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Synthese Library 388
Studies in Epistemology, Logic, Methodology,
and Philosophy of Science

Marcos Silva Editor

How Colours
Matter to
Philosophy
Synthese Library

Studies in Epistemology, Logic, Methodology,

and Philosophy of Science

Volume 388

Editor-in-Chief
Otávio Bueno, University of Miami, Department of Philosophy, USA

Editors
Berit Brogaard, University of Miami, USA
Anjan Chakravartty, University of Notre Dame, USA
Steven French, University of Leeds, UK
Catarina Dutilh Novaes, University of Groningen, The Netherlands
The aim of Synthese Library is to provide a forum for the best current work in
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together by an extensive editorial introduction or set of introductions if the volume
is divided into parts. An extensive bibliography and index are mandatory.

More information about this series at http://www.springer.com/series/6607

Marcos Silva
Editor

How Colours Matter

to Philosophy

123
Editor
Marcos Silva
Federal University of Alagoas
Maceió, Alagoas, Brazil

Synthese Library
ISBN 978-3-319-67397-4 ISBN 978-3-319-67398-1 (eBook)
DOI 10.1007/978-3-319-67398-1

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Endorsements

Questions about the nature of colour – how and what we know about them; how we
experience them; how they fit into nature – grip the philosophical imagination
as much today as they have done down the ages. This outstanding volume
collects together scholars with a deep understanding of the history of longstanding
epistemological and metaphysical puzzles and the history of debates associated
with colour. Yet it does much more, its addresses a gap in the market, by revealing
how learning lessons from both the analytic and continental philosophical traditions
can bear fruit in our thinking about colour. More than that, the collection showcases
how new research about the nature of colour figures can inform and be informed
by new thinking about language, mind, phenomenology, aesthetics, logic and
mathematics. Its chapters make clear why questions they examine matter to those
working in fields and disciplines outside of philosophy. In achieving all of this,
this book beautifully prepares the ground for the next steps in our research on and
philosophising about colour.
Daniel D. Hutto is Professor of Philosophical Psychology at the University of
Wollongong, Australia

It is not an overstatement to say that How Colours Matter to Philosophy is a ground-

breaking publication. It is the first edited volume on the philosophy of colour to
reach far beyond the core topics of metaphysics, epistemology and naturalistic
philosophy of mind, bringing within its purview logic, philosophy of mathematics
and aesthetics. There are also important contributions to the literature on relatively
under-explored historical topics, such as Wittgenstein’s theory of colour and
Goethe’s philosophy of the scientific method. The entries are methodologically
sophisticated and philosophically rewarding, and I expect that many of them will be
the starting point for future scholarly debates. The volume is essential reading for
advanced students and researchers seeking new perspectives on a perennial issue in
systematic philosophy.
Mazviita Chirimuuta is Associate Professor, History & Philosophy of Science at
the University of Pittsburgh, USA

v
vi Endorsements

The book assembles an impressive variety of contributions from different theoret-

ical approaches and philosophical traditions. The puzzling nature of colours, its
central common theme, unites chapters touching on a great number of domains
(philosophy of mind, philosophy of language, metaphysics, logic, philosophy of
science and history of philosophy). This diverse collection has the potential to
seduce readers into getting to know research areas they would otherwise have
ignored. It contains a number of rigorously argued and yet accessible original
papers. Anyone interested in philosophical issues about colour will find it highly
stimulating.
Martine Nida-Rümelin is Philosophy Professor at the Université de Fribourg,
Switzerland

Colour has intrigued and puzzled scientists and philosophers since antiquity.
Nowadays, the philosophy of colour is a highly active subfield in analytic phi-
losophy. While acknowledging the contemporary discussions and often directly
contributing to them, this collection of papers aims to also approach the philos-
ophy of colour from less well-travelled paths. Thus detailed attention is paid to
the history of thinking about colour, to phenomenology and mathematics, and
not all of this book’s authors take naturalism, or the standard forms of it, for
granted. The high-quality papers included in this anthology succeed admirably
in enriching current philosophical thinking about colour. The reader will find
refreshing treatments of familiar problems and will be guided to neglected and
novel philosophical questions that the elusive phenomenon of colour continues
to pose.
Erik Myin is Professor of Philosophy at the University of Antwerp, Belgium

This is certainly the most complete collection of philosophical essays on colours

ever published. Those interested in the history of the problem since antiquity or in
the famous debate opposing Newton and Goethe will find here new interpretations.
But there is much more. Colours can be approached from many different points
of view: logico-linguistic, phenomenological, aesthetic or metaphysical. It is hard
to believe that all these approaches can find their place in one single book. This
is Marcos Silva’s feat: for years, he worked hard and patiently to gather all
these essays and organized them by topics. The result is astonishing: what makes
our sensorial experiences so rich and lively is here, at least in part, deciphered.
If not all the philosophical enigmas associated with colours are resolved, this
collection of original and up-to-date essays is no doubt an important step in the
right direction.
André Leclerc is Professor of Philosophy at the University of Brasília, Brazil

This outstanding volume merges eighteen excellent paradigms of showing how

colours and their expressions matter to philosophy. Transcending traditional bound-
aries authors from twelve countries combine aspects of metaphysics, history
of philosophy, epistemology, philosophy of mind, aesthetics, psychology, lin-
guistics, logic and mathematics in a very inspiring manner. Anyone who is
interested in philosophical investigations on colours can learn a lot starting with
Endorsements vii

the ancient Greeks, continuing with Goethe, Newton, Husserl, Katz, Bühler,
Heidegger, Wittgenstein and ending up with advanced studies in the contexts
of vagueness, cognition and the four-colour theorem. All in all, this collection
represents a new milestone in the ongoing philosophical debate on colours and
colour expressions.
Ingolf Max is Professor of Analytic Philosophy at the University of Leipzig,
Germany
Contents

Part I History of Philosophy

Dispositionalism: Democritus and Colours by Convention. . . . . . . . . . . . . . . . . . 3
Barry Maund
Hue, Brightness & Saturation in Classical Greek Chroma Terms . . . . . . . . . 25
Ekai Txapartegi
How Many Colours? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Kirsten Walsh
Goethe contra Newton on Colours, Light, and the Philosophy
of Science . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Olaf L. Müller
On Color: The Husserlian Material a Priori . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Jairo José da Silva
Impossible Colours: Wittgenstein and the Naturalist’s Challenge . . . . . . . . . 107
Andrew Lugg

Part II Phenomenology and Philosophy of Mind

Colours – Wittgenstein vs (Katz & Bühler) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Kevin Mulligan
What the Mind-Independence of Color Requires . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Peter W. Ross
Explaining Colour Phenomenology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Nicholas Unwin
Dasein Is the Animal That Sorts Out Colors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
Bernardo Ainbinder

ix
x Contents

Subjectivity and Normativity in Colour-Distinctions . . . . . . . . . . . . . . . . . . . . . . . . 195

Pirmin Stekeler-Weithofer
Colors: Presentation and Representation in the Fine Arts . . . . . . . . . . . . . . . . . . 215
Otávio Bueno

Part III Philosophy of Language and Philosophy of Logic

Things Are Not What They Seem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Graham Priest
Vagueness, Hysteresis, and the Instability of Color . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Diana Raffman
Logic and Colour in Cognition, Logic and Philosophy . . . . . . . . . . . . . . . . . . . . . . 249
Dany Jaspers
A Chromatic Hexagon of Psychic Dispositions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273
Jean-Yves Beziau
Proofs Versus Experiments: Wittgensteinian Themes Surrounding
the Four-Color Theorem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289
Gisele Dalva Secco and Luiz Carlos Pereira
The Wonder of Colors and the Principle of Ariadne . . . . . . . . . . . . . . . . . . . . . . . . . 309
Walter Carnielli and Carlos di Prisco

Name Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 319

Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321

Contributors

Editor

Marcos Silva Federal University of Alagoas, Maceió, Alagoas, Brazil

Authors

Bernardo Ainbinder Instituto de Humanidades, Universidad Diego Portales, San-

tiago, Chile
Jean-Yves Beziau Federal University of Rio de Janeiro, Rio de Janeiro, Brazil
Otávio Bueno Department of Philosophy, University of Miami, Coral Gables, FL,
USA
Walter Carnielli Department of Philosophy, Centre for Logic, Epistemology and
the History of Science, State University of Campinas UNICAMP, Campinas, SP,
Brazil
Jairo José da Silva Researcher, CNPq, MCT, Brazil
Carlos di Prisco Instituto Venezolano de Investigaciones Científicas, Universidad
de Los Andes, Bogotá, Colombia
Dany Jaspers KU Leuven campus Brussels CRISSP, Brussels, Belgium
Andrew Lugg University of Ottawa, Ottawa, Canada
Barry Maund University of Western Australia, Crawley, Australia
Olaf L. Müller Institute for Philosophy, Humboldt University, Berlin, Germany
Kevin Mulligan University of Italian Switzerland, Lugano, Switzerland
University of Geneva, Geneva, Switzerland
xi
xii Contributors

Luiz Carlos Pereira Pontifical Catholic University of Rio de Janeiro, Rio de

Janeiro, Brazil
Graham Priest Departments of Philosophy, CUNY Graduate Center, New York,
NY, USA
The University of Melbourne, Melbourne, VIC, Australia
Diana Raffman University of Toronto, Toronto, ON, Canada
Peter W. Ross California State Polytechnic University, Pomona, CA, USA
Gisele Dalva Secco Federal University of Rio Grande do Sul, Porto Alegre, Brazil
Pirmin Stekeler-Weithofer University of Leipzig, Leipzig, Germany
Ekai Txapartegi University of Basque Country (UPV-EHU), Lejona, Spain
Nicholas Unwin Lancaster University, Lancaster, UK
Kirsten Walsh Philosophy Department, University of Nottingham, Nottingham,
UK
Introduction to “How Colours Matter to
Philosophy”

Colours induce us to philosophize. Colours pose problems and challenges to theories

of perception, of rule-following, and classical principles of logic; they serve as
illustrations of harmonic and holistic systems; they motivate phenomenological
arguments and modal systems of incompatibility. Colours encourage intricate
models in linguistics and mathematics, and they also represent common ground for
the Gestalt tradition as well as puzzles for some central accounts in the philosophy
of mind. Furthermore, colours are a favourite example in many central philosophical
arguments where they are used systematically in support of some theses and
as counterexamples in refutation of other theories. Indeed, discussions about the
nature of colours reside at the core of many classic disputes in metaphysics and
epistemology, such as those between Locke and Leibniz, Newton and Goethe as
well as Wittgenstein and himself. This book examines and discusses different and
seminal ways in which colours matter to a range of theorists, including philosophers.
The contributions in this volume advance several discussions in different areas of
contemporary philosophy, which represent a comprehensive platform for further
studies and collaboration programmes.
The present volume provides an international forum for philosophers from 12
different countries to interact across old disciplinary boundaries. Significantly, this
compilation approaches work on colour in a different way from previous books
such as Readings on Color edited by Byrne and Hilbert (1996) and Colors for
Philosophers authored by C. L. Hardin (1988). Both of these seminal works
are focused primarily in the naturalist tradition of analytic philosophy, surveying
colour science to rely upon empirical research and conceptualizing philosoph-
ical questions as continuous with enquiries in the natural sciences. This is a
legitimate and fruitful approach; however, neither of these works fully addresses
the breadth and depth of philosophical problems that have been reflected in
thinking about colours. How colours matter to philosophy brings these philosophical
problems to a broader audience by dealing with language, mind, phenomenology,
aesthetics, logic and mathematics, in addition to the traditional epistemological

xiii
xiv Introduction to “How Colours Matter to Philosophy”

and metaphysical problems of subjectivity and objectivity. Several contributions

focus chiefly upon highlighting the interrelations between analytic and continental
philosophical traditions.
Three significant innovations provided by the volume overall are noteworthy:
Firstly, the collection delivers a historically informed discussion of colours in the
Western philosophical canon. Secondly the collection offers to bridge continental
and analytic philosophy, whereby the extant literature based on the naturalistic
tradition is examined given the grounding of a conscientious historical perspective.
Finally, this volume presents discussions of the logical and linguistic aspects of
colours from various philosophical points of view. Overall, the volume stands as an
extensive resource for anyone doing philosophical work on colour.
Those focussing on the history of ideas more broadly construed may also be
interested in this book, as colours challenge important philosophical theses popular
in different eras. Scientists and psychologists looking for theoretical background for
their findings may also find the chapters absorbing. Accordingly, the audience for
this manuscript is not restricted to standard philosophy audiences but will attract
people working in aesthetics, phenomenology, conceptual semantics and logical
syntax, for inspiration will surely continue to be found in the eccentricities that
colours offer.
The book is divided into three parts, each containing six papers: (i) history of
philosophy, (ii) phenomenology and philosophy of mind and (iii) philosophy of
language and philosophy of logic.
Barry Maund’s contribution opens up the first part, devoted to the history of
philosophical colour problems. Maund supports the claim that dispositionalism
and eliminativism are actually compatible, by showing that the position of the
ancient Greek philosopher Democritus is more complex than is usually recognized.
Maund maintains that that there is a way of understanding Democritus’ position that
comprises a special form of dispositionalism, one that is compatible with one form
of eliminativism, which can be thought of as colour fictionalism. Ekai Txapartegi,
in his paper, defends Plato’s keen awareness of the distinction between saturation,
brightness and hue. Drawing on cross-language colour naming systems, Txapartegi
explains why classical Greek chroma terms might have referred to hue regions of
the colour space.
In her contribution, Kirsten Walsh explores Newton’s mathematico-experimental
methodology, which eschews ‘hypotheses’ in favour of ‘theories’. Concerning the
number and division of colours, Walsh investigates why Newton argued that the
number of different original colours was indefinite, in spite of an allegedly onto-
logical inflation. Olaf Mueller’s paper advocates Goethe’s objections to Newton’s
theory of light and colour as typically underestimated in the literature. According to
Mueller, by insisting that the step to theory is not forced upon us by the phenomena,
Goethe revealed our own free, creative contribution to theory construction. Thus, he
suggests that Goethe explored the problem of theory underdetermination a century
before Quine made Duhem’s arguments famous.
In his chapter, Jairo da Silva discusses Husserl’s concept of material a priori truth,
particularly with respect to colour concepts. Da Silva exposes misunderstandings
Introduction to “How Colours Matter to Philosophy” xv

and misconceptions in Schlick’s criticism of Husserl’s notion (as advanced in

Schlick’s ‘Is There a Material A Priori?’). Andrew Lugg’s contribution makes
a case for philosophy as conceptual analysis, against the background of the rise
of naturalistic philosophy. He then draws on Ludwig Wittgenstein’s account of
‘reddish green’ and ‘transparent white’ as grammatically excluded. Lugg defends
that while Wittgenstein’s view of such impossible colours is open to question, so
too is the naturalist’s alternative.
Part II of the book opens with Kevin Mulligan’s paper on phenomenological
issues concerning complex interactions between colour perception and light. Mulli-
gan shows that Wittgenstein’s everyday explorations of colours in ordinary light are
intended to illustrate his view that the world of colours is very complex and displays
no system, at least none rooted in the nature of colours themselves. Mulligan
compares Wittgenstein’s remarks with the psychological account of colour by David
and Buehler, defending their approach to solving puzzles concerning optics.
The paper which follows by Peter Ross explores an ‘argument from structure’
against the claim that colours are primary qualities. Ross addresses the problem
about the ordering of quality spaces as explained by mental processing which
conflicts with the standard assumption that primary qualities are mind independent.
Using colour perception as a leitmotif, he avoids this conflict by way of a model for
visual spatial perception according to which the ordering of determinates is in part
explained by mental processing while at the same time the determinate qualities are
mind independent.
Nicholas Unwin argues, in his contribution, that the hypothesis of inverted
colour qualia is of philosophical and scientific importance by attempting to colour
phenomenology. He argues against reductive explanations of colour phenomena,
preferring to show that we should instead try to show that colour hue inversions are
as problematic as other qualia inversions; this, he says, yields a more modest sort of
explanation which does not imply physicalism.
Bernardo Ainbinder deals with Heidegger’s conception of experience as a
normative conceptual one. Against this background, he claims that Heidegger’s
description of Umsicht, i.e. the way of coping with the world as a meaningful
whole, is not limited to typical forms of practice explicitly mentioned by Heidegger,
but should be extended to analyse other basic cases such as colour perception.
Ainbinder maintains that colour perception therefore illustrates the sense in which
conceptualism is true.
Stekeler-Weithofer holds that not only Kant’s Ding an sich but also subjec-
tive qualia must be understood as merely limiting concepts (Grenzbegriffe). He
maintains that colour words express plastic contrasts by which we (in many cases
successfully) split up a manifold and continuum of colour experiences into ‘discrete’
colours of surfaces. According to Stekeler-Weithofer, assertions about colours
are derived from and dependent upon a generic system of relations and modal
inferences. If this is true, then a merely classificatory understanding of colours as
one-place predicates or concepts is misguided.
Otavio Bueno closes the second part of the volume by offering a framework to
examine colours in the fine arts. He argues that despite the relatively uniform phe-
xvi Introduction to “How Colours Matter to Philosophy”

nomenology of colours, their meanings vary dramatically in painting, photographs

and films. According to Bueno’s account, this makes colours behave in far more
conventional ways than one might initially suppose (given our relatively stable
colour phenomenology) – that is, colours are represented to us via conventional
codes rather than via recognitional prompts.
Part III of the book is opened by the logician Graham Priest addressing some
solutions to the sorites paradox by defending a dialetheic solution. On this approach,
statements concerning a borderline area of a sorites sequence are both true and
false, whereby modus ponens fails. Priest is concerned with an application of this
approach to colour sorites. He introduces a logic of appearance to deal with the
suggestion that one can actually see contradictory states, though they may not appear
contradictory. In the paper following, Diana Raffman also addresses philosophical
problems about vagueness, but from a different perspective. She explores the
implications of some experimental data derived from psycholinguistic experiments
which counter objectivist views of how vague colour predicates identify physical
properties such as reflectance profiles. According to Raffman, dynamic patterns of
hysteresis and enhanced contrast in everyday speech are purely psychological and
give rise to subjective variation in subjects’ applications of vague predicates.
Dany Jaspers’ contribution makes a case for the observation that the four logical
oppositions (contradiction, contrariety, subcontrariety, entailment), as embodied in
Aristotle’s square and its extensions such as the Blanché hexagon, are inexact
with respect to the actual conceptual content of the lexical fields they organize.
According to Jaspers, they generalize over modal operators, propositional operators,
predicate calculus operators and tense operators. He maintains that the vertices of
the hexagon, for instance, could be abstracted from concrete incarnations, where a
homologous patterning among primary and secondary colour percepts is obtained.
Next Jean-Yves Beziau introduces a ‘chromatic hexagon of psychic dispositions’,
based on the extension of Aristotle’s square discussed by Jaspers, through which he
develops a theory of mental disposition and emotions inspired by colour theory.
Giselle Secco and Luiz Carlos Pereira’s contribution is devoted to exploring
the philosophical significance of the four-colour theorem proof, presented to the
mathematical community in a pair of papers by Appel and Haken in the late 1970s.
Against this background, Secco and Pereira formulate relevant questions regarding
some Wittgensteinian topics in the philosophy of mathematics such as surveya-
bility as a key criterion for distinguishing mathematical proofs from empirical
experiments. They hold that the ‘characteristic Wittgensteinian invention’ – i.e. a
controversially strong distinction between proofs and experiments – can shed light
on the conceptual confusions surrounding the four-colour theorem.
Walter Carnielli and Carlos di Prisco close the book by discussing the philo-
sophical interest of Ramsey’s theorem of finite combinatorics connected to colours
in mathematics and indicating some relevant philosophical open problems. This
theorem, thought to be a result in logic, states that in any colouring of the edges of a
sufficiently large complete graph, one will find monochromatic complete subgraphs.
Carnielli and di Prisco explore the idea of Ramsey’s theorem expressing a form of
unavoidable order, using colour system examples. They hold that it is not trivial to
Introduction to “How Colours Matter to Philosophy” xvii

completely clarify the role of colours in this statement: besides being hopelessly
unconstructive, the intuitive understanding of Ramsey’s theorem is that ‘complete
disorder is impossible’.
I would like to express my gratitude to Andrew Lugg, Andre Leclerc, Aparecida
Montenegro, Stekeler-Weithofer, Ingolf Max, Jean-Yves Beziau, Otavio Bueno,
João Marcos and Luiz Carlos Pereira for their decisive support of the ideas that first
motivated this colourful project and to Helen Lauer for helping with the preparation
of the manuscript.

Federal University of Alagoas Marcos Silva

Maceió, Alagoas, Brazil
 Part I
History of Philosophy
Dispositionalism: Democritus and Colours
by Convention

Barry Maund

The ancient philosopher and atomist, Democritus apparently wrote over 50 books,
none of which have survived, except for some intriguing fragments. One of the most
intriguing ones is very famous:
For by convention colour exists, by convention bitter, by convention sweet, but in reality
atoms and the void.

This is usually acknowledged as denying that there are colours in the natural
world or, as it is often put ‘in reality’. I think that this is right, but the view is
more subtle than is commonly thought. Understood properly, Democritus provides
an inspiration for a valuable account in the philosophy of colour.
There is a certain argument that have been attributed to him – by ancient
commenters, and repeated by modern historians/philosophers – an argument to
the effect that colours do not exist. Professor Burnyeat begins an important paper,
‘Conflicting Appearances’, with a quote from the ancient empiricist philosopher,
Sextus Empiricus:
From the fact that honey appears bitter to some and sweet to others Democritus concluded
that it is neither sweet nor bitter, Heraclitus that it is both. (p. 69)

As Burnyeat comments, this report testifies that arguments from conflicting

appearances came early to the repertoire of philosophy. This point is not just of
historical interest. A modern version of the argument from Conflicting Appearances,
we should note, forms a central role in the argument by Jonathan Cohen for a
Relationalist view of colour. Cohen sees his view of colour as incompatible with
Democritus’s response to the phenomenon of conflicting appearances. (In Maund
2012, I have a detailed critique of Cohen’s approach to conflicting appearances in
his argument.)

B. Maund ()
University of Western Australia, Crawley, Australia
e-mail: jbmaund@bigpond.com

© Springer International Publishing AG 2017 3

M. Silva (ed.), How Colours Matter to Philosophy, Synthese Library 388,
DOI 10.1007/978-3-319-67398-1_1
4 B. Maund

There is a related context in which Democritus’s view is important. R. Chisholm

has defended a dispositionalist view of colours. In exposition of this view, he
proceeds by means of a criticism of Democritus. Following the lead from Aristotle,
he attributes to Democritus the argument above, and accuses Democritus of
committing the fallacy of equivocation. In the first part of this paper, I shall argue
that Chisholm’s criticism is unfair to Democritus. His argument is more complex
and does not depend on such a fallacy.
In Part II of the paper, I argue more positively in favour of Democritus. I claim
that there is reason to think that a dispositionalist account of colour stands in need
of an error component, as Democritus suggests. (As it happens, these reasons will
also apply to Cohen’s theory.)
Having said all this, however, let me make a crucial qualification. It is not my
aim to defend a historical claim about what Democritus actually said, let alone what
he meant. My competence, such as it is, does not go so far as to allow that. My aim
is more limited. It is two-fold. First, I think that there is a possible interpretation
of what Democritus was arguing, an interpretation that is coherent, and is plausible
for him to have held. Second, whatever the truth of the first claim, the thesis is an
eminently defensible, and is a worthy candidate for providing an adequate account
of colour perception.

1 Part I

1.1 Colours by Convention

We have seen that commentators standardly take Democritus to be denying that

any external thing is black or white or red or sweet, or bitter, etc. That is to say,
Democritus is understood as being what is today called an Eliminativist, or Irrealist,
with respect to colours: holding that, contrary to common belief, there are no bodies
that have the colours. However, let us go back to the famous remark by Democritus,
in the fragment:
For by convention colour exists, by convention bitter, by convention sweet, but in reality
atoms and the void.

This remark suggests that his position is more complex than is usually supposed.
If it comprises eliminativism, it is not a bald eliminativism. For in the first part of the
fragment, he says “By convention colour exists, by convention sweet exists, : : : ” So,
he is not actually denying that colours exist. He is saying something more nuanced:
colours exist, but they exist by convention.
One might wonder whether the remark as a whole is consistent. On the one
hand, reality seems to comprise the atoms and the void, but on the other, there are
conventions and presumably people that make and accept the conventions. Let us
put that aside for one moment. Presumably, someone who says that colours exist by
convention is contrasting this sort of existing, with another sort. That is to say, it is
Dispositionalism: Democritus and Colours by Convention 5

implied, there are things that that exist but not by convention, or at least, not merely
by convention. Colours and tastes and odours, etc., do not belong to this class of
things.
Modern social theorists and philosophers commonly draw a distinction between
social reality and physical reality. The social reality is construed as a socially
constructed reality. John Searle has written extensively on the subject.1 He explains
the social reality as comprising things whose existence relies on agreement among
social beings (typically, humans). The agreement, we should note, may be implicit
or explicit.
It may be possible to argue that Democritus’s view should be understood in this
sense. I think it can be. Nevertheless, I shall argue, his view is more nuanced than
this thesis suggests.

1.2 Chisholm on Democritus

Chisholm attributes to Democritus, a similar argument to the one cited by Burnyeat,

the argument from conflicting appearances. Following Aristotle, Chisholm writes:
Oversimplifying slightly, we may say that Democritus reasoned in this way: “The wine that
tastes sweet to me tastes sour to you; therefore I do not perceive that it is sweet and you do
not perceive that it is sour, and the wine itself is neither sweet nor sour.” (Chisholm 1966,
p. 92)

However, this statement comes just after he has ascribed a more complex
argument to Democritus:
The appearances of things, he [Democritus] said change with the condition of our body and
the influences coming toward it or resisting it. The question as to whether any particular
thing will appear white, black, yellow and red, sweet, or bitter, he noted, cannot be answered
merely by reference to the nature of the thing; one must also refer to the nature of the person
or animal who is perceiving the thing. (p. 91)

Chisholm then goes on to say:

And from these premises, which are undeniable, Democritus then went on to infer (1) that
no one ever perceives any external thing to be white, black, yellow, red, sweet, or bitter, and
also (2) that no unperceived external thing is, in fact, white, black, yellow, red, sweet, or
bitter. (p. 91)

Chisholm’s evaluation of this argument is that it is fallacious: “we can accept the
premises that Democritus used and at the same time, reject his conclusions, for the
conclusions do not follow from the premises”. It seems to me, however, that, on
the contrary, the more complex argument has the seeds of a more promising line of
thought. This suggestion is strengthened when we consider, I argue, that it is highly
questionable that Democritus is guilty of the fallacy claimed.

1
See especially Searle 1995.
6 B. Maund

The argument only seems valid, Chisholm writes, since Democritus commits
an equivocation. In saying this, Chisholm agrees explicitly with Aristotle, who he
quotes as saying:
The earlier students of nature were mistaken in their view that without sight there was
no white or black, without taste no savour. The statement of theirs is partly true, partly
false. ‘Sense’ and ‘the sensible object’ are ambiguous terms; i.e., they may denote either
potentialities or actualities. The statement is true of the latter, false of the former. This
ambiguity they wholly failed to notice.2

Chisholm explains this passage as follows:

In suggesting that the terms “white” and “black” are ambiguous, Aristotle is taking note
of the fact that in certain uses, these terms are intended to refer to ways of appearing and
that in other uses they are intended to refer to properties or dispositions of physical things –
those properties or dispositions in virtue of which the things appear in the ways in which
they do appear. [My emphasis] (p. 93)

Chisholm writes:
Let us say of such terms as “white”, “black”, “yellow”, “red”, “bitter” and “sweet”, that
when they are used to refer to these properties or dispositions, [i.e., dispositions of physical
things], they have a dispositional use, and that when they are used to refer to ways of
appearing, to ways in which things may appear, they have a sensible use. [My emphasis]
(p. 93)

The way the fallacy is said to arise is as follows:

Aristotle is telling us, then, that the statement “Without sight, there is no white or black,
without taste, no savour” is true if the terms “white”, “black” and “savour” have a sensible
use, and false if they have a dispositional use. Democritus, therefore, seems to have
committed the fallacy of equivocation: Having established that the statement is true when
it is taken in the first of these two ways, he goes on to infer fallaciously that it is also true
when it is taken in the second.” (p. 93)

However, given the fuller argument that Chisholm and Aristotle attribute to
Democritus above – that is, given the background argument that precedes the
argument from conflicting appearances – it is not at all clear that they are being
fair to him. That argument cited is
The question as to whether any particular thing will appear white, black, yellow and red,
sweet, or bitter, he noted, cannot be answered merely by reference to the nature of the thing;
one must also refer to the nature of the person or animal who is perceiving the thing.

The thrust of this argument, it seems to me, is that, for the use of terms, “white”,
“black”, “yellow”, “red”, “bitter” and “sweet”, it is a conventional matter which
observers are normal and which conditions are standard. We should keep in mind
that to say a matter is decided conventionally is not to say it is an arbitrary matter,
that there is no good reason for adopting the convention. It is natural to assume that
Democritus is appealing to some such argument as the following:

2
De Anima, Bk III, Ch.2, p. 426a; also Metaphysics, Bk IV, Ch.5, 1010b.
Dispositionalism: Democritus and Colours by Convention 7

Given that the way things appear to a normal perceiver depends on the constitution of that
person, as well as on the qualities of the external bodies (the qualities of the atoms) then it
is easy enough for us to imagine that the constitution of those observers should change so
that things appeared quite differently.

In other words, what counts as “normal perceivers” depends on the constitution

of human beings, and that it is easy enough to conceive of possible worlds in which
this constitution is different for human beings, but everything else is the same. In
those worlds, either there are worlds in which no things are blue, yellow, purple etc.,
or different things are blue, yellow, purple, etc.
In the light of these considerations, we can make the response, on behalf of
Democritus, that to accept the dispositional account of colour is to grant him his
case. To say that objects have colours, in the sense of having dispositions to appear,
is to say that things have colours by convention. The reason for this is that if the
relevant disposition is disposition to appear, it must be a disposition to appear to
certain kinds of observers. This means that that the relevant class of observers will
be ones that have a certain nature/constitution. What Democritus is doing, we may
suppose, is to say that we can imagine that whatever that constitution involves, we
can easily enough imagine it being changed in such a way that rubies and tomatoes
and apples will look differently to people; they will no longer look red, and hence
will no longer be red. In that case, we will adopt a different convention for picking
out red objects.
We might strengthen these considerations by drawing upon modern arguments
presented by Averill, in Averill 1992 and 2005.3 His arguments are directed
at colour-dispositionalists and colour-realists, who both depend crucially on the
notions of normal perceivers and standard viewing conditions in exposition of their
theories. His arguments depend on difficulties that stem from trying to give a non-
arbitrary account of normal observers and standard viewing conditions. We can
easily suppose changes in either our eyes (and hence in normal observers) or in
standard viewing conditions, such that some objects that previously were yellow
would look red, and others would still look yellow—while remaining otherwise
physically unchanged.
As Averill points out, how colours appear (the colours things appear to have)
depends on a very complex structure of the visual system: they depend on (a)
response functions of cones in the retina; (b) an opponent-processing system,
consisting of channels that take as input responses of the cones, and deliver as output
stimulation of cells that deliver experiences of colour. As Averill further points out,
we can easily enough imagine these structures changing in a variety of ways: either
by natural evolution, genetic engineering, medical intervention, or alien invasion.
Some changes will be unproductive, but not all. We can imagine systematic changes
that can affect what will count as “normal perceivers”.

3
Averill, E. W., 1992, “The Relational Nature of Colour”, Philosophical Review, 101: 551–588;
Averill, E.W. (2005), ‘Toward a Projectivist Account of Color’, The Journal of Philosophy, 102,
217–234.
8 B. Maund

I am not saying that Democritus is anticipating the empirical discoveries that

Averill is drawing attention to. I am saying that Averill gives added strength to an
argument that has much to say for itself previously. I am suggesting that Democritus
has in mind the sorts of considerations that Jonathan Bennett appealed to in his
discussion of how secondary qualities such as colours differ from primary qualities,
in his Locke, Berkeley and Hume.4 Averill makes the case that much stronger.
We might ask ourselves what reasons do we have for having concepts of colour
and taste. Why do we classify certain things as sweet, bitter and tasteless – and
other sorts of things as white, blue, purple, orange, etc.? There seem to me to be two
major reasons. One is that the qualities of being sweet and being bitter are related
to the psychological capacities of humans and other sentient beings: they provide
motivations for actions. The second reason is that they work for us as signs of the
presence of other qualities. This reason is more important for colours, but it has
some place for tastes as well.
Given that this is so, we can see that the point of colour-vision is not, in the first
place, to tell us about perceiver-independent properties of objects: it is to tell us
about perceiver-dependent properties – ways things appear – which in turn tell us
about things in the world that are of special interest to human beings.
This point is reinforced by this consideration. There is a group of philosophers
who are colour-realists, who defend a physicalist reductionist theory of colour. They
hold that colours are perceiver-independent qualities of physical objects, and, ore
particularly, are light-related properties, e.g., for surface colours of physical objects,
a colour such as blue is a member of a special group of spectral reflectances.
Examples of such philosophers are Tye (2000), Byrne and Hilbert (2003), Smart
(1975/1997). Although all of these philosophers describe themselves as colour
realists, they also describe their view as Anthropocentric realism. The point of this
label is that although colours are identified as real properties, they are properties
that are mainly of interest only to human beings, and in particular, to human beings
with a certain type of colour vision. And this is to imply that the colour visual
system might change in important ways so as to identify a different group of “real”
qualities that are of special human interest. Again, on behalf of Democritus, we
might respond that on the view colours are real, but real by convention.

1.3 The Case for Equivocation

The passage in which Chisholm explains the ambiguity that Aristotle points to,
is very revealing. It implies that Aristotle’s argument depends on a fact that is
supposed to be available to all of us, presumably those of us who reflect on our
use of certain linguistic terms. Aristotle refers to how, in different uses, the term
are intended to refer either to dispositions or, alternatively, to ways of appearing.

4
J. Bennett, Locke, Berkeley and Hume, 1971, pp. 89–123.
Dispositionalism: Democritus and Colours by Convention 9

It ought to raise the question as to who intends the terms to refer in these ways.
Since the context is one in which Chisholm is describing our ordinary use of ‘looks’
expressions, he must mean that ordinary language-users, intend that the terms be
used in these two different ways. But, it strikes me, it is highly implausible that
ordinary language users intend “red” in “looks red” to refer to ways of appearing.
(It sounds suspiciously like something it would need a philosopher to say.) Not only
that, but it strikes me that it requires considerable philosophical acumen to detect
the ambiguity.
Consider these examples: ‘the lines in the Muller-Lyer illusion look unequal
(to S)’; ‘the straight stick partially immersed in water looks less bent than the one in
benzene’; ‘the cars in the rear vision mirror look quite distant, but are rather close’.
These would seem to be examples of the descriptive or phenomenal use of ‘looks
F’ that Chisholm relies upon. But here surely, the terms, ‘unequal’, ‘bent’, ‘quite
distant’, are used in their standard, everyday sense. It is because these terms have
their normal use, that we are struck by the illusions. It would seem that there is an
ordinary, everyday, use of ‘looks F’, which presupposes a contrast between looks F
and is F. Or, to take the example discussed by Chisholm, that of ‘looks centaurian”.
It is hard to believe that the term ‘centaurian’ here is not used in with its normal
meaning.
There is a further complication in Chisholm’s account.
If we are to speak more strictly, we should not say that “white” in its sensible use, always
refers to the way in which an object appears; it refers, rather, to the way in which one is
appeared to – whether or not an object appears. Or if we introduce an active verb, such
as sensing, or “experiencing” as a synonym for the passive “is appeared to”, we could say
that “white” in its sensible use, refers to the way in which a man may sense or experience.
(p. 94)

It seems even more implausible that ordinary language-users intend to use the
term in this kind of sensible use. It sounds even more like a concept that a philoso-
pher would introduce to give an account of the phenomenology of perception. It
seems to me that Chisholm is within his rights in producing such a theory. My
objection is to his claim that Democritus is engaged in a fallacy of equivocation.
The charge should be a different one: that he has the wrong philosophical theory.
But then Chisholm has to do far more to sustain this charge. He cannot rely on the
claim that Democritus is committing a fallacy, that of equivocation on the meaning
of terms in common use.
Before leaving this topic, I should acknowledge that in this area there are a
number of ambiguities, and it is important to distinguish between the different
types of equivocation that might arise. One important ambiguity is with the notion
of ‘looks’, which I discuss later (in II: 3 ‘Two Approaches to Dispositionalism’).
This concerns the threefold distinction that Chisholm and Jackson have both drawn
attention to. This ambiguity is different from the putative distinction that Chisholm,
following Aristotle, alleges to hold.
10 B. Maund

2 Part II

2.1 Modern Versions of Colour Dispositionalism

It might be thought that, in my defence of Democritus, I have gone too far. I have
emphasised the importance of the first part of his famous quote: that colours exist
by convention. I have argued that this claim is still true, even if it is true that our
colour terms designate dispositional properties, as has been argued by some. It may
be thought, however, that I have not given full attention to the second part of the
quote. For if all that he had in mind is compatible with Dispositionalism, why need
he bother with the remark that in reality there are (only) atoms and the void?
The point is that to explain the positive part of the remark, i.e., the claim that
colours exist by convention, we need some account of what might make this true.
The dispositionalist account supplies an answer. It is not the only answer. It could
be, as I suspect, Democritus would have thought that our ordinary colour terms were
thought of as designating perceiver-independent qualities of objects. He might have
understood his conventionalism as a form of fictionalism: the colour terms apply to
fictional properties. However, he might also have been a revisionist about our colour
terms and held that we should read the ascription of colour terms as if they applied
to relevant dispositional properties. Even so, I argue, there is good reason to think
that the theory requires some error element as a component. In this second part of
the paper, I wish to address this question. I propose to answer the question with
respect to modern versions of colour-dispositionalism.
Levin (2000) and McGinn (1983) are two of the most prominent defenders of
dispositionalist views of colour. (Later, McGinn 1996 revises his view). Both reject
the thought that dispositionalism requires granting a place to errors in the account of
perception. They both see their positions as being in the same tradition as Locke and
other early philosophers. In so doing, they take Locke to be committed to the view
that the dispositionalist view of colour is an account of our ordinary colour concepts.
In my view this gets Locke wrong. His dispositionalist account of colour should
be read as a proposal: that if we wish to adopt clear philosophical thinking about
colours, and other secondary qualities – as opposed to our thinking in our “vulgar”
moments – we should think of them in dispositional terms: secondary qualities will
be powers to induce ideas of secondary qualities. However, this revisionary proposal
is compatible with holding an error view about colour experience. (I argued for this
thesis in Maund 1996.)
In one sense, it does not matter what views Locke held. The views of disposi-
tionalists such as Levin and McGinn can be evaluated on their own merits. Still, as
I think Locke’s revisionary proposal is, in broad terms the right one, it is important
to bear it in mind. An important reason for this is that Levin and McGinn are keen
to avoid resort to an error element in their theories. I shall argue, however, that the
strongest versions of this theory are best construed as containing (happily) an error
component.
Dispositionalism: Democritus and Colours by Convention 11

2.2 Error Theories

I prefer the term “illusory theories”, “fictionalism” to “error theories”. These labels
are better in that “illusion” and “fiction” are more positive than “errors”. Illusions
and fictions can serve a number of useful purposes. Illusions can carry information.
One good example is that the bent-stick illusion carries information about the
refractive index of the media in which the stick is placed. (See Edmond Wright
1996).5
However, the term “error” is in widespread use, and as long as the dangers
of possible misrepresentation are heeded, no great damage should be done by its
continued use.
Even errors and falsehoods can carry information, especially if the errors are
systematic ones. The clock in my car always runs slow, and, as time passes, it
falls more and more behind. It is never accurate, except for 1 day at a time. Yet
it is nevertheless very useful. Governments and central banks depend on inflation
measures and unemployment measures. But these measure are never accurate – how
could they be? Or, if they are accurate, no-one knows whether they are. The point is
that it does not really matter. What matters is that we use the same measures and we
note the variations in the measures. As long as the errors are systematic errors they
can be very useful.
In the case of colours and most secondary qualities, there are two main reasons
why we are interested in knowing about them, i.e., in knowing whether an object
has a certain colour, or gives off a certain sound, or ejects a certain smell, etc. One
reason is that colours – and patterns of colour – are very good signs for the presence
of some thing or quality or another. Red is very good for picking out certain fruits.
Certain patterns of colours are good for identifying ripeness of fruits, their decay and
so on. The importance of colour cannot be over-estimated. Perception works almost
entirely through the recognition of objects and their qualities, and in effect, almost
all of this identification is through the appearance of objects. And central to the
appearance of objects is their colour, and patterns of colour. The importance of this
function is also illustrated in the conventional uses of colour, e.g., in color-coding: in
traffic-lights, colour of the coating of electric wiring, colour-coding in various kinds
of images. The significance of the fact that colours serve as signs is the following.
Being a sign for the presence of some other quality, the causal relevance of the sign
is not so much that it has causal powers, but rather, that it is a causal effect, and
hence it is an indicator of qualities that we want to know about.
A second function that colours have is in the service of broadly aesthetic effects:
in sexual attraction, in appreciation of nature (sunsets, landscapes, gardens, etc.,) in
clothing, in fashion, in bodily decoration, in habitat design, in art, and so on. These
effects are not solely for beauty: they can tell us of states of decay, of terror, of high
status, etc.

5
Wright 1996, p. 34.
12 B. Maund

With respect to both of these functions, the function is served just as well if colour
perception works through errors and illusions – provided of course that the errors,
(illusions, fictions) are systematic ones. On a properly constructed error theory, of
course, they are systematic errors, illusions, fictions. The point about colours is that
they serve their functions through the way they look, and an error theory does not
deny that things look blue, or look purple, or look light blue, etc. For serving the
major functions, it does not matter whether the objects really are blue, purple, light
blue, etc. what matters is that they look blue, look purple, look light blue. What
matters for delight in golden or red sunsets is that the sunset looks a specific colour.

2.3 Problems for Dispositionalism

Colour-Dispositionalism is usually understood in this sense: for X to be blue is

for X to be disposed to look blue to normal perceivers in standard conditions.
As it stands, this account has two problems: the circularity problem and the
phenomenological problem. McGinn (1983) and Levin (2000) both argue for
versions of dispositionalism that, they hold, overcome these problems.6
One of the difficulties with the accounts given by McGinn and Levin is that they
do not actually give any detailed account of what it is for anything to look any
way at all, nor, in particular, what it is to look blue, say. They take the notion for
granted as not needing explanation. Accordingly, any adequate defence of Colour-
Dispositionalism will require an adequate account of what it is to look blue, look
yellow, etc. This defect is crucial if it turns out that the relevant dispositional
property that constitutes colours are not dispositions to look blue, look yellow, etc.,
but dispositions to look something else. (Maund 2012 has attempted to re-frame the
arguments of Levin and McGinn on the basis of such an account.)
The circularity problem has been formulated in different ways. One way is to
say that the account of what it is to be blue depends on the notion of looks blue,
and would seem to require prior understanding of what it is to be blue. One way of
defusing this problem is to say that ‘blue’ has different meanings in the explanans
and the explanandum. (This seems to be implicit in Chisholm 1966). A more
satisfactory resolution of the problem is to say that ‘blue’ has the same meaning
in each, but the expression, ‘looks blue’, designates different properties in different
contexts. In some contexts, it designates a relationship to the property of being blue;
in other contexts, however, it does not. (It seems to me that Peacocke 1984/1997
defends a version of this response.)

6
Jonathan Cohen in his defence of Colour-Relationalism, appeals to Levin’s account in defence of
his theory against objections based on similar problems. Cohen’s theory may be thought of as a
relativised dispositionalist theory: colours are dispositions, but the dispositions are relativised to
classes of perceivers, and kinds of circumstances.
Dispositionalism: Democritus and Colours by Convention 13

There is another way to formulate the circularity problem which, perhaps, makes
even clearer how this response to the problem operates. It is one that Levin has
described. (She attributes it to McGinn 1996 as a paraphrase of his discussion):
If an object is red iff it’s disposed to look red (under appropriate conditions), then an object
must be disposed to look red iff it’s disposed to look to be disposed to look red : : : and so
on, ad infinitum.7

A solution to this way of formulating the problem is to say that the sense of
‘looks red’ is such that to look red is not a matter of looking to have the property
of being red. Several possibilities are open: (i) it is to look to have some other
property say red*; (ii) to look red is to look a certain way. On the second possibility,
the expression ‘looks red’ employs the term ‘red’ with its normal meaning, but the
expression ‘looks red’ designates rigidly a certain property – a property of appearing
a certain way – which does not involve any relation to the property of being red.
It seems to me, therefore, that the circularity problem has a solution, though it
comes at a price. The phenomenological problem is more difficult. One can admit,
with Levin, that coloured objects may look to have dispositional qualities – there
are examples of dispositional qualities that objects look to have, e.g., they may look
fragile or heavy or opaque or solid, etc. However, in such examples, it seems that
the object only looks to have the dispositional property by virtue of looking to have
intrinsic qualities and, on the face of it, these qualities are ones physical objects
don’t have.

2.4 Two Approaches to the Dispositionalist Theory

It is customary for philosophers to make a threefold distinction between three dif-

ferent types of expressions and concepts that apply in ‘looks : : : ’ contexts. The
three uses for the respective expressions (and concepts) are usually called the
comparative, perceptual-epistemic and phenomenological uses, respectively. They
are commonly represented by the respective expressions: ‘X looks to A the way
F-things look in such and such circumstances’; ‘X looks to A as if it is F’, and ‘X
looks F to A’. (For classical discussions of this three-fold distinction see Jackson
1977 and Chisholm 1957 and 1966.)
There has been, in recent times, controversy over the characterisation of the
phenomenological use and, indeed, of its validity. Maund (2012) presents a defence,
and characterisation, of this use. (He distances himself from both Jackson and
Chisholm, in their accounts of this use.) In this account, there are two aspects to
the phenomenological use:

7
Levin 2000, p. 163.
Another random document with
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genital plates is also pierced by the madreporic pores. Some
zoologists have separated the ocular and the genital plates under
the name of "calyx" from the rest of the corona, under a mistaken
idea that they are homologous with the plates of the body or calyx of
a Crinoid.

Fig. 229.—The peristome of Echinus esculentus. × 2. 1, Tube-feet of the lower

ends of the radii; 2, gill; 3, teeth; 4, buccal tube-foot; 5, smooth peristomial
membrane. (After Kükenthal.)

The periproct (Fig. 228, 4) is covered with small plates and bears a
few pedicellariae. The peristome (Fig. 229) is covered by flexible
skin with abundant pedicellariae; it terminates in a thick lip
surrounding the mouth, from which the tips of five white teeth are just
seen projecting. There are ten short tube-feet projecting from the
peristome—one pair in each radius—and each tube-foot terminates
in an oval disc and is capable of little extension, and each has
around its base a little plate. The presence of these tube-feet shows
that in Echinus the peristome extends outwards beyond the water-
vascular ring, whereas in Asteroidea it is contained entirely within the
ring. In the primitive Cidaridae (Fig. 235) the whole peristome down
to the lip surrounding the mouth is covered with a series of
ambulacral and interambulacral plates similar to those forming the
corona, though smaller and not immovably united, and the series of
tube-feet is continued on to it. It is thus evident that the peristome is
merely part of the corona, which has become movable so as to
permit of the extension of the teeth. In Echinus the peristome is
continued in each interradius into two branched outgrowths called
gills, the relation of which to the respiratory function will be described
later. These gills (Fig. 229, 2) are situated in indentations of the edge
of the corona called "gill-clefts" (Fig. 230, g).

Fig. 230.—The dried peristome of Echinus esculentus and the surrounding

portions of the corona. × 1. amb, Ambulacral plate; b.t, buccal tube-foot; g,
gill-cleft; inter, interambulacrum; per, peristome.

The most conspicuous plates in the peristome are those surrounding

the buccal tube-feet; besides these, however, there are in Echinus
esculentus, and probably in most species, a large number of thinner
irregularly-scattered plates (Fig. 230).

The term ambulacral plate, applied to the plate pierced by the pores
for the tube-feet, conveys a misleading comparison with the
ambulacral plate of an Asteroid. In Echinoids the ambulacral groove
has become converted into a canal called the "epineural canal," and
the ambulacral plates form the floor, not the roof, of this canal; they
may perhaps correspond with the adambulacral plates of the
Starfish, which one may imagine to have become continually
approximated as the groove became narrower until they met.
 Fig. 231.—Dissection of Echinus esculentus. × 1. The animal has been opened
by a circumferential cut separating a small piece of the skeleton at the
aboral end, which is turned outwards exposing the viscera on its inner
surface. The other viscera are seen through the hole thus made. amp,
Ampullae of the tube-feet; aur, auricle; b.v, so-called "dorsal blood-vessel";
comp, "compasses" of Aristotle's lantern, often termed "radii" by English
authors; comp.elv, elevator muscles of the compasses; comp.ret, retractor
muscles of the compasses; eph, epiphyses of the jaws in Aristotle's lantern;
gon, gonad; g.rach, genital rachis; int, intestine; oe, oesophagus; prot,
protractor of Aristotle's lantern; rect, rectum; ret, retractor of Aristotle's
lantern; siph, siphon; st, stomach; stone.c, stone-canal.

The internal organs of the Urchin can best be examined by making a

horizontal incision about one-third the distance from the mouth and
pulling the two parts gently asunder. A large amount of fluid escapes
from the exceedingly spacious coelomic cavity, the alimentary canal
being comparatively narrow.

The alimentary canal commences with a short vertical tube which

has been shown to be a stomodaeum; this is surrounded by the
upper ends of the teeth and their supporting ossicles, which are
collectively termed "Aristotle's lantern." The oesophagus leads into a
baggy, flattened tube, the stomach, which runs horizontally round the
animal, supported by strings of tissue from the coelomic wall, so that
it hangs down in a series of festoons. Having encircled the animal, it
bends directly back on itself and immediately opens into the
intestine, which is also a flattened tube, which runs round the
circumference of the animal, but in the opposite direction, the
festoons of the second circle alternating with those of the first. The
intestine opens into a short rectum which ascends vertically to open
by the anus. The stomach is accompanied by a small cylindrical tube
called the "siphon" (Fig. 231, siph), which opens into it at both ends;
this represents merely a gutter which has been completely grooved
off from the main intestine; it is lined by cilia, and its function is
believed to be that of keeping a stream of fresh water flowing
through the gut, so as to subserve respiration.

Echinus esculentus seems to feed chiefly on the brown fronds of

Laminaria and the small animals found thereon, which it chews up
with its teeth, but it may regale itself on the same diet as Brittle
Stars, as Allen[474] has shown to be the case in Plymouth Sound.
Dohrn[475] has described the Neapolitan Sphaerechinus granularis
attacking and capturing Crustacea such as Squilla.

The water-vascular system presents several features of great

interest. The ring-canal is situated at a considerable distance above
the nerve-ring, and is separated from it by the whole of the jaws and
teeth. It has five small interradial pouches on it, which apparently
correspond to Tiedemann's bodies in an Asteroid. The stone-canal
(Fig. 231) opens as usual into the ring-canal, and is accompanied by
the axial sinus and genital stolon. The name "stone-canal" is very
unsuitable in this order, for there are no calcifications in its walls; it is
a simple membranous tube of circular section. On reaching the
upper wall of the test it expands into an ampulla, into which the
numerous ciliated pore-canals traversing the madreporite open. The
radial canals, starting from the ring-canal, pursue a downward
course till they come into contact with the radial nerve-cords, and
they then bend upwards and run along the centre of the ambulacral
region, finally terminating in the small terminal tentacles. In the just
metamorphosed Echinoid these are well-developed tube-feet, each
with a well-developed sucker, in the centre of which is a conical
sensory prominence, but as development proceeds they become
enclosed in a circular outgrowth of the test, so that only the tip
projects in the adult.

The long extensible tube-feet are connected by transverse canals

with the radial canal. Instead of the pair of valves which in Asteroids
prevent the reflux of liquid into the canal, there is a perforated
diaphragm[476] with circular muscles, which by contraction close the
opening in the diaphragm, while when they are relaxed fluid can
return from the tube-foot. The ampulla is flattened, and is contracted
by muscular fibres called "trabeculae" stretching across its cavity.
These muscular strands are developed by the cells lining the
ampulla. The external portion of the tube-foot, as in Asteroids, is
provided with powerful longitudinal muscles, and there is the same
alternate filling and emptying of the ampulla as the tube-foot is
contracted and expanded. The tube-foot is connected by a double
canal with the ampulla, the object of which is to assist in respiration.
The cells lining it are ciliated, and produce a current up one side of
the tube-foot and down the other, and the double canal leading to the
ampulla separates these two currents and prevents them interfering
with one another. Thus water is continually transported from the
ampulla to the tube-foot, through the thin walls of which it absorbs
oxygen, and it is then carried back to the ampulla, and transfers its
oxygen to the fluid of the general body-cavity through the walls of the
ampulla. The disc of the tube-foot is supported by a calcareous plate
(Fig. 232, oss), a circumstance which enabled Johannes Müller to
recognise the Echinoid larva when the form of the adult was as yet
unrecognisable. Below the edge of the disc there is a well-marked
nerve-ring, from which two bundles of nerve-fibres go to the disc
itself, in the edge of which there is an abundance of sense-cells.

The buccal tube-feet (Fig. 229, 4) are much shorter than the rest,
and are provided with oval discs which are highly sensory. These
feet are not used for seizing, but for tasting food; when a piece of
food is placed near them they are thrown into the most violent
agitation.
 Fig. 232.—Diagrammatic transverse section of the radius of an Echinoid.
amb.oss, Ambulacral ossicle; amp, ampulla of the tube-foot; ep, epineural
canal; musc, muscles attaching spine to its boss; nerv, nervous ring in base
of spine; n.r, radial nerve-cord; oss, ossicle in sucker of tube-foot; ped,
tridactyle pedicellaria; perih, radial perihaemal canal; pod, tube-foot; wv.r,
radial water-vascular canal.

The nervous system has the same form as in an Asteroid, viz. that
of a ring surrounding the mouth and giving off radial nerve-cords
(Fig. 232, n.r), one of which accompanies each water-vascular canal
to the terminal tentacle, where it forms a nervous cushion in which
pigmented cells are embedded.

A large band-like nerve is given off from the radial nerve-cord to

each tube-foot. This pedal nerve, as it is called, contains bipolar
neurons, and is really an extension of the nerve-cord itself. Beneath
the sucker it branches out to form a sensory ring. From the base of
the pedal nerve, branches are given off which run to the ectoderm
and enter into connexion with the plexus there. Romanes[477]
scraped away the radial cords and found that the spines still
converged when a point on the ectoderm was stimulated, but that,
on the other hand, if definite locomotor movements were to be
carried out, the presence of these cords was a necessity; hence he
concluded that the superficial plexus sufficed for ordinary reflexes,
but that for purposeful movements the central nervous system was
necessary.

Von Uexküll[478] has made an exhaustive study of the physiology of

the nervous system in the Echinoidea. He points out that all the
organs controlled by the nervous system, spines, pedicellariae, tube-
feet, and (see below) Aristotle's lantern, give two opposite reactions
in response to the same stimulus according as it is strong or weak,
bending away from the point of stimulation when it is strong and
towards it when it is weak. This reversal of reaction can only be due
to the action of the neuron in altering the effect of the stimulus on the
muscles, and this Uexküll regards as its fundamental property. Thus
in Preyer's[479] experiments with Starfish the strong form of
stimulation is obtained by directly applying the stimulus to the radial
cord or to the tube-feet, the weak form by stimulating the back, when
of course the stimulus has to traverse a longer path before affecting
the tube-feet, and is consequently weakened. Von Uexküll also
introduces the conception of "tone" with regard to the nervous
system. This term has been used to denote the amount of chronic
contraction in a muscle, and it is to be distinguished from the fleeting
contractions which cause movement. The more tone there is in a
muscle the less responsive it is to stimuli tending to bring about
movement. As applied to the nervous system "tone" denotes a
condition when it is not receptive to small stimuli, but when it is
maintaining a condition of tone in a muscle by which of course its
own tone is measured. Tone in a neuron can therefore be measured
by the produced tone in the muscle, and the one is to be
discriminated from the other only by using stimulants, such as
caffeine, which have no direct action on muscle. Tone can also be
measured by the amount of stimulus necessary to irritate the neuron.
When muscles are stretched the tone is lowered, and this loss of
tone extends to the neuron controlling the muscle, and vice versa.
When the spines on being gently stimulated bend towards the point
of stimulation, this is due to the contraction of the muscles on the
side towards the point of stimulus, for if the superficial plexus of
nerve-fibres be cut through so that the stimulus has to pursue a
round-about course the spine will bend towards the direction from
which the stimulus comes. The bending of the spines away from the
stronger stimulus is likewise due to the muscles on the side towards
the stimulus. It is caused by a sudden fall of tone in these muscles,
which causes them to yield to the tone of the muscles on the
opposite side, and this fall of tone is due to a fall of tone in the
neurons, for it can be produced by chemicals, and the direct action
of all chemicals applied to muscle is to raise tone.

In Arbacia this form of reaction cannot be produced; the spines

respond to stimuli of all degrees of intensity by convergence towards
the point of stimulation.

When a general skin-irritant like dilute acetic acid, or even strong

light, is applied to the skin of a Sea-urchin the spines bend
alternately to all points of the compass, or, in a word, rotate. This is
due to the fact that the weight of the inclined spine stretches the
muscles of one side and so renders them more open to the general
stimulus; these muscles in consequence, contract, and so move the
spine to a new position in which other muscles are stretched, and a
similar result follows. A continuation of this process brings about
rotation.

When a piece of glass rod or other light object is laid on the spines of
a Sea-urchin, it naturally, by its weight, presses asunder the spines
and stretches their muscles on one side, thus lowering the tone. If
now the skin be stimulated at any point the piece of rod will be rolled
by the spines towards the point of stimulation. This is caused by the
fact that the muscles of the spines holding the rod are made more
receptive by being stretched, and therefore they contract more than
do the others in response to the stimulation, and so the rod is rolled
onwards on to the next spines, which then act in the same manner.
This passage of stimulus is entirely independent of direct nervous
connexion between the bases of the spines, for it will traverse at
right angles a crack going clean through the shell; it is merely the
result of the mechanical weight of the object and of the juxtaposition
of the spines.

If the stimulation be too violent the first spines affected diverge wildly
and strike their neighbours with vehemence, so arousing into activity
the block musculature of these. This causes them to stand rigidly up,
and so the path of the stimulus is barred.
Now the escape movements of the animal under strong stimulation
which Romanes[480] alludes to are just an example of this handing
on of stimulation from spine to spine, not by nervous connexion but
by mechanical touch only; the object in this case is the substratum
on which the animal lies, which is, so to speak, rolled towards the
point of stimulation, or putting it otherwise, the animal is rolled away
from it. Righting when upset is another example of the same
phenomenon; the aboral spines are stretched by the weight of the
animal, and the animal acts as if it were stimulated in the region of
the periproct. When a Sea-urchin is in its normal position and is
stimulated in the periproct (as for instance by a strong light), it would,
according to this rule, tend to move downwards, which is of course
impossible; but as the stimulus never affects all sides quite alike the
result is that the Urchin rotates, turning itself ever away from the
point of strongest stimulation. In the case of Strongylocentrotus
lividus when living on limestone, as on the west coast of Ireland, this
results in the animal excavating for itself holes in the rock, where it is
safe from the action of the breakers.[481]

But it may be objected that no account is taken in the above

description of the action of the "central nervous system," i.e. of the
ring and the radial cords, and yet Romanes found that when they
were removed the escape movements could not be carried out. The
answer is that the central nervous system is a store-house of tone,
not, as in higher animals, a controlling centre for co-ordinating the
movements of the spines. When it is removed at first the escape
movements can be carried out, but in a day or two all tone in the
spine-muscles is lost, and then, since the tone of all is equally low,
there is no tendency in those that are stretched to be more
responsive than others, and hence the escape movements cannot
be carried out. Sea-urchins kept in the tanks of an aquarium are apt
to lose the tone of their spines owing to the poisoning of the nervous
system.

The central nervous system is, however, the system which controls
the movements of the tube-feet. As we have seen, extensions of the
radial nerves run to the tip of each podium. Tube-feet are chiefly
used in ordinary progression; when this is quickened the spines
come into play exclusively. The extent to which these two organs of
locomotion are used varies from genus to genus. Thus
Centrostephanus uses its spines a good deal, Echinus and
Strongylocentrotus very little. The last-named genus sometimes
walks on its tube-feet entirely without touching the ground with its
spines.

The faculty of vision in its simplest form may be defined as

sensitiveness to light and shade. Now strong light acts on all Sea-
urchins as a general skin irritant. They fly from it towards the darkest
corner, and then if it continues the spines rotate. A number of little
violet spines on the aboral pole of Centrostephanus longispinosus
are especially sensitive to light, and hence are almost constantly in
rotation. This is due, according to Uexküll,[482] to a pigment of a
purple colour, which can be extracted by means of alcohol and which
is decomposed by light, the products of decomposition being
supposed to irritate the nerves. Centrostephanus when exposed to
light becomes darker in colour. This is due to the migration outwards
of amoebocytes, which carry a pigment which acts as a screen in
order to prevent the valuable visual purple being too rapidly
decomposed. Not all Sea-urchins, in fact very few of those living in
northern waters, give a reaction to shadow. C. longispinosus is one
of the few; it reacts to a shadow by converging its spines towards it.
A much larger number of species inhabiting tropical waters show this
reaction. It is entirely stopped if the radial nerve-cords be removed,
whereas the reaction to strong light continues. The reaction to shade
is strongest after a long previous exposure to light, hence Uexküll
has given the following explanation of it. The continued irritation due
to light, having spread to all the spines, eventually reaches the radial
cords and is there stored in the bipolar nerve-cells as tone. When the
light-stimulus is interrupted some of the stored tone spreads
upwards to the spines, causing the weak form of spine reaction, and
the spines converge.
 Fig. 233.—To show character and distribution of the sphaeridia in
Strongylocentrotus droëbachiensis. A, a portion of a radius, with sphaeridia,
and the adjoining edge of the peristome. p, Pair of pores for a tube-foot; per,
peristome; t, primary tubercle. B, an isolated sphaeridium. (After Lovén.)

It will be seen therefore that the so-called central nervous system of

Echinus does not act in any sense as a brain, as indeed might have
been guessed from the absence of any differentiation in it. As
Uexküll points out, when an animal is covered all over with similar
organs, such as spines and pedicellariae, capable of acting
automatically, a brain is not needed. The object of a brain is to direct
organs which are in a certain place to a danger which may come
from any quarter, but in the Sea-urchin any spine is as good as any
other spine, and such orientation is not needed. "In a dog the animal
moves its legs, in a Sea-urchin the legs move the animal." What the
Sea-urchin does need is a means to prevent its pedicellariae
attacking its own organs with which they may come into contact.
Thus it possesses an "autodermin," a chemical contained in the
ectoderm which paralyses the muscles of the pedicellariae, as may
be seen by offering to them a spine of the same animal. If, however,
the spine be treated with boiling water, and then offered, it is
viciously seized, showing that this substance can be dissolved out.

Just as in the case of the Starfish, when the nerve-ring is cut

through, the tube-feet in the various radii are no longer co-ordinated
with one another.

Besides the tips of the tube-feet the Urchin possesses another kind
of sense-organ, the sphaeridia (Fig. 233). These are minute glassy
spheres of calcareous matter attached by connective tissue to
equally minute bosses on the plates of the ambulacra, generally near
the middle line. They are in fact diminutive spines, and like the latter
are covered with a thick layer of ectoderm, beneath which is a
particularly well-developed cushion of nerve-fibrils. Only the layer of
muscles which connects a normal spine with its boss is wanting.
Although definite experimental proof is lacking, the whole structure of
the sphaeridia shows that they belong to the category of "balancing
organs." As the animal sways from side to side climbing over uneven
ground, the heavier head of the sphaeridia will incline more to one
side or to another, and thus exercise a strain on different parts of the
sheath, and in this way the animal learns its position with regard to
the vertical.

Intervening between the radial nerve-cord and the radial vessel is a

single radial perihaemal canal (Fig. 232, perih), representing the
two parallel canals found in the same position in the Asteroid. The
five perihaemal canals lead downwards to a space called the
lantern-coelom, surrounding the oesophagus.[483] Since the
skeleton of the corona is composed of plates immovably connected
together, muscles corresponding to the ambulacral muscles of the
Asteroids would be useless, and so the wall of the perihaemal canal
remains thin and the side of it turned towards the general coelom
develops no muscles, and that turned towards the nerve-cord no
nerve-cells. Where, however, the radial nerve enters the nerve-ring,
and on the ring itself, an inner layer of nerve-cells is developed from
the lantern-coelom which represents the lower or oral portions of the
radial perihaemal canals. These cells control the muscles moving the
teeth. These canals are originally parts of the lantern-coelom, but in
the adult they become closed off from it.
 Fig. 234.—Echinus esculentus dissected in order to display Aristotle's lantern, ×
2. The whole upper part of the shell has been cut away. 1, Upper growing
end of tooth; 2, outer forked end of one "compass"; 3, muscle joining
adjacent compasses and acting as elevator of these ossicles; 4, depressor
of the compasses; 5, lower end of jaw; 6, retractor of the whole lantern; 7,
protractor of the whole lantern; 8, auricle; 9, ampullae of the tube-feet; 10,
interambulacral plate; 11, lower part of tooth; 12, water-vascular ring; 13,
meeting-point of a pair of epiphyses; 14, so-called Polian vesicle, really
equivalent to Tiedemann's body in an Asteroid; 15, oesophagus; 16, so-
called ventral blood-vessel; 17, genital stolon; 18, stone-canal; 19, rectum;
20, aboral sinus. (Partly after Chadwick.)

In the outer wall of this space are developed the calcareous rods
forming Aristotle's lantern. These are first: five teeth (Fig. 234, 11),
chisel-shaped ossicles of peculiarly hard and close-set calcareous
matter, the upper ends (1) pushing out projections of the upper wall
of the lantern-coelom. These projections are the growing points of
the teeth, whose lower ends pierce the ectoderm and project into the
lower end of the oesophagus. Each tooth is firmly fixed by a pair of
ossicles inclined towards one another like the limbs of a V and
meeting below. Each ossicle is called an "alveolus," and taken
together they form a "jaw." Their upper ends are connected by a pair
of ossicles called "epiphyses" (13). These two epiphyses meet in an
arch above. The jaws and their contained teeth are situated
interradially. Intervening between successive alveoli are radial pieces
called "rotulae," which extend directly inwards towards the
oesophagus. Above the rotulae are pieces termed "radii" or
"compasses" (2), which are not firmly attached to the other pieces
but lie loosely in the flexible roof of the lantern-coelom.

The uses of the various components of this structure can be made

out from an inspection of the muscles which connect them together.

Overarching each radial perihaemal canal where it leaves the lantern

is a bridge of calcareous matter called the "auricula" (Fig. 234, 8).
This arises as two rods which meet each other in a pent-house over
the canal. It is the only part of the skeleton which can be compared
to the ambulacral ossicles of the Asteroidea, and like them it serves
as the point of insertion for important muscles. Thus we find (1)
protractor (Fig. 234, 7) muscles which arise from the upper ends of
the alveoli and are inserted in the auricula; when these contract they
tend to push the whole "lantern" outwards so as to expose the tips of
the teeth. (2) The retractor muscles (Fig. 234, 6) extend from the
auriculae to the lower ends of the jaws and restore the lantern when
it has been extruded to its original position. (3) The comminator
muscles connect adjacent jaws with one another: these on
contraction approximate the pair of jaws into which they are inserted,
and it will easily be seen that by the successive contraction of the
five comminator muscles a rotating movement of the teeth would be
produced which would cause them to exert an action something like
that of an auger; by their simultaneous contraction the teeth are
brought to a point. (4) The internal and external rotula muscles:
these are small muscles which connect the outer side of the
epiphysis with the rotula. There are two facets on the epiphysis,
which permit it to rock to and fro on the rotula under the action of
these muscles. This rocking action must greatly increase the cutting
power of the tooth. These muscles are controlled by the nerve-ring
and the incipient portions of the radial nerves, which, as we have
seen, have an inner layer of nerve-cells. If the nerve-ring be gently
stimulated on one side the upper end of the lantern bends away from
the spot, causing the lower end, i.e., the teeth, to move towards it;
but a stronger stimulation produces the opposite effect, just as is the
case with spines. But besides these masticatory muscles there are
others which have nothing to do with moving the teeth. These
muscles are attached to the rods called radii or compasses (Fig.
234, 2),[484] which lie in the upper wall of the lantern-coelom, and
may be termed the compass muscles. There are two sets:—(1) The
elevator muscles (Fig. 234, 3), which connect the inner ends of the
compasses with one another. When these contract, the radii tend to
bend upwards at the inner ends and thus raise the roof of the
coelom. (2) The depressor muscles (Fig. 234, 4), which run
downwards from the forked outer ends of the compasses to the
auriculae. Uexküll[485] has shown that the function of these muscles
and of the rods to which they are attached is respiratory. These
muscles are also controlled by the nerve-ring. If this be stimulated by
passing a pin-head into the oesophagus, the roof of the lantern
cavity is raised by the contraction of the elevator muscles. This is
followed by contraction of the depressor muscles lowering it; the
same result may be brought about by placing the animal in water
with excess of carbonic acid. The ten branched gills described on p.
514 are outgrowths of the lantern-coelom. When the roof of this
cavity is depressed the fluid contents are driven out into the gills,
which are thus expanded and then absorb oxygen from the
surrounding sea water. When, on the other hand, the roof is raised
the aerated water is sucked back into the lantern cavity, and the
oxygen passes easily through the thin walls of the lantern into the
fluid filling the main coelomic cavity. There are thus two independent
respiratory mechanisms in the Sea-urchin, the one being the
compass muscles, the other the cilia lining the interior of the tube-
feet.

The function of excretion is performed, as in Asteroidea, by the

amoebocytes floating in the general coelomic cavity. These in part
escape through the thin bases of the gills. In other parts of the body
they seem not to succeed in reaching the exterior at all, but to
degenerate and to form masses of pigment; the colour of the animal
is largely due to these excrementitious substances.

The reproductive system, as in the two preceding orders, consists

of a vertical pillar, the "genital stolon," and a circular "genital rachis"
giving off interradial branches from which the genital organs bud.
The genital stolon is developed from the wall of the general coelom
near the upper end of the axial sinus; it attains a great development
and ultimately completely surrounds the axial sinus, which then
appears like the cavity of a glandular tube, the walls of which are
constituted by the genital stolon. The compound structure consisting
of stolon and axial sinus was actually described as a nephridium by
the Sarasins[486] in the case of Asthenosoma. Its true nature,
however, is shown when the upper end is examined; it is then seen
to open into the stone-canal and to be in communication with the
ampulla, into which the pore-canals open. Lying alongside the upper
end of the axial sinus is the somewhat elongated "madreporic
vesicle," or right hydrocoele, which was described by Sarasin as the
accessory kidney (Nebenniere), since like the axial sinus it is partly
enveloped by the genital stolon. Leipoldt,[487] however, showed
clearly that it is a completely closed space.

The genital rachis springs from the upper end of the stolon, and as in
Asteroids, it lies in the outer wall of a space called the "aboral sinus"
(Fig. 234, 20) intervening between it and the test. In adult specimens
it seems to degenerate. The genital organs are situated at the ends
of five interradial branches of the rachis (Fig. 231, gon). Each is an
immense tree-like structure consisting of branching tubes, which are
lined by the sexual cells. So enormous do they become in the
breeding season that they form an article of food among fishermen.
The term esculentus is derived from this circumstance. Other
species are regularly sold for food as Frutta di Mare (Fruit of the
Sea) at Naples, and as "sea eggs" in the West Indian Islands. One
female Echinus esculentus will produce 20,000,000 eggs in a
season.
The so-called blood system is more distinctly developed in
Echinoidea than in Asteroidea and Ophiuroidea. There is an oral ring
of lymphoid tissue surrounding the oesophagus below the water-
vascular ring. From this are given off two strands, the so-called
"dorsal" (Fig. 231, b.v), and "ventral" vessels (Fig. 234, 16), which
run along the two opposite sides of the stomach or first coil of the
alimentary canal. The position of these strands suggests that like the
lacteals of the human intestine they are channels along which the
products of digestion exude from the stomach. The dorsal strand is
situated on the same side as the genital stolon, and from it branches
are given off which ramify on the surface of the stolon, on account of
which this organ, as in Asteroidea, was at one time regarded as a
"heart," but the distinction of the stolon from the strands is easily
made out. An aboral ring enclosing the genital rachis lies embedded
in the septum dividing the aboral sinus (Fig. 234, 20) from the
general coelom.

Classification of Echinoidea.
The Echinoidea are sharply divided into three main orders, which
differ from each other profoundly in their habits and structure. These
are: (1) The Endocyclica or Regular Urchins, of which the species
just described may be taken as the type. (2) The Clypeastroidea or
Cake-urchins, which are of extremely flattened form, and in which
the periproct is shifted from the apical pole so that it is no longer
surrounded by the genital plates, while some of the tube-feet of the
dorsal surface are flattened so as to serve as gills. (3) The
Spatangoidea or Heart-urchins, in which the outline is oval: the
periproct is shifted, as in the Cake-urchins, and the dorsal tube-feet
are similarly modified; but the Heart-urchins have totally lost
Aristotle's lantern, whilst the Cake-urchins have retained it. This
strongly-marked cleavage of the group was primarily due, as in all
such cases, to the adoption of different habits by different members
of the same group. Were we to term the three orders Rock-urchins,
Sand-urchins, and Burrowing-urchins, it would not be entirely true,
for secondary invasions of the other's territory on the part of each
order have undoubtedly taken place; but still the statement would
remain roughly true, and would give a fair idea of the differences in
habitat which have led to the differentiation of the group.

Order I. Endocyclica (Regular Urchins).

The principal variations concern (1) the peristome, (2) the periproct,
(3) the corona, (4) Aristotle's lantern and its appendages, (5) the
spines, (6) the pedicellariae, and lastly, (7) the tube-feet. We shall
consider these points in order.

Peristome.—In the vast majority of species this region is covered

only with flexible skin in which ten small plates are embedded,
pierced by pores for the buccal tube-feet; besides these there are
irregularly arranged thin plates. In the Cidaridae both the ambulacral
and the interambulacral series of plates are continued on it; these
plates differ from those of the corona in being movable on one
another. In Echinothuriidae only the ambulacral series of plates is
continued on to the peristome. In the case of both these families
there are a considerable number of tube-feet within the region of the
peristome which may be classed as buccal.

Periproct.—This area, which represents the whole dorsal surface of

Asteroidea, is very large in the Cidaridae, where, as in Echinus, it is
covered with leathery skin in which small plates are embedded. In
the Saleniidae it is covered with a single large sur-anal plate, in the
edge of which the anus is excavated; in the Arbaciidae it is covered
with four valve-like plates; whilst in the remaining species its
condition is similar to that described in the case of Echinus
esculentus.

Corona.—In Echinothuriidae all the plates are separated by slips of

membranous skin, so that the test is flexible. In all other families it is
an unyielding cuirass. In the Cidaridae the pore-plates remain
separate throughout life, and are therefore identical with the
ambulacral plates. These are small and placed in two vertical rows,
and so the ambulacra are exceedingly narrow. In Echinothuriidae
there is some tendency to adhesion amongst the pore-plates; these
are of different sizes, and usually one larger and one smaller adhere
to one another. In all other species regular ambulacral plates are
formed at least in the lower part of the radii near the peristome by
the adhesion of the pore-plates in groups of two, three, or more.
Sometimes as many as nine pore-plates may thus adhere.

When adhesion takes place between the pore-plates it is of course

preceded by crowding, and this interferes with their equal
development. Some which extend so far horizontally as to meet their
fellows of the opposite side of the radius are called primary plates;
others which are small and wedged in between the larger ones are
called demi-plates. Systems of classification have been built up
(chiefly by palaeontologists) in which great stress has been laid on
how the primaries and secondaries enter into the constitution of the
compound plate, but it does not seem to the present author as if this
were at all a satisfactory basis for classification. All the pore-plates
are primarily equivalent, and the question as to which are interfered
with in their growth so as to become secondary is trivial. The so-
called Arbacioid type consists of one primary with a secondary on
each side; the Diadematoid type of three primaries, with occasionally
a secondary between the aboral and the middle primary; and finally
the Triplechinoid type of two primaries, with one or more secondaries
between them.

Aristotle's Lantern.—Under this head we may consider the

auriculae and gills as well as the jaws and teeth. In Cidaridae
external gills appear to be absent, but from the lantern coelom large
radial pouches project upwards into the general coelom cavity.
These pouches are supposed to be respiratory, and are termed
internal gills or Stewart's organs.[488] They co-exist with external
gills in Echinothuriidae and in Diadematidae, though in the last family
they are present only in a vestigial form, two being found in each
radius. The auricular arch both in Cidaridae and in Arbaciidae is
composed of two pillars which do not meet, but in the last-named
family they are based, as in Echinidae, generally on the ambulacral
plates, whereas in Cidaridae they arise from the interambulacral
plates (the ambulacral plates being here very narrow). The
epiphyses are absent in Cidaridae and Arbaciidae, and are imperfect
in Diadematidae.

Spines.—These organs are extraordinarily variable, and usually

differ very much in species of the same genus. In the vast majority of
species there is a limited number of long spines called "primaries,"
amongst the bases of which a large number of much shorter
"secondaries" are distributed. In Cidaridae the primaries are very
long and thick and blunt at the ends, and the secondaries form small
circles around their bases. The primaries in Cidaridae and the tips of
the primaries in Arbaciidae and Echinothuriidae are covered with a
special investment of extremely close, hard, calcareous matter very
different from the loosely fenestrated material out of which the
bodies of the spines of all species are composed. In Colobocentrotus
and Heterocentrotus the primaries are very thick and triangular in
section, whilst the secondaries on the aboral surface have expanded
outer ends, which form a close-set pavement protecting the
ectoderm from the shocks of the breakers. In Echinothuriidae the
primaries are short and so delicate as to be termed silky.

Pedicellariae.—In Cidaridae only gemmiform and tridactyle

pedicellariae are found. In the gemmiform the glands lie inside the
grooved blades instead of outside as normally, and they are covered
internally by ingrowths of calcareous matter from the edges. In
Echinothuriidae only tridactyle and trifoliate are found in most
species, but rudimentary gemmiform are found in one species and
well-developed ophicephalous in another. In some species
(Centrostephanus longispinosus) there are found gemmiform
pedicellariae which have lost the jaws but retained the glands. These
are termed "globiferae." Mortensen[489] uses minute details in the
structure of the pedicellariae to discriminate species and even
genera, but in this the present author is not prepared to follow him.