3/10/03 14:01 Page xvii

Introduction:
The Astrobiology
Revolution and the
Rare Earth Hypothesis
n any given night, a vast array of extraterrestrial organisms fre-

O quent the television sets and movie screens of the world. From Star
Wars and “Star Trek” to The X-Files, the message is clear: The Uni-
verse is replete with alien life forms that vary widely in body plan, intelli-
gence, and degree of benevolence. Our society is clearly enamored of the ex-
pectation not only that there is life on other planets, but that incidences of
intelligent life, including other civilizations, occur in large numbers in the Uni-
verse.
This bias toward the existence elsewhere of intelligent life stems partly
from wishing (or perhaps fearing) it to be so and partly from a now-famous
publication by astronomers Frank Drake and Carl Sagan, who devised an es-
timate (called the Drake Equation) of the number of advanced civilizations
that might be present in our galaxy. This formula was based on educated
guesses about the number of planets in the galaxy, the percentage of those
that might harbor life, and the percentage of planets on which life not only

xvii
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R A R E E A R T H

could exist but could have advanced to exhibit culture. Using the best avail-
able estimates at the time, Drake and Sagan arrived at a startling conclusion:
Intelligent life should be common and widespread throughout the galaxy. In
fact, Carl Sagan estimated in 1974 that a million civilizations may exist in our
Milky Way galaxy alone. Given that our galaxy is but one of hundreds of bil-
lions of galaxies in the Universe, the number of intelligent alien species
would then be enormous.
The idea of a million civilizations of intelligent creatures in our galaxy
is a breathtaking concept. But is it credible? The solution to the Drake Equa-
tion includes hidden assumptions that need to be examined. Most important,
it assumes that once life originates on a planet, it evolves toward ever higher
complexity, culminating on many planets in the development of culture.
That is certainly what happened on our Earth. Life originated here about 4
billion years ago and then evolved from single-celled organisms to multicel-
lular creatures with tissues and organs, climaxing in animals and higher
plants. Is this particular history of life—one of increasing complexity to an
animal grade of evolution—an inevitable result of evolution, or even a com-
mon one? Might it, in fact, be a very rare result?
In this book we will argue that not only intelligent life, but even the
simplest of animal life, is exceedingly rare in our galaxy and in the Universe.
We are not saying that life is rare—only that animal life is. We believe that life
in the form of microbes or their equivalents is very common in the universe,
perhaps more common than even Drake and Sagan envisioned. However,
complex life—animals and higher plants—is likely to be far more rare than is
commonly assumed. We combine these two predictions of the commonness
of simple life and the rarity of complex life into what we will call the Rare
Earth Hypothesis. In the pages ahead we explain the reasoning behind this
hypothesis, show how it may be tested, and suggest what, if it is accurate, it
may mean to our culture.
The search in earnest for extraterrestrial life is only beginning, but we
have already entered a remarkable period of discovery, a time of excitement
and dawning knowledge perhaps not seen since Europeans reached the New
World in their wooden sailing ships. We too are reaching new worlds and are

xviii
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Introduction

acquiring data at an astonishing pace. Old ideas are crumbling. New views
rise and fall with each new satellite image or deep-space result. Each novel bi-
ological or paleontological discovery supports or undermines some of the
myriad hypotheses concerning life in the Universe. It is an extraordinary
time, and a whole new science is emerging: astrobiology, whose central focus
is the condition of life in the Universe. The practitioners of this new field are
young and old, and they come from diverse scientific backgrounds. Feverish
urgency is readily apparent on their faces at press conferences, such as those
held after the Mars Pathfinder experiments, the discovery of a Martian mete-
orite on the icefields of Antarctica, and the collection of new images from
Jupiter’s moons. In usually decorous scientific meetings, emotions boil over,
reputations are made or tarnished, and hopes ride a roller coaster, for scien-
tific paradigms are being advanced and discarded with dizzying speed. We
are witnesses to a scientific revolution, and as in any revolution there will be
winners and losers—both among ideas and among partisans. It is very much
like the early 1950s, when DNA was discovered, or the 1960s, when the con-
cept of plate tectonics and continental drift was defined. Both of these events
prompted revolutions in science, not only leading to the complete reorgani-
zation of their immediate fields and to adjustments in many related fields, but
also spilling beyond the boundaries of science to make us look at ourselves
and our world in new ways. That will come to pass as well in this newest sci-
entific revolution, the Astrobiology Revolution of the 1990s and beyond.
What makes this revolution so startling is that it is happening not within a
given discipline of science, such as biology in the 1950s or geology in the
1960s, but as a convergence of widely different scientific disciplines: astron-
omy, biology, paleontology, oceanography, microbiology, geology, and ge-
netics, among others.
In one sense, astrobiology is the field of biology ratcheted up to en-
compass not just life on Earth but also life beyond Earth. It forces us to re-
consider the life of our planet as but a single example of how life might work,
rather than as the only example. Astrobiology requires us to break the shack-
les of conventional biology; it insists that we consider entire planets as eco-
logical systems. It requires an understanding of fossil history. It makes us

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R A R E E A R T H

think in terms of long sweeps of time rather than simply the here and now.
Most fundamentally, it demands an expansion of our scientific vision—in
time and space.
Because it involves such disparate scientific fields, the Astrobiology
Revolution is dissolving many boundaries between disciplines of science. A
paleontologist’s discovery of a new life form from billion-year-old rocks in
Africa is of major consequence to a planetary geologist studying Mars. A sub-
marine probing the bottom of the sea finds chemicals that affect the calcula-
tions of a planetary astronomer. A microbiologist sequencing a string of
genes influences the work of an oceanographer studying the frozen oceans of
Europa (one of Jupiter’s moons) in the lab of a planetary geologist. The most
unlikely alliances are forming, breaking down the once-formidable academic
barriers that have locked science into rigid domains. New findings from di-
verse fields are being brought to bear on the central questions of astrobiol-
ogy: How common is life in the universe? Where can it survive? Will it leave
a fossil record? How complex is it? There are bouts of optimism and pes-
simism; E-mails fly; conferences are hastily assembled; research programs are
rapidly redirected as discoveries mount. The excitement is visceral, powerful,
dizzying, relentless. The practitioners are captivated by a growing belief: Life
is present beyond Earth.
The great surprise of the Astrobiology Revolution is that it has arisen in
part from the ashes of disappointment and scientific despair. As far back as
the 1950s, with the classic Miller–Urey experiments showing that organic
matter could be readily synthesized in a test tube (thus mimicking early Earth
environments), scientists thought they were on the verge of discovering how
life originated. Soon thereafter, amino acids were discovered in a newly fallen
meteorite, showing that the ingredients of life occurred in space. Radio-
telescope observations soon confirmed this, revealing the presence of organic
material in interstellar clouds. It seemed that the building blocks of life per-
meated the cosmos. Surely life beyond Earth was a real possibility.
When the Viking I spacecraft approached Mars in 1976, there was great
hope that the first extraterrestrial life—or at least signs of it—would be found
(see Figure I.1). But Viking did not find life. In fact, it found conditions hostile

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Introduction

Figure I.1 Percival Lowell’s 1908 globe of Mars. Some thought that the linear features were irri-
gation canals built by Martians.

to organic matter: extreme cold, toxic soil and lack of water. In many people’s
minds, these findings dashed all hopes that extraterrestrial life would ever be
found in the solar system. This was a crushing blow to the nascent field of as-
trobiology.
At about this time there was another major disappointment: The first se-
rious searches for “extrasolar” planets all yielded negative results. Although
many astronomers believed that planets were probably common around

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R A R E E A R T H

other stars, this remained only abstract speculation, for searches using Earth-
based telescopes gave no indication that any other planets existed outside our
own solar system. By the early 1980s, little hope remained that real progress
in this field would occur, for there seemed no way that we could ever detect
worlds orbiting other stars.
Yet it was also at this time that a new discovery paved the way for the
interdisciplinary methods now commonly used by astrobiologists. The 1980
announcement that the dinosaurs were not wiped out by gradual climate
change (as was so long thought) but rather succumbed to the catastrophic ef-
fects of the collision of a large comet with Earth 65 million years ago, was a
watershed event in science. For the first time, astronomers, geologists, and bi-
ologists had reason to talk seriously with one another about a scientific prob-
lem common to all. Investigators from these heretofore separate fields found
themselves at the same table with scientific strangers—all drawn there by the
same question: Could asteroids and comets cause mass extinction? Now, 20
years later, some of these same participants are engaged in a larger quest: to
discover how common life is on planets beyond Earth.
The indication that there was no life on Mars and the failure to find ex-
trasolar planets had damped the spirits of those who had begun to think of
themselves as astrobiologists. But the field involves the study of life on Earth
as well as in space, and it was from looking inward—examining this planet—
that the sparks of hope were rekindled. Much of the revitalization of astrobi-
ology came not from astronomical investigation but from the discovery, in
the early 1980s, that life on Earth occurs in much more hostile environments
than was previously thought. The discovery that some microbes live in sear-
ing temperatures and crushing pressures both deep in the sea and deep be-
neath the surface of our planet was an epiphany: If life survives under such
conditions here, why not on—or in—other planets, other bodies of our solar
system, or other plants and moons of far-distant stars?
Just knowing that life can stand extreme environmental conditions,
however, is not enough to convince us that life is actually there. Not only must
life be able to live in the harshness of a Mars, Venus, Europa, or Titan; it must
also have been able either to originate there or to travel there. Unless it can be

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Introduction

shown that life can form, as well as live, in extreme environments, there is lit-
tle hope that even simple life is widespread in the Universe. Yet here, too,
revolutionary new findings lead to optimism. Recent discoveries by geneti-
cists have shown that the most primitive forms of life on Earth—those that
we might expect to be close to the first life to have formed on our planet—
are exactly those tolerant life forms that are found in extreme environments.
This suggests to some biologists that life on Earth originated under conditions
of great heat, pressure, and lack of oxygen—just the sorts of conditions found
elsewhere in space. These findings give us hope that life may indeed be
widely distributed, even in the harshness of other planetary systems.
The fossil record of life on our own planet is also a major source of rel-
evant information. One of the most telling insights we have gleaned from the
fossil record is that life formed on Earth about as soon as environmental con-
ditions allowed its survival. Chemical traces in the most ancient rocks on
Earth’s surface give strong evidence that life was present nearly 4 billion years
ago. Life thus arose here almost as soon as it theoretically could. Unless this
occurred utterly by chance, the implication is that nascent life itself forms—
is synthesized from nonliving matter—rather easily. Perhaps life may origi-
nate on any planet as soon as temperatures cool to the point where amino
acids and proteins can form and adhere to one another through stable chem-
ical bonds. Life at this level may not be rare at all.
The skies too have yielded astounding new clues to the origin and dis-
tribution of life in the Universe. In 1995 astronomers discovered the first ex-
trasolar planets orbiting stars far from our own. Since then, a host of new
planets have been discovered, and more come to light each year.
For a while, some even thought we had found the first record of extra-
terrestrial life. A small meteorite discovered in the frozen icefields of Antarc-
tica appears to be one of many that originated on Mars, and at least one of
these may be carrying the fossilized remains of bacteria-like organisms of ex-
traterrestrial origin. The 1996 discovery was a bombshell. The President of
the United States announced the story in the White House, and the event
triggered an avalanche of new effort and resolve to find life beyond Earth. But
evidence—at least from this particular meteorite—is highly controversial.

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R A R E E A R T H

All of these discoveries suggest a similar conclusion: Earth may not be

the only place in this galaxy—or even in this solar system—with life. Yet if
other life is indeed present on planets or moons of our solar system, or on far-
distant planets circling other stars in the Universe, what kind of life is it?
What, for example, will be the frequency of complex metazoans, organisms with
multiple cells and integrated organ systems, creatures that have some sort of
behavior—organisms that we call animals? Here too a host of recent discov-
eries have given us a new view. Perhaps the most salient insights come, again,
from Earth’s fossil record.
New ways of more accurately dating evolutionary advances recognized
in the Earth’s fossil record, coupled with new discoveries of previously un-
known fossil types, have demonstrated that the emergence of animal life on
this planet took place later in time, and more suddenly, than we had sus-
pected. These discoveries show that life, at least as seen on Earth, does not
progress toward complexity in a linear fashion but does so in jumps, or as a
series of thresholds. Bacteria did not give rise to animals in a steady progres-
sion. Instead, there were many fits and starts, experiments and failures. Al-
though life may have formed nearly as soon as it could have, the formation of
animal life was much more recent and protracted. These findings suggest that
complex life is far more difficult to arrive at than evolving life itself and that
it takes a much longer time period to achieve.
It has always been assumed that attaining the evolutionary grade we call
animals would be the final and decisive step: that once this level of evolution
was achieved, a long and continuous progression toward intelligence should
occur. However, another insight of the Astrobiological Revolution has been
that attaining the stage of animal life is one thing, but maintaining that level is
quite something else. New evidence from the geological record has shown
that once it has evolved, complex life is subject to an unending succession of
planetary disasters that create what are known as mass extinction events.
These rare but devastating events can reset the evolutionary timetable and
destroy complex life, while sparing simpler life forms. Such discoveries again
suggest that the conditions hospitable to the evolution and existence of com-
plex life are far more specific than those that allow life’s formation. On some

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Introduction

planets, then, life might arise and animals eventually evolve—only to be

quickly destroyed by a global catastrophe.
To test the Rare Earth Hypothesis—the paradox that life may be nearly
everywhere but complex life almost nowhere—may ultimately require travel to
the distant stars. We cannot yet journey much beyond our own planet, and the
vast distances that separate us from even the nearest stars may prohibit us from
ever exploring planetary systems beyond our own. Perhaps this view is pes-
simistic, and we will ultimately find a way to travel much faster (and thus far-
ther), through worm holes or other unforeseen methods of interstellar travel,
enabling us to explore the Milky Way and perhaps other galaxies as well.
Let’s assume that we do master interstellar travel of some sort and begin
the search for life on other worlds. What types of worlds will harbor not just
life, but complex life equivalent to the animals of Earth? What sorts of plan-
ets or moons should we look for? Perhaps the best way to search is simply to
look for planets that resemble Earth, which is so rich with life. Do we have to
duplicate this planet exactly to find animal life, though? What is it about our
solar system and planet that has allowed the rise of complex life and nour-
ished it so well? Addressing this issue in the pages ahead should help us an-
swer the other questions we have posed.

RARE PLANET?

If we cast off our bonds of subjectivity about Earth and the solar system, and
try to view them from a truly “universal” perspective, we also begin to see as-
pects of Earth and its history in a new light. Earth has been orbiting a star
with relatively constant energy output for billions of years. Although life may
exist even on the harshest of planets and moons, animal life—such as that on
Earth—not only needs much more benign conditions but also must have
those conditions present and stable for great lengths of time. Animals as we
know them require oxygen. Yet it took about 2 billion years for enough oxy-
gen to be produced to allow all animals on Earth. Had our sun’s energy out-
put experienced too much variation during that long period of development

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R A R E E A R T H

(or even afterward), there would have been little chance of animal life evolv-
ing on this planet. On worlds that orbit stars with less consistent energy out-
put, the rise of animal life would be far chancier. It is difficult to conceive of
animal life arising on planets orbiting variable stars, or even on planets orbit-
ing stars in double or triple stellar systems, because of the increased chances
of energy fluxes sterilizing the nascent life through sudden heat or cold. And
even if complex life did evolve in such planetary systems, it might be difficult
for it to survive for any appreciable time.
Our planet was also of suitable size, chemical composition, and distance
from the sun to enable life to thrive. An animal-inhabited planet must be a
suitable distance from the star it orbits, for this characteristic governs
whether the planet can maintain water in a liquid state, surely a prerequisite
for animal life as we know it. Most planets are either too close or too far from
their respective stars to allow liquid water to exist on the surface, and al-
though many such planets might harbor simple life, complex animal life
equivalent to that on Earth cannot long exist without liquid water.
Another factor clearly implicated in the emergence and maintenance of
higher life on Earth is our relatively low asteroid or comet impact rate. The
collision of asteroids and comets with a planet can cause mass extinctions, as
we have noted. What controls this impact rate? The amount of material left
over in a planetary system after formation of the planets influences it: The
more comets and asteroids there are in planet-crossing orbits, the higher the
impact rate and the greater the chance of mass extinctions due to impact. Yet
this may not be the only factor. The types of planets in a system might also
affect the impact rate and thus play a large and unappreciated role in the evo-
lution and maintenance of animals. For Earth, there is evidence that the giant
planet Jupiter acted as a “comet and asteroid catcher,” a gravity sink sweeping
the solar system of cosmic garbage that might otherwise collide with Earth.
It thus reduced the rate of mass extinction events and so may be a prime rea-
son why higher life was able to form on this planet and then maintain itself.
How common are Jupiter-sized planets?
In our solar system, Earth is the only planet (other than Pluto) with a
moon of such appreciable size compared to the planet it orbits, and it is the

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Introduction

only planet with plate tectonics, which causes continental drift. As we will try
to show, both of these attributes may be crucial in the rise and persistence of
animal life.
Perhaps even a planet’s placement in a particular region of its home
galaxy plays a major role. In the star-packed interiors of galaxies, the fre-
quency of supernovae and stellar close encounters may be high enough to
preclude the long and stable conditions apparently required for the develop-
ment of animal life. The outer regions of galaxies may have too low a per-
centage of the heavy elements necessary to build rocky planets and to fuel
the radioactive warmth of planetary interiors. The comet influx rate may even
be affected by the nature of the galaxy we inhabit and by our solar system’s
position in that galaxy. Our sun and its planets move through the Milky Way
galaxy, yet our motion is largely within the plane of the galaxy as a whole,
and we undergo little movement through the spiral arms. Even the mass of a
particular galaxy might affect the odds of complex life evolving, for galactic
size correlates with its metal content. Some galaxies, then, might be far more
amenable to life’s origin and evolution than others. Our star—and our solar
system—are anomalous in their high metal content. Perhaps our very galaxy
is unusual.
Finally, it is likely that a planet’s history, as well as its environmental con-
ditions, plays a part in determining which planets will see life advance to an-
imal stages. How many planets, otherwise perfectly positioned for a history
replete with animal life, have been robbed of that potential by happen-
stance? An asteroid impacting the planet’s surface with devastating and life-
exterminating consequences. Or a nearby star exploding into a cataclysmic
supernova. Or an ice age brought about by a random continental configura-
tion that eliminates animal life through a chance mass extinction. Perhaps
chance plays a huge role.
Ever since Polish astronomer Nicholas Copernicus plucked it from the
center of the Universe and put it in orbit around the sun, Earth has been pe-
riodically trivialized. We have gone from the center of the Universe to a
small planet orbiting a small, undistinguished star in an unremarkable region
of the Milky Way galaxy—a view now formalized by the so-called Principle

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R A R E E A R T H

of Mediocrity, which holds that we are not the one planet with life but one
of many. Various estimates for the number of other intelligent civilizations
range from none to 10 trillion.
If it is found to be correct, however, the Rare Earth Hypothesis will re-
verse that decentering trend. What if the Earth, with its cargo of advanced
animals, is virtually unique in this quadrant of the galaxy—the most diverse
planet, say, in the nearest 10,000 light-years? What if it is utterly unique: the
only planet with animals in this galaxy or even in the visible Universe, a bas-
tion of animals amid a sea of microbe-infested worlds? If that is the case, how
much greater the loss the Universe sustains for each species of animal or plant
driven to extinction through the careless stewardship of Homo sapiens?
Welcome aboard.

xxviii
Another random document with
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at the upper part and terminating posteriorly at the lower part of the
body cavity.

In Fig. 63 we show the arrangement of some of the chief organs of

the body, with the exception of the muscular and respiratory
systems, and the fat-body. It is scarcely necessary to point out that
the figure is merely diagrammatic, and does not show the shapes
and sizes of the organs as they will be found in any one Insect.

Muscles.

The muscular system of Insects is very extensive, Lyonnet[31] having

found, it is said, nearly 4000 muscles in the caterpillar of the goat-
moth; a large part of this number are segmental repetitions,
nevertheless the muscular system is really complex, as may be seen
by referring to the study of the flight of dragon-flies by von
Lendenfeld.[32]

The minute structure of the muscles does not differ essentially from
what obtains in Vertebrate animals. The muscles are aggregations of
minute fibrils which are transversely striated, though in variable
degree. Those in the thorax are yellow or pale brown, but in other
parts the colour is more nearly white. The muscles of flight are
described as being penetrated by numerous tracheae, while those
found elsewhere are merely surrounded by these aerating tubules.

The force brought into play by the contractions of Insect muscles is

very great, and has been repeatedly stated to be much superior to
that of Vertebrate animals; very little reliance can, however, be
placed on the assumptions and calculations that are supposed to
prove this, and it is not supported by Camerano's recent researches.
[33]
Some of the tendons to which the muscles are attached are very
elaborate structures, and are as hard as the chitinous skeleton, so
as to be like small bones in their nature. A very elaborate tendon of
this kind is connected with the prothoracic trochantin in Coleoptera,
and may be readily examined in Hydrophilus. It has been suggested
that the entothorax is tendinous in its origin, but other morphologists
treat it, with more reason, as an elaborate fold inwards of the
integument.

Fig. 64.—Cephalic and ventral chain of ganglia: A, larva of

Chironomus; B, imago of Hippobosca. (After Brandt.)

Nervous System.

Insects are provided with a very complex nervous system, which

may be treated as consisting of three divisions:—(1) The cephalic
system; (2) the ventral, or ganglionic chain; (3) an accessory
sympathetic system, or systems. All these divisions are intimately
connected. We will consider first the most extensive, viz. the ventral
chain. This consists of a series of small masses of nervous matter
called ganglia which extend in the longitudinal direction of the body
along the median line of the lower aspect, and are connected by
longitudinal commissures, each ganglion being joined to that
following it by two threads of nervous matter. Each of the ganglia of
the ventral chain really consists of two ganglia placed side by side
and connected by commissures as well as cellular matter. In larvae
some of the ganglia may be contiguous, so that the commissures do
not exist. From the ganglia motor nerves proceed to the various
parts of the body for the purpose of stimulating and co-ordinating the
contractions of the muscles. The number of the ganglia in the ventral
chain differs greatly in different Insects, and even in the different
stages of metamorphosis of the same species, but never exceeds
thirteen. As this number is that of the segments of the body, it has
been considered that each segment had primitively a single
ganglion. Thirteen ganglia for the ventral chain can, however, be only
demonstrated in the embryonic state; in the later stages of life eleven
appears to be the largest number that can be distinguished, and so
many as this are found but rarely, and then chiefly in the larval stage.
The diminution in number takes place by the amalgamation or
coalescence of some of the ganglia, and hence those Insects in
which the ganglia are few are said to have a highly concentrated
nervous system. The modes in which these ganglia combine are
very various; the most usual is perhaps that of the combination of the
three terminal ganglia into one body. As a rule it may be said that
concentration is the concomitant of a more forward position of the
ganglia. As a result of this it is found that in some cases, as in
Lamellicorn beetles, there are no ganglia situate in the abdomen. In
the perfect state of the higher Diptera, the thoracic and abdominal
ganglia are so completely concentrated in the thorax as to form a
sort of thoracic brain. In Fig. 64 we represent a very diffuse and a
very concentrated ganglionic chain; A being that of the larva of
Chironomus, B that of the imago of Hippobosca. In both these
sketches the cephalic ganglia as well as those of the ventral chain
are shown.

Turning next to the cephalic masses, we find these in the perfect

Insect to be nearly always two in number: a very large and complex
one placed above the oesophagus, and therefore called the supra-
oesophageal ganglion; and a smaller one, the sub- or infra-
oesophageal, placed below the oesophagus. The latter ganglion is in
many Insects so closely approximated to the supra-oesophageal
ganglion that it appears to be a part thereof, and is sometimes
spoken of as the lower brain. In other Insects these two ganglia are
more remote, and the infra-oesophageal one then appears part of
the ventral chain. In the embryo it is said that the mode of
development of the supra-oesophageal ganglion lends support to the
idea that it may be the equivalent of three ganglia; there being at one
time three lobes, which afterwards coalesce, on each side of the
mouth. This is in accordance with the view formulated by
Viallanes[34] to the effect that this great nerve-centre, or brain, as it is
frequently called, consists essentially of three parts, viz. a Proto-, a
Deuto-, and a Trito-cerebron. It is, however, only proper to say that
though the brain and the ventral chain of ganglia may appear to be
one system, and in the early embryonic condition to be actually
continuous, these points cannot be considered to be fully
established. Dr. L. Will has informed us[35] that in Aphididae the
brain has a separate origin, and is only subsequently united with the
ganglionic chain. Some authorities say that in the early condition the
sub-oesophageal ganglion is formed from two, and the supra-
oesophageal from the same number of ganglia; the division in that
case being 2 and 2, not 3 and 1, as Viallanes' views would suggest.
The inquiries that are necessary to establish such points involve very
complex and delicate investigations, so that it is not a matter of
surprise that it cannot yet be said whether each of these views may
be in certain cases correct. The supra- and sub-oesophageal ganglia
are always intimately connected by a commissure on each side of
the oesophagus; when very closely approximated they look like one
mass through which passes the oesophagus (Fig. 66, A). The large
supra-oesophageal ganglion supplies the great nerves of the
cephalic sense-organs, while the smaller sub-oesophageal centre
gives off the nerves to the parts of the mouth. From the lower and
anterior part of the supra-oesophageal ganglion a nervous filament
extends as a ring round the anterior part of the oesophagus, and
supplies a nerve to the upper lip.[36] This structure is not very well
known, and has been chiefly studied by Liénard,[37] who considers
that it will prove to be present in all Insects.

Whether the two cephalic ganglia be considered as really part of a

single great ganglionic chain, or the reverse, they are at any rate
always intimately connected with the ventral ganglia. We have
already stated that the two cephalic masses are themselves closely
approximated in many Insects, and may add that in some Hemiptera
the first thoracic ganglion of the ventral chain is amalgamated into
one body with the sub-oesophageal ganglion, and further that there
are a few Insects in which this latter centre is wanting. If the cephalic
ganglia and ventral chain be looked on as part of one system, this
may be considered as composed originally of seventeen ganglia,
which number has been demonstrated in some embryos.

The anatomy of the supra-oesophageal ganglion is very complex; it

has been recently investigated by Viallanes[38] in the wasp (Vespa)
and in a grasshopper (Caloptenus italicus). The development and
complication of its inner structure and of some of its outer parts
appear to be proportional with the state of advancement of the
instinct or intelligence of the Insect, and Viallanes found the brain of
the grasshopper to be of a more simple nature than that of the wasp.

Fig. 65.—Brain of Worker Ant of Formica rufa. (After Leydig, highly

magnified.) Explanation in text.

Brandt, to whom is due a large part of our knowledge of the anatomy

of the nervous system in Insects, says that the supra-oesophageal
ganglion varies greatly in size in various Insects, its mass being to a
great extent proportional with the development of the compound
eyes; hence the absolute size is not a criterion for the amount of
intelligence, and we must rather look to the complication of the
structure and to the development of certain parts for an index of this
nature. The drone in the honey-bee has, correlatively with the
superior development of its eyes, a larger brain than the worker, but
the size of the hemispheres, and the development of the gyri
cerebrales is superior in the latter. In other words, the mass of those
great lobes of the brain that are directly connected with the faceted
eyes must not be taken into account in a consideration of the relation
of the size and development of the brain to the intelligence of the
individual. The weight of the brain in Insects is said by Lowne to vary
from 1⁄150 to 1⁄2500 of the weight of the body.

Figure 65 gives a view of one side of the supra-oesophageal

ganglion of the worker of an ant,—Formica rufa,—and is taken from
Leydig, who gives the following elucidation of it: A, primary lobe, a,
homogeneous granular inner substance, b, cellular envelope; B,
stalked bodies (gyri cerebrales), a, b, as before; C, presumed
olfactory lobes, c, inner substance, d, ganglionic masses; D, ocular
lobes, e, f, g, h, various layers of the same; E, origin of lateral
commissures; F, median commissure in interior of brain; G, lower
brain (sub-oesophageal ganglion); H, ocelli; J, faceted eye.

Fig. 66.—Stomato-gastric nerves of Cockroach: A, with brain in situ,

after Koestler; B, with the brain removed, after Miall and Denny:
s.g, supra-oesophageal ganglion; o, optic nerve; a, antennary
nerve; f.g, frontal ganglion; oe, oesophagus; c, connective; p.g,
paired ganglia; v.g, crop or ventricular ganglion; r, recurrent nerve.

Besides the brain and the great chain of ganglia there exists an
accessory system, or systems, sometimes called the sympathetic,
vagus, or visceral system. Although complex, these parts are
delicate and difficult of dissection, and are consequently not so well
known as is the ganglionic chain. There is a connecting or median
nerve cord, communicating with the longitudinal commissures of
each segment, and itself dilating into ganglia at intervals; this is
sometimes called the unpaired system. There is another group of
nerves having paired ganglia, starting from a small ganglion in the
forehead, then connecting with the brain, and afterwards extending
along the oesophagus to the crop and proventriculus (Fig. 66). This
is usually called the stomatogastric system. The oesophageal ring
we have already spoken of.

By means of these accessory nervous systems all the organs of the

body are brought into more or less direct relation with the brain and
the ganglionic chain.

Our knowledge of these subsidiary nervous systems is by no means

extensive, and their nomenclature is very unsettled; little is actually
known as to their functions.

Organs of Sense.

Insects have most delicate powers of perception, indeed they are

perhaps superior in this respect to the other classes of animals.
Their senses, though probably on the whole analogous to those of
the Vertebrata, are certainly far from corresponding therewith, and
their sense organs seem to be even more different from those of
what we call the higher animals than the functions themselves are.
We have already briefly sketched the structure of the optical organs,
which are invariably situate on the head. This is not the case with the
ears, which certainly exist in one Order,—the Orthoptera,—and are
placed either on the front legs below the knee, or at the base of the
abdomen. Notwithstanding their strange situation, the structures
alluded to are undoubtedly auditory, and somewhat approximate in
nature to the ear of Vertebrates, being placed in proximity to the
inner face of a tense membrane; we shall refer to them when
considering the Orthoptera. Sir John Lubbock considers—no doubt
with reason—that some ants have auditory organs in the tibia. Many
Insects possess rod-like or bristle-like structures in various parts of
the body, called chordotonal organs; they are considered by
Graber[39] and others to have auditory functions, though they are not
to be compared with the definite ears of the Orthoptera.

The other senses and sense organs of Insects are even less known,
and have given rise to much perplexity; for though many structures
have been detected that may with more or less probability be looked
on as sense organs, it is difficult to assign a particular function to any
of them, except it be to the sensory hairs. These are seated on
various parts of the body. The chitinous covering, being a dead, hard
substance, has no nerves distributed in it, but it is pierced with
orifices, and in some of these there is implanted a hair which at its
base is in connexion with a nerve; such a structure may possibly be
sensitive not only to contact with solid bodies, but even to various
kinds of vibration. We give a figure (Fig. 67) of some of these hairs
on the caudal appendage of a cricket, after Vom Rath. The small
hairs on the outer surface of the chitin in this figure have no sensory
function, but each of the others probably has; and these latter, being
each accompanied by a different structure, must, though so closely
approximated, be supposed to have a different function; but in what
way those that have no direct connexion with a nerve may act it is
difficult to guess.

The antennae of Insects are the seats of a great variety of sense

organs, many of which are modifications of the hair, pit and nerve
structure we have described above, but others cannot be brought
within this category. Amongst these we may mention the pits
covered with membrane (figured by various writers), perforations of
the chitin without any hair, and membranous bodies either concealed
in cavities or partially protruding therefrom.
 Fig. 67.—Longitudinal section of portion of caudal appendage of
Acheta domestica (after Vom Rath): ch, chitin; hyp, hypodermis;
n, nerve; h1, integumental hairs, not sensitive; h2, ordinary hair;
h3, sensory hair; h4, bladder-like hair; sz, sense-cell.

Fig. 68.—Longitudinal section of apex of palpus of Pieris brassicae:

sch, scales; ch, chitin; hyp, hypodermis; n, nerve; sz, sense cells;
sh, sense hairs. (After Vom Rath.)

Various parts of the mouth are also the seats of sense organs of
different kinds, some of them of a compound character; in such
cases there may be a considerable number of hairs seated on
branches of a common nerve as figured by Vom Rath[40] on the apex
of the maxillary palp of Locusta viridissima, or a compound organ
such as we represent in Fig. 68 may be located in the interior of the
apical portion of the palp.

The functions of the various structures that have been detected are,
as already remarked, very difficult to discover. Vom Rath thinks the
cones he describes on the antennae and palpi are organs of smell,
while he assigns to those on the maxillae, lower lip, epipharynx, and
hypopharynx the rôle of taste organs, but admits he cannot draw any
absolute line of distinction between the two forms. The opinions of
Kraepelin, Hauser, and Will, as well as those of various earlier
writers, are considered in Sir John Lubbock's book on this subject.
[41]

Alimentary and Nutritive System.

The alimentary canal occupies the median longitudinal axis of the
body, being situated below the dorsal vessel, and above the ventral
nervous chain; it extends from the mouth to the opposite extremity of
the body. It varies greatly in the different kinds of Insects, but in all its
forms it is recognised as consisting essentially of three divisions:
anterior, middle, and posterior. The first and last of these divisions
are considered to be of quite different morphological nature from the
middle part, or true stomach, and to be, as it were, invaginations of
the extremities of a closed bag; it is ascertained that in the embryo
these invaginations have really blind extremities (see Fig. 82,
p. 151), and only subsequently become connected with the middle
part of the canal. There are even some larvae of Insects in which the
posterior portion of the canal is not opened till near the close of the
larval life; this is the case with many Hymenoptera, and it is
probable, though not as frequently stated certain, that the occlusion
marks the point of junction of the proctodaeum with the stomach.
The anterior and posterior parts of the canal are formed by the
ectoderm of the embryo, and in embryological and morphological
language are called respectively the stomodaeum and proctodaeum;
the true stomach is formed from the endoderm, and the muscular
layer of the whole canal from the mesoderm.

Fig. 69.—Digestive system of Xyphidria camelus (after Dufour): a, head

capsule; b, salivary glands; c, oesophagus; d, crop; e,
proventriculus; f, chyle, or true stomach; g, small intestine; h,
large intestine; i, Malpighian tubes; k, termination of body.
The alimentary canal is more complex anatomically than it is
morphologically, and various parts are distinguished, viz. the canal
and its appendicula; the former consisting of oesophagus, crop,
gizzard, true stomach, and an intestine divided into two or more
parts. It should be remarked that though it is probable that the
morphological distinctions correspond to a great extent with the
anatomical lines of demarcation, yet this has not been sufficiently
ascertained: the origin of the proctodaeum in Musca is indeed a
point of special difficulty, and one on which there is considerable
diversity of opinion. In some Hemiptera the division of the canal into
three parts is very obscure, so that it would be more correct, as
Dufour says, to define it as consisting in these Insects of two main
divisions—one anterior to, the other posterior to, the insertion of the
Malpighian tubes.

It should be borne in mind that the alimentary canal is very different

in different Insects, so that the brief general description we must
confine ourselves to will not be found to apply satisfactorily to any
one Insect. The oesophagus is the part behind the mouth, and is
usually narrow, as it has to pass through the most important nervous
centres; extremely variable in length, it dilates behind to form the
crop. It may, too, have a dilatation immediately behind the mouth,
and in such case a pharynx is considered to exist. The crop is
broader than the oesophagus, and must be looked on as a mere
dilatation of the latter, as no line of demarcation can be pointed out
between the two, and the crop may be totally absent.

In some of the sucking Insects there is a lateral diverticulum, having

a stalk of greater or less length, called the sucking-stomach; it is by
no means certain that the function this name implies is correctly
assigned to the organ.

The gizzard or proventriculus (French, gésier; German, Kaumagen)

is a small body interposed in some Insects between the true
stomach and the crop or oesophagus. It is frequently remarkable for
the development of its chitinous lining into strong toothed or ridged
processes that look as if they were well adapted for the comminution
of food. The function of the proventriculus in some Insects is
obscure; its structure is used by systematists in the classification of
ants. The extremity of the proventriculus not infrequently projects
into the cavity of the stomach.

The true stomach, or chylific ventricle (Magen or Mitteldarm of the

Germans), is present in all the post-embryonic stages of the Insect's
life, existing even in the imagines of those who live only for a few
hours, and do not use the stomach for any alimentary purpose. It is
so variable in shape and capacity that no general description of it
can be given. Sometimes it is very elongate, so that it is coiled and
like an intestine in shape; it very frequently bears diverticula or
pouches, which are placed on the anterior part, and vary greatly in
size, sometimes they are only two in number, while in other cases
they are so numerous that a portion of the outside of the stomach
looks as if it were covered with villi. A division of the stomach into
two parts is in some cases very marked, and the posterior portion
may, in certain cases, be mistaken for the intestine; but the position
of the Malpighian tubes serves as a mark for the distinction of the
two structures, the tubes being inserted just at the junction of the
stomach with the intestine.

The intestine is very variable in length: the anterior part is the

smaller, and is frequently spoken of as the colon; at the extremity of
the body the gut becomes much larger, so as to form a rectum.
There is occasionally a diverticulum or "caecum" connected with the
rectum, and in some Insects stink-glands. In some Hemiptera there
is no small intestine, the Malpighian tubes being inserted at the
junction of the stomach with the rectum. The total length of the
alimentary canal is extremely variable; it is necessarily at least as
long as the distance between the mouth and anal orifice, but
sometimes it is five or six times as long as this, and some of its parts
then form coils in the abdominal cavity.
The alimentary canal has two coats of muscles: a longitudinal and a
transverse or annular. Both coexist in most of its parts. Internal to
these coats there exists in the anterior and posterior parts of the
canal a chitinous layer, which in the stomach is replaced by a
remarkable epithelium, the cells of which are renewed, new ones
growing while the old are still in activity. We figure a portion of this
structure after Miall and Denny, and may remark that Oudemans[42]
has verified the correctness of their representation. The layers below
represent the longitudinal and transverse muscles.

Fig. 70.—Epithelium of stomach of Cockroach (after Miall and Denny):

the lower parts indicate the transverse and longitudinal muscular
layers.

In addition to the various diverticula we have mentioned, there are

two important sets of organs connected with the alimentary canal,
viz. the salivary glands and the Malpighian tubes.

The salivary glands are present in many Insects, but are absent in
others. They are situate in the anterior portion of the body, and are
very variable in their development, being sometimes very extensive,
in other cases inconspicuous. They consist either of simple tubes
lined with cells, or of branched tubes, or of tubes dilated laterally into
little acini or groups of bags, the arrangement then somewhat
resembling that of a bunch of grapes. There are sometimes large
sacs or reservoirs connected with the efferent tubes proceeding from
the secreting portions of the glands. The salivary glands ultimately
discharge into the mouth, so that the fluid secreted by them has to
be swallowed in the same manner as the food, not improbably along
with it. The silk so copiously produced by some larvae comes from
very long tubes similar in form and situation to the simple tubes of
the salivary glands.
The Malpighian tubules are present in most Insects, though they are
considered on good authority to be absent in many Collembola and
in some Thysanura. They are placed near the posterior part of the
body, usually opening into the alimentary canal just at the junction of
the stomach and the intestine, at a spot called the pylorus. They vary
excessively in length and in number,[43] being sometimes only two,
while in other cases there may be a hundred or even more of them.
In some cases they are budded off from the hind-gut of the embryo
when this is still very small; in other cases they appear later;
frequently their number is greater in the adult than it is in the young.
In Gryllotalpa there is one tube or duct with a considerable number
of finer tubes at the end of it. There is no muscular layer in the
Malpighian tubes, they being lined with cells which leave a free canal
in the centre. The tubes are now thought, on considerable evidence,
to be organs for the excretion of uric acid or urates, but it is not
known how they are emptied. Marchal has stated[44] that he has
seen the Malpighian tubes, on extraction from the body, undergo
worm-like movements; he suggests that their contents may be
expelled by similar movements when they are in the body.

The functions of the different portions of the alimentary canal, and

the extent to which the ingested food is acted on by their mechanical
structures or their products is very obscure, and different opinions
prevail on important points. It would appear that the saliva exercises
a preparatory action on the food, and that the absorption of the
nutritive matter into the body cavity takes place chiefly from the true
stomach, while the Malpighian tubes perform an excretory function.
Beyond these elementary, though but vaguely ascertained facts, little
is known, though Plateau's[45] and Jousset's researches on the
digestion of Insects throw some light on the subject.

Respiratory Organs.

The respiration of Insects is carried on by means of a system of

vessels for the conveyance of air to all parts of the body; this system
is most remarkably developed and elaborate, and contrasts strongly
with the mechanism for the circulation of the blood, which is as much
reduced as the air system is highly developed, as well as with the
arrangement that exists in the Vertebrates. There are in Insects no
lungs, but air is carried to every part of the body directly by means of
tracheae. These tracheae connect with the spiracles—the orifices at
the sides of the body we have already mentioned when describing
the external structures—and the air thus finds its way into the most
remote recesses of the Insect's body. The tracheae are all intimately
connected. Large tubes connect the spiracles longitudinally, others
pass from side to side of the body, and a set of tracheae for the
lower part of the body is connected with another set on the upper
surface by means of several descending tubes. From these main
channels smaller branches extend in all directions, forking and giving
off twigs, so that all the organs inside the body can be supplied with
air in the most liberal manner. On opening a freshly deceased Insect
the abundance of the tracheae is one of the peculiarities that most
attracts the attention; and as these tubes have a peculiar white
glistening appearance, they are recognised without difficulty. In
Insects of active flight, possibly in some that are more passive,
though never in larvae, there are air-sacs, of more than one kind,
connected with the tracheae, and these are sufficiently capacious to
have a considerable effect in diminishing the specific gravity of the
Insect. The most usual situation for these sacs is the basal portion of
the abdominal cavity, on the great lateral tracheal conduits. In
speaking of the external structure we have remarked that the
stigmata, or spiracles, by which the air is admitted are very various in
their size and in the manner in which they open and close. Some
spiracles have no power of opening; while others are provided with a
muscular and valvular apparatus for the purpose of opening and
closing effectually.

The structure of the tracheae is remarkable: they are elastic and

consist of an outer cellular, and an inner chitinous layer; this latter is
strengthened by a peculiar spiral fibre, which gives to the tubes,
when examined with the microscope, a transversely, closely striated
appearance. Packard considers[46] that in some tracheae this fibre is
not really spiral, but consists of a large number of closely placed
rings. Such a condition has not, however, been recorded by any
other observer. The spiral fibre is absent in the fine capillary twigs of
the tracheal system, as well as from the expanded sacs. The mode
of termination of the capillary branches is not clear. Some have
supposed that the finest twigs anastomose with others; on the other
hand it has been said that they terminate by penetrating cells, or that
they simply come to an end with either open or closed extremities.
Wistinghausen[47] states that in the silk-glands the tracheal twigs
anastomose, and he is of opinion that the fine terminal portions
contain fluid. However this may be, it is certain that all the organs are
abundantly supplied with a capillary tracheal network, or arboreal
ramification, and that in some cases the tubes enter the substance of
tissues. Near their terminations they are said to be 1⁄30 to 1⁄60
millimetre in diameter.

Fig. 71.—Portion of the abdominal part of tracheal system of a Locust

(Oedipoda): a, spiracular orifices; b, tracheal tubes; c, vesicular
dilatations; d, tracheal twigs or capillaries. (After Dufour.)

We must repeat that such a system as we have just sketched forms

a striking contrast to the imperfect blood-vascular system, and that
Insects differ profoundly in these respects from Vertebrate animals.
In the latter the blood-vessels penetrate to all the tissues and form
capillaries, while the aerating apparatus is confined to one part of the
body; in Insects the blood-circulating system is very limited, and air
is carried directly by complex vessels to all parts; thus the tracheal
system is universally recognised as one of the most remarkable of
the characters of Insects. Many Insects have a very active
respiratory system, as is shown by the rapidity with which they are
affected by agents like chloroform; but the exact manner in which the
breathing is carried on is unknown. In living Insects rapid movements
of contraction and expansion of parts of the body, chiefly the
abdomen, may be observed, and these body contractions are
sometimes accompanied by opening and shutting the spiracular
orifices: it has been inferred that these phenomena are respiratory.
Although such movements are not always present, it is possible that
when they occur they may force the air onwards to the tissues,
though this is by no means certain. It is clear that the tracheal
system is the usual means of supplying the organisation with
oxygen, but it appears to be improbable that it can also act as the
agent for removing the carbonaceous products of tissue-changes. It
has been thought possible that carbonic acid might reach the
spiracles from the remote capillaries by a process of diffusion,[48] but
it should be recollected that as some Insects have no tracheal
system, there must exist some other mode of eliminating carbonic
acid, and it is possible that this mode may continue to operate as an
important agent of purification, even when the tracheal system is, as
a bearer of air to the tissues, highly developed. Eisig[49] has
suggested that the formation of chitin is an act of excretion; if so this
is capable of relieving the system of carbonic acid to some extent.
Others have maintained that transpiration takes place through the
delicate portions of the integument. Lubbock[50] has shown that
Melolontha larvae breathe "partly by means of their skin." The mode
in which the carbon of tissue-change, and the nitrogen of inspiration
are removed, is still obscure; but it appears probable that the views
expressed by Réaumur, Lyonnet, and Lowne[51] as to inspiration and
expiration may prove to be nearer the truth than those which are
more widely current. In connexion with this it should be recollected
that the outer integument consists of chitin, and is cast and renewed
several times during the life of the individual. Now as chitin consists
largely of carbon and nitrogen, it is evident that the moulting must
itself serve as a carbonaceous and nitrogenous excretion. If, as is
suggested by Bataillon's researches,[52] the condition accompanying
metamorphosis be that of asphyxia, it is probable that the secretion
of the new coat of chitin may figure as an act of excretion of
considerable importance. If there be any truth in this suggestion it
may prove the means of enabling us to comprehend some points in
the development of Insects that have hitherto proved very
perplexing.

Peyrou has shown[53] that the atmosphere extracted from the bodies
of Insects (Melolontha) is much less rich in oxygen than the
surrounding atmosphere is, and at ordinary temperatures always
contains a much larger proportion of carbonic acid: he finds, too, that
as in the leaves with which he makes a comparison, the proportion
of oxygen augments as the protoplasmic activity diminishes. Were
such an observation carried out so as to distinguish between the air
in the tracheal system and the gas in other parts of the body the
result would be still more interesting.

We know very little as to the animal heat produced by insects, but it

is clear from various observations[54] that the amount evolved in
repose is very small. In different conditions of activity the
temperature of the insect may rise to be several degrees above that
of the surrounding medium, but there seems to be at present no
information as to the physiological mode of its production, and as to
the channel by which the products—whether carbonic acid or other
matters—may be disposed of.

In the order Aptera (Thysanura and Collembola) the tracheal system

is highly peculiar. In some Collembola it apparently does not exist,
and in this case we may presume with greater certainty that
transpiration of gases occurs through the integument: in other
members of this Order tracheae are present in a more or less
imperfect state of development, but the tracheae of different
segments do not communicate with one another, thus forming a
remarkable contrast to the amalgamated tracheal system of the
other Orders of Insects, where, even when the tracheal system is
much reduced in extent (as in Coccidae), it is nevertheless
completely unified. Gryllotalpa is, however, said by Dohrn[55] to be
exceptional in this respect; the tracheae connected with each
spiracle remaining unconnected.

Water Insects have usually peculiarities in their respiratory systems,

though these are not so great as might à priori have been
anticipated. Some breathe by coming to the surface and taking in a
supply of air in various manners, but some apparently obtain from
the water itself the air necessary for their physiological processes.
Aquatic Insects are frequently provided with gills, which may be
either wing-like expansions of the integument containing some
tracheae (Ephemeridae larvae), or bunches of tubes, or single tubes
(Trichoptera larvae). Such Insects may either possess stigmata in
addition to the gills, or be destitute of them. In other cases air is
obtained by taking water into the posterior part of the alimentary
canal (many dragon-flies), which part is then provided with special
tracheae. Some water-larvae appear to possess neither stigmata nor
gills (certain Perlidae and Diptera), and it is supposed that these
obtain air through the integument; in such Insects tracheal twigs may
frequently be seen on the interior of the skin. In the imago state it is
the rule that Water Insects breathe by means of stigmata, and that
they carry about with them a supply of air sufficient for a longer or
shorter period. A great many Insects that live in water in their earlier
stages and breathe there by peculiar means, in their perfect imago
state live in the air and breathe in the usual manner. There are, in
both terrestrial and aquatic Insects, a few cases of exsertile sacs
without tracheae, but filled with blood (Pelobius larva, Machilis, etc.);
and such organs are supposed to be of a respiratory nature, though
there does not appear to be any positive evidence to that effect.

Blood and Blood-Circulation.

Owing to the great complexity of the tracheal system, and to its

general diffusion in the body, the blood and its circulation are very
different in Insects from what they are in Vertebrates, so that it is
scarcely conducive to the progress of physiological knowledge to call
two fluids with such different functions by one name. The blood of
Insects varies according to the species, and in all probability even in
conformity with the stage of the life of the individual. Its primary office
is that of feeding the tissues it bathes, and it cannot be considered
as having any aerating function. It is frequently crowded with fatty
substances. Graber says: "The richness of Insect blood in
unsaponified or unelaborated fat shows in the plainest manner that it
is more properly a mixture of blood and chyle; or indeed we might
say with greater accuracy, leaving out of consideration certain
matters to be eliminated from it, that it is a refined or distilled chyle."
Connected in the most intimate manner with the blood there is a
large quantity of material called vaguely the fat-body; the blood and
its adjuncts of this kind being called by Wielowiejski[56] the blood-
tissue. We shall return to the consideration of this tissue after
sketching the apparatus for distributing the refined chyle, or blood as
we must, using the ordinary term, call it.

There is in Insects no complete system of blood-vessels, though

there is a pulsating vessel to ensure distribution of the nutritive fluid.
This dorsal vessel, or heart as it is frequently called, may be
distinguished and its pulsations watched, in transparent Insects
when alive. It is situate at the upper part of the body, extending from
the posterior extremity, or near it, to the head or thorax, and is an
elongate tube, consisting as it were of a number of united chambers;
it is closed behind, except in some larvae, but is open in front, and
has several orifices at the sides; these orifices, or ostia, are
frequently absent from the front part of the tube, which portion is also
narrower, being called the aorta—by no means a suitable term. Near
the lateral orifices there are delicate folds, which act to some extent
as valves, facilitating, in conjunction with the mode of contraction of
the vessel, a forward movement of the blood. The composition of the
tube, or series of chambers, is that of a muscular layer, with internal
and external membranous coverings, the intima and adventitia. Olga
Poletajewa states[57] that in Bombus the dorsal vessel consists of
five chambers placed in longitudinal succession, and not very
intimately connected, and that there is but little valvular structure. In