Rupert Glasgow CONCIUSNESS PDF
Rupert Glasgow CONCIUSNESS PDF
Rupert Glasgow CONCIUSNESS PDF
R.D.V. Glasgow
Julius-Maximilians-Universität Würzburg
Würzburg University Press
Universitätsbibliothek Würzburg
Am Hubland
D-97074 Würzburg
www.wup.uni-wuerzburg.de
The present book originally formed a rather long final chapter of a PhD
dissertation entitled The Minimal Self. Once again, I owe a great debt of
gratitude to Roland Borgards, Martin Heisenberg and Karl Mertens of
Würzburg University for their generous guidance and encouragement
throughout that project. Together with Bertram Gerber of the Leibniz
Institute of Neurobiology in Magdeburg, moreover, it was their idea that I
should turn that long, final chapter into a book in its own right. Given the
stamina that would have been required of readers of The Minimal Self to
reach that long, final chapter, I think that the advice was absolutely spot-on.
As with The Minimal Self, special thanks go to Bertram, who has provided
invaluable criticism and feedback as well as support and friendship. It was
Bertram who first encouraged me to pursue my thinking on ‘selfhood’, and
without his influence I doubt whether the book would have even been
conceived (at least by me), let alone written. It goes without saying that I
bear full responsibility for the mistakes and shortcomings.
Again, I should like to thank the organizers and participants in the vari-
ous courses and talks I have given on the ‘The Phylogeny of the Self ’. In
particular, I thank (in alphabetical order) Mirjam Appel, Achim Engelhorn,
Alex Gómez Marín, Apollonia Heisenberg, Manos Paissios, Alois Palmet-
shofer, Teiichi Tanimura and Ayse Yarali. Many thanks also go to Christina
Nath for contributing the beautiful illustrations, to the team at Würzburg
University Press for their great care and skill in preparing the book and to
Christina Kolbe for helping me to track down some of the research material.
I am very grateful, once more, to Bertram Gerber for making the illustra-
tions possible.
A special mention also goes to my lovely friends in Zaragoza and else-
where. Carmen Canales, Las Peñas Altamira and Los Helechos, and Winni
6
Schindler and Sarah Lothian have all coped admirably with my even
more than usually idiosyncratic behaviour, as have Barbara Glasgow in
Berkhamsted and Faith Glasgow and Tony and Mattie Doubleday in Stoke
Newington.
RDVG
Zaragoza, Spain
Contents
I.
A Brief Introduction to Minimal Selfhood
Minimal Selfhood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
II.
Consciousness: Preliminary Considerations
III.
From Motionlessness to Directed Motility
IV.
Appetite and Tacit Selfhood
V.
Where Consciousness is Superfluous
VI.
Limits to Claims about Rudimentary Consciousness
VII.
Epilogue: Consciousness in Simple Animals
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
List of Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
Endnotes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243
I.
A Brief Introduction
to Minimal Selfhood
Minimal Selfhood
(the free-living cell), but also to more complex organisms insofar as they
too are characterized by all three forms of intrinsic reflexivity. Unicellular
organisms such as bacteria and protozoa represent a paradigm of minimal
selfhood: they maintain themselves through an ongoing process of meta-
bolic self-production; they reproduce themselves, for example by period-
ically splitting into two; and they contain themselves by generating their
own membrane and forming functionally integrated bodies. By analogous
arguments, animals ranging from sponges and placozoans to cetaceans and
primates can also be regarded as embodying minimal selfhood.12 It goes
without saying that this is not an exhaustive description of the selfhood of
humans or complex animals. However, more sophisticated forms of animal
or human selfhood are grounded in and presuppose this foundation.
In particular, the parallelisms and correspondences between their
respective forms of self-containment – their physical boundaries and func-
tional integration – cast light both on free-living single-celled organisms
and on multicellular selves such as animals and plants. In neither the uni-
cellular nor the multicellular case is self-containment merely a matter of
enclosure or autonomy. Self-contained though it may be, any particular self
is always reliant on the environment that sustains it with a flow of energy
and matter. The boundary implied by self-containment must thus always
allow nutrients in and waste out, while keeping the internal environment
strictly separate from the external environment. Indeed, there is a sense in
which a boundary or limit is in essence ambiguous, for it cannot help but
link what it separates (in this case self and non-self). To the extent that the
boundary represents the point at which self coincides with non-self, it is
necessarily Janus-faced, infringing the law of identity, and the notion of
self-delimitation or self-containment is inextricably bound up with that of
self-transcendence. Given this inherent ambiguity, a self-containing self is
also a self-transcending self. This is crucial to the possibility of directed
self-movement and, by extension, consciousness.
The best-known illustration at the unicellular level is perhaps provided
by the model bacterium Escherichia coli, where this ‘openness’ specifically
takes the form of transmembrane receptor proteins that can be conceived
as gateways or windows traversing the boundary between self and non-
self. As we shall see below, these membrane-spanning receptors ‘recognize’
certain molecules (amino acids or toxins) on the outside of the cell and in
turn activate the biochemical circuitry on the inside.13 This is what makes
it possible for the cell to direct its locomotion towards what is good for
18 A Brief Introduction to Minimal Selfhood
itself and away from what is bad for itself. Activated by the transmembrane
protein, a protein-based signal pathway involving the addition or removal
of phosphates from a small protein called CheY regulates the rotary motor
activity of the bacterium’s locomotory flagella, causing the cell either to
‘tumble’ (i.e. change direction in search of food) or to continue swimming in
the same direction. E. coli has some ten thousand such membrane-spanning
receptors clustered mainly at one end of the organism, thanks to which it
inhabits a chemical universe comprising over fifty attractants and repellents.
As a consequence of this sensitivity, even the ‘slightest whiff ’ of its preferred
amino acid will elicit a change in its swimming activity.14
It is this openness to its environment that permits a hungry organism
not just to stumble or grope around at random, but to engage in intentional,
directed behaviour, i.e. to go after what is good (for itself). By sensing
environmental non-self, the organism can be said to reach out beyond its
boundaries. To the extent that a self-containing self thus also transcends
itself, it is able to behave (i.e. to move itself) in a manner that conforms with
its interests and fosters its self-perpetuation. Such self-transcendence can
also be conceived in terms of indexicality. While the concept of ‘self-con-
tainment’ entails that I am always here now (this is tautologically true of the
first-person perspective associated with selfhood: here is where I always am;
now is the time it always is), my self-transcendence allows for me to be there
and then (i.e. to be intentionally directed towards wherever my next meal
is likely to be in the future).
In a multicellular context, this openness manifests itself immediately in
our sense of touch. Our skin is not just a container that holds us in, but
one of the ‘main sensory portals’15 by which we open onto the world. The
human dermis contains a variety of specialized receptor cells that commu-
nicate with the central nervous system about the external world and the
state of the skin. These include mechanical pressure receptors, temperature
receptors and diverse pain sensors that alert us to the presence of potentially
harmful physical stimuli and to inflammation and injury, prompting us to
recoil from what might threaten our bodily integrity and to protect areas
where damage has already occurred. Yet even the single-celled Paramecium
has something akin to a sense of touch, reacting to a bump from behind by
speeding up its swimming and to a thump at the front by swimming off in
a new direction. These responses are mediated by movements of charged
ions through special channels in the cellular membrane that cause changes
in the beating of the organism’s cilia. Such ion channels, it is thought,16
Minimal Selfhood 19
Consciousness: Preliminary
Considerations
Consciousness and Behaviour
The aim of this brief treatise is to show that such minimal selfhood
– as embodied in the threefold intrinsic reflexivity of self-maintenance, self-
reproduction and self-containment23 – grounds the possibility of con-
sciousness. It will seek to do so first by arguing that the minimal selfhood
of single-celled protozoa such as amoebae, ciliates and flagellates24 may in
certain (but not all) cases beget an elementary or rudimentary25 form of
consciousness. It will then proceed to apply this same analysis to a number
of the simplest multicellular animals. The converse of these claims is that
the rudimentary consciousness of these protozoans and simple metazoans
– like the more complex forms of consciousness for which such rudimentary
consciousness provides the foundation – in turn presupposes minimal self-
hood. In experiential terms, this will be seen to manifest itself as what has
been termed ‘tacit’ selfhood (or sometimes ‘pre-reflective self-awareness’).
This refers to an implicit bodily self-familiarity: the appetites and aversions,
drives26 and dispositions that shape and structure a selfish perspective on
the world.
The argument, however, is not that all minimal selves exhibit conscious-
ness all the time, and the underlying task will be to distinguish selfhood
that does from selfhood that does not. Certainly, a conception of selfhood
in terms of the intrinsically reflexive relationship that a self has to itself also
implies an intrinsic relationship to the non-self to which it is structurally
coupled.27 The deep dependence of any metabolic entity upon meaningful
non-self for energy and nutrition thus prompted the philosophical biologist
Hans Jonas to propose that ‘world’ is there from ‘the earliest beginning’,28
thereby including passively drifting micro-organisms or sedentary plants
within the realm of sentience. There is a sense indeed in which all living
selves – and even merely ‘self-like’ phenomena such as self-organizing dis-
sipative structures – must necessarily be open or responsive to the changing
environmental non-self around them in order to perpetuate themselves.
The self-concern or care that is essential to selfhood is wed to the presence
of a ‘world’ upon which the self depends for energy and sustenance but that
constantly threatens, one way or another, to put an end to its striving to
maintain itself.
This striving may manifest itself as various forms of self-adaptation,
ranging from homeostatic self-regulation in the face of environmental fluc-
tuation to the phenotypic plasticity shown by organisms able to modify their
internal physiology or morphology so as to cope with variations in external
conditions. The form of self-adaptation that is relevant to consciousness,
Consciousness and Behaviour 25
distinguished man from the rest of the animal kingdom, whose movements
– whether microscopic or macroscopic – were performed mechanically and
without the aid of a ‘mind’.
More recently, philosopher John Searle has espoused what he calls ‘the
principle of the independence of consciousness and behavior’, according
to which there is no ‘conceptual or logical connection between conscious
mental phenomena and external behavior’.52 This principle is grounded in
the perfectly licit realization that certain sorts of behaviour, such as pain
behaviour, can be feigned, whereas conversely certain sorts of mental
states, such as genuine pain, can be concealed. Yet both simulation and
dissimulation are themselves categories of behaviour. The fact that behav-
ioural strategies may be highly complex (especially once they incorporate
the dimension of intersubjectivity, i.e. the capacity of one self to view itself
from the perspective of another self, with the concomitant opportunities for
deceiving that other self)53 does not invalidate the close logical connection
between mental phenomena and the behaviour through which they come
to expression. It simply means that the correlation between them may not
always be straightforward, and considerable interpretative expertise may
be required to decipher the mental phenomenon underlying any particular
behaviour. By the same token, pathologies such as locked-in syndrome
(where there is consciousness more or less completely without movement)
in no way disprove that consciousness is conceptually or causally bound up
with its behavioural manifestations. The fact is simply that behaviour can
break down.54 It will be argued in the following pages that appropriately
directed, self-caused self-movement (i.e. behaviour in the sense of ‘action’)
and the consciousness it both presupposes and expresses exist in such a
tight logical embrace that neither can be conceived without the possibility
of the other.
A Scale of Consciousness
Yet there are other arguments that can be drawn upon to deny conscious-
ness to unicellular organisms. Perhaps the most extreme such argument
is that consciousness does not exist anyway. This was a claim made by the
psychological behaviourist John Watson,55 who held the behaviour of all
living things to be a product merely of conditioning. Maintaining that we
do not consciously act but merely react to stimuli, Watson extended to the
whole of the human and non-human animal kingdom the sort of argument
normally reserved for micro-organisms or ‘lower’ animals. Indeed, his dec-
laration that the behaviourist recognized ‘no dividing line between man and
brute’56 could have been rephrased to proclaim the continuity between ‘man
and amoeba’. Watson’s asseveration that consciousness was a metaphysical
fiction might have been felt by some to be liberatingly egalitarian in divest-
ing the ‘high’ and the ‘low’ alike of any pretensions to mindedness. Yet it is
of course a metaphysical claim in itself57 and a supremely counterintuitive
one at that.
A more common approach to denying the possibility of single-celled
consciousness has been to regard the phylogenetic ‘scale’58 as in some sense
coincident with a ‘scale’ of consciousness. As we descend towards the ‘low-
lier’ forms of life, there comes a point in the hierarchy where the ‘light’ of
consciousness flutters and is eventually extinguished. Philosopher of mind
Michael Tye thus opens his paper ‘The Problem of Simple Minds’ with a
sequence of questions: ‘Are frogs conscious? Or fish? What about honey
bees? Do paramecia have experiences? Somewhere down the phylogenetic
scale consciousness ceases. But where?’59 Not surprisingly, Tye’s answer is
A Scale of Consciousness 33
that consciousness has already ‘ceased’ long before we reach the depths
represented by paramecia, which are considered bereft of all flexibility in
their behaviour. ‘Tropistic’ organisms, which here seem to include the ranks
of plants, protozoa and rigidly phototactic animals such as caterpillars, ‘feel
and experience nothing’; they are ‘full-fledged unconscious automata or
zombies’.60 Yet Tye’s argument, like Fodor’s, relies on the assumption that
all the behaviour of all such ‘lower’ organisms can be interpreted as a mere
reflex, a tropistic or tactic response to a stimulus. It can accordingly be
refuted by showing that the behavioural repertoire of certain organisms
includes just one mode of activity that cannot be reduced to reflex or taxis.
This is what the present argument will attempt.
However, the notion of a ‘scale’ – perhaps of body size, brain size or com-
plexity – remains appealing to common sense. The idea that paramecia or
dinoflagellates are philosophical ‘zombies’61 is certainly less disconcerting
to the modern-day sensibility than the idea that one’s lover, child or dog is
a zombie. Cetaceans and great apes also seem privileged with subjectivity,
whereas amphibians and fish – expressionless as they are – appear to reside
somewhere in between.62 Most people, I suspect, would not hesitate long in
flushing insects and arachnids down the plughole of zombiehood, whereas
rotifers and nematode worms, if given a second thought, would surely be
deemed too small to harbour a mind (after all, where would they keep it?).
But what is so special about size? Certain large unicellular organisms are
bigger than a whole range of small multicellular animals. Ciliates such as
Stentor or Neobursaridium may attain sizes of around a millimetre, overlap-
ping in range with metazoans such as rotifers, gastrotrichs, nematodes and
tardigrades.63 Amoebae may be similar in size, dwarfing the more Lillipu-
tian of insects: not only the diminutive parasitic wasps known as fairyflies,
which are blind and wingless and are scarcely likely to require much in the
way of consciousness anyway, but even the 170 µm-long hymenopterans of
the genus Megaphragma, which exhibit behaviours such as flight, feeding
and the ability to search for hosts for oviposition.64 H. S. Jennings drew
attention to the size chauvinism implicit in our customary dismissals of
amoeban consciousness, noting that if an amoeba were the size of a whale
we would not think twice about granting it states such as pleasure and pain,
hunger and desire. We would be foolish to do otherwise in the presence of
a whale-sized ‘beast of prey’.65
Equally, what is so special about complexity? The philosophers M. R.
Bennett and P. M. S. Hacker describe our tendency to be led astray by the
34 Consciousness: Preliminary Considerations
partly a result of certain philosophers being misled by the fact that when-
ever they reflect on their consciousness, their consciousness necessarily
assumes the form of a reflective consciousness, couched in the form of an
internalized dialogue.79 In reflecting on this reflective process, such philoso
phers have thus shown a recurrent tendency to forget that consciousness
is not primarily an attribute of reflective philosophers. By contrast with
such higher-order theories, a first-order theory of the sort I am advocating
holds that mental states or processes can be described as ‘conscious’ not on
account of the subject’s higher-order awareness of them, but because the
states or processes in themselves make the subject aware of – and able to
behave in – the external environment.80
In fact, not all ‘higher-order’ theories of consciousness require linguistic
thought. A divisive issue among the proponents of such models has been
whether the higher-order, meta-mental states that transform an unconscious
first-order mental state into a conscious one should be considered thought-
like or perception-like.81 On the latter view, perceptions and feelings become
conscious if they are the object not of a higher-order linguistic thought, but
of some form of higher-order inner sense or perception. Although this view
is not necessarily so radical in its exclusivity (in that it does not presuppose
linguistic ability), in both cases consciousness is understood as arising
when the mind directs its attention upon its own states and processes. Both
cases, however, overlook the logical problem raised by a scenario in which
one mental state takes another, different mental state as its object, where
this implies the occurrence of two mental states that are necessarily distinct
from one another – and yet the one has to ‘recognize’ the other as pertain-
ing to the same ‘self ’.82 The unanswered question is how to account for the
self-recognition that such models require for a mind to be able to focus on
its own mental states as its own.
Meta-mental theories fail to get to the bottom of consciousness because
any such recognition of self by self logically presupposes and is thus derivative
upon the intrinsic reflexivity of a self that is in some sense already familiar
with itself, the tacit selfhood or pre-reflective self-awareness broached above.
In the course of the present treatise, we shall look more closely at some of
the diverse forms of this non-representational, non-reflective self-awareness
that provides the logical underpinning for consciousness. We shall examine
below how it expresses itself, for example, in the appetites, emotions, pains
and immediate self-familiarity of the body that one is. Appetite in particular
will be seen to reside at the very heart of elementary consciousness.
Consciousness, Meta-Mental States and Tacit Selfhood 39
Thompson’s point is itself indicative of the other major cut-off in the ‘ascent’
that leads to human consciousness as its most glorious expression. This is
the distinction between multicellular animals – endowed with a brain – and
‘brainless’ single-celled organisms. Of course, the cut-off is not a clean one.
Not all animals have brains. Ancestral metazoans such as sponges and jelly-
fish either do without neurons altogether or have diffuse neural nets rather
than brains, whereas other metazoans seem to have evolved and then ‘lost’
their brain in the process of natural selection.90
Again, however, our tendency to privilege the brain makes intuitive good
sense, given the close relationship between consciousness and the brain in
humans (evidenced by the effects of brain damage and the workings of drugs)
and the remarkable structural and biochemical similarities between how
human and non-human brains respond to environmental contingencies.
Such similarities are lacking in our relationship with single-celled organ-
isms, which are decidedly more alien to us. In his account of the evolution
of mind, neurophysiologist Rodolfo Llinás thus writes that ‘neurons arose
within the space between sensing and moving: this space mushroomed to
become the brain’,91 endowing us with flexibility and adaptability rather
than restricting us to an invariant spatial relay of information between
stimulus and response. Yet unicellular prokaryotes and eukaryotes are
likewise equipped with a logical space between sensing and self-movement:
in this case it is the reversible phosphorylation of proteins that allows these
proteins to act as molecular ‘switches’, guiding cellular processes in one
Consciousness and Brains 43
the two hemispheres of their brain surgically severed.105 One might also
cite the ‘perspectival drift’ and ‘attentional disturbance’ characteristic of
schizophrenia, a ‘failure to stay anchored within a single frame of reference,
perspective, or orientation’.106 Like auditory hallucinations and passivity
experiences, such episodes are believed to have their origin in disorders of
tacit selfhood affecting mechanisms of corollary discharge or proprioception
that are normally taken for granted.107 It is above all in such contexts that
fragmented forms of consciousness are conceivable, i.e. as a consequence of
fragmented forms of selfhood.
The question of cognitive binding is one of seven ‘easy’ problems famously
identified by David Chalmers in an influential essay on consciousness. In
designating these problems ‘easy’, Chalmers means that they are ‘directly
susceptible to the standard methods of cognitive science, whereby a phe-
nomenon is explained in terms of computational or neural mechanisms’.108
As well as the integration of information by a cognitive system, these
‘easy’ problems include the ability to discriminate, categorize, and react to
environmental stimuli; the reportability of mental states; the ability of a
system to access its own internal states; the focus of attention; the deliberate
control of behaviour; and the difference between wakefulness and sleep. It
is noteworthy that while one of these problems (the reportability of mental
states) relies upon the possession of language, and another (internal access)
falls back on a ‘meta-mental’ model of consciousness, the five other easy
problems are all – as I hope will become clearer below – in some degree
pertinent to single-celled organisms.
Chalmers contrasts the ‘easy’ problems of consciousness with what he
calls the ‘hard’ problem, which he associates with the subjective or experien-
tial nature of consciousness, the notion of it being like something to be a con-
scious organism. This hard problem is considered so intractable that it has
led some thinkers to conclude that a theory of consciousness will be forever
beyond our grasp. Chalmers’ own response is to suggest that consciousness
can only be understood in non-reductive terms as something fundamental,
in other words as something that – like mass or space-time – cannot be
explained in terms of anything simpler. It is the notion of ‘information’ that
provides this explanatory bedrock. Chalmers’ self-confessedly speculative
proposal is what he calls the ‘double-aspect’ theory of information, which
is based on the observation that information has both a physical and a
phenomenal aspect.109 Among a raft of other questions,110 however, one is
Consciousness and Brains 47
again left wondering who or what the information is for? Surely information
only has a phenomenal aspect when it is being used by a self for that self in
the pursuit of interests that matter to that self.
In fact, questions such as Nagel’s111 about what it is like to be a bat are
not necessarily so intractable. For a start, they generally only make sense if
a context is specified. The question thus needs rephrasing as: what is it like
to be such-and-such an organism in such-and-such a context or performing
such-and-such an activity? In more concrete terms, one might rephrase
it as: what is it like to be such-and-such an organism in such-and-such a
context having just eaten copiously, as opposed to undergoing a longer-
than-usual period of food deprivation? The difference between the two
scenarios can be empirically observed in the conduct (no words are needed)
of a hungry as opposed to a satiated human, but also in the behaviour of
other, non-human and arguably even microscopic predators. The ascription
of hunger in one scenario (food deprivation) and not in the other (satiety)
implies an empathetic bridge that may span the width separating us from
the most phylogenetically remote of organisms. Question: what is it like to
be a food-deprived bat or amoeba in the presence of, or within perceptual
range of, a potential prey? Answer: it is like being hungry.112 So how far
are we prepared to go in attributing appetites and aversions to non-human
organisms?
I shall argue below that a capacity to experience various degrees of appe-
tite, aversion, motivation, pleasure and discomfort is not only something I
have in common with other human beings and bats, but also perhaps the
only thing – or one of only a very few things – that I have in common with
amoebae, euglyphids and dinoflagellates. Concepts such as appetite, moti-
vation and pleasure lay the groundwork for a potentially informative answer
to the question of what it is like to be an organism other than a human.
III.
From Motionlessness to
Directed Motility
Non-Movers and the Almost Immobile
made a lifestyle ‘choice’, finding a particularly stable niche that relieves them
of the need to ‘live freely’ (i.e. move) or have much, if any, awareness of their
surroundings. The various other strategies of unicellular or multicellular
self-adaptation are neither more nor less ‘sophisticated’ than motility.
Such adaptability may take the form, for example, of what is known as
phenotypic plasticity, which allows micro-organisms to grow and flourish
in a wide variety of physiological conditions rather than necessarily having
to go and find more favourable conditions if a key nutrient is missing. The
most intensely studied example is the ability of E. coli to change to using
lactose as a source of nutrition in the absence of the more habitual glucose.
As soon as lactose is encountered in a glucose-free environment, a genetic
switch is thrown and the appropriate lactose-digesting enzymes are pro-
duced, a process that occurs even if the bacteria have never previously come
across lactose.115 In such conditions, the genetically controlled fine-tuning
of metabolism enables the bacteria to stay put and renders locomotion
superfluous. Human beings benefit in particular from the phenotypic plas-
ticity exhibited by symbiotic gut bacteria such as Bacteroides thetaiotaomi-
cron, which are endowed with a remarkable capacity to vary the digestive
enzymes they produce according to cues in the intestinal environment.
This bacterial versatility redounds to our advantage in that it permits us to
degrade otherwise indigestible plant polysaccharides and thus broaden the
range of carbohydrates we can assimilate.116
Perhaps the most versatile of all micro-organisms is a rod-shaped
bacterium called Rhodopseudomonas palustris, which belongs to the class
of ‘purple non-sulphur bacteria’.117 The remarkable biochemical flexibility
and nutritional adaptability of this single cell, endowed with a genome of
just 5.5 Mb, enables it to swap between the four fundamental modes of
metabolism,118 getting by as required with or without oxygen, adopting
an ancestral, anoxygenic form of photosynthesis or feeding on organic or
inorganic compounds. By way of a rough comparison, it is as though we
humans were able to switch, when necessary, from powering ourselves
heterotrophically on organic compounds (as we do in our capacity as ani-
mals) to doing so autotrophically by means of photosynthesis (like plants),
before in turn changing to inorganic energy sources such as hydrogen sul-
phide, elemental sulphur or molecular hydrogen, while still keeping another
nutritional option (‘photoheterotrophy’) up our metabolic sleeves for the
lean times to come. Biochemically and metabolically, R. palustris is the most
complete jack of all trades, resisting the tendency to specialize to which
Non-Movers and the Almost Immobile 53
other, possibly less ancient lineages have yielded. The R. palustris way is to
leave its metabolic options completely open. Such biochemical versatility
makes directional locomotion unnecessary insofar as the creature in ques-
tion is likely to be able to extract energy from the environment wherever
it happens to be. Exempt from the need to search for nutrition elsewhere
(wherever ‘elsewhere’ may be), consciousness of any ‘elsewhere’ will thus
be surplus to requirements. Yet in metabolic terms, writes microbiologist
Harald Brüssow, ‘we are dwarfs with respect to this bacterium’.119
Another form of self-adaptation available to bacteria occurs not at
the level of the individual cell, but rather that of the collective or lineage.
Specifically, the phenomenon of hypermutation or adaptive mutagenesis
confers upon bacteria an increased chance of communally coming across a
beneficial mutation that will help them to cope with variations in local envi-
ronmental circumstances. Such bacterial group intelligence is augmented
by the ability of bacteria to engage in the apparently purposive exchange of
genetic information by means of non-genealogical horizontal gene transfer
(HGT), possibly with phages (i.e. bacterial viruses) serving as a manner
of ‘genetic repository’120 that can be used to optimize their efficiency in
self-defence and self-maintenance.
Accomplishments of this sort have led bacterial geneticist James Shapiro
to describe bacteria as ‘sophisticated natural genetic engineers’, concluding
that ‘even the smallest cells are sentient beings’.121 Biological physicist Eshel
Ben-Jacob referred to the bacterial genome as ‘an adaptive cybernetic unit
with self-awareness’.122 Such claims raise not necessarily straightforward
questions relating to whether self-adaptation that is essentially communal
can be associated with any manner of ‘agency’ on the part either of the
individual cells or the genome. Above all, a multicellular logic seems to be at
work, and it perhaps makes more sense to interpret this genomic adaptabil-
ity as the property of a collective self rather than of the individual bacteria.
Nor does it seem necessary to propose the existence of consciousness in
such a context. Although there is no doubt that the process of collective
self-adaptation involves the cognitive processing of informational input,
the scenario has more in common with the metabolic self-adaptation of a
bacterium responding to a shortage of glucose. It does not entail directed
self-movement so much as non-locomotive self-modification.
To reiterate: the basic idea underlying this approach to elementary
consciousness is that a capacity for conscious experience of the world can
be ascribed to an intrinsically reflexive being (a self) precisely to the extent
54 From Motionlessness to Directed Motility
in question not only throws up logical conundrums (in that the ‘movement’
is in fact the displacement of an identical replica) but fails to comply fully
with the criterion of genuine self-causation. Although the retroelement
encodes the enzymes needed for getting itself moved, it is crucially depend-
ent on a highly specific cellular environment that provides the requisite
chemistry and energy. In short, the ‘jumping gene’ does not perform work
in hopping from A to B; its movement is made possible merely by its own
largely invariant configurational features within the framework provided
by the metabolic activity of a self-maintaining host cell. It can jump simply
by virtue of remaining as it is rather than engaging in any form of activity.
Selfish though such genetic material may be, the lack of a self-maintaining
metabolism of its own can be taken to debar mobile DNA from full minimal
selfhood. The same applies to viruses. The reflexivity of a virus or phage is
thus indirect rather than direct, manifesting itself as a capacity to get itself
replicated, assembled, coated and above all moved. Although it is essential
for a recently assembled virion to find, recognize and penetrate a new host
in order to perpetuate the lineage, this movement is not a consequence
of behaviour or action on the part of the virion in question, which relies
instead on environmental contingency, with random encounters assisted
by diffusion and convection currents, as well as possible additional help
from self-encoded ‘movement proteins’.129 Most importantly, there is again
no investment of energy or performance of work: no drive or reward. When
the E. coli bacteriophage T4 is described as using its ‘host cell recognition
sensors’ to ‘recognize’ a potential host and then attach to the cell surface,130
this sensory activity – for all its complexity131 – can be fully understood
without any need to invoke consciousness: the phage ‘recognizes’ and
‘senses’ its host, but it is not conscious of it.132 Here we may indeed speak of
philosophical zombie-hood.
In some instances it is the viral factory – a protective structure inside
the host cell within which viral replication and assembly take place – that is
thought to provide a better expression of viral ‘selfhood’ than the infective
virion.133 On this view, the viral ‘self ’ is confined to an intracellular existence,
again ruling out genuine movement and dispensing with the possibility or
need for consciousness. In this sense, such viral factories are reminiscent
of bacterial endosymbionts such as Buchnera aphidicola and ‘Candidatus
Tremblaya princeps’,134 which enjoy a life of intracellular plenitude but
forfeit (almost) all behavioural or locomotive versatility.135 In the case of
these cells-within-cells, moreover, it is not just that they do not have to
56 From Motionlessness to Directed Motility
move anywhere to find their nutrition; they do not even have to operate a
mouth. Such cells acquire their nutrients by means of an ongoing process
of predominantly passive exchange determined by chemical concentration
gradients rather than any activity on their own part.
More generally, a distinction can thus be drawn between osmotrophy,
which refers to the uptake and assimilation of dissolved organic compounds
across the cell membrane by means of osmosis,136 and phagotrophy, which
involves the active engulfment of food in particulate form. Phagotrophy
may take the form of phagocytosis by pseudopodia in the case of amoe-
bae, in which case the amoeba as a whole is its mouth (and the amoeba is
described as ‘pantostomate’). Alternatively, ingestion may be restricted to
a specific part of the cell, a mouth or groove known as a cytostome that is
specialized to perform the task of engulfment, as in some ciliates. Further
variants exist. Among dinoflagellates three categories of feeding mechanism
are known, including not only direct engulfment of intact prey, but also the
deployment of some form of tube (usually a cytoplasmic ‘peduncle’) to suck
the contents out of prey, or a ‘pallium’ (a type of pseudopod formerly known
as a ‘feeding veil’) to envelop the prey and digest it extracellularly.137
In all these cases, work is required, performed in the main by the cellular
cytoskeleton, and there are clear parallels between the work involved in
locomotion (a search or pursuit) and that required for the ingestion of
prey (the self-contortions of engulfment). Nor is the extent of this work
to be underestimated. Rapacious ciliates such as Didinium feast on prey
(often Paramecium) that may be much bigger than the predator, requiring
an expandable cytostome to engorge their quarry. Dinoflagellates too are
renowned for devouring whole prey that may be up to five times their own
size.138 The operation of a mouth can thus be regarded as a significant form
of self-movement. The work involved suggests appetite, motivation and
the possibility of reward. To the extent that osmosis is purely passive, by
contrast, effort is unnecessary and motivation is superfluous. No reward is
needed.
The Sessility of Plants
but the first of which are present in plants.143 As living beings endowed with
souls, moreover, plants are characterized by what Aristotle terms ‘entelechy’,
an intrinsically reflexive term that implies the inherent possession of an
‘end’. Plants are ends in themselves, in other words, existing for their own
sake (i.e. as selves) rather than merely for the sake of humans.144 Aristotle’s
pupil Theophrastus, widely known as the father of botany, goes further in
his appreciation of plants as autonomous, intentional selves that actively
strive to pursue their own interests.145 This contrasts starkly with subsequent
thinkers such as Hegel, who divests plants of selfhood and in the process
justifies their instrumentalization for human purposes: by means of fruits,
he writes, ‘the silent essence of self-less (selbstlos) Nature … offers itself to
life that has a self-like (selbstisch) nature’.146
Yet plants are selves: they maintain themselves and reproduce themselves,
and although the element of self-containment may be more indeterminate
(resulting in uncertainty about the relevant unit of selfhood), it is not
absent.147 Moreover, plant growth may be considered a form of self-adap-
tation, i.e. a mechanism by which each organism modifies itself according
to environmental circumstances in such a way as to maximize its wellbeing
and fitness. Modern-day botanists refer to the ‘phenotypic plasticity’ of
plants, meaning their capacity to exhibit flexibility not only in physiology
– i.e. rates of photosynthesis and transpiration – but also in morphological
development in a way that presupposes both an assessment of the external
conditions and the selection of the best response. The question is whether
such growth and plasticity imply the presence of plant consciousness.148 The
answer depends on whether self-adaptation of this sort can indeed be classi-
fied as a mode of self-movement. There are some good arguments to suggest
that it can. As plant scientists Anthony Trewavas and František Baluška149
and philosophical botanist Matthew Hall have argued, it is the roots that
play a pivotal role in plant movement and cognition:
Rich soil patches are exploited by increased plastic root branching and
root growth. In the presence of few nutrients, root growth has been
found to accelerate in order to facilitate the detection of new, more
nutritious patches of soil in other locations. There is clear and active
perception of the resources available, which ... involves the construc-
tion of a ‘three dimensional perspective’ of the local space. Here plants
display their behavioural intelligence with an ongoing assessment of
the costs and benefits involved in exploiting the resources that exist
in the soil. ... Root plasticity allows plants to make choices about the
The Sessility of Plants 59
soil patches they feed in – to the extent that plants have been referred
to in ecological studies as ‘foragers’.150
If Jonas is right, the constant contiguity or adjacency of the root and its
nutrition means that there is no space for the emergence of appetite, desire
and thus the fulfilment of a goal. Whereas the plant is immersed in the
immediate satisfaction of its organic needs, therefore, the ‘great secret of
animal life lies precisely in the gap which it is able to maintain between
immediate concern and mediate satisfaction, i.e. in the loss of immediacy
corresponding to the gain in scope’.155 One might query whether it even
makes sense to speak of a search for environmental nutrients – and thus
the possible awareness of such nutrients – if there is no gap between the
searcher and the sought-for. Expressed in epistemological terms, conscious-
ness presupposes a degree of separation between subject and object.156
Plant-like and Animal-like Unicellulars
the use of either existent or specially generated water currents from which
food particles and suspended matter are strained; diffusion feeding involves
the use of bodily extensions such as axopods with which approaching prey
collide and to which they subsequently adhere; raptorial feeding involves the
pursuit and active engulfment of prey.163 Requiring hunt and capture, it is
the latter that is most suggestive of a creature armed with an awareness of
its surroundings, and we shall return to this strategy in subsequent sections.
Diffusion feeding is typified by radiolarians and heliozoans, etymologi-
cally known as ray animalcules and sun animalcules respectively,164 whose
delicate cytoplasmic axopods protrude radially from a central body to
capture the prey that inadvertently gets stuck to them. It is also illustrated
by the suctorians, sessile ciliates provided with sticky, extended tenta-
cle-mouths to which passing protozoa adhere before having the cytoplasm
sucked out of them.165 Although the strategy is largely passive, some work is
required, for the prey has to be incorporated or engulfed by the predator’s
body and then transported – by cytoplasmic flow – to the inner part of the
cell for digestion; one might perhaps speak of internalized self-movement
rather than the externalized self-movement of locomotion. Nevertheless,
there seems little need to posit consciousness of a ‘world’ on the part of the
predatory organism. An item of prey only becomes relevant to the organism
once it is already physically adjacent and adhering to it and on its way to
being materially assimilated. A similar strategy is employed by animals
such as the Gorgon’s Head basket star, a type of echinoderm that perches
in a prominent position, extends its arms as though to create a Medusan
bouffant, and ensnares small crustaceans that venture too close.
In itself, filter feeding also suggests a broadly sessile or passive life-
style and a non-discriminatory process of ingestion that renders sensory
awareness superfluous. Yet there are great variations in the amount of work
performed by different sorts of filter feeder. Within the slightly broader
category of ‘suspension feeding’,166 a distinction has been drawn between
active and passive forms according to whether ambient water currents
are exploited to drive water across the filtering apparatus or whether the
organism’s own metabolic energy produces the flow.167 Such currents may
be generated by appendages such as cilia but also by the movement of the
animal as a whole, rather in the manner of the filter-feeding baleen whales
that swim through areas rich in zooplankton, gulping or gaping while their
baleens sieve prey from the water. As in the case of animals, there are some
unicellular filter feeders such as Paramecium that likewise pursue their prey
Plant-like and Animal-like Unicellulars 63
in an actively motile manner (sweeping water with bacteria and algae into
their oral groove as they move along), whereas others such as Vorticella
tend rather to be sessile, relying on environmental currents to bring the sus-
pended material to them. Among the flagellates, filter feeding is generally
carried out by sessile organisms.
Even the lack of discrimination sometimes imputed to bacterivorous
filter-feeding ciliates may not be as all-embracing as once thought. It was
initially assumed that the feeding of such ciliates was predominantly ‘automa
tized’ and that they were unable to differentiate between different kinds
of particles except in terms of their geometrical properties.168 Although
filter-feeding ciliates may indeed lack the ‘discernment’ of raptorial predators
attacking individual prey items, and certainly the shape and size of potential
prey are a ‘first-order’ determinant in particle selection,169 there is evidence
that biochemistry – the ‘taste’ or ‘smell’ – plays a role as well. Researchers
have shown that prey items of similar morphology ‘are not always ingested
from mixtures with equal alacrity by filter-feeding ciliates’, putting such
selectivity down to ‘biochemical differences perceived by the grazer’.170 In
addition to being able to differentiate among morphologically similar prey
types in mixed assemblages, such ciliates can distinguish live from dead
bacteria, discriminate between living cells and inert microspheres, and on
occasion actively reject unattractive prey items in the course of processing
them.171
The aversive behaviour exemplified in particular by the filter-feeding
ciliate Stentor, the trumpet animalcule (illustration page 160), likewise sug-
gests that the life of a filter feeder need not be a matter merely of meek and
undiscriminating quiescence. In a famous experiment conducted by H. S.
Jennings, a specimen belonging to the species Stentor roeseli was bombarded
with a stream of potentially harmful carmine particles. In response to this
noxious chemical stimulus, Stentor went through a whole sequence of adap-
tive measures, first ingesting the particles, then trying (several times) to
bend away from the source, then reversing the motion of the cilia around its
mouth so as to drive the particles away, then trying (several times) to contract
its whole body towards its point of attachment to the substrate, before even-
tually – when harassed persistently enough – making a violent contraction
of its whole body and swimming away. Once it had settled down again, Sten-
tor proceeded to ‘explore’ the local terrain, or so it seemed, until it had come
across a suitable site less bothersome to it, where it duly embarked upon a
series of elaborate measures to establish and construct a new tube to live
64 From Motionlessness to Directed Motility
in.172 The persistence of Stentor’s endeavours to avoid what was ‘bad for self ’
– and the intricacy of its behavioural response – represent a marked contrast
with more genuinely sessile or plant-like ways of life.
An underlying distinction among feeding strategies is thus between those
that require no energy expenditure on the part of the organism (involving
sessility or the random dispersal to which Brownian motion may subject
small free-floating bacteria) and those that entail active locomotion. This
in turn may take the form of either directional or non-directional move-
ment. As a manner of indiscriminately ‘aroused’ behaviour, undirected
locomotion does not in itself suggest consciousness, precisely because it
does not presuppose discernment between one location and another, or
between directions that are ‘better’ and ‘worse’ for self. Such behaviour was
traditionally ascribed to the voracious hunter-ciliates of the genus Didin-
ium, which were believed to move around at random eating whatever they
bumped into and were physically capable of ingesting. Jennings described
the ‘process of food-getting’ in Didinium as ‘one of trial of all sorts of things’.
Denying that it perceived its prey at a distance or took a decision to attack
certain organisms, he posited that Didinium simply tried everything out
and held fast to what was good.173 More recent research has suggested that
Didinium is not in fact a ‘random contact hunter’ but shows discrimination
and chemotactic sensitivity.174 As shown by the motile filter-feeding ciliates,
however, the strategy undoubtedly makes sense to the extent that – given
certain size requirements175 – even haphazard movement increases the
likelihood of encounters with potential prey.
A corresponding phenomenon in the plant world is nastic movement,
such as the so-called thigmonasty of carnivorous plants, an unvarying and
undirected reaction to touch or vibration. The non-directional movement
characteristic of the Venus flytrap (the trap merely closes) makes it logically
superfluous to posit any ‘awareness’ of the prey on the part of the plant,
which does not need to perceive where the prey is located in order to be
able to trap and consume it. The Venus flytrap simply produces an invariant
mechanical response to what is in effect the pulling of a trigger.
Chemokinesis is another class of non-directional response to a stimulus.
Like chemotaxis, this refers to chemically induced movement, but it is dis-
tinguished from chemotaxis by the random orientation of the locomotion,
as when the presence of a particular substance causes an organism to speed
up, slow down or engage in aleatory movements such as turning. If the
organism finds itself in a favourable location (characterized, for example,
Plant-like and Animal-like Unicellulars 65
has now been shown to be capable of true taxis (suggesting at least the
possibility of further such species). The species in question is a sulphur-oxi-
dizing bacterium called Thiovulum majus, a spherical cell between five and
ten micrometres in diameter and largely covered in flagella, which propel
it to swimming velocities of up to 600 micrometres a second. This is the
highest speed known for bacteria197: a hundred cell lengths per second (and
through the equivalent of treacle at that). These cells display various strat-
egies aimed at keeping their preferred position within a gradient of oxygen
concentrations.198 One strategy is to attach themselves to a solid surface by
means of a mucous stalk. Alternatively, they may remain free-swimming
and use chemical signals to form narrow bands at their preferred oxygen
concentration. This they achieve on the one hand by a mechanism of
‘steered turning’, a phobic – and basically non-directional – response that
consists in changing direction whenever they stray from their comfort zone.
The other trick they use is known as helical klinotaxis, which exploits the
natural tendency of cells to swim in a helical trajectory.199 Sampling the
chemical concentration at successive points in time as they follow a helical
pathway along their preferred isopleth, Thiovulum cells are sensitive to peri-
odic changes in concentration resulting from the helical geometry of their
course. In response to undesirable deviations, they regulate the activity of
their flagella, modulating the rotational and translational components of
their helical motion – the parameters of the helix – in such a way as to
maintain their pathway.200
The adherence of Thiovulum to an oxygen isopleth is notably lacking in
the ‘distance’ presupposed by consciousness. It makes little sense to distin-
guish the informational from the energetic function of the oxygen, and the
bacterium remains firmly ensconced within the immediacy of metabolic
need fulfilment. Yet although we may be disinclined to see the truly chemo-
tactic locomotion of Thiovulum as grounding anything like consciousness in
itself, what it announces is that genuinely directional self-propulsion cannot
be denied on principle to prokaryotes.
Motile Protozoa: Oxyrrhis marina
and Company
lectins on the surface of the predator cell that bind to specific prey-associ-
ated ligands such as the carbohydrate mannose.215 In addition to the direct
cell-cell contact of mannose-binding lectins, the flagella too are considered
possible mediators of prey recognition.216 An element of randomness seems
to play an important part in the capture of prey; the absence of visual acuity
makes the foraging process appear ‘clumsier’. Accordingly, it might be
asked whether the goal of the chemotactic search is an individual item or
merely a productive prey patch (an area rich in nutrients). Having sniffed
out a hunting ground, the predators might be imagined ‘groping’ about in
a microscopic game of blind man’s buff, where it is contact chemoreception
(i.e. ‘cell-surface recognition’217 or ‘taste’) as opposed to distance chemore-
ception (‘smell’) that gets the results.
This in turn raises a further question, namely whether a nutrient-rich
prey patch signalled by a ‘gradient’ of infochemicals really amounts to a
‘target’. To the extent that the gradient is gradual (which, etymologically
speaking, is exactly what a gradient is) and each minuscule ‘step’ (gradus)
towards the prey patch constitutes a statistical improvement in the chance
of bumping into an item of prey, the movement might be interpreted not as
motion towards an objective, but simply as an immediate chase from ‘good’
to ‘better’ to ‘even better’ in probabilistic terms. Whether this represents a
gradual progression up a gradient of what is ‘better for self ’ or the inten-
tional pursuit of a target at a distance is perhaps best deemed undecidable.
Indeed, this ambiguity may well have been the logical legerdemain by which
intentionality itself was first made possible, i.e. as a product of the seamless
transformation of an incremental sequence of micro-steps (towards or away
from what was directly contiguous according to whether it was statistically
more likely to be better or worse for self) into the all-or-nothing targeting of
a goal. One might speculate that this was the mechanism by which minimal
selfhood was first ‘manipulated’ into moving itself directionally for a reward
yet to come.
Even though directional chemotaxis towards a prey patch followed by
random hunting at close quarters is doubtless a perfectly viable strategy
for snatching a meal, chemotaxis can in fact be used to target individual
items even at a unicellular scale. A striking example of this occurs in the
realm of mate attraction rather than nutrition acquisition and features the
small (10-µm-long) flagellate Chlamydomonas allensworthii,218 the females
of which release a pheromone while the males pursue the resulting gradient
to find the individual female with which to fuse. Perhaps unsurprisingly,
Motile Protozoa: Oxyrrhis marina and Company 77
observations on its predatory habits. The authors note that ‘the intensely
interesting sight of an amoeba after numerous trials gradually sliding its
pseudopods around a feeding paramecium, throwing a cover over it, closing
the pseudopods, and gradually squeezing the struggling victim down to
a rounded mass can hardly be described without using anthropomor-
phic terms’.225 Nicholas Money has more recently described how the cell
‘embraces its microbial quarry in a pseudopodial hug, casts the morsel into
a vacuole, and showers the terrified bacterium with digestive enzymes. ...
Later, any waste materials are voided by the reverse of the feeding process at
the posterior end of the amoeba. No mouth, no anus, but the essence of all
animal life is there’.226 Collectively, in fact, amoebozoans are not restricted
to phagocytosis of this classic sort. In a phenomenon known as trogocytosis,
the parasitic protozoan Entamoeba histolytica has been found to take bite-
size chunks out of intestinal cells as it grazes through the human gut.227
More than 100,000 people are killed each year by the amoebic dysentery to
which it gives rise.
Also like O. marina, A. proteus is selective in what it consumes. As Gibbs
and Dellinger comment, the amoeba ‘shows distinct food preferences: with
diatoms and unicellular algae, it takes algae, but when feeding on algae it
will leave them to “pursue” ciliates. In the presence of large paramecia, some
amoebas leave algae and ciliates to catch these larger forms. Amoeba eats
nothing dead’.228 As a rule, it also avoids eating members of its own species,
although it is observed to eat amoebae belonging to other species. Indeed,
A. proteus is known to emit an identity marker – a peptide called A-factor –
that reveals its presence not only to its clone mates (thus preventing clonal
cannibalism), but also to various species of potential prey, allowing certain
ciliates of the genus Euplotes to beat a hasty retreat and avoid amoeba’s
pseudopodial hug. While reducing prey uptake by letting more ciliates
escape than otherwise, this form of collective ‘self-recognition’229 presuma-
bly pays its way in preventing collective self-consumption.
Whereas the locomotion of O. marina involves the coordinated use of its
two flagella, A. proteus and amoebozoans in general use their cytoskeleton
to move themselves. In spite of the connotations of the term ‘skeleton’, the
cytoskeleton is far from being a fixed framework, but is a self-assembling
structure that dynamically and adaptively responds to cues from the
environment mediated by scores of interacting regulatory proteins.230 The
protein actin, which constitutes up to a tenth of the total protein comple-
ment of mobile cells, plays a pivotal role in the dynamic functioning of
Hunger, Satiety and Siesta 83
away waste particles) and whose tireless activity had led some investigators
to believe that protozoa ‘never rest’. The rhythm of work and rest, it was
thought, ‘was only gradually evolved with the more complex forms of life’.236
Unicellular A. proteus proves otherwise, for the work of the chase and the
feed is followed by what might best be described as a post-prandial nap. As
Gibbs and Dellinger construe the phenomenon, ‘the period of rest appears
to be simply the result of organic satisfaction, or a period of recuperation.
It suggests the lowest form of sleep; for this tendency to rest, to sleep, as a
food reaction is illustrated by the higher animals’.237
We shall return below to consider the question of ‘sleep’. Another inter-
pretation might be to dismiss this period of non-movement as a purely ‘bio-
mechanical’ phenomenon determined by changes in body structure. Indeed,
the explanation for an amoeba’s ‘satiety’ could simply be that the contents
of its food vacuoles, when full, are too ‘massive’ or ‘bulky’ for the creature
to be energetically or metabolically capable of dragging itself around in
pursuit of further nutrition. A possible analogy from the metazoan world
is the female mosquito, whose weight increases fourfold after it has drunk
its fill of (usually vertebrate) blood and for whom rest and digestion thus
become an urgent priority.238 Motion proves to be an energetic extravagance
if one’s weight has quadrupled, and a newly sated mosquito is unlikely to
go far. Whether an organism’s post-prandial immobility is biomechanically
or biochemically ‘enforced’, however, is perhaps secondary to the present
argument. The point here is simply that – unlike the indefatigably239 feeding
ciliate Vorticella – the amoeba is sometimes disposed to perform the work
required to capture and ingest an item of prey, and sometimes not. Its dispo-
sition to move depends not exclusively on external circumstances (say, the
presence of prey), but also on the internal state of the amoeba in question
(how many it has already eaten).
This clearly contravenes an understanding of unicellular self-move-
ment as mere taxis or tropism. To provide a full explanation of the causes
of such locomotion it is not enough to cite the stimulus or chemical cue
emanating from the prey; we also need to know the internal state of the
amoeba. Manifest as a disposition to behave or move itself to such-and-
such an end or not, an internal state of this sort is indistinguishable from
what is commonly referred to as a ‘mental’ or ‘psychological’ state or, more
commonly still, how an organism ‘feels’. To the extent that mental states can
only be differentiated in terms of the behaviour that they cause or as which
Hunger, Satiety and Siesta 85
There is something charming and quaint about the paper by Gibbs and Del-
linger, dating back as it does to 1908. But presumably there are other, more
recent studies that might cast light on the phenomenon just in case Gibbs
and Dellinger cooked the whole thing up (which of course they did not).
The search for such corroboration proves fruitful but raises a number of
doubts. The underlying question is how to recognize hunger in a free-living
eukaryotic cell. I have been suggesting that to do this requires us to be able
to distinguish between different behaviours (different degrees or forms of
self-movement) in otherwise similar external circumstances.
A rather fundamental doubt relates to how far the proposed distinction
between hunger (as manifest in the pursuit of prey) and satiety (as manifest
in non-pursuit) overlaps in practice not only with a distinction between
behaviour and non-behaviour but by extension with that between wakeful-
ness and sleep. A variation on this theme comes to light in the behavioural
patterns of the haptorid ciliate Pseudomonilicaryon anser, formerly known
as Dileptus anser.243 Measuring as much as a millimetre in length, this vora-
cious predator is armed with a trunk-like proboscis that sweeps through the
surrounding waters to increase its chances of hitting upon prey organisms,
which it then disables by firing toxic trichocysts.244 Groups of this protozoan
have been observed to start feeding shortly before dawn and to terminate
abruptly, four hours later.245 This cut-off varies in time from one day to the
next but always takes place more or less in unison. A group engaged in
finding and ingesting prey switches to a state of complete quiescence within
a matter of minutes, with little or no more feeding occurring throughout the
Recognizing Hunger: Some Doubts 87
rest of the 24-hour cycle (during which time cell division takes place and
broken proboscises are regenerated). The precise nature of this biological
‘clock’ remains uncertain,246 as does the extent to which it provides the foun-
dation for a possible distinction between wakefulness (used for hunting and
ingesting prey) and a sleep-like state (used for bodily renovation). During
their limited period of feeding activity, can the dileptids meaningfully be
described not only as ‘hungry’, but also as ‘awake’?247Are the two terms
synonymous?
Similar concerns are raised by O. marina, which has been shown to
exhibit periodic rhythms in its feeding behaviour, manifest as higher rates
of ingestion during the day than at night.248 These rhythmic activities of
O. marina seem very likely to be circadian in nature, for they are not deter-
mined solely by diel light cycles but persist when the cells are subjected to 24
hours of darkness. This suggests that, although the cycle may be entrained by
environmental cues, it is governed by endogenous biochemical processes.249
Given such 24-hour periodicity, it is tempting indeed to extrapolate from
human experience to propose that O. marina feels both ‘hungrier’ and more
‘awake’ during the daytime. However, other flagellates are nocturnal in their
habits. Diel variations in feeding rates are also exhibited by bacterivorous
nanoflagellates, yet some of these tiny protists graze primarily in the day
whereas others graze primarily at night.250 Perhaps the deeper question
unearthed by these differences among protozoans is whether their periodic
reduction in activity is itself caused by the organism having fed to satiety
or merely coincident with a cessation of feeding brought on by exogenous
(light-related) and/or endogenous (biochemical) rhythms. The diversity in
behaviour seems both to substantiate the existence of some sort of parallel-
ism between satiation and a sleep-like state while also calling into question
any over-tidy correlation.
The possible equivalence of satiety with a condition akin to sleep may also
be felt to have other logical ramifications for the question of consciousness.
To the extent that it is not ‘like anything’ to be in a state of sated sleep (there
is no reason for it to be like anything in that no directional self-movement
is undertaken), the converse state of ‘wakefulness’ or ‘hunger’ lacks a foil, or
point of contrast. If my experiential world consists solely of either feeling
hungry or not feeling anything at all, i.e. of alternating between ‘hunger’
and ‘nothing’, then ‘hunger’ is my only experience, and in the absence of
any counterpoint, it becomes as empty an experience as it is in the case of
Vorticella.
88 Appetite and Tacit Selfhood
Hunger and appetite have at times been described as emotions. The neuro-
physiologist Derek Denton, for example, uses the term ‘primal’ or ‘primor-
dial’ emotion to refer to what he calls the ‘subjective element of instinctive
behaviour’, which ‘subserves control of the vegetative systems of the body’.268
Also referred to as ‘homeostatic’ emotions,269 primal emotions such as thirst
and hunger monitor and regulate the constancy of the body’s internal con-
ditions, prompting behaviour whenever this is necessary to counter an inner
imbalance. According to Denton, such primordial emotions represent the
origins of consciousness.270 The neuroscientist E. T. Rolls, by contrast, opts
to use the word ‘emotion’ to denote ‘states elicited by rewards and punishers’,
where a reward is ‘anything for which an animal will work’ and a punisher
is ‘anything that an animal will work to escape or avoid’.271 In these terms,
‘emotion’ corresponds more to the state that arises from the satisfaction of
an appetite, or to that associated with a failure to avert harm or homeostatic
disequilibrium.
Yet the use of the term ‘emotion’ in the context of appetites and their sat-
isfaction is not uncontroversial. To be sure, both emotions and appetites are
feelings in the sense that it ‘feels like something’ to be angry or hungry. Sub-
tler light is thrown on the matter, however, by the dissection of the concept
of a ‘feeling’ – in all its bewildering breadth – undertaken by Bennett and
Hacker.272 This includes not only affections such as emotions (feeling love
or hatred, fear or jealousy) and moods (feeling happy or gloomy), but also
feelings in the sense of bodily sensations (feeling pain or pleasure, feeling
ill or well, feeling warm or cold), feelings in the sense of tactile perceptions
Appetite, Motivation and Pleasure 93
More recently, this notion of the adaptive utility of pleasure has been
taken up by the physiologist Michel Cabanac, who has highlighted the
homeostatic nature of pleasure: when we feel cold, we are motivated to seek
warmth, and this warmth feels good; when the warmth becomes too much
for us, we look for a cooler place. When we are hungry, we search for food,
and once we have found it, eating it is acutely enjoyable. Once we have sated
our appetite, our enjoyment rapidly declines, and before long we stop.280
Cabanac indeed describes pleasure as the ‘common currency’ by which the
relative strengths of motivational drives can be compared and appropriate
action undertaken.281 In other words, when there are competing or clashing
motivations (such as food intake versus thermoregulation; pursuit of prey
versus inactivity), pleasure is the yardstick by which an organism ranks
its priorities and optimizes its behaviour. In Cabanac’s view, the ‘algebraic
summation’ of pleasure and displeasure occurs ‘not only within one sensory
modality such as taste, but across different modalities of perception and
experience’.282 An example of such cross-sensory assessment might relate
to how much cold an organism is willing to tolerate, or how much work
it is willing to perform, in order to attain a nutritious treat, the resultant
behaviour thus representing a disposition to maximize the sum of pleasure.
In the field of behavioural ecology, the actions of organisms are often
regarded as corresponding to what is optimal on a cost-benefit curve: the
‘common currency’ enables the animal to weigh up rewards and costs and
then choose the behaviour with the maximal net reward or minimal aver-
sive outcome. Although pleasure thus serves as the common currency of
decision-making, there is no implication that the ensuing decision need be
a product of conscious reasoning or deliberation. What such decision-mak-
ing does presuppose is a creature that is conscious of the world and can
choose rationally among its behavioural options on the basis of expected
outcomes.283 ‘Rationality’ here refers not to logical argument, therefore, but
simply to behaviour that promotes the well-being of the organism itself284; it
is the rationality of a ‘selfish’ or ‘self-interested’ self that (by definition) has
reason to pursue its own interests.285 Once endowed with the gift of moti-
vated, directional self-movement, a self thus becomes an entity that has not
only interests but also the capacity to behave or take action in accordance
with those interests. It is with motivated locomotion that ‘interests’ in a
genuinely intrinsically reflexive sense – interests as gauged and pursued by
a self rather than ascribed by an ‘other’ (i.e. an observer) – come into being.
Defined in terms of appetites and pleasures, true self-interest co-emerges
with the motivated self-movement or behaviour that permits its pursuit.
98 Appetite and Tacit Selfhood
with the consumption of intoxicants, which are said to hijack the reward
pathways designed to motivate us to eat and to mate.287 Intoxication is dan-
gerous and maladaptive,288 and animals with a predilection for the pleasures
conferred by psychoactive plants tend to be more accident-prone, vulner
able to predators, and neglectful of their offspring. Another, more debatable
illustration of the apparent non-coincidence of pleasure and utility is the
archetypically ‘non-functional’ pleasure of play, which is often defined as
activity that is autotelic or an ‘end in itself ’. In the case of play, however,
longer-term benefits are generally considered likely to compensate for the
hazards it may occasion and the energetic extravagance it represents.289
The connection between what is pleasurable and what is in one’s interests
is perhaps most graphically undermined by the timelessly well-attested
fact that most children need to be force-fed what is ‘good’ for them. In the
popular imagination ‘healthy’ and ‘fun’ tend to be regarded as more or less
antonymous. Again, however, a widespread proclivity to stuff our face with
sugary treats of dubious nutritional value should probably be understood
as harking back in evolutionary terms to a time where our sugar intake took
the form not of milk chocolate and caramel goo but of ripe fruit loaded
with energy, minerals and vitamins, and a sugar-rich fruit-based diet was of
unquestionable benefit.
Although the link between pleasure and what is good for self is certainly
flawed, therefore, such flaws are derived or secondary effects in complex
systems where sub-optimal ‘viability’ will in most cases do the trick. Impor-
tantly, an amoeba’s sense of what is good for itself can be presumed to be
reliable enough, statistically speaking, to ensure that it passes on its genes
to the next generation. Jennings hits the nail on the head, once again, in
discussing the concept of ‘choice’: in its regulatory sense, he notes, choice
‘is not perfect … in either lower or higher organisms. Paramecium at times
accepts things that are useless or harmful to it, but perhaps on the whole
less often than does man’.290
Persistent sceptics will question whether the satisfaction of an appetite
really is a sufficient condition to speak of ‘pleasure’. A distinction has
been drawn in psychology between ‘liking’ and ‘wanting’,291 where the key
feature of ‘liking’ is that it is reflected in positive behavioural reactions to
the ‘immediate hedonic impact of pleasurable events’, whereas ‘wanting’
is not considered a sensory pleasure or an inherently ‘hedonic’ state and
does not ‘potentiate positive affective reactions to pleasure’.292 Pleasure-less,
compulsive drives and addictions are spotlighted as cases of wanting rather
100 Appetite and Tacit Selfhood
than liking. Surely, one might suppose, the ‘hunger’ of an amoeba is a case
of pleasure-less wanting rather than hedonic liking. This impression is rein-
forced by the failure of amoebae to exhibit any ‘positive affective reactions’
when they engulf paramecia. After all, they have no lips to lick, no eyes to
roll, no vocal cords with which to emit hyperbolic groans of delight; they are
not even equipped with the taste buds that would enable them to enjoy their
meal. Can there be gastronomic pleasure without taste buds?
Two related points need to be made. The reason an amoeba neither has
nor needs taste buds is that – amongst many other things – it is its taste buds.
The amoeba’s ‘tasting’ or ‘smelling’ of the chemical cues that emanate from
potential prey organisms is precisely what alerts it to the nearby presence
of something desirable. Moreover, the nature of the relationship between
pleasure and the expression of pleasure is itself a matter of contention. Many
mammals exhibit positive and negative ‘affective reactions’ to taste in the
form of facial and gestural expressions. These resemble human reactions
and lend themselves to hedonic interpretation. Monkeys and apes repeat-
edly stick out their tongue if offered something sweet; rats likewise show
rhythmic tongue protrusions. Bitter tastes elicit a triangular gape, head
shakes and arm shakes.293 The ‘yuck!’ displays shown by mammals or birds
on interacting with distasteful food items have been found to put observing
conspecifics off similar items, signalling unpalatability and fostering the
social acquisition of avoidance behaviours.294 Yet if a non-social animal,
in consuming its preferred prey, failed to produce a hedonic response but
simply gobbled up its meal with no public expression of delectation, would
we withhold the ascription of pleasure on these grounds? The absence of
an overt expression of pleasure should not be confused with the absence of
pleasure. ‘Watch someone eating’ suggests Jonathan Balcombe.295 ‘Especially
if they are alone, you won’t find facial expressions or other conclusive signs
that the food is pleasurable’.
The social component in the expression of pleasure is likely, therefore, to
rule out any such expression on the part of the solitary amoeba. Imagine the
hypothetical case of an amoeba endowed with a flagellum-like appendage
which it wagged vigorously on perceiving, capturing or ingesting a para
mecium.296 What could such a signal possibly be for? Would we be any more
justified in attributing pleasure to the creature merely on the basis of this
wagging? The basalmost behavioural sign of pleasure is not lip-licking or
tail-wagging, but the fact that a self-interested self will perform work in
Appetite, Motivation and Pleasure 101
order to reach the target or attain the gratification in question (not always, but
sometimes: i.e. until it is sated or has had enough, or until the pleasure has
turned stale).297
Yet this still fails to rule out the possibility that zombie-like ‘wanting’
may be the only form of appetite available to expression-less amoebae, or
any other expression-less creatures. Proponents of this view would insist
that less mentalistic terms such as ‘utility’ or ‘reward’ are perfectly adequate
to describe what is going on when an organism makes the requisite behav-
ioural adjustments to optimize its performance.298 It is superfluous, they
would say, to posit any explicit awareness of pleasure. But this is just the
point. Explicit self-awareness is not implied by the above account of appetite
and the concomitant possibility of pleasure, which are conceived not as
objects of attention (so-called ‘intentional’ objects) but as part of the tacit
or pre-reflective selfhood of a self-concerned self. The claim, in short, is not
that the organisms in question have a capacity to reflect on their pleasure.
What they experience is not reflectively conscious pleasure.299 They do
not think: ‘boy, this is pleasurable!’ or ‘life just doesn’t get any better than
ingesting juicy paramecia!’ Pleasure in this elementary sense is merely the
pre-reflective experiential corollary of a successfully pursued reward. It is
the pleasure-giving world – not the mental state – that is the object of the
predator’s attention.
Further Aspects of Tacit Selfhood:
Pain and Emotion
Consciousness of the world – even of a world that consists of little more than
paramecia, diatoms and lesser morsels – is always structured and shaped
by the tacit selfhood of the self-interested self that is conscious. In this
respect, consciousness is necessarily tendentious, involving a perspective300:
namely, from me, here, now, hungry or otherwise, perhaps also anxious
or angry, drowsy or distressed. Incorporating appetites, motivations and
possibly emotions, this tacit selfhood is the precondition and foundation
for the possibility of our consciousness of the world, enabling an organism
to focus its attention on meaningful non-self and to perceive prey as prey,
and predators as predators. In answer to the question whether the organism
‘knows’ that it is hungry or is ‘aware’ of its pleasure in satisfying its hunger,
an appeal might be made to a time-honoured distinction between disposi-
tional and propositional knowledge. Though not propositionally explicit, its
appetite is what tacitly guides its disposition to move itself appropriately to
where food is. The organism embodies and thus is its appetite, together with
the attendant implicit anticipation of reward. This is what structures both its
(tendentious) awareness of the world and its choice of action.
The conception of tacit or pre-reflective self-awareness is a crucial feature
of phenomenological thought, going back to Husserl’s insight that ‘to be a
subject is to be in the mode of being aware of oneself ’.301 For Heidegger too,
experience of the world from a subjective perspective is indissolubly bound
up with co-disclosure of the self. On this view, self-awareness is not some-
thing derived, secondary or ‘higher-level’ that may be optionally ‘added on’
to supplement a more basal form of consciousness; rather, it accompanies
Further Aspects of Tacit Selfhood: Pain and Emotion 103
and shapes consciousness from the outset, albeit in a tacit or implicit form.
This is not to deny that – in humans at least – these tacitly present appetites,
pleasures, pains, moods or emotions may on occasion become the objects
of explicit awareness. Although we do not need to be expressly conscious
of our interoceptive signals (of hunger or satiety) for these to organize our
perception of the world, we may become conscious of them, for example if a
problem is encountered, if an obstacle needs to be overcome, or if our stom-
ach rumbles obtrusively. While they may contribute to the phenomenology
of hunger, however, our stomach contractions and the accompanying
cacophony are no more the cause of hunger than a dry throat is the cause
of thirst.302 In both cases, it is a homeostatic imbalance that generates the
appetite, the generalized feeling of which may be augmented by specific
bodily sensations that subsequently enter our consciousness.
Appetites such as hunger, thirst or sexual drive are just one aspect of tacit
selfhood among many others that shape our consciousness of the world. In
the case of human and non-human animals, further crucial factors include
pain and an array of emotions and moods. The present subchapter will look
briefly at these aspects of pre-reflective self-awareness and their possible
presence in protists, while leaving out of account other, more complex
aspects such as corollary discharge and proprioception303 which seem less
likely – if only for anatomical reasons – to have analogues in the unicellular
realm. In particular, it will focus on pain as a basalmost response to the
world (or to what is harmful in the world) that is commonly thought to
underlie and structure consciousness in a manner akin to an appetite, albeit
resulting in movement away rather than towards.
The parallels between pain and hunger are striking. Philosopher Sydney
S. Shoemaker famously argued304 that one’s awareness that one is in pain
does not involve a kind of perception of oneself, conceived on the model
of one’s sensory perception of the external world. Nor does my sense of
being in pain involve an act of identification (I do not have to recognize
myself as the person of whom pain is predicated); pain is thus immune
from the possibility of mis-identification. By contrast with the ascription of
pain to other people, the self-ascription of pain is not based on criteria and
does not require evidence. As Bennett and Hacker put it, no more grounds
– whether behavioural or introspective – are required to say that one has a
headache when one has a migraine ‘than a groan of pain needs grounds’.305
Philosophers of mind have traditionally mused on the existence of a sort
of privileged ‘inner access’ or a logically private ‘inner sense’ that a person
104 Appetite and Tacit Selfhood
has to his or her own pain,306 but this leaves unanswered the question how
I recognize my inner sense as my inner sense unless I already have some
prior sense of myself as myself. This is where our tacit selfhood steps in. As
with hunger and other, similar psychological attributes, self-recognition is
superfluous to the extent that a pain is something I not only have but am:
the reason I cannot misidentify myself as the bearer of a particular pain is
that I am that pain. There is no logical separation between the knower and
what is known.307 When I have a toothache my discomfort and my desire for
it to stop structure and shape my entire perception of the world.
We have seen that Romanes and other thinkers locate pleasure and pain
at the very origins of consciousness, regarding them as the most rudimen-
tary manifestation of mind or subjectivity. As neurologist Antonio Damasio
points out, however, pleasure and pain are not mirror images of one another,
but ‘asymmetric physiological states’, belonging to ‘two different geneal
ogies of life regulation’.308 Indeed, there is a sense in which the opposite of
‘pleasure’ is not ‘pain’ but ‘less pleasure’ or ‘discomfort’, and it is by no means
obvious in what sense, if at all, the lower rungs of this scale of pleasure
coincide with pain. This in turn raises the question of whether pain really is
linked to consciousness in the same primordial way as pleasure, i.e. whether
pain and pleasure are equally basal components of consciousness.309
For a start, a distinction is frequently drawn between suffering and
nociception, where the latter refers to a reflex to withdraw from something
harmful or damaging. Inflexible and to a certain extent non-directional
withdrawal responses to stimuli such as heat, electrical currents, noxious
chemicals and mechanical interference occur across the spectrum of motile
organisms from bacteria and protozoa to insects and vertebrates. When
poked with a fine needle, for example, paramecia modulate the rate at which
their cilia beat – by means of changes in the electrical properties of the cell
membrane – so as to take evasive action and avoid the mechanical insult.310
If ‘irritated’ by water turbulence or sudden changes in acidity, cryptomonads
may be seen to zigzag rapidly away, performing a non-directional, random
escape movement or ‘jump’ triggered by the rapid expulsion of a pent-up
protein ribbon known as an ‘ejectisome’.311 As an invariable reflex, nocicep-
tion implies not that an organism is conscious of an unpleasant experience
or its cause, but simply that its body registers a harmful stimulus and is
immediately prompted to recoil from it. By contrast with appetite, indeed,
a painful stimulus is one that is already present.312 Accordingly, there is no
call for anything like a self-guided search or pursuit involving perception
Further Aspects of Tacit Selfhood: Pain and Emotion 105
activity. Whereas insects and arachnids do not seem to ‘mind’ their pain
(if such they have), vertebrates certainly give the impression of doing so.
Analogy with humans suggests that vertebrates in general have a capacity
to feel bad or even ‘miserable’.318 What is awful about pain, it often seems, is
an associated emotional state. The notion of ‘not minding’ a pain may seem
counterintuitive or even contradictory to the extent that a pain is something
that we by definition do mind having. Nonetheless, the distinction between
‘having’ and ‘minding’ a pain is known to exist in certain cases of excruci-
ating human pain, as shown by the emotional transformation that occurs
when specific parts of the frontal lobes are operated on in people with the
condition trigeminal neuralgia, or tic douloureux.319 Damasio thus draws
a distinction between ‘pain sensation’ and ‘pain affect’, pointing out how
certain drugs (analgesics) can block the sensory awareness of pain, while
others such as Valium or beta-blockers do not affect the signal transmission
of tissue damage but blunt the emotion and thus do away with the suffer-
ing that would have otherwise accompanied the pain.320 Human pain is a
multidimensional phenomenon, involving a whole nexus of affective and
cognitive factors that may include stress, anxiety and depression.
There is a sense, therefore, in which the attendant emotions may repre-
sent a large part of what is bad about pain, or what turns pain into suffering.
Insofar as the pain undergone by non-human animals is dissociated from
emotion (less burdened, perhaps, by time-dependent factors such as
anticipation and the anxiety to which this gives rise), it may be felt that
their suffering is in some sense less severe than human suffering. However,
there are clear indications that mammals such as rats scarcely differ from
humans in this regard, displaying a number of bodily corollaries of fear in
anticipation of pain.321 Moreover, animal suffering seems to be connected
with other affections and states of mind instead of or as well as pain. Animal
scientist Temple Grandin accordingly claims that for animals fear is ‘worse’
than pain,322 while Hans Jonas associates animal suffering not directly with
pain, but with the exigencies of unsated appetite: ‘the suffering intrinsic
in animal existence’, he writes, ‘is thus primarily not that of pain (which is
occasional and a concomitant) but that of want and fear, i.e., an aspect of
appetitive nature as such’.323
*
Further Aspects of Tacit Selfhood: Pain and Emotion 107
such affections – indeed to anything more than mere appetite, desire, and
states of greater or lesser pleasure. Can we attribute even the most primitive
of emotions, such as fear, to a retreating amoeba that has one of its more
sizeable conspecifics hot on its pseudopodial heels? In the case of fear the
answer will depend, at least in part, on whether the escape response is
considered a mere reflex. If a chemical cue emitted by a predator invariably
produces the same non-directional reflex withdrawal, there is presumably
no need to posit any further ‘awareness’ of the predator, any ‘emotional’
response to its presence, or any subjective sense of it ‘being like anything’ to
react in this way. The explosive ‘jump’ of a cryptomonad in reply to a chem-
ical or mechanical disturbance should not be taken to imply an affective
correlate. The same goes for the insect-like, wingless hexapods known as
springtails, which are equipped with a specialized jumping organ, or fur-
cula, that catapults them out of the clutches of potential predators and into
the air in a spectacular but uncontrolled leap.328 Insofar as escape strategies
need to be modulated (escape is not always just AWAY!), however, a merely
invariable reflex may not be enough. Some capacity to discriminate relevant
features of the environment may be required. In animals there is frequently
a rather sophisticated mixture of directional and non-directional factors in
operation.329
The question of unicellular fear was indeed a matter of 19th-century
controversy. Whereas Romanes declined to attribute emotions to protists,
considering the most ancestral emotions such as fear to have first found
expression in worms,330 Binet countered forcefully that ‘there is not a single
ciliate Infusory that cannot be frightened, and that does not manifest its fear
by a rapid flight through the liquid of the preparation’. If a drop of acetic
acid is added to a preparation containing plentiful Infusoria, he pointed
out, these will flee in all directions ‘like a flock of frightened sheep’.331 Binet
may appear to have succumbed hook, line and sinker to the temptations
of anthropomorphism,332 yet he remained agnostic on the question of con-
sciousness, thus implying the possibility of unconscious fear.
To the extent that Romanes is right to deny unicellulars even the most
ancestral of emotions, and to the extent that Binet’s sheep-like ciliates are
in fact displaying a merely invariant, non-directional aversive response, it
would seem that protozoans lead a life animated only by appetite and the
possibility of its pleasurable satisfaction or less pleasurable non-satisfaction.
However, there is one category of affection that may yet be of relevance to
the tacit selfhood of a unicellular predator: namely agitation in the form of
varying degrees of arousal, wakefulness, alertness or attention.
Attention, Arousal and Their Absence:
Sleep and Anaesthesia
A capacity for attention implies the selective spotlighting that makes spe-
cifically appropriate action possible. What this means is that a particular
signal or set of signals stands out from a manifold of other signals, acquiring
meaningfulness through its salience. The importance of being able to focus
in this way again comes to light in the paper by Gibbs and Dellinger:
are in large measure endogenous but also entrained by external cues such
as daylight – pervades the living world. It is found not only in plants and
animals, but also in fungi, protozoa and cyanobacteria. Evolutionarily
ancient, highly conserved light-sensing molecules such as cryptochrome
regulate the day-night cycle in plants and fruit flies alike, determining the
opening and closing of a plant’s leaves and the daily patterns of activity and
inactivity, responsiveness and unresponsiveness, studied in Drosophila.347
DNA sequence comparisons yield an age of over 3,500 million years for the
primordial rhythm gene of cyanobacteria.348 This suggests that biological
clocks had evolved long before multicellularity entered the scene and long
before plants and animals branched into two distinct kingdoms, presumably
permitting early micro-organisms to shield light-sensitive processes such
as DNA replication from the deleterious effects of UV radiation or perhaps
serving to partition mutually incompatible metabolic functions (such as
oxygenic photosynthesis and nitrogen fixation) between daytime and
night.349
The circadian rhythms of cyanobacteria regulate the diurnal patterns
of expression for most of their genes, giving rise to daily fluctuations in
metabolic rates, nitrogen fixation and reproduction.350 In autotrophic
dinoflagellates, such rhythms govern crucial physiological functions such as
cell division, photosynthesis, phototaxis and bioluminescence, with mitosis
usually taking place in the dark phase.351 In plants the effects of the circadian
clock are witnessed not only in leaf movements, but also in rhythms of
growth, germination, enzymatic activity, gas exchange, photosynthesis and
the opening and closing of flowers.352 Yet the presence of circadian rhythms
in such photosynthetic organisms raises the question of how far these
rhythms can be identified with cycles of wakefulness and sleep and how far
this distinction between wakefulness and sleep can in turn be equated with
that between consciousness and its absence. Although circadian rhythms
are certainly responsible for regulating an organism’s levels of metabolic
and physiological activity, not all such activity necessarily coincides with the
specific, directional forms of self-movement associated with consciousness.
The activity of an organism need not be ‘action’ in the sense outlined above.
A circadian ‘sleep-like’ state has thus recently been attributed to the
upside-down jellyfish Cassiopea, a cnidarian and as such one of the most
ancestral metazoans.353 Not only does Cassiopea exhibit periods of rapidly
reversible night-time quiescence marked by delayed responses to stimula-
tion, but the day after being deprived of this quiescent period it is less active
114 Appetite and Tacit Selfhood
an otherwise motile organism does not in itself provide proof of a shift from
consciousness to unconsciousness. As we shall see, consciousness might
not be truly abolished by immobilization (it might persist in an immobile
body).357 In an organism whose locomotion is restricted to taxis, it might
not have been present in the first place. Yet these conundrums are not in
principle different at a unicellular from a multicellular level.
The whole issue of anaesthesia is beset with controversy. One of its most
remarkable features is that chemically highly diverse gases can induce it,
ranging from nitrous oxide, ether, chloroform and isoflurane to the inert
element xenon.358 This has traditionally fostered the notion of a unitary
molecular mechanism common to the various chemical agents in spite of
their structural and pharmacological diversity. The focus has generally been
on either of two main features of cells that are known to be affected by
volatile anaesthetics, namely the membrane and the cytoskeleton.
The original unified theory of general anaesthesia was the ‘lipid hypoth-
esis’, which propounded that anaesthetics act by non-specific interference
with the lipid plasma membranes of neurons, thus preventing conduction
of electrical impulses, or action potentials. More recently, the emphasis
has been on ion channels and neurotransmitter receptors, but here too the
effect resides in the capacity of anaesthetics to inhibit the transmission of
signals among neurons. In particular, the breakdown in communication
among neurons has been associated with a disruption of the ‘functional
connectivity’ deemed necessary for consciousness, i.e. a failure to integrate
information among various cognitive networks that are thus left uncou-
pled and uncoordinated.359 Indeed, circuits involved with the integration
of information are thought to be among those most directly linked with
anaesthesia, which has been attributed, for example, to a loss of connectivity
within the corticothalamic network. As argued above,360 however, connec-
tivity and the integration of information are best regarded as constitutive
not of consciousness itself, but of the unitary selfhood or self-containment
that grounds consciousness. This line of investigation may thus be taken to
imply that it is by undermining minimal selfhood that anaesthesia extin-
guishes consciousness.
To the extent that they have highlighted the disruption of signal trans-
mission and connectivity within nervous systems, membrane-centred
interpretations have tended to limit themselves to the multicellular anaes-
thesia of animals and disregard the case of single-celled organisms.361 This
is less so in the instance of the other main centre of attention in studies of
116 Appetite and Tacit Selfhood
The claim of the present tract is neither that all selfhood displays conscious-
ness nor even that all self-moving selfhood entails consciousness. An indis-
pensable part of any attempt to demonstrate that some micro-organisms
may in certain respects be endowed with consciousness is to show other,
related respects in which they are almost certainly not. The underlying idea
is that consciousness, in its logical origins, enables a self-concerned self to
guide itself successfully towards what is ‘better’ or away from what is ‘worse’
for itself. In these terms, a self ’s consciousness of what is relatively ‘good for
self ’ in the world around it (say, potential nutrients) is logically inextricable
from the capacity and disposition to behave or take action, i.e. to move itself
to wherever this relative ‘goodness’ may be.
This has already been seen to render consciousness logically superfluous
where the organism is not big enough to resist the disorganizing effects of
Brownian motion, the random jostling that prevents it from maintaining
a steady orientation and thus pursuing its interests by means of guided
self-propulsion.369 Consciousness is also pointless where the acquisition of
nutrition is by osmotrophy – the passive assimilation of dissolved organic
compounds – or by other forms of passive capture such as the diffusion
feeding of radiolarians, heliozoans and suctorians. Little or no locomotive
work is presupposed by such feeding strategies370; nor is appetite required
to motivate the performance of work for a reward yet to come; nor, again,
is consciousness of a world required to guide the organism appropriately to
where the nutritional reward may be found. Similar considerations suggest
that fish have no reason to be conscious of water: unlike terrestrial reptiles
122 Where Consciousness is Superfluous
and mammals, they can be assumed neither to experience thirst nor to pos-
sess water-seeking activity within their behavioural repertoire.371 Awareness
of the location of nutrition within the environment is equally redundant
for modes of non-specific restlessness or non-directional movement such
as chemokinesis – presumably including the biased random walk of E. coli
– and the thigmonastic behaviour of carnivorous plants.
At the same time, consciousness is only required if the locomotion of the
self-concerned self is genuinely self-caused. This means, for a start, that the
movement of the hungry predator should come from ‘within’ rather than
having its causal roots in a purely physical force exerted by the prey. Hans
Jonas thus visualizes the case of a target-seeking torpedo that is impelled by
direct magnetic attraction between itself and its target, comparing this with
a missile that uses magnetism merely as a signal to guide it:
What constitutes the difference in the two cases, assuming that mag-
netic principles operate in both, is that in the self-steering torpedo the
magnetic factor does not itself provide the power for the acceleration
of the entity whose steering arrangement it affects, and the effect on
the latter is not a function of the quantity of the magnetic force acting
on it. Given sufficient sensitivity, this force may be as small as you
please, and given efficient coupling and sufficient motor resources,
the effect in terms of power may be as large as you please. The torpedo
is not attracted but is steered toward the target – in response, to be
sure, to an influence emanating from it, but this influence is of the
order of ‘message’ and not of acceleration.372
On a human level, Jonas adds, this amounts to the claim that purposive
behaviour involves perception. Yet while the conscious pursuit of prey may
require an influence from the prey in the form of information (a cue or set
of cues that is registered by the predator), the force comes from the predator.
It is the predator’s energy that drives the pursuit rather than a force exerted
by prey on predator. The predator has to work for its meal; without work it
is not truly self-movement.
In fact, magnetotaxis has itself been described as ‘dumb’.373 Magneto-
tactic bacteria such as Magnetospirillum magnetotacticum contain within
themselves an organelle known as a magnetosome, enclosing magnetite
crystals upon which the Earth’s magnetic field exerts a torque.374 In this
way, the bacterium’s body is oriented according to the local magnetic field
lines, and the cell swims in the direction it happens to be pointing. This
Taxis and Reflexes 123
Tye concludes that there is no more reason ‘to attribute phenomenal con-
sciousness to a caterpillar on the basis of how it moves than to an automatic
door’.387 Just as a door responds in a purely mechanical way to the applica-
tion of pressure to a plate in the ground in front of it, the lowly caterpillar
responds in a purely mechanical way to the presence of light.
126 Where Consciousness is Superfluous
cold, mechanical shock or chemicals, all of which may cause them to swim
downwards.390 Such versatility implies that information is being processed
rather than merely transmitted, a procedure equally possible in nervous
systems or in the phosphoprotein circuits of single-celled organisms, where
neural or protein ‘switches’ are capable of performing logical operations
and thus generating a diversity of behaviours that may vary according to
circumstance. Yet despite the cognitive sophistication engendered by pro-
cessing of this kind, it still implies unidirectional causality insofar as it is the
environment – albeit a multi-modal environment – that wholly determines
the behaviour of the organism. The genuinely self-caused movement of a
conscious self presupposes that endogenous signals, i.e. signals generated
within the organism itself, can also be integrated into the informational
circuits, which thus accommodate interoceptive cues relating to the body’s
own chemical state (its need for food or water, its temperature, etc.).391
In vertebrates, such integrative processes tend to be centralized in the
brain, whereas the control of movement in itself – and rhythmic motion in
particular – need not be. Aspects of the motor activity of the frog such as
its stereotypical reflex reaction to remove irritants from the skin are known
not to require any part of the brain, and this sort of brain-independent
reflex can be assumed to be no more related to consciousness than the taxis
of microbes.392 In arthropods such as insects and crustaceans, the presence
of other relatively autonomous concentrations of neurons (ganglia) in
addition to the brain allows for even more spectacular feats of brainless,
and presumably unconscious, movement. Headless cockroaches have been
shown to be capable of learning how to avoid an electric shock. Brainless
fruit flies can stand up if knocked over and perform a grooming reflex if
prodded. As Greenspan drily points out, however, flying ‘requires a head’.393
Indeed, most (if not all) of the flexibly directional locomotion performed
by animals endowed with a brain involves the use of that brain, an organ
that evolved as the intermediary between sensory input and motor output
meticulously attuned to the particular needs of the respective self. The
question of whether all animal consciousness demands an anatomical brain
will re-emerge in the Epilogue.
The analogy with metazoans has suggested to some that a similar, ‘brain-
like’ centralization of the information-integrating function might be present
in unicellular organisms. On a speculative note, Dennis Bray thus posits the
presence in amoebae of a ‘single executive complex’ – an integrative protein,
perhaps – that would ‘receive relevant signals about the cell’s internal state
128 Where Consciousness is Superfluous
*
140 Limits to Claims about Rudimentary Consciousness
The latter was perfectly spherical and very easily moved, so that
when the anterior edge of the Amoeba came in contact with it the
cyst merely moved forward a little and slipped to one side (the
left). The Amoeba thereupon altered its course so as to follow the
cyst. … The cyst was shoved forward again and again, a little to the
left; the Amoeba continued to follow. This continued until the two
had traversed about one-fourth the circumference of a circle; then …
the cyst, when pushed forward, rolled to the left quite out of contact
with the Amoeba. The latter then continued forward with its broad
anterior edge in a direction which would have taken it past the cyst.
But a small pseudopodium on its left side came in contact with the
cyst. The Amoeba thereupon turned again and followed the rolling
cyst. At times it sent out two pseudopodia, one on each side of the
cyst … , as if trying to enclose the latter, but the ball-like cyst rolled
so easily that this did not succeed. At other times a single very long,
slender pseudopodium was sent out, only the tip of which remained
in contact with the cyst. Then the body of the Amoeba was brought
up from the rear and the cyst pushed further.427
A World of Objects 141
Eventually the Euglena cyst was whisked away by the ciliary current of a
passing infusorian, ‘one of those troublesome disturbers of the peace in
microscopic work’.428 The amoeba continued its pursuit for a short time,
before reversing its course and heading in a new direction.
Jennings is struck by the resemblance between the amoeba and
‘immensely higher organisms’: one ‘seems to see’, he writes, that ‘the Amoeba
is trying to obtain this cyst for food, that it puts forth efforts to accomplish
this in various ways, and that it shows remarkable pertinacity in continuing
its attempts to ingest the food when it meets with difficulty’.429 Elsewhere, in
describing the pursuit of one amoeba by another, he notes that ‘it is difficult
to conceive each phase of action of the pursuer to be completely determined
by a simple present stimulus’.430 What Jennings is almost bashfully431 insin-
uating here is the influence of some form of – admittedly very short-term
– memory trace (or ‘representation’) that keeps the predator focused on its
prey even when the latter temporarily slips from its clutches.432 The question
is whether the predator ever really does leave the sphere of chemoattraction
exerted by its prey, be it the ‘smell’ of something distant or the ‘taste’ of what
is contiguous. Given the presumed limitations of unicellular learning (to be
discussed below), an explanation based on the prey’s ongoing ‘presence’ is
perhaps more parsimonious than one involving an amoeba ‘remembering’
or ‘representing’ its prey as it fleets into and out of its presence. But the jury
is still out.
Choice and Freedom
Amoeba proteus seems not to inhabit a world of lasting objects, which for
some people may exclude it from membership of the exclusive club of truly
conscious entities. Yet the space in which it lives is structured by the differ-
ential desirability that motivates it not only to move towards what it likes (to
eat) and avoid what is harmful, but also to distinguish between things on the
basis of how much it likes (eating) them: in other words, it has preferences
and makes choices.433 This has already come to light in its predilection for
paramecia over other ciliates, and for ciliates over diatoms and other algae.
A great deal of work has also been done to establish the impressive prey
selectivity of O. marina, both between species and within species.434 As we
have seen, moreover, selectivity may depend on the physiological state of
the predator, whose relative hunger or satiety conditions how ‘choosy’ it
is.435 Nor are a protozoan’s preferences immune to modification over the
course of its own life-history. The effects of ‘dietary imprinting’ have been
demonstrated in the predator Didinium nasutum, which will preferentially
ingest the stock of Paramecium bursaria on which it has been reared. It can
even be ‘trained’ to feed on specimens containing mutualistic algae (called
zoochlorellae) that are otherwise ‘distasteful’ to it, although this training
is quickly overcome if it is subsequently presented with ‘bleached’ cells.436
Preferences also come to light in the amount of work a predator is willing
to undertake in order to obtain an item of prey. The basic principle is that
the more motivated an organism is to procure an item, the harder it will
work to do so, just as a hungry organism is expected to work harder to
get food than one that is satiated. The field of ‘consumer demand’ studies
Choice and Freedom 143
circumstances. In this respect protists cannot be free either, for they too are
necessarily shackled by the requirements of their bodily constitution.
Alternatively, it might be felt, ‘true’ freedom encompasses the possible
subjugation of these basic needs, for example by deferring or postponing
their gratification in ways we shall encounter in the following section. At a
deeper level, it might take the form of a capacity to overcome conatus and the
instinct of self-preservation, and thus a capacity to overcome selfhood itself.
Conceived in such terms, freedom may be understood to manifest itself
in the Jainist sallekhanā or ‘fast unto death’ by which the devout embrace
their own imminent demise,446 or in the sacrifice of one’s own selfhood for
a higher-order collective good, a behavioural option available (at the very
least) to humans, hymenopterans and the social amoebae. To the extent
that freedom is understood as an expression of selfhood, of course, the
renunciation of selfhood – though it may be a final act of freedom or even
its consummation – is also, paradoxically, a renunciation of freedom.447
At a most elementary level, perhaps, freedom may best be seen to inhere
in how one meets one’s needs, ‘basic’ or otherwise. If there are no alterna-
tives (as in conditions of stress or poverty), there is indeed no freedom. It
can thus be characterized as an ability to choose and pursue one’s own way of
maximizing pleasure,448 i.e. of seeking what is good for self, be it in the form
of a paramecium or a pizza, the titillations of a life of luxury or the delights
of an ascetic, altruistic or selfless lifestyle.
Future and Past
– and thus seem to act against their own interests as individuals – may be
explained by a collective logic dictating that the multigenerational benefits
of their husbandry are reaped by existing kin groups.461 Individual aware-
ness of the future is perhaps made unnecessary by this multicellular logic.
Whatever the explanation, the consequence of the ‘selflessness’ shown by
the amoeban farmers is that they either ‘lose’ their appetite or are some-
how able to ‘overcome’ it. Their motivation to eat transforms itself into a
motivation to hoard, and not just into a lack of motivation to eat, as in
satiety. The precise molecular mechanisms that produce this subsumption
of short-term individual appetite within long-term collective well-being are
not currently known.
A sense of time may manifest itself not only in the deferral of gratification,
but also in a willingness to initiate an action (directed self-movement) in
the absence of the anticipated reward. A striking example of an animal satis-
fying an appetite by a strategy that seems to involve planning and temporal
awareness is the behaviour observed in African elephants, which trek vast
distances to specific salt ‘mines’ such as Mount Elgon on the Kenya-Uganda
border in order to meet their need for salt. After a lengthy expedition, herds
of salt-hungry animals descend 100 metres into the pitch-black depths of
the extinct volcano, where they use their tusks to gouge lumps of sodium
sulphate rock from the cave walls. Ian Redmond, who has studied the ele-
phants, believes that their knowledge of the caves and the salt they contain
has been passed down from mother to calf over generations for perhaps
hundreds of thousands of years.462 The salt-appetite of the elephants, in
conjunction with individual and possibly cultural memory, thus allows
the animals to undertake a targeted ‘pursuit’ of the food they need – as
opposed to a merely random search – in spite of the lack of any direct
sensory stimulation from the target in question. The elephants set out on
their journey far from the mountain, following established trails through
the Kenyan forest long before any outward cue could betray the proximity
of the salt on which they depend to restore their homeostatic equilibrium.
Motivation and memories are what get the elephants to where they want
and need to be.
Future and Past 151
and in pursuing its own interests in general. Such an organism should like-
wise not be understood to have ‘self-knowledge’ in the sense of an explicit
apprehension of its own identity as the subject of a sequence of experiences
or as an autobiographical self with a life-story. There are no grounds for
believing that a protozoan is blessed with a concept of itself as itself. But it
does have the pre-reflective self-awareness that has been examined above,
embodied in its appetites, its motivations and its bodily self-presence: what
we have called its ‘tacit selfhood’.
To the extent that the consciousness displayed by predatory protozoans
is logically prior to the use of concepts, moreover, such micro-organisms
cannot be ascribed a capacity for thought. Oxyrrhis and Didinium (with all
due respect) are not ‘thinkers’. Again, the terms in question require further
specification. The notion of a ‘concept’ has itself been understood in differ-
ent ways by different people. Some regard the possession of concepts as a
linguistic capacity that consists in an ability to use words correctly in diverse
situations according to a meaning established by convention. This ‘strong’
understanding of concepts unequivocally excludes any creature without a
language.482 A ‘weaker’ sense of the term equates concept-possession with
an ability to recognize or distinguish different categories of entity. Yet this
sense seems rather too flimsy, leaving it unclear how the possession of
concepts is to be distinguished from mere perception, in that to perceive
is already to perceive as.483 In general usage, something more is demanded
of a concept.
One option is to characterize concepts as enabling us to represent some-
thing in its absence, a faculty seemingly denied to protozoans such as A.
proteus. Or we might expect the possession of concepts to involve an ability
to combine them – rather like building blocks – into meaningful composites,
or thoughts.484 In contrast with the structural complexity of much human
thought, we might concede that the structure of non-human thought need
consist of no more than concept A (or B or C) in conjunction or association
with concept a (or b or c). In the case of the salt-hungry elephants, for exam-
ple, we might imagine a concept of ‘salt’ (A) being combined with a concept
of a particular spatial location (a) to produce the thought (Aa). The fact that
the concept of ‘salt’ (A) could, in theory at least, be combined with concepts
of other spatial locations (Ab or Ac), just as the particular spatial location
(a) could be combined with other resources or activities such as dust baths
or underwater pools (Ba or Ca), is what turns ‘salt’ and the particular spatial
location into concepts, and the composite of the two of them into a thought.
Knowledge, Thought and Morality 157
it is itself an ‘other’ to other selves, producing our awareness that we are our-
selves objects of perception to those who view us (and judge us) from their
own ‘selfish’ perspective. These two factors, empathy and shame, have been
decisive in the development of selves that are motivated – or regard it as in
their interest – to behave in the interests of selves other than themselves, i.e.
to act in ways that may be held to be moral.
Figure 2: Dugesia tigrina
typically measuring ca. 5 –10 mm in length
Epilogue: Consciousness in
Simple Animals
Three Questions
The argument of this book has been that the minimal self, appropriately
defined, provides an explanatory foundation for the most elementary forms
of consciousness. The idea is that consciousness is logically entailed by cer-
tain directional modes of self-moving selfhood. This has been seen to allow
for the ascription of consciousness to some free-living protozoans some of
the time (though by no means all or always), but it also has similar impli-
cations for metazoans that engage in similar activities in similar contexts.
If the preceding argument is on the right lines, there are three sets of
questions that need to be asked in order to ascertain whether or not the
attribution of consciousness to a particular entity is appropriate. The first set
of questions relates to whether the entity in question is a self, as defined in
terms of intrinsic reflexivity. This is not merely a matter of whether it shows
a capacity to maintain itself, i.e. whether it is a metabolic system that uses an
influx of energy or fuel to keep itself going. Equally significant, as we shall
see, is whether it displays the intrinsic reflexivity of self-containment, mani-
fest in a capacity to separate self from non-self and in the functional unity
and interdependence of its constituent parts.488 The second set of questions
concerns whether the entity genuinely moves itself and, if it does, whether
this movement is guided or oriented by external cues: in other words, is it
truly directional self-movement as opposed to random locomotion? The
third set of questions asks whether the entity’s self-propulsion represents
an invariable reflex that always occurs in response to a particular external
stimulus or whether it depends upon a variable internal state (e.g. hunger
as opposed to satiety). Such a dichotomy raises the possibility of it being
166 Epilogue: Consciousness in Simple Animals
themselves from their own multicellular body and join forces with a nearby
grex that wanders too close. Any particular grex can be split into two, and
two of them can fuse with one another. In this respect, the individual amoe-
bae do not fully subsume their selfhood within an indivisible collective self,
and it is as discrete cells that they may respond to attractants and repellents
in the environment around them. In spite of the adaptive appropriateness
of its coordinated self-movement, therefore, the multicellular Dictyostelium
grex lacks the requisite self-containing unity to be considered a potential
candidate for consciousness, at least as a grex.
Sponges and Other Filter Feeders
Many species appear all the more plant-like because individual cells host
photosynthetic endosymbionts such as cyanobacteria or green algae, which
share the energy they harvest from the sun in return for protection and a
cosy home. Like plants, moreover, they are able to respond appropriately to
a range of environmental stimuli, closing the oscula to stop or diminish the
flow of water through the aquiferous system if the suspended particles are
too large or densely concentrated, or secreting toxic mucus to ensure that
the oscula are not overgrown by corals. Fleeing is not an option in the face
of predators, but sponges are afforded special protection by calcareous or
siliceous needles and an array of chemicals that are poisonous to animals
with a nervous system.494
Sponges were traditionally thought to be filter-feeding detritivores, her-
bivores or bacterivores, ‘restricted in their aggressive activities to waging
chemical warfare in substrate competition with other sessile organisms and
in anti-predatory defence against mobile animals’.495 This is now known
not necessarily to be the case. Deep-sea sponges such as the harp sponge
Chondrocladia lyra have dispensed with the system of filtering bacteria and
microalgae from a throughflow of water, instead adopting a carnivorous
lifestyle that involves ensnaring copepods and other small crustaceans
that happen to bump into their branching appendages. The captured prey
are then gradually engulfed and broken down within a secreted digestive
membrane. Clearly distinct from filter feeding, such a strategy recalls the
diffusion feeding of unicellular radiolarians, heliozoans and suctorians.
Although the manipulation and ingestion of the prey requires work, no
genuine locomotion is called for, and as with the above-mentioned protists
it seems logically superfluous to posit any consciousness on the part of the
predator, whose strategy is essentially a passive assimilation of whatever
comes its way.496
According to the present argument, the sedentary nature of both dif-
fusion-feeding carnivorous sponges and the filter-feeding bacterivorous
variety renders consciousness unnecessary. Yet the phenomenon of filter
feeding in particular is diverse in its manifestations. As in the case of unicel-
lulars, there are variations in the amount of work and movement demanded
of different types of filter feeder. One relevant factor is whether it is ambient
currents that bring the food to be filtered out or whether the flow of water
is produced by the organism’s own metabolic energy. The flow through the
internal chambers of sponges, for example, is at least partially generated
by the beating flagella of specialized cells called choanocytes. Sea lilies
Sponges and Other Filter Feeders 171
– which belong to the phylum of echinoderms – are sessile animals that sit
on a stalk, holding five arms outstretched to funnel sinking detritus into
their mouth.497 A less passive strategy is deployed by salps, chordates like
us, which vigorously contract their barrel-shaped gelatinous body, using a
kind of jet propulsion to power water through themselves and trap plankton
and organic matter as it traverses a series of internal meshes.498 Another fil-
ter-feeding chordate, the simple, fish-like lancelet (or amphioxus) burrows
into the sand in shallow waters, with just its head protruding to sift food
from ambient currents.499
Varying degrees of metabolic work notwithstanding, none of these filter
feeders requires any form of directed self-movement for the acquisition of
food, meaning that their eating habits do not in themselves provide a founda-
tion for consciousness. This is not to say that an animal such as the lancelet
is never conscious; it simply rules out food-seeking, directional locomotion
as the root of amphioxus consciousness, if such there is. It is possible that
consciousness might emerge for other purposes involving sporadic direc-
tional behaviour, such as steering clear of predators or searching for a mate
or a suitable place to bury itself in the sediment.
That filter feeding need not be incompatible with the possible occur-
rence of consciousness is highlighted by examples such as the basking shark,
Cetorhinus maximus, the second largest of all extant fish. As with unicellular
filter feeders such as Paramecium, the currents that are filtered are actively
generated by the motion of the animal as a whole. In theory, the movement
could be non-directional to the extent that the feeder simply sweeps through
an area rich in nutrients. However, the survival of these gentle leviathans
hangs crucially upon their ability to pinpoint the richest possible patches
of plankton, and this presupposes sensory and decision-making apparatus
that dictates not only where to go but how long to stay, in short how to
maximize intake and minimize risk. Observations of the feeding behaviour
of basking sharks have shown that the time they spend in a particular prey
patch is proportional to the density of zooplankton found there. Movements
between these patches take one or two days and involve distances of several
kilometres. The sharks apparently follow temperature gradients and tidal
flows, possibly also sensing the weak electric fields generated by copepod
muscle activity or the dimethyl sulphide released when zooplankton graze
on phytoplankton.500 In such circumstances, food-oriented consciousness
seems more than plausible, as well as that associated with the range of other
activities in which basking sharks engage.
The Non-Directional Movement
of Placozoans
that is offered to it but will remain quietly attached to the substrate, only
displaying its feeding response if stimulated with an unusually high con-
centration of glutathione.512 Once its appetite has returned, it will exhibit a
different set of behaviours, contracting and stretching its body and tentacles
in new directions as though ‘seeking’ a quarry. With time (and increasing
hunger), the contractions and stretchings increase in frequency and vigour.
Eventually, the hydra will move off to a new location, gliding along on the
amoeboid cells of its base or performing a ‘somersault’ that involves bend-
ing over, attaching its tentacles to the substrate, loosening its base and then
swinging over like a tiny gelatinous gymnast.513 Such displacements may
be random in orientation or in some cases minimally directional, taking
advantage of the hydra’s photosensitivity to aim for better-lit areas where
nutrition tends to be more abundant. In other cases, the hydra simply allows
itself to float freely with the currents.
Leaving aside any residual gelatinophobia, one thus wonders whether
the sporadic positive phototaxis of a ‘hungry’ hydra provides grounds for
the attribution of elementary consciousness. Is it ‘like’ anything to be such
a hydra as it heads towards what is better for self, i.e. the light that signals a
more probable presence of food? To the extent that its behaviour is different
from the behaviour it shows when it is in a different physiological state
(‘sated’), the ascription of ‘hunger’ makes sense. But the recurrent uncer-
tainty is whether we are justified in divesting the ‘hunger’ and ‘satiety’ of
their scare quotes.
The present argument is that true appetite, in this basalmost context,
is what tacitly structures and shapes the consciousness presupposed by
flexibly directional self-movement, i.e. by genuine behaviour in the sense of
‘action’. If there is no action but only random or invariant locomotion, con-
sciousness and the tacit selfhood that structures it are logically unnecessary:
no genuine ‘being like’ need be inferred, and the attribution of ‘hunger’ or
‘satiety’ must remain metaphorical. In the case of our hydra, of course, the
motion may be considered directional precisely to the extent that it includes
phototaxis alongside sessile and non-directional strategies. Even granting
the occasional occurrence of guided locomotion, however, movement up a
gradient towards a potential prey patch (i.e. an area that is statistically more
likely to yield nourishment) perhaps falls short of movement towards an
individualized prey item, bringing us back to the logical puzzle raised on
page 76. We may still be a step or two away from elementary consciousness.
‘Hungry’ Cnidarians 177
Other cnidarians are more motile, yet not necessarily more likely to be
endowed with consciousness. The ‘slow swimming’ of the feeding jellyfish
Aglantha digitale, for example, is based upon an initial process of upward
propulsion that involves the rhythmic contraction of a conical sheet of
muscles and the expulsion of water from within its gelatinous bell. This
is followed by a stage of drifting passively downwards and thereby ‘fishing’
for whatever items of prey happen to be in the way.514 At the bottom of
this descent, the jellyfish realigns itself by means of a set of gravity-sensing
statocysts akin to the otoliths of the vertebrate vestibular system to which
we owe our sense of balance. Having righted itself in this manner, Aglantha
can duly propel itself upwards again. Noteworthy is that if its statocysts
are removed, the swimming movement of the jellyfish becomes haphazard
and aimless. This shows just how reliant the successful feeding movement
of Aglantha is upon the implicit orientation associated with its statocysts,
the prerequisite for an invariant form of ‘dumb’ but highly effective taxis.
Insofar as the directionality of its movement is pre-established and thus
unvarying with respect to environmental contingency, consciousness is not
in itself entailed by such a strategy.
Still other free-swimming cnidarian medusas appear to possess a greater
degree of discernment of their environment. The Caribbean box jellyfish (or
cubozoan) Tripedalia cystophora (illustration page 161) has aroused particu-
lar interest on account of an elaborate visual system comprising 24 eyes of
four different kinds.515 Eight of these eyes are complex, lens-bearing sensory
structures morphologically similar to the vertebrate eye, sparking a certain
amount of controversy and speculation on their capacity for image-forma-
tion and ‘crude’ forms of vision.516 Even though the retina of T. cystophora’s
lensed eyes is commonly thought to lack the resolution needed to generate a
focused image, however, the sophistication of the animal’s visual system – in
conjunction with its ability to modulate the dynamics of bell contraction
and to regulate and channel bell outflow – permits it to adjust both the
speed and direction of its swimming to suit its needs.517 By means of rapid
phototactic locomotion, it is thus able to navigate towards the vertical light
shafts that tend to be populated by the dense swarms of copepods on which
it habitually preys. Once located within a light shaft, it maintains its position
there by non-directional modifications of its swimming behaviour, reducing
its speed, increasing its turning rate, and pivoting abruptly should it happen
to stray away. The result is that Tripedalia cystophora can discern and target
178 Epilogue: Consciousness in Simple Animals
the light shaft even though it does not see the copepods themselves, its
swimming behaviour within the shaft remaining largely unaffected by the
presence or absence of its prey.518
There remain important questions pertinent to the possible ascription
of consciousness to Tripedalia. Is its movement towards light in any way
modulated by levels of ‘hunger’ or ‘satiation’? If not, it may be dismissed as
an automatic feeder or ‘eating machine’, its navigation towards light little
more than an invariant taxis reminiscent of unicellular Paramecium, and its
persistence within the light shaft just non-directional kinesis. As with hydra,
we may continue to harbour residual doubts about the targeting of a prey
patch as opposed to an individual item.519 Yet the box jellyfish is perhaps the
most likely cnidarian candidate for consciousness on account of the greater
motility and more varied behavioural repertoire that set it apart from the
largely sedentary hydra. Its multifaceted visual system makes guided loco-
motion possible in situations other than feeding, enabling it (for example) to
avoid obstacles in the water and use visual cues to navigate back – if washed
away – to its preferred habitat among the mangrove roots.520 Tripedalia has
even been found to exhibit diurnal activity patterns (foraging in the day and
resting at night) that may imply a condition in some respects resembling
sleep.521 It would be precipitate to rule out rudimentary consciousness on
principle or simply because it is made of jelly and lacks a brain.
Two Worms
under stones. During the next 24 h, the planarians did not react to
a new portion of food. However, [after just] 2 days, when the food
appeared again, the planarians repeated, with rare exception, the
whole repertoire of their feeding behavior.530
The varying ‘inner state’ of the planarian can easily be distinguished on the
basis of the variation in its behaviour in otherwise similar circumstances:
when hungry, it performs the work of approaching its prey and activating its
pharynx; when sated, it ignores – or fails to perceive – the food and remains
quiescent. The satiety of Dugesia, which lasts two days, may be associated
with the temporary transformation of its intestines into a syncytium,531
this process possibly signalling repletion and inhibiting feeding behaviour.
Subsequently, the intestine structure is restored and the worm regains its
appetite. A further consideration is that the planarian lacks a through-gut,
its pharynx doubling as an anus.
After decapitation, by contrast, Dugesia no longer approaches the mos-
quito grub,532 and decerebrate planarians generally show a reduction in
guided, organized locomotion. Yet there are some species, such as Planocera
gilchristi, that continue to engage in feeding activity – albeit lacking in
coordination – provided that direct contact is made with the prey item.533
In these cases the manipulation and ingestion of food can occur even in the
absence of a brain. Significantly, satiety no longer inhibits further feeding.
So although the distinction between hunger and satiety holds for an intact
animal, decerebration seems to turn certain planarians – within their
mechanical limitations – into ‘eating machines’.534 An implication of this is
that, for planarians at least, the presence or absence of a brain makes the
difference between the possible occurrence or otherwise of rudimentary
consciousness. Simple rhythmic motions may be feasible without a brain,
but not complex, coordinated behavioural sequences attuned to the inner
state of the animal.535
Other behaviours and abilities evinced by flatworms are also closely
associated with consciousness. Though not in themselves sufficient for con-
sciousness to be inferred, such behaviours and abilities add dimensions that
may enrich its phenomenology. For a start, planarians periodically display a
state of rapidly reversible behavioural quiescence and reduced responsive-
ness to stimuli that ticks all the boxes to be classed as sleep.536 The quiescence
exhibited by planarians is not always a function of satiety, therefore, but
may conform to circadian rhythms, occurring in this form predominantly
182 Epilogue: Consciousness in Simple Animals
during the day (not surprisingly, given their aversion to light).537 Like sleep,
this underlying rhythm has been found to persist in conditions of continu
ous darkness, implying an endogenous origin, and it is homeostatically
regulated, with sleep deprivation having to be compensated by a subsequent
decrease in activity. Such quiescence usually assumes the form of a typical
contracted posture. As in other animals such as humans, moreover, it is
regulated by the hormone melatonin, pointing to an evolutionarily shared
origin. This distinction between wakefulness and sleep – between endog-
enously generated rhythms of activity and quiescence – provides further
evidence of a differential ‘inner’ state that has recognizable behavioural or
locomotive ramifications, manifesting itself as a disposition to move oneself
or not (in a given context).
A capacity for learning, as we have seen, is also commonly identified with
consciousness, although it is not essential to its most elementary form. Here
too planarians prove to have certain abilities. Dugesia japonica, for example,
shows non-associative modes of learning such as habituation, responding
to water turbulence by coming to a halt, but ceasing to react in this way if
the turbulence is recurrent.538 In the field of associative learning, Dugesia is
usually deemed rather a dullard, apparently requiring scores of training ses-
sions to relate a particular stimulus (a flash of light) with another stimulus
(an electric shock) that follows immediately afterwards.539 However, there
are other contexts in which planarians do demonstrate more of a memory. It
has been established that worms that have been fed in a particular environ-
ment will subsequently be quicker to start eating in this ‘familiar’ context
than others that have never previously been exposed to the feeding area.540
Such concepts of environmental familiarity or unfamiliarity clearly imply
that the planarian has some kind of relationship – albeit implicit – to its
relatively distant past and is not wholly swallowed up in its ‘present’.
Flatworms also have ‘preferences’. Their robust negative phototaxis
dictates that they generally eschew well-lit areas. If an enticing drop of liver
extract is placed in the middle of an illuminated area, therefore, the planar-
ian must ‘overcome’ its natural aversion to light – as well as its tendency to
hug the edges of containers – in order to venture out to procure its coveted
meal.541 This brings us back to the question of how much discomfort an
animal is willing to undergo in order to gain a reward, the phenomenon of
‘pleasure’ again emerging as the common currency by which motivational
drives – attractions and aversions – can be weighed up and a decision taken.
Experiments on the planarian Dugesia japonica have attempted to ascertain
Two Worms 183
If the arguments I have put forward are valid, it would be churlish not to
admit planarians such as Dugesia to the club of (sometimes) conscious
organisms. Given that the planarian is the only unequivocally successful
metazoan candidate so far, this might be taken to imply that consciousness
in animals requires a brain, dependent as it is upon coordinated and direc-
tional self-movement attuned to an internal state, say, of hunger or satiety.
This in turn would seem to rule out nematodes such as Caenorhabditis
elegans, a diminutive worm comparable in size to many protozoans (roughly
1.3 mm in length and 80 µm in diameter), the adult hermaphrodite of which
has a nervous system with just 302 neurons and some 7,000 connections.545
184 Epilogue: Consciousness in Simple Animals
But perhaps this hurried dismissal of what has been described as the ‘hydro-
gen atom of systems neuroscience’546 is unwarranted. Possessed of neither a
circulatory nor a respiratory system, the free-living soil-dweller C. elegans is
certainly capable of adapting its locomotion in a coordinated and integrated
manner to a wide range of sensory stimuli, including odorants, touch, light,
temperature and vibration. The head or neck region is equipped with a pair
of conspicuous sensory organs known as amphids, each of which contains
various chemosensory, mechanosensory and perhaps also thermosensory
neurons, making appropriately targeted self-movement feasible at least.547
At the same time, the first of our three criteria is certainly met. C. ele-
gans is robustly self-containing, in terms both of structural integrity548 and
functional integration. By contrast with planarians, it has only very limited
powers of regeneration. Apoptosis, or programmed cell death, is not only a
normal feature of its development, but a crucial part of the immune system
whereby infected or stressed cells ‘sacrifice’ themselves for the greater good
of the organism as a whole.549 It is clearly a self. Granted the selfhood of
each individual C. elegans organism, the aim in the remaining pages is to
cast a glance at the other two questions relevant to the possible presence of
consciousness: 1) does C. elegans display genuinely directional locomotion
in its pursuit of nourishment? 2) is its self-movement duly attuned to an
inner state, say of hunger or satiety?550
Some features of the nematode may suggest that the answer to the first
question is no. For a start, C. elegans is a bacterivorous filter feeder, taking
water with suspended food particles (bacteria) into its pharynx, trapping
the particles and then expelling the liquid.551 In itself, this suggests a meas-
ure of passivity and non-discernment in its feeding behaviour insofar as the
worm does not strictly ‘choose’ – i.e. individually pick out and pursue – the
food that ends up in its pharynx. To be sure, it is a self-propelling filter feeder,
which means that the work of locomotion is involved. Like the basking
shark considered above, in fact, C. elegans has the work both of propelling
itself through a prey patch shovelling up whatever food it can, and of finding
that prey patch in the first place. Yet it is clear that we are not dealing with a
predator that perceives and identifies its prey on a one-by-one basis.
A second doubt is whether the nematode moves by orienting itself in the
direction of a gradient (the target being where its next meal is) or adopts
an essentially non-directional mode of locomotion, i.e. kinesis. The latter
has already been seen to subsume a distinction between orthokinesis and
klinokinesis. These strategies involve modulation of the animal’s speed and
Two Worms 185
By contrast with the biased random walk, however, it has been argued
that the pirouette functions not by randomizing the animal’s orientation, but
by correcting its course. If this is so, a form of error compensation may be
achieved by correlating (albeit weakly) the size of the turn with the degree to
which the worm has veered off course immediately prior to the pirouette.565
Nematologists have proposed that C. elegans pirouettes may thus represent
‘a transitional stage between biased random locomotion and continuous
alignment with the direction of the gradient’.566 In fact, it seems likely that
the course-correction and weathervane techniques may function in concert,
complementing and reinforcing one another.567
These different techniques – run-and-tumble, the pirouette and ‘weath-
ervane’ continuous alignment – incorporate different levels of randomness
and directionality, bringing to light once again that the elementary con-
sciousness with which guided locomotion is associated is unlikely to be
an all-or-nothing phenomenon. Even though the question of whether the
nematode’s self-propulsion is genuinely directional is best left as undecided,
however, the question of its attunement to an inner state is much less
ambivalent.568 We have already noted that the reduction in speed shown
by C. elegans on being presented with a meal of bacteria is modulated by its
hunger state. Whereas recently fed worms exhibit a ‘basal’ slowing response,
food-deprived worms show a slowing response that is recognizably
‘enhanced’. Whether the nematodes are hungry or satiated thus determines
differential behaviour in the face of external stimuli and can be identified
as such.569
This is not all. A state of satiety in C. elegans has been found to express
itself as a behavioural ‘quiescence’ that consists in decreased movement
and a gradual cessation of food intake.570 This quiescence bears a close
resemblance to satiety in other animals. For a start, it is brought on by
high-quality food,571 which, unlike low-quality food, induces the worm
gradually to stop eating and moving about. It also depends on the worm’s
nutritional status. Mutants deficient in ingestion (pharyngeal pumping) or
nutrient absorption show less quiescence, which is thus assumed to require
the appropriate signals from the intestine. Thirdly, worms that have fasted
prior to feeding are more likely to reach this state of satiety, a previous
period of enforced abstinence thus augmenting subsequent quiescence.572
Intriguingly, mutants lacking a particular kinase573 (a protein that among
other functions determines locomotor states and body size control) do not
show quiescence at all, but constantly move and feed. Failing to attain satiety,
188 Epilogue: Consciousness in Simple Animals
Allo-: preface signalling the opposite of auto-, i.e. by ‘non-self ’ or ‘other’: e.g. if auto-
poiesis denotes self-creation, allopoiesis is creation by non-self or other; in the present
context allo-containment is used to denote containment by non-self or other, as opposed
to self-containment
Arthropod: member of the phylum Arthropoda (‘animals with jointed legs’), which
includes insects, spiders, scorpions, crabs and the extinct trilobites; they have an exo-
skeleton, or external skeleton
ATP: adenosine triphosphate: a molecule that stores and transports the energy required
to power living processes
Autotroph: an organism that can produce complex organic compounds from simple in-
organic compounds, for example by using the energy from sunlight (photosynthesis)
Bilateria: the clade of animals characterized by bilateral symmetry at some stage in their
life cycle, i.e. possessing a front and a back, a top and a bottom, and a right and a left; it
includes the protostomes and the deuterostomes (q.v.), but not sponges or cnidarians
such as corals or jellyfish
Chordate: member of the phylum Chordata, which includes vertebrates, tunicates (such
as salps and sea squirts) and cephalochordates (whose extant representative is the lance-
let, also known as amphioxus); they are characterized, among other things, by posses-
sion of a notochord for at least part of their life cycle
Ciliate: group of protists whose cell surface features hair-like extensions called cilia; like
flagella (q.v.), cilia serve as organelles of motility but they are generally shorter and more
numerous than eukaryotic flagella
Cnidaria: a phylum of exclusively aquatic animals that include corals and sea anemones,
jellyfish and box jellyfish, freshwater hydras and venomous siphonophores such as the
Portuguese man o’ war
Cytoplasm: the contents of a cell that are outside the nucleus (if there is one) but within
the cell membrane
Glossary 193
Cytoskeleton: dynamic protein structure in the cytoplasm of cells, fulfilling a wide range
of functions including the maintenance of cell shape, locomotion, intracellular trans-
port and cell division
Diatoms: group of mainly unicellular photosynthetic algae endowed with glass tests or
shells known as frustules; they rank among the world’s most abundant aquatic organ-
isms
Endosymbiont: an organism that lives inside the body or cells of another organism in a
mutually beneficial relationship of endosymbiosis
Epithelium: type of animal tissue, taking the form of one or more layers of cells that
cover a body’s outer surface and line its cavities
Euglena: a genus of flagellate protist, most species of which have chloroplasts and can
photosynthesize but are also able to feed by phagocytosis (q.v.)
Eukaryote: an organism whose cells contain their genetic material within a mem-
brane-bound nucleus
Foraminifera: class of mainly marine protists with pore-studded tests or shells from
which microtubule-reinforced projections emerge
Haptorid: a class of ciliates that includes Didinium and Dileptus, which capture or kill
their prey mainly by firing toxic filaments known as trichocysts
HGT, horizontal gene transfer (also: lateral gene transfer): transfer of genes between
organisms other than by parent-offspring transmission; it contrasts with vertical gene
transfer, which is by sexual or asexual reproduction
Interneuron: a class of neuron that conveys information from one neuron to another
Isomerization: process by which one form of a molecule can change into another form
with the same atoms but arranged in a different configuration
Isopleth: a line connecting points that register the same amount or ratio of some meas-
urable variable
Kinase: an enzyme that catalyses the addition of phosphate groups to proteins, thus
activating protein function; the enzymes that catalyse the removal of phosphate groups
and thus the de-activation of protein function are called phosphatases
Glossary 195
Lipid: any of a large and diverse group of organic compounds that are insoluble in water
but soluble in organic solvents such as alcohol, ether and chloroform; examples include
fats, oils, waxes and steroids; they are among the main constituents of plant and animal
cells
Macrophage: a type of white blood cell whose functions include the phagocytosis (q.v.)
of pathogens as part of the immune system
Mitochondrion: organelle in eukaryotic cells that uses energy from aerobic respiration
to synthesize ATP (q.v.), commonly termed the ‘powerhouse’ of the cell
Myxobacteria: (also known as ‘slime bacteria’) a group of bacteria that includes the
model organism Myxococcus xanthus; noted for travelling as ‘swarms’ and aggregating as
‘fruiting bodies’ when nutrients are scarce
Osmotrophy: a form of nutrition that involves the uptake of small organic molecules by
osmosis
196
Phagocytosis: process by which a cell engulfs a solid food particle; phagocytes are im-
mune cells that ingest potentially harmful non-self in this way
Phagosome: a vesicle (q.v.) formed during phagocytosis (q.v.) when a food particle or
prey item is engulfed and internalized by the cell membrane; the phagosome subse-
quently fuses with a lysosome (q.v.) for digestion to take place
Phospholipid: one of a class of lipids (q.v.) that includes both a phosphate group and one
or more fatty acids
Placozoa: one of the most basal groups of Metazoa, generally considered to consist of a
single species, Trichoplax adhaerens
Polymer: a molecule that consists of many repeated elements (monomers): e.g. proteins
are polymers of amino acids; DNA and RNA are polymers of nucleotides; polymerization
is the process by which polymers are formed
Proteobacteria: a major group of Gram-negative bacteria that includes the genera Rick-
ettsia, Escherichia, Salmonella and Buchnera as well as myxobacteria such as Myxococcus
Protostome: a superphylum of bilaterian animals that include the arthropods (q.v.), mol-
luscs, annelid worms and nematode worms
Radiolaria: major class of protozoa (q.v.) characterized by a delicate silica skeleton and
long thin projecting pseudopods known as axopods
Retroelement, retrotransposon: a genetic element that can multiply its presence within
a genome by using an enzyme to turn RNA copies of itself back into DNA, which may
then be inserted into the genome
Social amoebae: (also known as ‘cellular slime moulds’) a class of amoeboid protists that
includes the model organism Dictyostelium discoideum
Suctoria: a class of ciliate that is sessile in its developed stage and feeds by means of
projecting tentacles
Taxis: directional movement of a motile organism towards or away from a stimulus such
as light (phototaxis), chemicals (chemotaxis), physical contact (thigmotaxis) or gravity
(gravitaxis); it is characterized as positive or negative depending on whether it is ‘to-
wards’ or ‘away from’ the stimulus; it is distinguished from kinesis, which is a non-direc-
tional response to a stimulus
Trophont, trophic stage: the feeding stage of an organism, especially of a ciliate; in other
contexts, the term trophozoite is sometimes used
Vacuole: a type of vesicle (q.v.), a membrane-bound space within the cytoplasm of a cell,
often used for storing food, water or waste
Virion: a complete virus particle comprising both the genetic material and the protective
capsid
Viroid: a short, circular, infectious, single-stranded RNA molecule that does not code for
any proteins and does not have a protective capsid
1 Glasgow (2017); the book can be downloaded for free from the publisher via the following link:
https://nbn-resolving.org/urn:nbn:de:bvb:20-opus-145252.
2 Realistically, the aim is thus not necessarily to serve up all the right answers in this analysis of
rudimentary consciousness. Our empirical understanding of (for example) the locomotion of
single cells will undoubtedly be fine-tuned with time, and this is likely to have a bearing on some
of the conclusions proposed. Rather than providing answers, the objective is – given a particular
understanding of ‘selfhood’ based on intrinsic reflexivity – to unearth the questions that are most
relevant in considering whether consciousness can be ascribed to a particular form of life. I owe this
interrogative approach to W. D. Glasgow, who believed strongly that the purpose of philosophy is not
to produce ready-made answers, but to learn how to ask the right questions.
3 The other main approach to the concept of ‘selfhood’ derives from the emphatic use of the pronoun.
On this view, your self is what you are, or perhaps what you really are. The implication tends to be
that selfhood denotes some kind of timeless essence. Intrinsically reflexive selfhood does not imply
an essence, but nor does it preclude the possibility of one.
4 The attribute ‘reflexive’, based on the term’s grammatical use, is to be distinguished from that of
‘reflective’, which denotes the process of reflection or thought and is here employed primarily in the
context of ‘pre-reflective self-awareness’, a form of awareness that is prior to explicit thought processes
or mental representations. ‘Reflexive’ is also to be distinguished from the notion of a reflex in the
sense of an automatic bodily response to a stimulus. Here the adjective used is ‘reflex’, as in a ‘reflex
reaction’.
5 On the contrast between self-organization and self-assembly see Glasgow (2017), 63– 64.
6 In fact, the formalism in question is of a rather special kind. Intrinsically reflexive processes are
constitutive of a type of entity that in some sense forms itself. As proposed in Glasgow (2017; 24), the
coinage self-formalism captures an important feature of intrinsic reflexivity in marrying formalism
with process and change. One might say that – like Aristotle’s ‘soul’ – a self is not just a form, but a
202
form constantly engaged in and constituted by the process of sustaining itself as the form that it is, as
well as adapting this form (within limits) to changing circumstances. This is suggested by Aristotle’s
notion of entelechy, understood as denoting the work of self-maintenance.
7 On the selfish or self-like nature of energy flows and genetic material, see Glasgow (2017), chapters
II and III respectively. On the selfhood inherent in organisms and biological individuals, see chapter
VI. Forthcoming works, I hope, will look at the selfhood of super-organisms, biospheres and universes.
8 For a discussion of whether ‘reproduction’ really is ‘self-reproduction’ and can thus aptly be
described in terms of intrinsic reflexivity – in other words, the unanswerable logical quandary thrown
up by the question of whether progenitor and progeny really are ‘the same’ – see Glasgow (2017),
38–45.
9 See Maynard Smith and Szathmáry (1999), 3, according to whom natural selection presupposes
a population of entities endowed with the properties of multiplication (or, in intrinsically reflexive
terms: self-multiplication), heredity and variation.
10 On the relationship between selfhood and selfishness see Glasgow (2017), 53–56, 101– 4.
11 By contrast, the individual cells that make up multicellular bodies such as animals and plants are
deemed not to be selves. For an explanation, see the subchapter below on ‘Metazoan Cells’.
12 As noted above, the lack of one or more of the three fundamental categories of intrinsic reflexivity
results in something that is merely selfish or self-like. The absence of self-maintenance, for example, is
typified by the non-metabolic selfishness characteristic of viruses or virus-like entities. The absence
of self-multiplication rules out heredity, variation and the concomitant possibility of evolution
by natural selection among a population of entities. In the absence of self-containment, intrinsic
reflexivity takes the form of the dissipative structures exemplified by self-organizing flow patterns or
self-propagating forest fires, which are unable to channel their energetic flow into the performance
of (constructive) work.
13 See Bray (2009), 89–93.
14 Ibid., 6.
15 Jablonski (2006), 17.
16 On Paramecium, see Greenspan (2007), 5 –21.
17 See Dawkins (2004), 398.
18 On the shadow reflex of barnacles and the molecular basis of photoreception in general see
Greenspan (2007), 23– 40; on the evolution of sight see Lane (2009), 172–204.
19 Halobacterium, for example, uses bacteriorhodopsin to capture light energy and produce a proton
gradient across its cell membrane. This proton gradient is then employed to synthesize the energy-
carrying molecule ATP and thus power cellular activity.
20 On warnowiids such as Erythropsidinium, see Gómez et al. (2009).
21 The restoration of balance is generally not immediate, since time is required for it to take effect.
However, it is anticipated by bodily signals of ingestion. See page 95.
22 This is not to say that such ‘imprisonment’ can never be to the advantage of the ‘prisoner’, as in
Endnotes 203
the case of the protection provided by endosymbiosis, which makes self-movement largely pointless.
On the endosymbiotic existence of formerly free-living bacteria such as Buchnera aphidicola, which
are now (more or less) permanent inhabitants of specialized cells in their greenfly hosts, see Glasgow
(2017), 182– 86.
23 It may be objected that rudimentary consciousness does not in fact presuppose self-reproduction.
Indeed, there is a slight asymmetry in the relationship of consciousness to self-maintenance and its
relationship to self-reproduction, in that the original function of consciousness is to enable a hungry
self to get from ‘here’ (no food) to ‘there’ (food) in order to fuel itself and thus keep itself going. But this
is far from being the only plausible reason for guided self-propulsion. Once sexual reproduction has
emerged, it is logically feasible that nutrient uptake might occur purely passively and so necessitate
neither movement nor consciousness; consciousness, meanwhile, might arise from a periodic need
to find a mate rather than prey, i.e. to get from ‘here’ (no mate) to ‘there’ (mate). Even in this scenario,
however, the directed self-propulsion here deemed necessary for consciousness always presupposes
energetic self-maintenance, since the self must have fuelled itself and must have energy surplus to its
own immediate metabolic requirements for movement to be possible. If this is so, then consciousness
does not logically require full minimal selfhood (i.e. all three forms of intrinsic reflexivity), but merely
the quality of being self-like (i.e. two out of the three modes of intrinsic reflexivity). As it occurs on our
planet, of course, consciousness does seem to coincide with full minimum selfhood, and it is difficult
to conceive of the requisite self-containment and self-transcendence occurring in a self that is not
itself the product of a process of evolutionary self-transformation through natural selection among a
population of self-reproducing selves. Whether this is merely an empirical contingency or reflects a
deeper logical limitation may simply be unanswerable at present. An example of a self-like entity that
has not yet reproduced itself (as far as we know) is the biosphere of our planet. Would we be willing to
ascribe consciousness to the biosphere as an entity if we were aware of it ‘behaving’, i.e. moving itself
directionally in relation to non-self, say, by planet-hopping? As it happens, such planet-hopping on
the part of the biosphere might even coincide with reproduction.
24 Protozoa may be categorized as flagellated, ciliated or amoeboid, depending respectively
on whether they move and feed using flagella, cilia or transient extensions of the cell body called
pseudopodia. However, these broad categories may overlap: pelobionts such as Mastigella, for
example, are amoeboid cells equipped with a flagellum; heteroloboseids such as the facultative
pathogen Naegleria fowleri have a life cycle incorporating both an amoeboid and a flagellate stage.
25 Throughout this work the terms ‘rudimentary’ and ‘elementary’ will be used interchangeably to
describe consciousness in its logically minimal or most simple manifestation, whether applied to
single-celled or multi-celled organisms.
26 The term ‘drive’ does not enjoy the best of reputations; it has often been dismissed as speculative,
metaphysical or simply vacuous. I here use the term as an alternative to ‘appetite’, especially in contexts
where the object relates to reproduction (or sex) rather than nutrition. Like ‘appetite’, the concept
implies both mechanical and purposive elements, suggesting both a physiological imbalance and
204
directedness towards a goal object. For a critique of the concept of ‘drive’, see Peters (1958), 93 –129.
27 On the notion of ‘structural coupling’, see Glasgow (2017), 52; see also Thompson (2007), 45.
28 See Jonas (1966; 2001), 84.
29 See Ari Berkowitz (2016; 85 –90) and Rodolfo Llinás (2001; 5 –7), who cite in particular the work
of the influential physiologist T. Graham Brown on cats in the early part of the 20th century. In most
vertebrates, it is the spinal cord rather than the brain that generates such rhythms, functioning rather
like a pacemaker.
30 Notably, it is only skeletal muscles – not the cardiac muscles responsible for pumping blood or the
smooth muscles responsible for gut movements – that are under voluntary control.
31 The underlying point is that consciousness is presumed, through the evolutionary eons, to have
fulfilled a function rather than exist as a mere epiphenomenon, a functionless by-product of cognitive
processes or nervous activity. Epiphenomenalism raises the question of why consciousness – if it
serves no function – should be associated with nervous or cognitive events and yet not with other
physical processes such as photosynthesis or combustion. See Morsella (2005), 1001.
32 Aristotle’s notion of ‘soul’ or psyche is akin to a ‘principle of animation’ by virtue of which a
thing can be said to be living. ‘Mind’ is a much more problematic term, but happens to be the more
common of the two in post-Cartesian philosophy. In what follows, I shall assume that anyone who is
prepared to claim that a particular entity has a mind will also be willing to ascribe the possibility of
consciousness to that entity. I shall treat the term ‘consciousness’ as synonymous with ‘awareness’, but
opt for ‘self-awareness’ over ‘self-consciousness’ on account of the distracting associations of the latter
with undue diffidence or bashfulness.
33 See Glasgow (2017), 89 – 90.
34 Aristotle, De Anima, (Book II.3), 414a –b. See also Aristotle (1986), 162.
35 See Schloegel and Schmidgen (2002), 621.
36 For an account of the ‘cellular psychology’ of Haeckel, Verworn, Binet and others, see ibid., 622 –33.
37 Jennings (1906), 336.
38 See Thompson (2007), 161.
39 This is not the same as what is known as psychological behaviourism, which is a method for
the study of human beings and animals and which we shall come across below. However, both
behaviourisms share an emphasis on what is accessible to public observation, and the former has
been interpreted as providing philosophical vindication for the latter. For an overview of logical
behaviourism see Priest (1991), 35 – 64; see also Shaffer (1968), 14 –17.
40 For a start, philosophical functionalism is generally treated as a theory of mind, not of consciousness.
Yet in spite of their origins in different realms of discourse, there is substantial concurrence between
the two sorts of functionalism in the account I am offering. The adaptive function of consciousness
consists precisely in the production of behaviour, where ‘behaviour’ is understood as a particular sort
of guided self-movement undertaken by a self-maintaining self. A functionalist theory of mind also
interprets mind in terms of the production of behaviour.
Endnotes 205
between consciousness and behaviour. A particular disposition to move may fail to be realized; the act
of speaking may have to be substituted by a complex pattern of eye movements or by the movement
of a cursor on a computer screen. On locked-in syndrome see for example Gazzaniga (2008), 338 –39.
55 See Priest (1991), 36.
56 Quoted in Gregory (ed.) (1987), 72.
57 The claim that ‘consciousness does not exist’ is as metaphysical as the claim that ‘God does
not exist’ and certainly inconsistent with the methodology of psychological behaviourism. Logical
behaviourism, by contrast, might have contended that claims about consciousness – including whether
or not it exists – are meaningless. This is more in line with Watson’s later claim that ‘consciousness is
neither a definite nor a usable concept’; see Watson (1930; 1970), 2.
58 To the extent that it incorporates a metaphor of ‘height’ with humans on top, the idea of a scale
of phylogeny is utterly and irrevocably misleading. It also tends to be harmfully anthropocentric. All
extant organisms constitute a ‘pinnacle’ on the time axis simply by virtue of being extant.
59 Tye (1997), 289.
60 Ibid., 301–3.
61 In effect, a zombie in the philosophical sense is a being that is capable of normal self-movement
or behaviour but is bereft of consciousness.
62 To Michel Cabanac (1999), for example, the signs of emotional tachycardia and elevated
body temperatures elicited by gentle handling in reptiles but not in amphibians suggest that the
phylogenetic emergence of emotion – which is sometimes taken as a proxy for consciousness – lies
between amphibians and early reptiles. However, the narrative of a phylogenetic progression from
emotion-less to emotionally endowed animals is problematic. There is increasing evidence of the
capacity of fish – though ostensibly more ‘primitive’ in phylogenetic terms than amphibians and
reptiles – to suffer and feel emotion; see Braithwaite (2010), Balcombe (2006), 185 –91. On the
relationship between emotion and consciousness, see below, chapter IV.
63 On the overlapping size ranges of protozoans and metazoans, see McMahon and Bonner (1983),
1–5, also Patterson and Hedley (1992; 1996), 27–28.
64 For an illustration of Megaphragma see Polilov (2012), 30; on the fairyfly see Mockford (1997), 115 –20.
65 Jennings (1906), 336 –37: ‘If [the amoeba] were as large as a whale, it is quite conceivable that
occasions might arise when the attribution to it of the elemental states of consciousness might save the
unsophisticated human being from the destruction that would result from the lack of such attribution’.
66 Bennett and Hacker (2003), 303. Of course, the question of ‘complexity’ is of some relevance to
consciousness. A ‘complex’ brain may provide increased opportunities for behavioural complexity
and flexibility; it may thus have a decisive influence on the nature of consciousness.
67 Ibid., 304.
68 Jonas (1966; 2001), 2.
69 Glisson held that the body’s natural energy is transformed by irritability into sensation. See
Giglioni (2008) for an account of why Glisson’s theory of irritability promptly fell into oblivion even
Endnotes 207
though the concept of irritability was to flourish under the less speculative influence of Albrecht
Haller. For Haller irritability was merely a manifestation of mechanical elasticity.
70 Schelling was also influenced by the work of the 18th-century Scottish physician John Brown on
‘excitability’.
71 See Binet (1889), 64, 105.
72 Ibid., 106.
73 For an example of this usage, see Llinás (2001), 113: ‘we know that single-cell “animals” are
capable of irritability, that is, they respond to external stimuli with organized, goal-directed behavior’.
Elsewhere (212), Llinás defines ‘irritability’ as an ability to produce a behavioural response to stimuli
consisting of ‘either moving away from, or approaching an object or another cell’. This is tantamount
to taxis.
74 Bray (2009), 25 – 26.
75 Some single-celled eukaryotes do indeed show phototactic behaviour. We shall return below
(chapter VI) to the question of how far this may be taken to imply vision.
76 For an example of such a theory see Rolls (1999), 244– 65.
77 On the term ‘feeling’, see below, chapter IV.
78 Donald Griffin, cited in Balcombe (2006), 28.
79 See note 4 above on the distinction between ‘reflexive’ and ‘reflective’. Of course, the act of reflecting
upon oneself reflecting is itself reflexive insofar as the grammatical subject (the thinker) is the same
as the grammatical object (the thinker). In fact, it is doubly reflexive: the thinker is reflecting upon
himself, and the reflections are ‘about’ themselves. Yet in neither respect is the reflexivity intrinsic. The
thinker, even qua thinker, is not constituted by this reflexivity, for he can think about other things and
he can stop thinking and do other things. Equally, the sequence of reflections may cease to be ‘about’
the act or process of reflection and instead focus on the thinker’s upcoming social commitments, a
persistent itch, or the sounds of connubial disharmony from next door. Sometimes the rather vague
word ‘self-reflection’ is used, suggesting both reflection and reflexivity. It is in the nature of reflection
that the reflexive relationship between the reflection and what is reflected is extrinsic.
80 See Lurz (2009), 9.
81 See Zahavi (2005), 18.
82 For an exemplary discussion see Zahavi (2005), 17– 30.
83 On corollary discharge, see Crapse and Sommer (2008). One of the classic papers is Von Holst
and Mittelstaedt (1950), who proposed that the organism sends to the sensory pathway a copy
(designated an ‘efference copy’) of the motor command issued to an effector such as a muscle. This
motor copy would then be combined, juxtaposed or in some way compared with the input from the
sensory receptors.
84 Merker (2005).
85 Ibid., 93.
86 Of course, one might imagine the consciousness of an absolutely stationary self that simply records
208
its surroundings. But what would such consciousness be for? How could consciousness emerge from
such a purely passive process?
87 On the mechanism of corollary discharge in the sea-slug Pleurobranchaea, see Crapse and
Sommer (2008), 589 – 90.
88 Thompson (2010), 162.
89 Ibid.
90 Of the four known phyla of deuterostomes, only chordates such as vertebrates have a centralized
nervous system. Echinoderms (which include starfish, sea lilies, sea urchins and sea cucumbers),
hemichordates (which include acorn worms and pterobranchs) and worm-like Xenoturbella have
radial or net-like systems. If the last common ancestor of all deuterostomes had a central nervous
system, which is believed to be the most parsimonious option, these lineages must have reverted
from a central nervous system to a diffuse nervous system. This was presumably because it was better
at keeping them alive long enough to reproduce. The deuterostomes constitute the sister clade to the
protostomes, which include arthropods (insects, spiders and crustaceans), nematodes, annelids and
molluscs. On the phylogeny see Moroz (2009), 184.
91 Llinás (2001), 78.
92 On the capacity of protein networks to transmit and process information, see Bray (2009), 71– 88.
93 Wittgenstein (1953), § 281.
94 So what is implied by the distinction between ‘naked’ amoebae and the ‘testate’ amoebae that
have thrown – or whose ancestors have thrown – a cranium-like exoskeleton around much of their
brain-cum-body?
95 See Crick and Koch (1990), esp. 270 –72.
96 Penrose (1995), 348 – 92.
97 Tononi (2004) likewise focuses on the thalamocortical system, arguing that ‘the fact that
consciousness as we know it is generated by the thalamocortical system fits well with the information
integration theory, since what we know about its organization appears ideally suited to the integration
of information’.
98 See Morsella (2005), 1010.
99 Chalmers (1995), 204 –5.
100 Morsella (2005; 1015) himself notes that ‘although intersystem integration could conceivably
occur without something like phenomenal states (as in an automaton or in an elegant “blackboard”
neural network with all of its modules nicely interconnected), such a solution was not selected in our
evolutionary history’. Yet this seems to sever consciousness from any logical connection to its function.
101 As was shown in the final chapter of Glasgow (2017), this ‘within’ need not imply spatial
boundaries, but rather ‘belonging to’ or being ‘a functional part of ’ the system in question.
102 Tononi (2004), by contrast, equates consciousness with ‘the ability of a system to integrate
information, whether or not it has a strong sense of self, language, emotion, a body, or is immersed in
an environment, contrary to some common intuitions’. The divorce between consciousness and any
Endnotes 209
sort of environment rather contravenes the thermodynamics of information processing, which relies
on an input of energy.
103 We tend to be led astray, again, by the metaphor of mental representations or images. The
implication is that they are something the brain has to assemble or put together from a multiplicity of
informational components. Yet an amoeba perceiving a paramecium does not sense a mental image
of a paramecium; it senses the paramecium. I do not perceive a mental representation of the nut roast
on my plate; I perceive the nut roast.
104 See Berkowitz (2016), 121–30, for a fascinating account of how this alignment is brought about
in a barn owl.
105 For a discussion of this operation, known as commissurotomy, see Glover (1988), 32– 35; see
also Gazzaniga (2008), 289 – 92.
106 See Sass and Parnas (2003), 435 (italics omitted).
107 On schizophrenia and proprioception see ibid., 427– 44; on schizophrenia and corollary
discharge see Blakemore et al. (2000).
108 Chalmers (1995), 200.
109 Ibid., 215 –17.
110 Ibid. Perhaps the most urgent question is whether all information has a phenomenal aspect. If it
does, then the ubiquity of information entails the ubiquity of consciousness.
111 Nagel (1974).
112 Given the behavioural parallels, the burden of proof for the counterclaim that it is not like
anything lies with the counterclaimant. As we have seen, this might be achieved by showing that the
movements of a predator, be it macroscopic or microscopic, do not genuinely count as ‘behaviour’
or ‘action’.
113 See Glasgow (2017), 36 – 37.
114 On Buchnera and other such endosymbionts see ibid., 182– 86.
115 On the functioning of such genetic circuits see Bray (2009), 179 – 81.
116 See Xu and Gordon (2003).
117 On Rhodopseudomonas palustris see Brüssow (2007), 180 – 82.
118 Broadly speaking, the four main modes of metabolism are photoautotrophy (deriving energy
from light and carbon from carbon dioxide), photoheterotrophy (acquiring energy from light but
carbon from organic compounds), chemoautotrophy (using inorganic compounds for energy and
carbon dioxide for carbon), and chemoheterotrophy (deriving both energy and carbon from organic
compounds). See ibid., 180.
119 Ibid., 181.
120 See Turney (2015), 222– 23.
121 Shapiro (2007), 814, 807.
122 Ben-Jacob (1998), 58.
123 The focus is here on appetite rather than aversion. More generally, one should perhaps say that it
210
with gustatory selection: having bumped into an item of food at random, the ‘decision’ to proceed
or not with the work of operating one’s mouth might depend on whether what one happened upon
tasted ‘good’ or ‘bad’. Would we be unwilling to ascribe consciousness to such a creature? What about
a microscopic carnivore such as the tardigrade Macrobiotus richtersi, a tactile predator that detects its
nematode prey by touching them with the sensory area around its mouth (the circumoral field)? In
the absence of eyes, it is uncertain whether chemical cues also provide information to guide it in its
nematode harvesting (see Hohberg and Traunspurger (2009)). Or what about the much-discussed
star-nosed mole, which likewise makes scarce use of teleception (it has a singular ability to smell
underwater but is more or less blind) and whose perception of the world, or at least of its food, appears
to be basically tactile or gustatory? This creature’s remarkable star-shaped nose is used primarily not
for olfaction but for ‘touching’ its prey as it moves along, employing a mechanical detection system that
is more rapid than any known optical system. Believed to be the fastest-eating mammal on the planet,
the star-nosed mole takes under a quarter of a second to identify, process and ingest individual food
items, its brain deciding in a matter of milliseconds whether they are edible or not. Few would deny
some form of consciousness to this breathtakingly efficient forager, especially given that foraging is far
from being its only activity. It is thought to live socially in loose colonies within networks of tunnels
of its own making, and it partakes in the range of activities associated with sexual reproduction. But
would we be so generous with the tiny tardigrade Macrobiotus, which has a similarly tactile feeding
strategy but may otherwise be more limited in its behavioural repertoire? My feeling is that a system
involving only random encounter in juxtaposition with gustatory or mechanical contact-based prey
selection would not in itself provide the foundation for consciousness. On the foraging speed of the
star-nosed mole, see for example Catania and Remple (2005), who describe how, ‘in an astounding
flurry of star movements and prey captures’, a mole locates and eats eight separate prey items in under
two seconds (521).
157 Harold (2001), 20.
158 See Rothschild (1989), esp. 280 – 90. A third term proposed for the microscopic kingdom was
‘Protoctista’, coined by the naturalist John Hogg and meaning ‘first created beings’. In the present
context I have often opted for the informal category of ‘protozoa’ to denote unicellular eukaryotes
capable of locomotion.
159 In their book Free-Living Freshwater Protozoa, Patterson and Hedley (1992; 1996) thus define
plankton in terms not of movement but of where they live, namely in the water column (i.e. above the
sediment). Planktonic protists, they point out, may either swim or drift (203).
160 Barnes (ed.) (1998), 63 – 64.
161 Ibid., 59 – 60.
162 Money (2014), 61: the phylum of dinoflagellates likewise includes many species considered
photosynthetic (about half of those that are known), but also many others that have either lost or
never had a chloroplast and that feed on diatoms or other living organisms. Many both possess a
chloroplast and actively ingest prey. These are known as mixotrophs.
Endnotes 213
163 See for example Verity (1991); Jürgens and DeMott (1995); Montagnes et al. (2008).
164 See Barnes (ed.) (1998), 97– 100; see Glasgow (2017), 274.
165 See Patterson and Hedley (1992; 1996), 177-79. In the case of the suctorians, each tentacle-
cum-mouth terminates in a knob that houses an ‘extrusome’, i.e. an organelle whose contents can
be extruded so as to hold on to the prey as its cytoplasm is extracted. In some cases, the prey may
be released (alive) once the suctorian has had its fill. Many suctorians are supported on a stalk and
contained within an extracellular lorica (shell).
166 The category of suspension feeding includes filter feeding or sieving on the one hand but also
various mechanisms of what is known as ‘aerosol’ filtration on the other. Aerosol forms of particle
capture involve filters that are additionally characterized by adhesive properties. See LaBarbera
(1984), 76 – 81.
167 Ibid., 71 – 72.
168 See Fenchel (1980).
169 Verity (1991), 70.
170 Ibid.; see Snyder (1991) on the chemoattractive response shown by various bacterivorous ciliates
to compounds normally found on the surface of prey cells, suggesting an ability to sense dissolved
substances derived from their prey.
171 Verity (1991), 70.
172 See Jennings (1906), 174 –79. For accounts of Jennings’ work on Stentor, see also McDougall
(1923), 66 – 67; Bray (2009), 14 –17.
173 Jennings (1906), 185 – 86.
174 Berger (1980), 402-3.
175 Dusenbery (2009; 152 –53) shows that – on account of the laws of small-scale hydrodynamics –
random movement does not increase the nutrient encounter rate for a medium-sized bacterium with
a diameter of ca. 1 µm feeding by osmotrophy in a uniform environment. In itself, swimming is not
worth the trouble. A bacterium of such proportions may as well just rely on encountering nutrition by
diffusion. According to Dusenbery’s calculations, an organism has to be more than 10 µm in diameter
for random locomotion to pay its way by enhancing nutrient uptake.
176 Harold (2001), 28; for a comparison of prokaryotes and eukaryotes see 25 –30.
177 The Reynolds number is defined as ‘the fluid density times a characteristic length times a
characteristic speed divided by the fluid viscosity’. In a given context this means it is a ratio ‘between
inertial and viscous forces per unit volume’. On the Reynolds number see McMahon and Bonner
(1983), 89 –97; see also Dusenbery (2009), 41– 49.
178 Dusenbery (2009), 44 – 45.
179 See McMahon and Bonner (1983), 197– 98, quoting the physicist Edward Purcell.
180 Ibid., 195 – 96: ‘the stopping distance, expressed as a fraction of the cell’s diameter, is one-eighteenth
of its Reynolds number based on its diameter and its initial speed. For a bacterium 2 micrometers long
swimming at a Reynolds number of 10– 6, the stopping distance is about 10– 7 micrometers’.
214
181 A further determinant is the reversibility of flow at low Reynolds numbers, as a result of which
self-propulsion cannot be achieved simply by moving a rigid, oar-like appendage back and forth, even
varying the speed in the two halves of the cycle. This is possible at high Reynolds numbers, endowing
bivalve molluscs such as scallops with a rather crude form of aquatic locomotion based on their
ability to open their shells slowly and then snap them shut. At a low Reynolds number, however, this
would get the scallop nowhere, but merely shift it back and forth. On how flagella and cilia overcome
the challenge posed by reversible microscopic flow, see Dusenbery (2009), 198 – 214.
182 The mechanical efficiency of rotating bacterial flagella has been estimated to be slightly more
than 1 %. By comparison, fish are calculated to attain efficiencies of up to 80 % at high Reynolds
numbers by pushing against water with their bodies; see ibid., 213 –14.
183 Ibid., 4.
184 Ibid., 231.
185 See ibid., 239 – 42; 248.
186 See ibid., 20, on how the size range for each genus was measured. Dusenbery stresses the
strength of this correlation, calculating that if motility and size were distributed independently, the
probability that 19 % of the non-motile genera were smaller than the smallest motile genus would be
less than two in a million (ibid., 22).
187 See Gitai (2005), 577–78: for example, FtsZ is a bacterial homologue of eukaryotic tubulin, and
MreB is a homologue of eukaryotic actin.
188 Norris et al. (1996), 197, define the enzoskeleton as ‘the ensemble of proteins (principally
enzymes) that interact with one another and with membranes and nucleic acids to form extended
structures within the cell’.
189 This is possible because the cellular circuitry consists of proteins which – by means of processes
such as phosphorylation and methylation – can flip between two distinct states in accordance
with circumstance, functioning as molecular switches able to channel the cellular processes in one
direction or another depending on whether they are in state a1 or a2.
190 Bray (2009), 117, describes the chemotactic pathway, for example, as being ‘small and relatively
independent of other processes... but the operative word here is relatively. You just have to scratch
the surface to uncover a multitude of links to all kinds of other cell processes. The kinase that drives
the cascade of chemotactic signals, for example, also plays a part in other signaling pathways, such
as responses to osmotic changes and the detection of glucose. Just making the proteins of this
pathway and positioning them in the membrane and cytoplasm requires numerous other molecular
interactions to be performed’. Ultimately, Bray believes, it would be possible ‘to trace connections
between chemotaxis and every other function of the cell: no circuit is an island unto itself ’.
191 See ibid., 94 – 96; see also Grebe and Stock (1998).
192 Dusenbery (2009; 124 –27) draws an extremely useful distinction between ‘causal inputs’ and
‘informational inputs’: causal inputs, he writes, ‘are inherently important because of the chemical or
physical effects that they exert and the work that they can do. In contrast, informational inputs are
Endnotes 215
important only because they are associated with some causal input and can be used to predict the
occurrence of the causal input at another place or a later time’.
193 Ibid., 125 –26; see also Adler (1969).
194 Adler (1969), 1590 – 93.
195 See Thar and Fenchel (2001; 3299) on the difference between true taxis and chemokinesis.
196 Ibid.
197 Ibid.
198 See ibid.
199 See Dusenbery (2009; 214 –16) on the ‘fundamental physical tendency for free-swimming objects
to move in a helical path’. This can be considered the default option unless sufficiently prejudicial. As
Dusenbery writes, ‘even if the flagellum does not rotate, the body may be forced into rotation. If
a swimming microorganism generates its propulsive thrust along a line that does not pass exactly
through its center of frictional resistance, it will cause rotation of the body around a perpendicular
axis. … Since no organism (or machine) is constructed perfectly, some rotation around such an axis
is inevitable, unless some other force or feedback system is included to correct for deviations from a
particular orientation’ (214 –15).
200 Thar and Fenchel (2001), 3300.
201 In fact, there is such genetic and phenotypic diversity among the different strains of O. marina
that it has been suggested they may represent different species. For the sake of simplicity I shall refer
throughout to O. marina as a single species. See Calbet et al. (2013).
202 Roberts et al. (2011 a), 604.
203 Ibid.
204 See Guo et al. (2013), 39: roughly two per cent of O. marina populations are believed to fall back
on cannibalism in food-depleted environments.
205 Calbet et al. (2013; 68) note that it is generally found in intertidal pools, salt marshes and
embayments, rarely inhabiting open ocean waters.
206 See ibid., 73, 78.
207 Roberts et al. (2011a), 609, 604. See also Guo et al. (2013), 38 – 40. O. marina has further been
reported to possess a proton-pump-type rhodopsin, possibly acquired from bacteria by means of
horizontal gene transfer. This photosensitive receptor protein may well generate energy directly from
light.
208 Roberts et al. (2011 a), 609 –10.
209 See Breckels et al. (2011).
210 Again, this difference is anything but absolute: we have seen that bacteria too may be ‘misled’ by
a chemical gradient, even though the information in principle coincides with the nutrition.
211 Jonas’s description is couched in terms of animal existence, yet applies verbatim to active
protozoan hunters. ‘In the motions of animals’, he writes, ‘we have activity made possible by the
surplus from previous metabolism and directed toward safeguarding its future, but itself a free
216
expenditure dissociated from the continuing vegetative activity, and thus action in a radically new
sense. … The outward motion is an expenditure to be redeemed only by the eventual success. But this
success is not assured. The external action, in order to be a possibly successful one, must be such that
it also can go wrong’. By contrast with the sedentary life of plants, therefore, motile animals and (one
might add) active protozoan predators lead an existence that is precarious, exposed and committed
to ‘wakefulness and effort’. With this active consciousness of one’s surroundings there emerges the
possibility of pleasure and discomfort: ‘pursuit itself may end in the disappointment of failure. In
short, the indirectness of animal existence holds in its wakefulness the twin possibilities of enjoyment
and suffering, both wedded to effort’. See Jonas (1966; 2001), 104 –5.
212 See Boakes et al. (2011), 646 – 47.
213 This brief account is indebted in large measure to the description of how dinoflagellates use their
flagella to swim by Tom Fenchel (2001). Again, microbes are now known to swim in a helical path by
default ‘because at low Reynolds numbers almost any type of swimming will lead to a rotation as well
as a translation of the cell’ (330).
214 A more linear path is used when prey concentrations are high, whereas at reduced concentrations
the helical component is increased. It is thought that such ‘helical walks’ optimize random foraging
where prey densities are low. See Roberts et al. (2011 a), 608.
215 See Montagnes et al. (2008); Roberts et al. (2011 a), 610; Roberts et al. (2011 b), 834.
216 Breckels et al. (2011), 635.
217 Roberts et al. (2011b), 834.
218 On Chlamydomonas allensworthii see Starr et al. (1995); see also Dusenbery (2009), 305 –7, who
points out that the use of pheromones in mating also occurs in two species of ciliates (Blepharisma
intermedium and Euplotes octocarinatus) but that here the attraction is to groups of organisms rather
than to individuals.
219 See Ralt et al. (1994). In the plant kingdom, the occurrence of chemotaxis in ferns is well known.
Ancient lineages of gymnosperms (non-flowering seed plants) such as ginkgoes and cycads also
have sperm that are flagellated and therefore motile. Renouncing the self-propulsion of sperm, more
derived plants rely on the random transportation of pollen by wind and passing animals.
220 Buskey (1997), 77.
221 Ibid. See also Hansen and Calado (1999), 386, who note: ‘when in the presence of a large
organism, the movement extends from end to end of the prey, as if the dinoflagellate were sizing the
potential food item’.
222 See Fenchel (2001), 335, describing the behaviour in Crypthecodinium: ‘every time the dinoflagellates
happen to swim away from the particle, they tend to make a turn and so move back towards the particle.
The behaviour is similar to what has previously been described for certain phagotrophic dinoflagellates
that ingest large prey. … In these cases, it superficially appears as if the dinoflagellates gauge the size of
their prey before attempting to engulf it; in reality the motile chemosensory behaviour simply forces
them to encircle the prey cell until their ventral side makes physical contact to its surface by chance’.
Endnotes 217
223 Echolocation and electroreception may be assumed to perform a similar function in the
identification and localization of objects.
224 See the subchapter ‘A World of Objects’.
225 Gibbs and Dellinger (1908), 240.
226 Money (2014), 11: a certain anthropomorphism or rather zoomorphism (or perhaps simply
poetic licence) might be detected in the attribution of terror to a ‘writhing’ bacterium. See the
subsequent considerations on fear in unicellular organisms.
227 Ralston et al. (2014).
228 Gibbs and Dellinger (1908), 240.
229 See Kusch (1999), 715 –18, who describes the contrasting case of Lembadion – ‘a fascinating
ciliate with a huge oral cavity’ – which likewise produces a self-recognition signal to inhibit clonal
cannibalism, but limits synthesis of the signal in conditions of starvation (so cannibalism can go
ahead if needs must).
230 On the cytoskeleton see Pollard (2003); Fletcher and Mullins (2010).
231 For a critique of models of amoeboid motility based solely on dendritic actin polymerization at
the leading edge of the cell, see Kay et al. (2008), who suggest that other factors such as hydrostatic
pressure may also play a part. The precise mechanisms involved do not alter the present argument.
232 Harold (2001), 137. Assuming a length of 300 μm, a speed of 5 μm per second would mean that
the amoeba is crawling the distance of one body length per minute. In these relative terms, bacteria
are very much faster.
233 See Kay et al. (2008), 455, who describe the chemotaxis of crawling cells such as amoebae and
neutrophils as ‘an extremely slow form of movement’. These authors attribute top speeds of just 10-
40 µm per minute to neutrophils, which are amoeboid, phagocytic cells belonging to the immune
system: ‘a neutrophil that moves at 10 µm per minute covers its own body length in approximately 1
minute, which is equivalent to a human taking an hour to cover 100 metres’.
234 See McMahon and Bonner (1983), 200 –1, who point out that most ciliated organisms tend to
swim at the same speed on account of the consistency in the length of their cilia and beat frequency.
Flagellates too tend to have the same swimming speed, irrespective of their size.
235 Gibbs and Dellinger (1908), 233.
236 Ibid., 236 –37. For the Vorticella study see Hodge and Aikins (1895), who note that although the
cell never normally seems to rest, it does undergo periods of ‘enforced’ rest in the form of encystment
in unfavourable environmental conditions. It also, of necessity, interrupts its ciliary activity during
stalk contraction (an aversive reaction), although this does not last long enough to count as a ‘period
of rest’ (ibid., 529).
237 Gibbs and Dellinger (1908), 237.
238 See Brüssow (2007), 689 – 90, who points out that it is only the egg-laying females that need such
a nutritious diet; male mosquitoes, assigned the relatively low-energy business of sperm-production,
remain ‘lifelong vegetarians’.
218
239 Or at least Vorticella is seemingly indefatigable in its feeding behaviour; see note 236. Two further
provisos should be made. If suspension-feeding ciliates are subjected to centrifugation or transferred
to water at a different temperature or pH level, they may ‘lose their appetite’. It may be minutes or even
hours before they recover it, and for this time it is as though they are off their food. Fenchel (1980, 2)
observes that in such conditions the ciliates may ‘refuse to feed altogether’. The second qualification
is that, like many sessile protozoans, Vorticella has a motile life stage. Known as telotroch larvae or
swarmers, Vorticella daughter cells are concerned solely with finding a place to settle, and no feeding
occurs in this phase: nourishment only comes once they have completed their search and settled
down. So in this sense, too, there is indeed a contrast between feeding and non-feeding states. This
raises questions that will be addressed in the next section. On Vorticella see Patterson and Hedley
(1992; 1996), 113 –14.
240 As Jennings points out (1906; 11, 20), citing the work of the zoologist Ludwig Rhumbler,
amoebae can also be brought to interrupt their meal by subjection to strong light, which in general
has the effect of interfering with their activities and in this context makes them ‘lose their appetite’.
Mechanical stimuli such as the shaking of the substrate or contact with a sharp needle can likewise
cause amoebae to withdraw their pseudopods and remain immobile – and thus indifferent to the
presence of prey – for varying periods of time. In such cases too, the amoebae ‘lose their appetite’. See
Schönborn (1966), 29, who describes the effects of such disturbances on various testate amoebae such
as Difflugia lobostoma and Nebela gracilis. Whereas the former moves off more or less at once, the
latter takes 20 minutes to start moving again.
241 See notes 236 and 239.
242 Quoted in Bray (2009), 136. Darwin suggests that only people with asthma are truly aware
of the value of breathing. Mountaineering and diving are activities that foster such awareness. The
extremely rare condition known as Ondine’s Curse is a failure of the normally automatic regulation of
our respiration, resulting in apnoea usually but not always during sleep.
243 The species was classified in the genus Dileptus until 2012, when it was moved to its new genus.
See Wikipedia: https://en.wikipedia.org/wiki/Pseudomonilicaryon_anser. Both genera belong to the
family of dileptids.
244 Patterson and Hedley (1992; 1996), 168.
245 Miller (1968), 313.
246 Ibid., 315.
247 Of course, a systematic answer to this question would require an analysis of the dileptid’s hunting
strategy, in particular the degree to which its prey are ‘targeted’ as opposed to merely bumped into.
248 See Jakobsen and Strom (2004), 1918.
249 See ibid., 1915: according to Jakobsen and Strom, circadian rhythms are biological rhythms
that are characterized by 24-hour periodicity, endogenously generated (though environmentally
modulated), and can be entrained by naturally occurring environmental cycles that also show 24-
hour periodicity.
Endnotes 219
265 Miller (1968), 316. Of course, this too may have a mechanical rather than an internal biochemical
explanation: it could be that successful prey handling is in some way physically hampered by the
organism‘s state of physical repletion.
266 Matsuoka et al. (2000), 85. This contrasts with the cannibalism shown by various species of the
normally bacterivorous filter-feeding ciliate Blepharisma. In the latter case, cannibalism is associated
with gigantism, and its occurrence seems to depend not on starvation but simply on whether the
predator’s mouthpart is large enough to engulf its conspecifics; see Giese (1973), 123 – 34.
267 Guo et al. (2013), 39.
268 Denton (2005), 7.
269 Ibid., 8, citing A. D. Craig.
270 Ibid., 7.
271 Rolls (1999), 60 – 61.
272 Bennett and Hacker (2003), 199 – 200.
273 Ibid., 200.
274 As we have seen, Rolls defines what he refers to as ‘emotions’ – but what I would refer to as
‘appetites’ – in just such terms.
275 Craig (1918), 91– 92, 93.
276 Denton (2005), 127. The equivalent in a unicellular context is a protozoan displaying behaviour
associated with satiety as soon as its food vacuoles are full rather than waiting until its inner
biochemical balance is restored. Vacuolar distension thus functions as a signal for an imminent
return to homeostatic equilibrium. This signal may be misleading, as shown by bacterivorous ciliates
fed to satiation on latex beads, which behave in the same way as ciliates that have been sated on
nutritious bacteria. See Snyder (1991), 210.
277 The implication that reproduction is good for self may rankle to the extent that reproduction is often
considered to entail the death of the progenitor self and the birth of a new, different self. On the logical
implications of viewing self-reproduction as a form of self-perpetuation, see Glasgow (2017), 39 – 41.
278 Balcombe (2006), 9: Balcombe’s work provides a marvellous account of the presence of pleasure
in the animal kingdom.
279 Romanes and Darwin (1885), 110 –11.
280 See Cabanac (1992), 174.
281 The logical need for a ‘common currency’ in order to choose which of various behavioural
options is ‘the best’ (for oneself) and should thus be pursued was already recognized by Aristotle. See
De Anima (Book III.11) 434 a; also Aristotle (1986), 216.
282 Cabanac (1992), 174, 176.
283 The role of outcome expectations further implies a temporal dimension that is absent in reflex
behaviour.
284 Ibid., 175, 193: Cabanac points out that it is a matter of ‘rationality’ not in the philosophical but in
the economic sense: ‘Maximization of pleasure may thus be the link between physiology and behavior
Endnotes 221
and give the key to the problem of physiological optimization without the implication of the animal’s
knowledge and rationality about its physiological state. A working man does not have to know his body
temperature, blood oxygen and glucose, muscle glycogen and lactic acid, etc. to take an occasional but
necessary break. He just has to “listen to” his sensations and maximize the algebraic sum of pleasures’.
285 On the relationship between selfhood and selfishness see Glasgow (2017), 51–56, 101– 4.
286 Balcombe (2006; 37) quotes the veterinarian Jerrold Tannenbaum on the unrelenting harshness
of animal life.
287 See Balcombe (2006; 161) for the effects of catnip on felids: ‘Under its spell, cats chase and paw at
non-existent mice and phantom butterflies. The plant contains nepetalactone, a chemical compound
akin to a pheromone in the urine of sexually receptive cats. The solicitous rolling about of female cats
under catnip’s influence suggests that sexual arousal is a side effect. A cat who has discovered a catnip
plant will return to it daily. Ditto cougars, lions, jaguars and leopards’.
288 See ibid., 161–64.
289 It is plausible that the two most rudimentary forms of play – locomotor play and object play – in
fact have their evolutionary roots in the pleasures of hunting, searching, foraging and the pursuit of
prey or mates, activities that have come to be positively evaluated in themselves in the light of the
subsequent reward. In other words, it is not just the final consummation of an appetite that is positively
valenced, but the whole process of pursuit and capture. Such ‘play’ activities may thus seem to be
non-functional to the extent that they involve self-movement without the subsequent consummation.
However, their occurrence can be explained not only in terms of the proximate causation of sheer
pleasure – what the German language refers to as Funktionslust – but also the ultimate causation of
increased ‘fitness’ deriving from enhanced locomotive or manipulative skills. As suggested by the
autotelic nature of play, play exists in a deep relationship with selfhood, which is likewise autotelic by
definition. For an exhaustive account of the phylogeny of play see Burghardt (2005).
290 Jennings (1906), 330.
291 Berridge (2003), 108.
292 Ibid., 113-14.
293 See Balcombe (2006), 93; Berridge (2003), 109.
294 See Thorogood et al. (2018). Social transmission of aversions has been observed, for example, in
vervet and tamarin monkeys, house sparrows, domestic chicks and great tits.
295 Balcombe (2006), 102.
296 In fact, some protozoans do combine the characteristic features of amoebae and flagellates; see
note 24. The so-called pelobionts are amoeboid in shape but also sport a beating flagellum. Mastigella
and Mastigamoeba are two such genera. However, neither of these is known to ‘wag’ its flagellum
specifically in excited anticipation of pelobiont chow. On the pelobionts, see Patterson and Hedley
(1992; 1996), 56 – 57, 100.
297 If my intention in making these claims had been to explain motivation by reference to (implicitly
anticipated) pleasure, the argument would here be circular in that it affirms that pleasure is ultimately
222
recognized by the occurrence of motivated behaviour. However, the aim here is not explanation but
conceptual analysis. Appetite, motivation and pleasure are viewed as logically or conceptually linked.
298 See McFarland (2008), 118 – 29, comparing the ‘hedonic view’ of Cabanac with the ‘automaton
view’ of opponents, who conceive an organism as a type of self-regulating machine.
299 Anyone who believes that a feeling of pleasure necessarily incorporates a conscious thought
about this feeling of pleasure will be justified in refusing to ascribe such a state to unicellulars. Yet
even though humans are endowed with this capacity to reflect on their pleasures, it does not follow
that – in order to experience a pleasure – one needs to be able to reflect on or think about this
experience. It goes without saying that the specifically human capacity to contemplate, remember and
anticipate one’s pleasures adds extraordinary new dimensions to the experience of pleasure.
300 Of course, concepts such as ‘perspective’ or ‘point of view’ are misleadingly visual in their
implications; the ‘point of smell’ is more relevant for many animals.
301 Zahavi (2005), 11, quoting from Husserl, Zur Phänomenologie der Intersubjektivität II.
302 See Denton (2005), 124 –26.
303 As described above, corollary discharge gives rise to our normally unquestioned capacity to
distinguish self-caused sensory input from sensory input caused by non-self. Our proprioceptive and
kinaesthetic self-awareness is a pre-reflective sense of fluctuating bodily ‘hereness’ in animals with
individuated limbs that can shift in relation to one another. Other aspects of tacit selfhood include
equilibrioception (i.e. our sense of balance, mediated in most mammals by the vestibular system),
and what is known as self-body-size perception (our dispositional ability to cross gaps and negotiate
openings such as doors). To the extent that directional self-movement is what grounds the possibility
of consciousness, these factors – coordinating and synchronizing the self-movement of a highly
complex body – silently structure consciousness itself. Given the necessarily speculative nature of
attempts to ascribe these aspects of tacit selfhood to unicellular organisms, the present discussion
will leave these questions unexplored. Yet even though amoebae lack permanent limbs (perhaps
unlike ciliates, which have cilia) and a constant body size, their movements are still governed by an
overriding need for cohesiveness and coordination.
304 Shoemaker (1968).
305 Bennett and Hacker (2003), 97.
306 For a full discussion of traditional misconceptions regarding pain and its ‘privacy’, see ibid., 88 – 97.
307 There is logical redundancy, therefore, in the claim ‘I know that I am in pain’; it is enough to say
‘I am in pain’. By contrast, there is no logical redundancy in the assertion ‘I know that I am RDVG’
(I may suffer sporadic amnesia and need to fall back on empirical reassurance to convince either
myself or someone else who I am). This is because such an assertion pertains to my narrative or
autobiographical self.
308 Damasio (2000), 77, 78.
309 The human affliction known as congenital insensitivity to pain, or congenital analgesia, hints
that the connection between pain and consciousness is far from absolute. This genetic disorder,
Endnotes 223
which affects fewer than one in a million people, is not associated with any form of cognitive disorder
except an inability to feel pain, leaving sufferers with a diminished awareness of – and an extreme
vulnerability to sustaining – bodily damage to themselves. See for example Lane (2009), 250.
310 Smith (1991), 26. For a general discussion of invertebrate and insect pain see also Sherwin
(2001), Eisemann (1984).
311 Ammermann et al. (2012). It is also conjectured that the discharged extrusome might prevent
the cryptomonads from being ingested by a potential predator, entangling the feeding apparatus.
312 See Craig (1918), 91, who provides a lucid account of the asymmetry in the relationship of
appetite and aversion, the former caused by the absence of an ‘appeted’ stimulus, the latter caused by
the presence of a ‘disturbing’ stimulus.
313 See Eisemann (1984; 166) on the absence of pain guarding in invertebrates: ‘No example is
known to us of an insect showing protective behavior towards injured body parts, such as by limping
after leg injury or declining to feed or mate because of abdominal injuries. On the contrary, our
experience has been that insects will continue with normal activities even after severe injury or
removal of body parts. … Among our other observations are those on a locust which continued to
feed whilst itself being eaten by a mantis; aphids continuing to feed while being eaten by coccinellids;
a tsetse fly which flew in to feed although half dissected; caterpillars which continue to feed whilst
tachinid larvae bore into them’.
314 Sherwin (2001), S113.
315 On octopus pain see Godfrey-Smith (2017), 102, 224, describing recent work by Jean Alupay
and her colleagues.
316 See chapter VI.
317 Non-directional learned avoidance is presumably insufficient to suggest consciousness. This has
been found to occur in single isolated ganglions of decapitated cockroaches; see below note 473.
318 On the case of fish as the ancestral vertebrate, see Braithwaite (2010).
319 Damasio (2000; 74 –75) reports on the effects in one such case. After the operation the patient
was a new person, relaxed and happy. When asked about the pain, he replied that ‘the pains were
the same,’ but that he felt fine now. The operation had done nothing to alter the sensory patterns
corresponding to the local tissue dysfunction, notes Damasio, but it had abolished the emotional
reaction: the suffering had gone.
320 Ibid., 75 – 76.
321 See Gerber et al. (2014), especially 240.
322 Grandin and Johnson (2005), 189 – 93. This may sound rather a bold claim. What is meant is that
fear has a greater potential to incapacitate an animal: animals in considerable pain can still function,
acting as though nothing were the matter; animals in a state of panic cannot function at all (190).
323 Jonas (1966; 2001), 105.
324 As with consciousness in general, there has been a tendency to posit a ‘cut-off ’ point in the
phylogenetic ‘scale’ beneath which it is inappropriate to ascribe emotions to animals. It has been
224
argued, for example, that amphibians, unlike reptiles, lack emotions in that they do not exhibit
changes in heart rate or core body temperature in response to handling stress. The idea has a certain
appeal given the apparently expression-less inscrutability of fish and amphibians, deprived as they
are of our mammalian dexterity in pulling faces and gesticulating. Although it may seem plausible,
however, it can be queried from two angles. First, it is not obvious that there is a straightforward
correlation between such autonomic responses and ‘emotions’. Second, an increasingly substantial
body of evidence suggests that even fish are indeed highly susceptible to strain and stress: it has been
shown, for example, that it is 48 hours before their hormonal levels return to normal following rough
handling. See Balcombe (2006), 186-87; Braithwaite (2010).
325 This classification is indebted to Bennett and Hacker (2003), 199 – 202, who not only distinguish
emotions, agitations and moods, but describe how they may shade off into attitudes that are not
emotions (such as liking or disliking) and into character traits (such as benevolence or irascibility).
The boundaries between such categories, as the authors concede, are far from sharp.
326 Dennett (1991), 101.
327 For his analysis of Stimmung (albeit as a specifically human phenomenon), see Heidegger
(1929/30; 1983), 89 –103. Heidegger was already developing similar thoughts in his 1919 – 20 lectures
on The Fundamental Problems of Phenomenology, where he speaks of the Irgendwie (the somehow)
and the Wiegehalt (the how-content) of experience. The phenomenal world is encountered not just
as a manifold of objects, but always in some way or other, funnelled through my own particular
Selbstwelt (self-world). In this fascinating early work Heidegger also elaborates the concept of Sich-
Selbst-Haben (having-oneself) as a form of pre-reflective self-intimacy (Vertrautheit mit sich selbst).
See Heidegger (1919/20; 1993).
328 The bristletails, another class of primitively wingless hexapods, also resort to a directionless
escape jump. Other insects incorporate varying degrees of directedness into their jumps. Depending
on urgency, fruit flies choose between two escape strategies in the face of a ‘looming’ stimulus. Both
these strategies exhibit directionality, involving a sequence of at least four sub-behaviours: freezing,
adjusting directional posture, elevating the wings and taking off. The two strategies differ above all
in the time spent on wing elevation prior to take-off. The ‘short’ mode produces a less stable initial
flight, but is quicker than the ‘long’ mode, and tends to be elicited by looming that is more rapid. See
von Reyn et al. (2014).
329 The complexities are illustrated by the fast escape responses of fish and crayfish (which are
not fish, but freshwater crustaceans). As the neurobiologist Ari Berkowitz reports (2016; 36 –38),
considerable attention has been paid to a neuron called the Mauthner cell (or M cell) in fish and
to the lateral giant neuron in crayfish. These huge neurons, provided with thick axons adept in the
expeditious transmission of information, resemble ‘command neurons’ in that a single neuronal
spike is enough to trigger a fast escape movement, a speedily executed tail flip in the case of crayfish
or a ‘turn and swim’ response in fish. Such mechanisms are ‘like a push button’ (and presumably call
for little in the way of consciousness). However, there are other neuronal circuits that come into
Endnotes 225
play if the M cell is put out of action, for example, generating a slower, less stereotypical response.
These smaller neurons also contribute to the later stages of normal escape behaviour, ‘fine-tuning
its trajectory’. In crayfish, such circuits are used when the emergency is somewhat less pressing,
producing a sequence of flips more tailored to the specific situation: ‘a giant-neuron circuit produces
a very stereotypical flip, whereas the nongiant circuits produce a flexible set of slightly different
flips, through which movement direction can be better controlled. So if you have some time to
collect information and deliberate, you might be better off using a nongiant circuit to make sure
you’re going off in the best direction, rather than using the eject button’. What initially seems to be
an invariant response, therefore, proves on second examination to include a much more adaptable
mechanism that may well entail directionality and thus some sort of awareness of the surroundings.
It may also be modulated by factors such as ‘social status’ (43 – 47), moreover, as determined by the
winning or losing of fights against conspecifics. Fight winners have been found to be more likely to
employ the non-directional escape triggered by the lateral giant; fight losers tend to make use of the
adjustable form of getaway.
330 Romanes and Darwin (1885), 342.
331 Binet (1889), vi.
332 Binet was himself aware of his liability to the charge of anthropomorphism, as shown by his
discussion of unicellular choice (see ibid., 62). For his doubts regarding consciousness see ibid., 61.
333 Gibbs and Dellinger (1908), 240 – 41.
334 Bray (2009), 18; on attention in this sense see also Jennings (1906), 331.
335 Jennings (1904), 198.
336 Ibid. Such behaviour seems to be a more general strategy in weaker gradients, where ‘cells do not
orientate pseudopodia directly up the chemotactic gradient, but instead produce new pseudopodia
by splitting existing ones with little reference to the gradient. … Cells then steer by favouring the
daughter pseudopod that is up-gradient, which then becomes dominant, and the unsuccessful
daughter is retracted’. See Kay et al. (2008), 456.
337 Bray (2009), 18.
338 See, for example, Zahavi (2005), 62– 64.
339 As Alva Noë has shown, this phenomenal background is closely associated with various aspects
of our tacit self, namely our dispositional sensorimotor skills and our proprioceptive or kinaesthetic
self-awareness. My sense of the perceptual presence of the world in all its detail does not consist in
a ‘representation’ of the minutiae of the world that I encounter, but rather in a selective awareness of
certain salient features, together with my possible sensorimotor access to further details by controlled
self-movements (a turn of the head, a shift of my eyes, a step forward) and in conjunction with my
tacit kinaesthetic self-awareness and understanding of how these movements would produce changes
in sensory stimulation. See Noë (2004), 49 – 66.
340 See Bennett and Hacker (2003), 244: ‘Transitive consciousness is a matter of being conscious of
something or other, or of being conscious that something or other is thus or otherwise. Intransitive
226
361 But see Sonner (2008), 849 –50, who suggests that the capacity to modulate ion channels in
the unicellular micro-organisms that preceded the emergence of multicellularity may have originally
had a beneficial defensive function, preventing the entry of deleterious ions that might have induced
‘spurious motile responses’ in the cell. Sonner’s proposals seek to provide an evolutionary narrative
explaining the reaction of present-day organisms to inhaled anaesthetics. His conjecture is that
‘organisms today respond to inhaled anesthetics because their ion channels are sensitive to inhaled
anesthetics by virtue of common descent from ancestral, anesthetic-sensitive ion channels in one-
celled organisms (i.e., that the response to anesthetics did not arise as an adaptation of the nervous
system, but rather of ion channels that preceded the origin of multicellularity)’. Sonner’s membrane-
centred hypothesis thus accounts for the continuity between unicellular and multicellular anaesthesia.
362 See Hameroff (2006), 406.
363 See Kaech et al. (1999).
364 Ibid. As the authors point out, volatile anaesthetics seem not to exert a direct influence on actin
dynamics (for example by binding to a hydrophobic site on the actin molecule and interfering with its
assembly into filaments), but to have an indirect effect, possibly by interacting with a component of
one of the pathways that influence polymerization (10436).
365 See Damasio (2000), 357– 58 (endnotes): Damasio describes curare as mimicking locked-in
syndrome.
366 Dennett (1978), 209 –10.
367 Ibid.: as Dennett points out, ‘patients administered our compound, curare-cum-amnestic, will
not later embarrass their physicians with recountings of agony, and will in fact be unable to tell in
retrospect from their own experience that they were not administered a general anesthetic. Of course
during the operation they would know, but would be unable to tell us. At least most of our intuitions
tell us that curare-cum-amnestic would not be an acceptable substitute for general anesthesia, even
if it were cheaper and safer. But now how do we know that general anesthetics in use today are not
really curare-cum-amnestic?’ (210).
368 Applewhite and Gardner (1971), 287.
369 See above, page 67.
370 Of course, the work of phagocytosis is required in diffusion feeders, and some prey manipulation
may be needed.
371 Denton (2005), 121–23. Having failed to evolve such a strategy since moving onto land,
amphibians do not search for water even when severely dehydrated. It seems to have been the first
wholly terrestrial vertebrates (reptiles) that evolved an ‘appetite’ for water, i.e. a capacity to discern its
presence and a disposition to undertake motivated behaviour to find and drink it.
372 Jonas (1966; 2001), 116.
373 See Turner (2000), 43 – 45. Some thinkers suggest that the term ‘taxis’ is misleading to the
extent that the process is not a stimulus-response mechanism but a purely passive phenomenon.
See Dusenbery (2009), who notes that ‘even microbiologists do not call the movement a taxis when
228
bacteria sink under the force of gravity’ (383; see also 164 – 65). Dusenbery prefers to call such
bacteria ‘magnetic’ rather than ‘magnetotactic’.
374 For an illuminating account of magnetotaxis in a philosophical context, see O’Malley (2014), 25 – 41.
375 Dusenbery (2009), 164 – 65.
376 As Dusenbery explains, ‘the magnets reduce the movements of the bacteria from three dimensions
to one, and … reduction of dimensionality makes for more efficient guidance’. This permits magnetic
bacteria to extend the duration of time over which ‘they can maintain their orientation without
attaching to the surface of a larger object. This allows them to take as much time as desirable to obtain
directional information, without the limitation imposed by rotational Brownian motion. They can
then swim forward and backward along this line, in response to chemical stimulation’ (ibid., 166 – 67).
377 On this distinction see Turner (2000), 44. The distinction partly reflects the uncertainty of
human investigators as regards the benefits of magnetic orientation, which are not as obvious as
the benefits of, for example, phototactic orientation. It is perfectly feasible, indeed, that there is
an adaptive explanation for magnetosomes that has nothing to do with locomotion or motility. It
has been suggested that magnetosomes might serve as a repository for excess iron or play a role in
the detoxification of reactive oxygen species such as hydrogen peroxide. On the various possible
functions of the magnetosome, see O’Malley (2014), 31.
378 Of course, it is self-caused insofar as it depends on the work performed by the organism and thus
on the organism being alive and metabolically active rather than dead or in a state of cryptobiosis.
Assuming the aliveness of the creature, however, the response is basically fixed or unvarying.
379 By contrast, other ciliates are known to regulate their phototaxis in accordance with how well-
fed they are. When undernourished, Chlamydodon exhibits positive phototaxis; when satiated, it
shows negative phototaxis. Though generally avoiding light and preferring the darkness of the depths,
that is, hungry specimens may swim to the water surface in order to feed on the phototrophic prey
that frequently accumulate there. See Jékely (2009), 2799, 2802.
380 Villarreal (2009), 111 – 12.
381 See McDougall (1923), 60 – 61. A very similar principle – though dispensing even with
photoreception – underlay the original conception of phototaxis developed by Jennings’s
contemporary Jacques Loeb and others. This envisaged the stimulus acting directly on the locomotor
organs of the creature (e.g. the cilia of a paramecium), causing those on the more strongly stimulated
side to contract either to a greater or lesser extent than those on the opposite side. Jennings (1904)
showed that the explanation for the movements of ciliates such as paramecia was not as simple as
Loeb and colleagues had proposed. The phototactic paramecia and euglenas he observed moved by
a process more akin to ‘trial and error’, which involved turning and trying out a new direction until
one was found that worked.
382 See Jékely (2009), who pinpoints four necessary and thus universal features required of an
organism for three-dimensional phototaxis to be possible: ‘(i) polarity and a fixed shape; (ii) spiral
swimming with cilia; (iii) photosensory molecules and a phototransductory cascade that affects ciliary
Endnotes 229
beating; and (iv) a shading or refractive body that ensures the orientation-dependent illumination of
the photopigments during axial rotation’ (2802). As Jékely notes, the relatively frequent conjunction
of these features has resulted in the evolution of phototaxis eight times independently in eukaryotes.
Prokaryotes, by contrast, are not able to discriminate the direction of light. Instead, they measure the
intensity of light and use a biased random walk to negotiate a light-intensity gradient (2795).
383 See Thar and Kühl (2003), 5751. More specifically, the flagella at the end with a rising oxygen
concentration increase their rotation speed, whereas those at the end with a falling concentration
decrease their speed.
384 See ibid., 5752. The reasonably large size of the bacterium (6 µm) is likely to ensure, note the
authors, that ‘the flagellar bundles at either end of the cell are controlled only by the local sensor
regions’.
385 See Glasgow (2017), 96 – 98.
386 Tye (1997), 303.
387 Ibid.
388 In fact, the ‘satiety’ of caterpillars is more associated with the stages of their life-cycle than
with reversible, post-prandial ‘fullness’, recalling to mind some of the questions raised in the section
‘Recognizing Hunger: Some Doubts’. Caterpillars generally only cease to eat just before shedding their
skin or in the time prior to pupation.
389 See page 30.
390 For other examples of conflicting stimuli, see Jennings (1906), 92 – 99.
391 On interoception see Damasio (2000), 149 – 53; 150: ‘Under no normal condition is the brain
ever excused from receiving continuous reports on the internal milieu and visceral states, and under
most conditions, even when no active movement is being performed, the brain is also being informed
of the state of its musculoskeletal apparatus. … The internal milieu and visceral division is in charge
of sensing changes in the chemical environment of cells throughout the body. The term interoceptive
describes those sensing operations generically’.
392 Llinás (2001), 133 –34.
393 On cockroaches see Rose (1992), 172; on fruit flies see Greenspan (2007), 93.
394 Bray (2009), 239 – 40.
395 Bray (ibid.) proposes the centrosome as a possible candidate in animal cells, while acknowledging
the conjectural nature of his proposal. The centrosome, or microtubule organizing centre, he writes,
‘acts as a seed for microtubules that grow out to other parts of the cell, including its membrane, and
thereby influence both the shape of the cell and its movements’. It also plays a central role in mitosis.
See also Penrose (1995; 360 – 61) on the centrosome as the ‘focal point’ of the cytoskeleton; it is a
structure that ‘apparently controls the cell’s movements and its detailed organization’.
396 Penrose (1995), 358.
397 Hanahan and Weinberg (2000), 61; see Ishizaki et al. (1995), 1443.
398 Both cadherins and integrins are known to be altered in metastatic cells. No longer tethered to
230
the cells around them, metastatic cells thus reacquire the motility that enables them to spread through
the body. See Hanahan and Weinberg (2000), 65.
399 See King (2004), 314.
400 Basal metazoans such as sponges, by contrast, exhibit a more flexible relationship between the
individual self of their component cells and the collective self of the organism as a whole. See Epilogue.
401 See Penrose (1995), 371; also 406 –7. Penrose remains uncommitted on whether such ‘apparent
absurdities’ will one day come to be accepted. However, he regards the question as scientifically
legitimate, its answerability depending on improvements in our understanding of physical nature.
402 Llinás (2001), 218.
403 It is as though one were to deduce the wetness of water molecules from the wetness of water.
404 This is not to mention mobile germ cells and the occurrence of sperm chemotaxis. However,
spermatozoa can be deemed behaviourally inflexible enough to exclude consciousness, in that their
locomotive repertoire consists merely in reaching the oocyte. There is no question of whether or not
to stop off for a quick snack on the way.
405 Equally intriguing is the peregrination that even neurons and other bodily cells have to
undertake in the course of the development of the embryo. The migration of the embryo’s cells is
guided, in part at least, by a range of different chemical signals emanating from different tissues in
the developing organism. The eventual settling and maturation of a migrating cell is in turn induced
by proteins produced by tissues at its final destination. Many cancer cells revert to a migratory mode
during metastasis. Unlike animal cells, plant cells – held immobile by their rigid cellulose wall – do
not migrate at all. By the same token, plant cancers are not vulnerable to metastasis; nor do plants
have mobile phagocytes. For a fascinating account of embryonic cell migration see Davies (2014),
92 –105; on plants see Hallé (2002), 130.
406 We have also encountered caterpillars as ‘eating machines’, although the question of whether
they genuinely live up to their reputation for insatiability was left open. A further question mark was
left hanging over so-called trophonts, whose non-stop feeding is just a stage in an organism’s life cycle.
Even the ceaseless feeding of Vorticella is just part of its life cycle; see above, note 239.
407 According to the description by Sompayrac (1999, 2008; 16 –17), macrophages certainly seem to
have a fluctuating appetite: in their ‘resting’ state, they are mainly garbage collectors, responsible for
ingesting dead cells; when ‘primed’ or activated by cytokines such as interferon gamma, they upgrade
their activity levels; they may further be ‘hyperactivated’ by direct bacterial signals such as LPS or
mannose. In such circumstances, they grow larger, step up their rate of phagocytosis, and increase
their digestive capacity (i.e. the number of lysosomes they harbour). Opsonisation is also described as
boosting the ‘appetite’ of a macrophage. It is notable, however, that these changes in ‘appetite’ depend
on signals that are external to the macrophage itself, though internal to the superordinate system of
which it forms a part. Perhaps, after all, the macrophage simply does what the system dictates.
408 At the same time, any such residual selfhood must be tightly reined in; it must not be allowed to
result in an increased risk of cancer.
Endnotes 231
450 We have seen that – by contrast with the dinoflagellate Oxyrrhis marina – for E. coli the chemical
gradient is only ‘metaphorically’ distanced from the reward, insofar as the gradient from less to more
glucose or aspartate comprises both the nutrition and the information signalling the nutrition. This
may be taken to imply immediacy. Yet it is the information, not the nutrition, that spurs E. coli to
undertake its pursuit.
451 Jennings (1906), 19, citing observations by Rhumbler.
452 Hansen and Calado (1999), 386 – 87.
453 Gazzaniga (2008), 388.
454 See Baumeister et al. (1998): the paper, which is entitled ‘Ego depletion: is the active self a limited
resource?’, argues that ‘because much of self-regulation involves resisting temptation and hence
overriding motivated responses, this self-resource must be able to affect behavior in the same fashion
that motivation does. Motivations can be strong or weak, and stronger impulses are presumably more
difficult to restrain; therefore, the executive function of the self presumably also operates in a strong
or weak fashion, which implies that it has a dimension of strength. An exertion of this strength in self-
234
control draws on this strength and temporarily exhausts it’ (ibid., 1253). In one of the experiments,
people who had resisted the temptation to eat chocolates but instead made themselves eat radishes
were shown subsequently to be much quicker to give up in their attempts to solve a brain-teaser.
455 See also Kahneman (2011), 43.
456 See Gazzaniga (2008), 314.
457 See Emery and Clayton (2004), Emery and Clayton (2001).
458 Kabadayi and Osvath (2017).
459 Balcombe (2006), 223: if the hens in question took a food reward straightaway, no more was
given to them, but if they waited for 22 seconds they got the ‘jackpot’: they were found to hold out for
the jackpot 90 % of the time. See Abeyesinghe et al. (2005).
460 See Brock et al. (2011). On Dictyostelium discoideum in general, see Bonner (2009), also Glasgow
(2017), 244 – 47. See ibid., 256, for a brief account of their ‘agricultural’ activities.
461 Brock et al. (2011), 393.
462 See Denton (2005), 75 – 80.
463 Quoted in ibid., 49: Koch continues: ‘if the organism’s performance is only marginally affected
by the delay, then the input must have been stored in some sort of intermediate short-term buffer,
implying some measure of consciousness. If the subject can be successfully distracted during this
interval by a suitable salient stimulus (e.g. flashing lights) it would reinforce the conclusion that
attention was involved in actively maintaining information during the delay period’.
464 Neuser et al. (2008) have shown that Drosophila melanogaster indeed possesses what is termed a
‘spatial working memory’: during locomotion, in other words, flies can be distracted from a target for
several seconds and yet subsequently resume their former trajectory.
465 Crick and Koch (1990; 265, 269 – 70) stress the intimate connection between consciousness and
working memory and point out that no case has ever been reported of a person ‘who is conscious but
has lost all forms of short-term memory’ (270). However, human consciousness is not to be equated
with consciousness in its minimal or basalmost manifestation. If there is absolutely no memory,
of course, the pursuit of prey seems reduced to a ‘mindless’ ascent of a gradient, yet even here the
organism has to perceive this gradient and it has to be motivated to ascend it given that the reward is
not immediate.
466 According to the present argument, the concept of a philosophical zombie (a hypothetical being
that is physiologically and behaviourally indistinguishable from a human being but without any sort
of consciousness) is logically incoherent. Non-philosophical zombies of the sort that lumber through
horror films can perhaps be conceived as ‘eating machines’, which therefore know neither satiety nor
hunger. In fact, however, they may be ‘choosy’, preferring the taste of some anatomical parts over others.
467 See Hofstadter (1980), 360.
468 See note 151.
469 A third category is noise, although here the concept of ‘variability’ rather than ‘flexibility’ seems
more apt, since flexibility connotes variability that is adaptive as opposed to random. Variations in
Endnotes 235
responses to sensory stimuli may thus result from the noise that is inevitable in sensory systems
operating near detection thresholds, and noise may affect signal transmission at any point between
sensory input and motor output. See Globus et al. (2015), 215.
470 For Rolls (1999; 266 – 67), by contrast, all behaviour that does not involve operant learning is
mere taxis, with only ‘a fixed response available for the stimulus’.
471 As Dennett points out (1996; 88), such trial-and-error learning is just fine as long as an early
error does not lead to an early demise. More complex forms of mentality make it possible for the
trials to be carried out ‘in one’s head’, permitting – in Karl Popper’s words – ‘our hypotheses to die in
our stead’.
472 See Bermúdez (2000), 194 – 95.
473 Strictly, the foundation for associative learning is provided not by consciousness (which
presupposes directional self-movement) but by value. This is why even a single isolated insect
ganglion – as demonstrated in decapitated adult male cockroaches – is capable of avoidance learning.
Non-directional movement can thus be learnt without consciousness needing to be posited. On the
avoidance learning shown by an insect ganglion see Eisenstein and Cohen (1965).
474 Berger (1980).
475 Berger (1980; 401) himself notes the possibility of dietary imprinting persisting ‘through
successive asexual generations’.
476 See Jennings (1906), 174 –79, who concludes: ‘the same individual does not always behave in
the same way under the same external conditions, but the behavior depends upon the physiological
condition of the animal. The reaction to any given stimulus is modified by the past experience of the
animal, and the modifications are regulatory, not haphazard, in character. The phenomena are thus
similar to those shown in the “learning” of higher organisms, save that the modifications depend
upon less complex relations and last a shorter time’ (179).
477 See Applewhite (1979).
478 Applewhite and Gardner (1971), 285: the ciliate is found to habituate to a series of mechanical
shocks ‘so that it no longer contracts to the stimulus; after several minutes the organisms have
returned to their pre-stimulus level of sensitivity’.
479 Applewhite (1979), 347.
480 Armus et al. (2006).
481 McClintock (1993), 193.
482 For a discussion see Bennett and Hacker (2003), 339 – 41.
483 One possible distinction might be that concepts involve more abstract categories of entity than
do percepts. Honey bees, for example, have been shown to be able to differentiate not only patterns,
but also distinct classes of pattern, discriminating symmetrical from asymmetrical patterns or even
patterns that are ‘the same’ from ones that are ‘different’. Such bees might thus be said to have a
‘concept’ of symmetry or sameness. On the capacity of honeybees for abstract categorization see
Giurfa et al. (2001); see also Greenspan (2007), 132 –36.
236
484 This paragraph and the next are indebted to a reading of Carruthers (2009), who establishes
the following constraints on the possession of concepts: ‘in order to count as having concepts, a
creature needs to be capable of thinking. And that means, at least, that it must possess distinct belief
states and desire states, which interact with one another (and with perception) in the selection
and guidance of behavior. In addition, the belief states need to be structured out of component
parts (concepts) which can be recombined with others to figure in other such states with distinct
contents’ (90 – 91). Carruthers argues that many invertebrates, including honey bees, satisfy these
constraints, as suggested by their ‘flexible use of spatial information in the service of a multitude
of goals’.
485 See ibid., 97.
486 ‘Slime bacteria’ or myxobacteria are noted for their cooperative activity. This includes travelling in
‘swarms’ and forming ‘fruiting bodies’ when nutrients are scarce; on the model organism Myxococcus
xanthus see Strassmann (2000).
487 Much has recently been written on the theory of mind. On empathy, for example, see Thompson
(2007), 382 – 411; on shame see Rochat (2009).
488 For the question of whether the third form of intrinsic reflexivity required for full minimal
selfhood (i.e. self-reproduction) is also presupposed by consciousness see above, note 23.
489 See Glasgow (2017), 96 – 98.
490 Ibid., 93 – 94: in these terms, the intrinsic reflexivity of their ‘self-reproduction’ can be understood
to be indirect insofar as successful models get themselves reproduced (by the humans whose needs they
satisfy). The marketplace is the driver of ‘natural selection’.
491 See ibid., 94-96, 322. On small drones see Hambling (2015), esp. 126-37.
492 On Dictyostelium discoideum as a compromise between unicellular and multicellular forms of
self-containment, see Glasgow (2017), 244-247; in general see Bonner (2009).
493 On sponge self-containment see Glasgow (2017), 236, 265 – 66.
494 On the feeding biology of sponges see for example Brüssow (2007), 330 – 36.
495 Lee et al. (2012), 259.
496 On the carnivorous sponges see Vacelet and Boury-Esnault (1996), Vacelet (2006), Lee et al. (2012).
497 On sea lilies see Brüssow (2007), 381.
498 On salps see ibid., 428.
499 On lancelets see ibid., 363 – 64.
500 On basking sharks see ibid., 431.
501 On Trichoplax see Schierwater (2005), Schierwater et al. (2010), Wenderoth (1986), Smith et al.
(2015), Ueda et al. (1999) and Kuhl and Kuhl (1966). Despite such work, placozoans remain relatively
little studied by comparison with other animals. Future studies of Trichoplax may throw up surprises
yielding as yet unsuspected insights into the question of consciousness in simple animals.
502 Smith et al. (2015).
503 Wenderoth (1986).
Endnotes 237
504 Smith et al. (2015) suggest that such behavioural coordination might be attributed to either
of two mechanisms: the release of neuropeptides by secretory ‘gland cells’ also found in the ventral
epithelium or possible electrical signalling among fibre cells.
505 I could just as well have chosen to focus on the phylum Ctenophora, commonly known as the
comb jellies. This phylum is believed by some biologists to be phylogenetically even more ancestral
than the cnidarians. Yet it is a much smaller phylum, comprising just a few hundred species, and
so offers rather less in the way of behavioural variety. Ctenophores were traditionally regarded as
automatic feeders or ‘eating machines’ and thus not susceptible to states of hunger or satiety. In fact,
they are now known to exert a remarkable degree of control over their prey capture and ingestion,
adjusting the area of their food-collection surfaces in accordance with their nutritional state. Over a
period of six hours, starved ctenophores have been found to ingest about twice as much as previously
fed ctenophores when exposed to the same food density. According to Reeve et al. (1978; 744), ‘the
initial feeding rate of starved animals can be five times higher at first, decreasing to the pre-fed rate
gradually over six hours. This pattern is also affected by the duration of starvation’.
506 On the relationship between nematocyst discharge and prey capture and ingestion in sea
anemones see Thorington et al. (2010).
507 Ibid., 123.
508 Ibid., 130.
509 Glauber et al. (2010).
510 Pagán (2014), 127– 28.
511 Loomis (1955), 221.
512 Ibid., 219.
513 On the hydra see Buchsbaum et al. (1987).
514 On Aglantha digitale see Greenspan (2007), 41–58. Greenspan compares the slow, rhythmic
swimming behaviour characteristic of feeding with the much faster escape response triggered by
contact with a predator or a bite from a mackerel.
515 On the visual capacities of the cubozoan Tripedalia cystophora see for example Buskey (2002),
Garm et al. (2007) and Petie et al. (2011).
516 See Buskey (2002), 225 – 26, 230.
517 Petie et al. (2011).
518 See Buskey (2002), 230.
519 Andrew B. Barron and Colin Klein (2016; 4905) cite the decentralized organization of the
cubozoan behavioural control system as a key argument for refusing to grant them any capacity for
subjective experience. I am not convinced that the centralized organization of information-processing
is a significant factor. Information from both internal and external sources must be integrated, of
course, but it is not obvious why one cannot take the unitary self as a whole (i.e. the organism insofar
as it behaves coherently and in its own interests) to be the relevant unit of integration. Barron and
Klein dismiss box jellyfish as ‘nothing more than simple decentralized stimulus-response systems’
238
(4905), yet this is only the case if there is no interoception, i.e. no internalized distinction between
starvation and satiety. If their appetite is variable, however, the jellyfish will sometimes do one
thing (move towards a shaft of light) and sometimes another (say, remain stationary or drift about
at random). Barron and Klein also pinpoint a capacity for ‘representation’ in the sense of a spatial
simulation or model of the world as an ineluctable criterion for the possibility of consciousness,
which is why they rule out consciousness in the nematode Caenorhabditis elegans. Though a key
feature of human consciousness and crucial to our conception of our own consciousness, it is not self-
evident that ‘spatial simulation’ is a defining characteristic of consciousness as such.
520 Petie et al. (2011), 2809.
521 See Garm et al. (2012). Whereas T. cystophora is active during the day and inactive at night,
another species of box jellyfish, Copula sivickisi, shows the converse behaviour, resting during the day
and actively foraging and mating at night. This difference between the two species reflects the separate
ecological niches they occupy, each corresponding to the activity cycles of their respective prey. Other
species of box jellyfish are believed to be active both day and night. Significantly, the activity pattern of
T. cystophora seems to follow the light regime rather than obeying an endogenous circadian rhythm.
522 All worms are characterized by a body exhibiting bilateral symmetry with a head at one end,
yet the distinct groups of worms are only distantly related. Today the three main groups are the
segmented annelids, the flatworms or platyhelminthes, and the roundworms or nematodes. The
former two groups are more closely related to molluscs, whereas nematodes are positioned nearer to
arthropods on the ‘tree’ of life. Other phyla include Priapulida (penis worms), Xenacoelomorpha (see
following note) and Hemichordata, which comprise acorn worms and semi-sessile, tube-dwelling,
mainly colonial animals called pterobranchs.
523 The phylum Platyhelminthes was for a long time thought to include members of what is
now regarded as distinct phylum, Xenacoelomorpha, which comprises the classes Acoela and
Nemertodermatida and the species Xenoturbella. These simple animals lack a brain, a circulatory
system and a through-gut (i.e. an anus), and are fascinating in their own right. The question of just
how ‘primitive’ they are, phylogenetically speaking, is hotly contested. I shall leave them out of
account partly out of ignorance and partly to avoid making the final chapter longer than the rest of
the book together. For an overview see Achatz et al. (2013), Nakano (2015), Nakano et al. (2013) and
Cannon et al. (2016).
524 On planarian powers of regeneration see Pagán (2014), 127– 32.
525 Ibid., 102 –7.
526 Ibid., 160.
527 See Buttarelli et al. (2008; 400), who list six features that distinguish a brain from a ganglion, also
including the presence of a greater number of interneurons than primary motor or primary sensory
neurons and a predominance of multisynaptic over monosynaptic circuits.
528 Sarnat and Netsky (1985), 297.
529 See Inoue et al. (2015), 2.
Endnotes 239
549 See Arvanitis et al. (2013): for example, intestinal cells infected by Salmonella have been shown
to engage in apoptosis; C. elegans also has a p53-like tumour suppression protein that triggers cell
death in response to DNA damage.
550 For the sake of simplicity and brevity I shall here focus on solitary rather than social behaviour
in nematodes. Social feeding is thought to be a response to stressful conditions such as crowding or
a shortage of food. On social feeding in C. elegans see, for example, de Bono et al. (2002), Sokolowski
(2002), Artyukhin et al. (2015).
551 Avery and Shtonda (2003). Most filter feeders separate nutrients from medium by passing
the suspension though a mesh that traps the food particles. In the case of C. elegans, there is no
obvious filter. Instead, it seems to be by means of the differential timing of various pharyngeal muscle
contractions that the nematode succeeds in sweeping the food particles backwards into its intestine
while ejecting the liquid from its mouth.
552 Sawin et al. (2000). By contrast, C. elegans does not modify its velocity in response to gradients of
chemoattractants such as salt; see Iino and Yoshida (2009), 5373. Also noteworthy is that this slowing
response in the presence of bacteria is exhibited by solitary strains only. Social strains of C. elegans
do not decelerate in this way; see de Bono et al. (2002), 899: ‘natural isolates of the soil nematode
C. elegans feed on bacteria either alone or in groups. Solitary feeders such as the standard N2 strain
reduce locomotory activity and disperse on encountering bacterial food. By contrast, social feeders
such as strain CB4856 continue moving rapidly on food and aggregate together’.
553 Sawin et al. (2000).
554 See Hills et al. (2004), 1219.
555 This strategy is sometimes referred to as ‘area-restricted search’; see also Hills (2006). It is
thought to be optimally adaptive when resources are distributed in a ‘clumpy’ or ‘patchy’ as opposed
to a homogeneous way. Since ‘clumpiness’ tends to prevail in biological environments, the best place
to look for resources is in the close vicinity of where they have just been found.
556 Ibid.
557 Ibid., 9.
558 Of course, possible variations in the degree to which the randomness of the turns is constrained
or controlled in response to external factors may suggest varying gradations of directionality in the
animal’s movement.
559 Nuttley et al. (2002), 12449; see also Bargmann and Horvitz (1991), 729: ‘C. elegans can chemotax
toward the peak of a gradient of a number of small molecules, including positively charged ions such
as Na+ and K+, negatively charged ions such as Cl- and SO42-, basic pH, and several small organic
molecules, including cAMP, cGMP, lysine, cysteine, and histidine’.
560 See Lockery (2011).
561 On nematode chemotaxis see Ward (1973), 820.
562 Ibid., 821.
563 Pierce-Shimomura et al. (1999), 9568.
Endnotes 241
564 Ibid., 9561: a pirouette may refer either to a bout of two or more sharp turns in quick succession
or to a single sharp turn: it is defined as ‘a series of one or more sharp turns separating consecutive runs’.
565 Ibid., 9568. Iino and Yoshida (2009), by contrast, regard the pirouette strategy as a form of
‘biased random walk’. However, this klinokinetic strategy, they agree, is combined with the klinotactic
strategy of weathervane orientation. Computer simulations suggest that neither the pirouette
mechanism nor the weathervane mechanism alone achieves the efficiency attained by real worms,
but the two mechanisms together combine to produce effective chemotaxis.
566 Pierce-Shimomura et al. (1999), 9568.
567 Ibid.: ‘We cannot rule out the possibility … that a weak weather vane strategy operates in parallel
with the course-correction model, making runs better oriented than they might otherwise have been.
… We also cannot rule out the possibility that the weather vane strategy exists as a default mechanism
when the course-correction strategy is inactivated or when animals are in steeper gradients’. See also
Iino and Yoshida (2009); Lockery (2011).
568 See for example Globus et al. (2015), who trace some of the neural interactions by which
sensory information may be integrated within the underlying dynamics of the interneuronal network,
generating ‘different behaviors under different feeding states’ (223).
569 See Sawin et al. (2000), 627.
570 On the quiescence of nematodes, see You et al. (2008).
571 You et al. (2008; 249) define ‘food quality’ operationally in terms of an ability to support growth.
The E. coli strain HB101 is considered high-quality, whereas the strain DA837 is regarded as low in
quality.
572 Ibid., 250.
573 Ibid., 252: the kinase in question is the protein kinase G (PKG), EGL-4.
574 Ibid., 251.
575 See You et al. (2008), Ben Arous et al. (2009).
576 Ben Arous et al. (2009).
577 Ibid.
578 Gallagher et al. (2013), 9716.
579 See Trojanowski and Raizen (2016) on the arguments for referring to lethargus as ‘sleep’ rather
than merely ‘sleep-like’. In addition to the developmentally timed sleep of lethargus, C. elegans also
exhibits a non-circadian ‘stress-induced’ sleep in response to environmental trauma. This likewise
manifests itself as a cessation of feeding and locomotion and reduced responsiveness. Though
behaviourally similar, lethargus and stress-induced sleep are two physiologically distinct sleep states.
580 On lethargus see also Raizen et al. (2008), Iwanir et al. (2013).
581 Again, the criterion of ‘rapid reversibility’ is required to be able to differentiate the state from
pathological forms of quiescence.
582 See Trojanowski and Raizen (2016): the C. elegans gene lin-42 is a homologue of the gene that
regulates circadian rhythms in other animals such as mammals; the lin-42 gene product cycles its
242
expression in phase not with circadian time but with moulting time.
583 On the homeostatic self-regulation of sleep see Raizen et al. (2008), 570; Iwanir et al. (2013), 393.
584 See Nuttley et al. (2002).
585 Wen et al. (1997).
586 On aversive olfactory learning in C. elegans see Zhang et al. (2005).
587 Ibid., 179.
588 See Pujol et al. (2001) on how the Toll signalling pathway enables the nematode to recognize
pathogenic S. marcescens and then undertake the appropriate strategy of behavioural avoidance,
leaving the bacterial lawn. Such recognition may come about when a protein akin to the Toll-like
receptors in vertebrate innate immunity binds to the pathogen’s characteristic LPS structure or its
bacterial flagellin, the primary protein component of the flagellum (818).
589 Zhang et al. (2005), 179.
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*
Minimal Selfhood and the Origins of Consciousness
In Minimal Selfhood and the Origins of Consciousness,
R.D.V. Glasgow seeks to ground the logical roots of
consciousness in what he has previously called the
‘minimal self ’. The idea is that elementary forms
of consciousness are logically dependent not, as is
commonly assumed, on ownership of an anatomical
brain or nervous system, but on the intrinsic reflexi-
vity that defines minimal selfhood. The aim of the
book is to trace the logical pathway by which mini-
mal selfhood gives rise to the possible appearance of
consciousness. It is argued that in specific circum-
stances it thus makes sense to ascribe elementary R. D.V. Glasgow
consciousness to certain predatory single-celled or-
ganisms such as amoebae and dinoflagellates as well
as to some of the simpler animals. Such an argument
involves establishing exactly what those specific
circumstances are and determining how elementary
consciousness differs in nature and scope from its
more complex manifestations.
R.D.V. Glasgow