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Nature’s Patterns
This page intentionally left blank
Nature’s
Patterns
A Tapestry in Three Parts

Philip Ball
Nature’s Patterns is a trilogy composed of
Shapes, Flow, and Branches

1
 3
Great Clarendon Street, Oxford OX2 6DP
Oxford University Press is a department of the University of Oxford.
It furthers the University’s objective of excellence in research, scholarship,
and education by publishing worldwide in
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With offices in
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South Korea Switzerland Thailand Turkey Ukraine Vietnam
Oxford is a registered trade mark of Oxford University Press
in the UK and in certain other countries
Published in the United States
by Oxford University Press Inc., New York
# Philip Ball 2009
The moral rights of the author have been asserted
Database right Oxford University Press (maker)
First published 2009
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
without the prior permission in writing of Oxford University Press,
or as expressly permitted by law, or under terms agreed with the appropriate
reprographics rights organization. Enquiries concerning reproduction
outside the scope of the above should be sent to the Rights Department,
Oxford University Press, at the address above
You must not circulate this book in any other binding or cover
and you must impose the same condition on any acquirer
British Library Cataloguing in Publication Data
Data available
Library of Congress Cataloging in Publication Data
Data available
Typeset by SPI Publisher Services, Pondicherry, India
Printed in Great Britain
on acid-free paper by
Clays Ltd., St Ives plc

ISBN 978–0–19–923796–8

1 3 5 7 9 10 8 6 4 2
Shapes

T
HERE is beauty to be found in regularity: the same element
repeating again and again, typically with geometric order.
There is no better example than the honeycomb, a miracle of
hexagonal perfection. This sort of pattern is a lattice. On a leopard’s
pelt, the lattice melts but a pattern remains: spots spaced at more or
less equal distances but no longer in neat rows. There is a comparable
order in stripes and concentric bands: a succession rather strictly
enforced on the angelfish, but more loosely applied in the meandering,
merging stripes of the zebra or of sand ripples. The means by which
these natural patterns are constructed may tell us something about
how the far more complicated forms of animals and plants are created
by a progressive division and subdivision of space, orchestrated by
nothing more than simple physical forces.
This page intentionally left blank
Contents

Preface and acknowledgements ix

1: The Shapes of Things 1

Pattern and Form

2: Lessons of the Beehive 33

Building with Bubbles

3: Making Waves 103

Stripes in a Test Tube

4: Written on the Body 151

Hiding, Warning, and Mimicking

5: Rhythms of the Wild 200

Crystal Communities

6: How Does Your Garden Grow? 226

The Mathematics of a Daisy

7: Unfolding the Embryo 257

The Formation of Body Plans

Appendices 287

Bibliography 295

Index 303
This page intentionally left blank
Preface and
acknowledgements

A
F T E R my 1999 book The Self-Made Tapestry: Pattern Formation
in Nature went out of print, I’d often be contacted by would-be
readers asking where they could get hold of a copy. That was
how I discovered that copies were changing hands in the used-book
market for considerably more than the original cover price. While that
was gratifying in its way, I would far rather see the material accessible to
anyone who wanted it. So I approached Latha Menon at Oxford Uni-
versity Press to ask about a reprinting. But Latha had something more
substantial in mind, and that is how this new trilogy came into being.
Quite rightly, Latha perceived that the original Tapestry was neither
conceived nor packaged to the best advantage of the material. I hope
this format does it more justice.
The suggestion of partitioning the material between three volumes
sounded challenging at first, but once I saw how it might be done,
I realized that this offered a structure that could bring more thematic
organization to the topic. Each volume is self-contained and does not
depend on one having read the others, although there is inevitably
some cross-referencing. Anyone who has seen The Self-Made Tapestry
will find some familiar things here, but also plenty that is new. In
adding that material, I have benefited from the great generosity of
many scientists who have given images, reprints and suggestions.
I am particularly grateful to Sean Carroll, Iain Couzin, and Andrea
Rinaldo for critical readings of some of the new text. Latha set me
more work than I’d perhaps anticipated, but I remain deeply indebted
to her for her vision of what these books might become, and her
encouragement in making that happen.
Philip Ball
London, October 2007
This page intentionally left blank
The Shapes of Things
Pattern and Form
1

A
R RIV IN G on Earth, the aliens approach the first thing they see
and utter the familiar words: ‘take me to your leader’ (Fig. 1.1).
Like many jokes, this one offers a damning critique. It under-
mines the venerable and serious scientific quest to find life on other
worlds, exploding the question of ‘how would we know if we found it?’
by answering that we tend to imagine it will look like us.
Now, let me assure you that astrobiologists (as scientists who study
aliens are called nowadays) are not really that foolish. They do not
imagine for a moment that when we touch down on another inhabited
world, we will be greeted by envoys who look like Leonard Nimoy.
Indeed, if there is life in those parts of our own solar system that
seem at all habitable (such as the subsurface seas of Jupiter’s icy
moon Europa), it is most unlikely to warrant the description ‘intelli-
gent’. And we may have to look hard and long to find it, precisely
because we don’t know what we’re looking for. Yet even if we know it
is not going to be Dr Spock, we have trouble shaking the conviction that
it will look something like the forms of life we have seen before.

Fig. 1.1: Do we inevitably expect life to ‘look’ like us?

2 j NATURE’S PATTERNS: SHAPES

Even if it did, that already makes the challenge of identifying extra-

terrestrial life bad enough. Take a look at life on Earth today, and you’ll
see such a bewildering variety of shape and form that you could be
forgiven for imagining anything is possible (Fig. 1.2). But scientists have
a rather more sophisticated view of life (although they still cannot agree

Fig. 1.2: Living organisms on Earth come in a bewildering variety of shapes and sizes. (Photos: a,
carolsgalaxy; b, Keenan Pepper; c, Sarah Nichols; d, twoblueday; e, Ed Schipul; f, Doug Bowman.)
 THE SHAPES OF THINGS j 3

on a universal definition of it), which gives them hope of distinguishing

it from its inorganic context. They recognize some attributes of living
systems that go beyond mere physical appearance, such as the fact that
life tends to destroy the chemical equilibrium of its environment. I’ll
explain later what I mean by that, but let’s say for now that it’s rather
like watching a film in which all you can see are balls being juggled: you
know that there is something out of frame that is keeping them in
motion. It’s true that some geological and astrophysical processes
that don’t involve life at all can also induce this disequilibrium—but
nonetheless, searching for disequilibrium as a potential fingerprint of
life seems a lot better than looking around for a loitering humanoid
alien to whom you can say ‘Take me to your leader’.
Nevertheless, old habits die hard. Meteorite ALH84001 is a potato-
shaped lump of Mars that was blasted from the Red Planet a few billion
years ago in an asteroid or meteorite impact and subsequently found its
way through space to Earth. It was discovered in 1984 in the snows of
Antarctica. Scientists who made a detailed study of this cosmic intruder
claimed in 1996 that it contains a ‘possible relic’ of Martian life. In support
of that claim, an image was broadcast around the world that seemed to
show worms crawling across the mineral surface (Fig. 1.3). These ‘worms’

Fig. 1.3: These microscopic structures found in Martian meteorite ALH84001 have been interpreted as
evidence of ancient bacterial life. Might they be the fossilized remnants of tiny organisms? (Photo: NASA.)
4 j NATURE’S PATTERNS: SHAPES

were mineral too, and so tiny that they could only be seen in the electron
microscope; but the suggestion was that they could be the fossilized
remains of Martian bacteria that once infested this chunk of stone.
The researchers who investigated ALH84001 admitted that this con-
clusion was tentative, and they didn’t make it lightly. These wormy
forms were by no means the sole evidence—and after all, the scientists
acknowledged, they were much smaller than earthly bacteria tend to
be. All the same, these structures didn’t look like inorganic forms: it was
hard to explain them as microscopic rock features formed by physical
forces alone. And so the researchers stuck out their necks and used
shape, pattern, form—what scientists tend to call morphology—as the
partial basis for inferring a possible signature of life.
That doesn’t seem an unreasonable thing to do, does it? Surely, after all,
we can distinguish a crystal from a living creature, an insect from a rock?
Well, you might think so. But take a look at Fig. 1.4. At the top are the
shells of marine creatures called diatoms (which we will meet again
shortly). Below are microscopic mineral formations created in a test

Fig. 1.4: Can we tell

apart inorganic
structures formed by
chemistry alone and
those wrought by
biology? The shapes in a
are the shells of marine
micro-organisms called
diatoms; but while those
in b have a similar
complexity that seems
also to speak of the
agency of living
organisms, they are the
product of a purely
chemical process.
(Photos: a, Rex Lowe,
Bowling Green State
University, Ohio. b,
Geoffrey Ozin, University
of Toronto.)
 THE SHAPES OF THINGS j 5

Fig. 1.5: Chemistry alone can also produce microscopic surface

patterns rather similar to those seen in meteorite ALH84001 (Fig. 1.3).
(Photo: Geoffrey Ozin, University of Toronto.)

tube, entirely without the agency of life. Would you trust yourself to say
which is a ‘living’ form, and which is not? Now look at Fig. 1.5, in which
the microscopic patterns are a product of much the same chemical
process that made those in 1.4b. Does this remind you of anything?
What is it that encourages us to typecast some forms as those made
by life, and others as the products of the non-living world? A tree, a
rabbit, a spider have rather little in common when considered as mere
shapes—and yet we don’t hesitate to see them as examples of living
morphology. Why? Perhaps we sense a kind of purpose, of design, in
these forms? They are ‘complex’, certainly, but what does that mean?
They may have some regularity or symmetry—the bilateral symmetry
of the animals, the repeated branching of the tree—but that can’t be all
there is to it, for surely it is often in a high degree of regularity and
symmetry, in the onset of crystallinity, that we might imagine we
discern life’s absence. Even if we can’t say exactly what living form is,
we’d like to think we know it when we see it. But do we?
In the late 1990s, a group of NASA researchers decided this was a
problem that, like many others, is best left to a computer. They believed
that artificial intelligence would be more likely than we are to distin-
guish the living from the non-living, and so they hoped to ‘train’ com-
puters to recognize life from morphology alone, using all the examples
6 j NATURE’S PATTERNS: SHAPES

they could muster from our own planet. The machine would sense
and assimilate the subtle characteristics of living form, and would
then look for these signatures on missions to Mars or other potentially
life-supporting worlds. They intended this to be another of the compu-
tationally intensive projects like that which analyses radio signals for
signs of extraterrestrial intelligence,* in which volunteers’ home com-
puters analyse the data—in this case, running the computationally
intensive learning process—during spare time. This distributed system
would feed its output into the central brain of the operation, a com-
puter that the NASA researchers proposed to call the D’Arcy Machine.
Now that seems truly ironic, for the name is inspired by the man who,
to my mind, did more than anyone else to undermine the notion that life
has characteristic forms that distinguish it from non-living systems. This
man was the Scottish zoologist D’Arcy Wentworth Thompson, whose
classic book On Growth and Form, first published in 1917, provides the
first formal analysis of pattern and form in nature. Thompson’s book was
deeply erudite, beautifully written, and like nothing that had gone before.
It was also several decades ahead of its time—and this, coupled with its
undoubted idiosyncrasy, has meant that On Growth and Form never
sparked the emergence of ‘morphology’ as a scientific discipline. For a
long time, no one really knew what to make of Thompson’s tome.
On Growth and Form presents a challenge to the naive view that non-
living systems can produce only ‘simple’, often classically geometric,
shapes and forms—the prismatic shapes of crystals, say, or the sterile
ellipses of planetary orbits. Physics apparently teaches us that the basic
laws of nature are simple and symmetrical, and so it seems natural to
expect that their manifestations will share that characteristic. By the
same token, we tend to imagine that life is a complicated and infinitely
plastic influence, generating complex shapes that geometry struggles to
encompass or describe.
D’Arcy Thompson argued that, on the contrary, life’s forms may often
echo those of the inorganic world, and both of them may be rather simple,
or conversely, delightfully subtle and complex. Many of them, moreover,
have an undeniable beauty, whether that be the elegant, Platonic beauty
that we seem to perceive in symmetry and regularity or the dynamic,
organic beauty of living nature. The ambitions of the D’Arcy Machine
notwithstanding, life leaves no characteristic signature in natural form.

*This project, called SETI@Home, is an example of so-called distributed computing.

 THE SHAPES OF THINGS j 7

What makes Thompson’s book especially appealing is that there is

nothing at all recondite about the things he discusses. These forms are
ones we see all around us: the spirals of snail shells or in a sunflower
head, the baroque whorls of flowing water, the lacework of clouds, the
stripes of a zebra, the orderly perfection of a honeycomb. In these three
books I shall consider many other examples which Thompson barely
touched on, if at all, such as the designs of a butterfly wing, the
undulations of sand dunes, the branching of trees and rivers. Science
now possesses the tools and concepts to unravel the processes that
create these things, and it has vindicated Thompson’s approach of
searching for universal physical causes of pattern and order, offering
us a glimpse of a kind of natural harmony that pervades and structures
the whole world.
The puzzle of how these structures arise has solutions that are both
surprising and inspiring, and that foil our intuitions about how com-
plex form and pattern are made. Many of the most striking examples
that we encounter around us are evidently the products of human
hands and minds—they are patterns shaped with intelligence and
purpose, constructed by design. Brickwork tiling schemes, the hori-
zon-spanning stepped terraces of Asian rice fields, the monotonous
regularities of the built environment, the delicate traceries of micro-
electronic circuitry (Fig. 1.6)—all bear the mark of their human makers.
The unconscious message that we take away from all this artifice is that
patterning the world, shaping it into forms that please us or do useful
things, is hard work. It requires dedicated effort and skill. Each piece of
the picture must be painstakingly put into place, whether by us or, in
the wider world, by the forces of nature: the sparrow building its nest,
the individual plants weaving together into a hedgerow. This, we have
come to believe, is the way to create any complex form.
So when the natural philosophers of ages past found complexity in
nature, it is scarcely surprising that many of them decided they were
gazing on God’s handiwork and artistry. Most famously, the English
pastor William Paley argued in his book Natural Theology that the
contrivances found in the living world, in the forms of animals and
plants, are so exquisite that they cannot but be the products of a
guiding intelligence. The shabby tatters of this idea survive today in
the Intelligent Design movement, but Paley’s argument was in its time
much more defensible and indeed more coherent than that. For it was
not until the latter half of the nineteenth century that Darwin’s theory of
8 j NATURE’S PATTERNS: SHAPES

Fig. 1.6: Most of the

complex patterns that
we create are the
products of
painstaking labour,
each element of the
pattern having been
put in position ‘by
hand.’ Here I show a
wall pattern from a
Korean palace (a), a
paddy field in China
(b), an apartment
block in Benidorm (c)
and circuitry on a
microprocessor chip
(d). (Photos: a, Craig
Nagy; b, McKay
Savage; c, Ross
Goodman; d, Digital
Equipment
Corporation.)
 THE SHAPES OF THINGS j 9

evolution by random mutation and natural selection furnished an

explanation of how apparent ‘design’ might arise in nature without a
designer.
Contrary to the usual perception, Darwinism did not wholly answer
Paley, for the reverend also considered that the principles of astronomy
show signs of God’s wisdom, for example in the physical layout of the
solar system and the stability, simplicity and ponderous pace of plan-
etary orbits. His arguments here are not terribly convincing, although it
has to be acknowledged that modern cosmology is now even more of a
happy hunting ground for those who hope to discern evidence of God’s
design. But Paley could have availed himself of many suggestive ex-
amples of ‘design’ in nature that did not come from the living world,
and it is rather surprising that he did not. Anyone looking at the world
with Paley’s convictions about a Divine Architect would surely have to
conclude that He possesses an irrepressible urge to create beauty for
beauty’s sake. Why else, for instance, is the snowflake—a mere ice
crystal—wrought with such extravagance? Why are clouds dashed and
dotted across the sky in orderly ranks? And can it be mere coincidence
that we see the same forms and patterns again and again in places that
apparently have nothing in common, not even the characteristic of
being alive? Why should river networks resemble our veins and arteries,
or whirlpools look like galaxies, if these are not the motifs chosen by
God?
It is unfair, I know, to create a new argument for Paley only with the
intention of demolishing it. But that is not really my aim. To my mind,
the fact that these patterns can arise, and that they share certain
elements, quite without any need for a Great Patterner, is much more
remarkable and exciting than the notion that they are merely the
products of a cosmic artisan weaving nature’s tapestry. For that is just
what happens. We would not expect to be able to make patterns such as
those in Fig. 1.6 by, say, letting dyes spontaneously unmix into labyrin-
thine patterns, or trusting soil and pebbles to arrange themselves into
terraces ready to take water and seed. But equivalent things really do
happen, and in that way nature weaves its own designs. And what is
more, we find that some of nature’s patterns recur in quite different
situations, as though this tapestry is woven from an archetypal design
book, its themes echoing one another in different parts of the fabric. We
shall see that nature’s artistry is spontaneous, but is not arbitrary.
10 j NATURE’S PATTERNS: SHAPES

The boundaries of evolution

The quixotic flavour of On Growth and Form stems not only from the
fact that Thompson was attempting to explain many things for which
he lacked the right instruments. He was also taking a stand against what
he considered to be a stifling orthodoxy imposed by those who, at the
beginning of the twentieth century, sought explanations for form and
pattern in the living world. For although Darwinism obviated the need
for Paley’s natural theology, it seemed at times in danger of conducting
the same conjuring trick by another name. Instead of proclaiming
‘God’s work’ in response to every example of apparent design in nature,
there was a tendency to exclaim ‘evolution’s work!’ instead.
Darwin argued that, given enough time, small random changes in the
forms of organisms could carry them towards those shapes that were
best adapted to the demands that their environment made on them,
because the struggle for survival weeded out those changes that made
survival harder while conferring a reproductive advantage on those
individuals who, by sheer luck, acquired a beneficial mutation.
This means that, in biology, it is natural to expect that form follows
function: that the shape and structure of a biological entity (which
could be a molecule, a limb, an organism, even a whole colony) is
that which best equips the associated organism for survival. Biologists
are still divided about whether the selective pressure that dictates such
forms acts primarily at the level of the individual genes responsible for
that characteristic, or of the whole organism. But either way, the impli-
cation is that form is naturally selected from a palette of possibilities. A
form that confers evolutionary advantage tends to stick.
This is an extraordinarily powerful idea, and no serious biologist
doubts that it is basically correct in explaining how organisms evolve
and adapt over time. But as an explanation for the forms of life, it is not
entirely satisfying—not because it is wrong, but because it says nothing
about proximate mechanism, about causes that operate not over evo-
lutionary time but in the here and now, in the shaping of each individ-
ual organism. There is a tendency to imagine that science boasts an
answer to every question about the material world, but the truth is that
many questions can be given several different kinds of answer. It is like
asking how a car gets from London to Edinburgh. One answer might be
‘You get in, switch on the engine, and drive up the M1.’ That is not so
much an explanation as a narrative, and Darwinian evolution is a bit
 THE SHAPES OF THINGS j 11

like that—it provides a narrative that rationalizes how we got to where

we are today. (As a result, it says rather little about where we might go in
the future.) A chemical engineer might offer a different scenario: the car
got to Edinburgh because the chemical energy of the petrol was con-
verted to kinetic energy of the vehicle, along with a fair amount of heat
and acoustic energy. That is a correct answer too, but is perhaps a little
vague and abstract for some tastes. Why did the car’s wheels go round?
To answer this we must go instead to the mechanical engineer, who
explains how they are connected to the engine via a crankshaft . . . and
before long you are getting into an account of the mechanics of the
internal combustion engine.
So what, we might ask, are the mechanics that create biological form?
The standard answer, from what D’Arcy Thompson might have called
the adaptionists, is that we must reason a posteriori, seeking to under-
stand what we observe in terms of its functional efficiency and thus its
adaptive value. He had no objection to this in principle, but com-
plained that the Darwinists don’t follow through with that conviction:
they are happy to assume that a particular form must be the most
efficient one, and do not bother (or do not know how) to demonstrate
that this is so. For that reason, Thompson said, the Darwinian morph-
ologist is apt to forget about the mechanics altogether, and to become
obsessed with comparing this or that feature of different organisms
without thinking how their shape is determined by their mechanical
function in the organism as a whole. He argued that it was absurd to
think, as comparative anatomists did, that individual bones are separ-
ately moulded by evolutionary forces, when what matters is how the
skeleton as a whole functions as an efficient mechanical structure—
something that can be understood using the same principles as those
used by engineers to design bridges.
But this was simply an accusation that Darwinian morphologists
were apt to lose the plot. The bulk of On Growth and Form was devoted
to a much more fundamental challenge to the Darwinists’ ‘principle of
heredity’. Thompson’s contemporary zoologists would not have
objected (at least, one hopes not) to the idea of invoking physics and
mechanics to explain biological form, using them to rationalize it in
terms of efficiency and thus adaptive advantage. But this assumes that
there is a shape or structure that can be identified as the ‘best’ within a
particular environment, and that biology can always find a way of
making it. Taken to the naive extreme, this view holds that everything
12 j NATURE’S PATTERNS: SHAPES

is possible: that nature has at its disposal an infinite palette, and that it
dabbles at random with the options, occasionally (oh, so rarely!) hitting
on a winning formula with which it then tinkers to make minor vari-
ations on a theme. The torpedo-and-fins theme works for fish, say, and
the four-legs-and-muscle design is just the ticket for land predators.
Given the diversity of living forms evident today (Fig. 1.2), which is
after all only a fraction of that which has been exhibited over geological
time, this assumption of an infinite palette is understandable. But as
D’Arcy Thompson reminded his readers, ‘Cell and tissue, shell and
bone, leaf and flower, are so many portions of matter, and it is in
obedience to the laws of physics that their particles have been moved,
moulded and conformed.’ Evolution operates within physical con-
straints which insist that not everything really is possible. Are the zebra’s
stripes really the ‘best’ form of camouflage, or just the best that nature
can come up with given the limitations imposed by physical law?
That might seem a minor quibble, since if the whole of nature
operates within the same constraints then all we’re apparently saying
is that Darwinian evolution is a matter of finding the most advanta-
geous forms out of those available. But by insisting on those limits,
D’Arcy Thompson brought to the fore the issue of exactly how such
forms come about through the action of physical forces. It wasn’t just a
question of ensuring that evolutionary biology obeys physical and
chemical laws; he felt that these laws play a direct, causative role in
determining shape and form in biology. Thus he insisted that there
were many forms in the natural world that one could, and indeed
should, explain not by arguing that evolution has shaped the material
that way, but as a direct consequence of the conditions of growth or the
forces in the environment.
What, he felt, could be more unnecessary than invoking millions of
years of selective fine-tuning to explain the curving shape of a horn or
shell when one could invoke a mathematically simple growth law,
based on proximate, physical causes, to account for it? The sabre-like
sweep of an ibex horn need not have been selected from a presumed
gallery of bizarre and ornate alternative horn shapes. We can assume
merely that the horn grows at a progressively slower rate from one side
of the circumference to the other, whereupon, hey presto, you have an
arc. In this case, then, an evolutionary argument is redundant, or at best
ancillary, because the forms of horns are inevitable. Either a horn grows
at the same rate all around its circumference, in which case it becomes
 THE SHAPES OF THINGS j 13

a straight cone, or there is an imbalance from one side to the other,

giving a curved cone. It just did not make sense to invoke other shapes:
nature’s palette contains just these two. Even the more elaborate spiral
form of a ram’s horn (Fig. 1.7) comes simply from ramping up the
asymmetry of growth rates, causing the horn’s tip to swing through
several complete revolutions. By the same token, biological forms such
as the shapes of amoeba can no more be regarded as ‘selected for’ than
can the spherical form of a water droplet; rather, they are dictated by
physical and chemical forces.
With Darwinism in its first flush, banishing to the dustbin of pseudo-
science the teleology of Paley, it is easy to see how Thompson’s ideas,
with their apparent exhortation of biological predestination governed
by physical law, looked dangerously close to heresy. Thompson was
conscious of that, admitting that On Growth and Form sometimes
‘undoubtedly runs counter to conventional Darwinism.’ Where that
was so, he said, ‘I do not rub this in, but leave the reader to draw the
obvious morals for himself.’
They did, and often unfavourably. At the University College of Dun-
dee in Scotland, Thompson felt marginalized and neglected, and found
himself in his sixth decade without having published anything of note.
When he dared suggest, as he did in 1894 at the meeting of the British

Fig. 1.7: Many animal horns, such as that of this Dall’s sheep, have
spiral shapes that can be explained with a simple growth law.
(Photo: Brian Uhreen.)
14 j NATURE’S PATTERNS: SHAPES

Association for the Advancement of Science, that there were ‘some

difficulties with Darwinism’, he was met with the response that, as he
put it, ‘there were no difficulties in Darwinism . . . to any sensible man in
those days.’ Nonetheless, in the 1910s he began to draw his ideas
together in what he intended to be a ‘little book’. When it appeared in
1917 it was anything but that; yet the formidable On Growth and Form
met with a largely positive reception, and it secured Thompson’s repu-
tation at last. But plans for a revised edition in the 1920s descended into
farce as Thompson failed repeatedly to meet his publishers’ deadlines,
and the second edition, an immense tome that had to be split in two,
did not appear until 1942. In some ways that delay served him poorly,
for the book remained an emphatically Victorian piece of scholarship.
It is illuminating to contrast it with the physicist Erwin Schrödinger’s
groundbreaking book What is Life?, published in 1944—an altogether
more forward-looking work that some regard as presaging the science
of molecular genetics and the notion of biology as an information
science. Thompson’s wartime edition, in contrast, gave the impression
that little had changed in the three decades separating it from its
predecessor.
And it is true that some of On Growth and Form has aged poorly,
while some is plainly wrong. All the same, the book’s central message
remains relevant, and in its breadth, audacity and ambition it continues
to inspire in generations of scientists (and others too) a sense of awe
and wonder about the natural world. Not the least of its virtues is that it
is so exquisitely written; the English biologist Peter Medawar has called
it ‘beyond comparison the finest work of literature in all the annals of
science that have been recorded in the English tongue’. On Growth and
Form provided the first glimpse of the landscape that we shall explore
in this series. Today we can see the geography considerably more
clearly, for we have the tools to map it and to interpret what we see.
But we are latecomers; D’Arcy Thompson was the pioneer.

The genetic black box

D’Arcy Thompson’s thesis, then, was that biology cannot afford to neglect
physics, and in particular that branch of it that deals with the mechanics
of matter. (He was far less concerned with chemistry, the other pillar of
the physical sciences, seemingly because he did not consider it to be
 THE SHAPES OF THINGS j 15

sufficiently mathematical. Today there is much in the field of chemistry

that would have served him well, as we shall see.) His complaint was
against the dogma of selective forces as the omnipotent answer to bio-
logical questions. For him, this did not address the question of cause, but
merely relocated it. A physicist, on the other hand, ‘finds ‘‘causes’’ in what
he has learned to recognize as fundamental properties . . . or unchanging
laws, of matter and of energy’.
Thompson seemed to consider that this neglect of physics in biology
produced an almost vitalistic streak in his contemporary Darwinists.
‘The zoologist or morphologist’, he wrote,

has been slow. . . to invoke the aid of the physical or mathematical sciences; and
the reasons for this difference lie deep, and are partly rooted in old tradition and
partly in the diverse minds and temperaments of men. To treat the living body as
a mechanism was repugnant, and seemed even ludicrous, to Pascal; and Goethe,
lover of nature as he was, ruled mathematics out of place in natural history. Even
now the zoologist has scarce began to dream of defining in mathematical
language even the simplest organic forms. When he meets with a simple geo-
metrical construction, for instance in the honeycomb, he would fain refer it to
psychical instinct, or to skill an ingenuity, rather than to the operation of
physical forces or mathematical laws; when he sees in snail, or nautilus, or tiny
foraminiferal or radiolarian shell a close approach to sphere or spiral, he is prone
of old habit to believe that after all it is something more than a spiral or a sphere,
and that in this ‘something more’ there lies what neither mathematics nor
physics can explain. In short, he is deeply reluctant to compare the living with
the dead, or to explain by geometry or by mechanics the things which have their
part in the mystery of life. Moreover he is little inclined to feel the need of such
explanations, or of such extension of his field of thought.

Strange that this was written nearly a century ago, for I have a feeling
that I’ve met this ‘morphologist’ and spent some time in frustrated
debate with him. He has a point: life’s mechanisms are seldom simple,
and (contrary to common belief) biology does not always do things by
the most economical means imaginable, but can become encumbered
with the legacy of history, on which physics must remain silent. More-
over, physicists can be as arrogant as biologists are stubborn. But a
tradition that has rendered ‘theoretical biology’ almost a term of abuse
does not seem likely to be very productive of theories that explain
(rather than describe) its subject.
Of course, today’s biologists have a more sophisticated answer to
questions of form than the magic word ‘adaptation’. They call on gen-
etics, the ‘microscopic’ basis of Darwinism. It is tempting to imagine,
16 j NATURE’S PATTERNS: SHAPES

from what has been said and written about genes, that they are where all
biological questions end. One hears about genes responsible for this
or that illness or trait or feature—for cancer, for intelligence, for the
development of flies’ wings or of blue eyes. The climate of the culture in
molecular biology (if not the expressed belief of all its practitioners) is
that, by understanding the roles of genes and the mutual interactions of
the protein molecules they encode, we will understand life.
That attitude underpinned the Human Genome Project, the inter-
national effort to map out every one of the 30,000 or so genes in the 23
chromosome pairs of the human cell. The first draft of this map was
completed in 2000 (most of the holes have been filled in subsequently),
and to judge from some of the hyperbole it elicited, you would think
that it has provided us with a complete instruction manual for the
human body. But it does not do that at all. The Human Genome Project
has created a bank of genetic data that is sure to be of immense medical
value, and which contains a great deal of information about how our
cells work. But for biological questions that have a genetic component
(and not all of them do), the respective genes are just the beginning of
an answer. Most of these genes encode the chemical structures, and
thus the chemical functions, of proteins. The issue is how the produc-
tion (or absence) of a particular protein affects the network of biochem-
ical processes in the cell, and how this gives rise to the particular
physiological consequences that we are studying. Identifying the gene
‘responsible’ for this or that trait is like discovering which door we need
to go through in order to enter this network and find out where it leads.
(And most answers can be accessed through several doors.)
Elucidating precisely how genes work is the really hard part of the
matter, which is why so much of genetics operates at the ‘black box’
level: we know that the presence or lack of a gene in the genome is
linked to a certain manifestation at the level of the whole organism, but
we do not know why. In the same way, our computers are blank boxes
(available in a range of colours more tasteful than black) that we know
will respond in certain ways when poked, even though most of us have
not the faintest idea of why this happens.
But in any event, organisms are not just genes and the proteins made
from them. There is all kinds of other stuff in the cell: sugars, fatty acids,
hormones, small inorganic molecules like oxygen and nitric oxide,
salts, and minerals such as those in bone and tooth. None of these
substances is encoded in genes (that is, in the structure of our DNA),
 THE SHAPES OF THINGS j 17

and you would never guess, by looking at the genome, that they were
required at all, let alone what roles they play. And yet these substances
tend to be highly organized and orchestrated in their interactions and
their structures at the level of the cell (and at larger scales too). Where
does that structure come from? Proteins often play a role in building it,
but so too do physical forces, such as surface tension, electrical attrac-
tion, fluid viscosity. Gene-hunting tells us nothing about such things.
In short, questions in biology of a ‘how?’ nature need more than
genetics—and frequently more than a reductionist approach. If nature
is at all economical (and there is good reason to suppose that is often
the case, though not invariably so), we can expect that she will choose
to create at least some complex forms not by laborious piece-by-piece
construction but by harnessing some of the organizational and pattern-
forming phenomena we see in the non-living world. Evolution, via
genetics, can exploit, tame and tune such phenomena; but it does not
necessarily generate them. If this is so, we can expect to see similarities
in the forms and patterns of living and purely inorganic systems, and to
be able to explain them both in the same manner.
I should add a cautionary note. It is true that biology has been
somewhat resistant to the idea that crude, general physical principles
might sometimes be capable on their own of explaining aspects of
biological form. To biologists, this seems too risky and uncontrollable,
like driving a car with no hands on the wheel and hoping that friction
and air resistance will somehow conspire to guide the vehicle along the
winding road. But one can go too far to the other extreme. It is popular
in some circles to denounce the so-called reductionist science of mo-
lecular biology and imply instead that the universe is somehow imbued
with a creative potential that operates in a ‘holistic’ way. The fad for the
notion of ‘complexity’, which shows that sophisticated forms and pat-
terns may emerge spontaneously from a miasma of interactions, may
sometimes veer towards a kind of neo-vitalism in the way that it
invokes a cosmic creativity. Worst of all is the tendency to make moral
distinctions, so that ‘holistic science’ becomes good and ‘reductionist
science’ meretricious. While I applaud a perspective that broadens the
horizons of ‘black-box’ biology and argues for a role of spontaneous
pattern formation in the living world, there is no getting away from the
fact that most of biology, particularly at the molecular level, is hid-
eously complicated. In distinction from complex, this means that the
details really do matter: leave out one part of the chain of events, and
18 j NATURE’S PATTERNS: SHAPES

the whole thing grinds to a halt. In such cases, one gains rather than
loses understanding by turning up the magnifying power of the micro-
scope. Until we get reductionistic about, say, the body’s immune re-
sponse, we won’t know much about it, let alone develop the potential to
tackle pathological dysfunctions such as AIDS. A reductionist view
won’t necessarily provide an explanation of how it works, but without
it we might not know quite what needs to be explained. Reductionism
can be aesthetically unattractive, I know, but it is wonderfully useful.

What is pattern and what is form?

These concepts are, after all, my topics, and yet I am afraid that I cannot
offer a rigorous definition of either, nor make a rigid distinction be-
tween them. If it makes you feel any better, remember that neither can
scientists offer a rigorous definition of life. (They have tried often
enough, but the very attempt is ill-advised, for the word is colloquial
rather than scientific. You might as well try to define the word ‘love’.)
What is clear is that ‘pattern’ is a supremely plastic word, and evi-
dently it implies quite different things to, say, a psychologist and to a
wallpaper designer. My definition, such as it is, is much closer to the
latter than the former. The best I can do is to say that a pattern is a form
in which particular features recur recognizably and regularly, if not
identically or symmetrically. And while I shall occasionally mention
patterns of a temporal nature—events that repeat more or less regu-
larly, such as the beating of a heart—on the whole I shall be talking in
spatial terms, and so the image of a pattern on wallpaper or a carpet is a
useful one to bear in mind. In those cases, however, the repeating units
are generally identical. My concept of pattern will not necessarily be so
demanding. The repeating elements may be similar but not identical,
and they will repeat in a way that could be called regular without
following a perfect symmetry. Yes, I know it’s vague—but I believe we
usually know such patterns when we see them. One such is made up
from the ripples of sand on a wave-lapped beach or in a windswept
desert (Fig. 1.8). No two of these ripples are identical, and they are not
positioned at exactly repeating intervals. But we can see at once that
there are elementary units (ripples) that recur throughout space. We see
the pattern. Indeed, we are remarkably good at seeing the pattern,
which, because it is not mathematically perfect, is typically harder for
 THE SHAPES OF THINGS j 19

Fig. 1.8: Ripples in sand clearly constitute a repetitive pattern even though no two parts of the pattern
elements are identical. (Photo: Nick Lancaster, Desert Research Institute, Nevada.)

a machine to recognize. It has become a cliché to say that the human

brain is a pattern-recognizing instrument while the electronic com-
puter is a data-crunching instrument; but like most clichés, it has
taken hold for good reason. Sand ripples are a relatively simple pattern,
but I think we can discern something pattern-like too even in rather
irregular structures, such as the peaks and valleys of a mountain range
or the skeleton of a tree in winter.
Patterns, then, are created from groups of features. Form is a more
individual affair. I would define it loosely as the characteristic shape of a
class of objects. Just as our brains allow us to organize a field of similar
shapes into a pattern, so they are adept at somehow discerning common-
alities of form between diverse objects—although we find it equally hard
to explain exactly why. Objects with the same form need not be identical,
or even similar in size; they simply have to share certain features that we
can recognize as typical, even stereotypical. The shells of sea creatures are
like this: those of organisms of the same species tend to be identifiably
akin even to the untrained eye. The same is true of flowers and of the
shapes of mineral crystals. You might say that the ‘form’ of these objects is
a rather Platonic concept—that which remains after we have averaged
away all the slight variations between individuals.
Patterns, then are typically extended in space—they go on and on—
while forms are bounded and finite. But take this as a guideline, not a rule.
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