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Biological Journal of the Linnean Society, 2014, 112, 261–267.

Between holism and reductionism: a philosophical

primer on emergence
MASSIMO PIGLIUCCI*

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Philosophy Program, The Graduate Center, City University of New York, 365 Fifth Ave., New York,
NY 10016, USA

Received 14 December 2012; revised 17 January 2013; accepted for publication 17 January 2013

Ever since Darwin a great deal of the conceptual history of biology may be read as a struggle between two
philosophical positions: reductionism and holism. On the one hand, we have the reductionist claim that evolution
has to be understood in terms of changes at the fundamental causal level of the gene. As Richard Dawkins famously
put it, organisms are just ‘lumbering robots’ in the service of their genetic masters. On the other hand, there is a
long holistic tradition that focuses on the complexity of developmental systems, on the non-linearity of gene–
environment interactions, and on multi-level selective processes to argue that the full story of biology is a bit more
complicated than that. Reductionism can marshal on its behalf the spectacular successes of genetics and molecular
biology throughout the 20th and 21st centuries. Holism has built on the development of entirely new disciplines
and conceptual frameworks over the past few decades, including evo-devo and phenotypic plasticity. Yet, a number
of biologists are still actively looking for a way out of the reductionism–holism counterposition, often mentioning
the word ‘emergence’ as a way to deal with the conundrum. This paper briefly examines the philosophical history
of the concept of emergence, distinguishes between epistemic and ontological accounts of it, and comments on
conceptions of emergence that can actually be useful for practising evolutionary biologists. © 2013 The Linnean
Society of London, Biological Journal of the Linnean Society, 2014, 112, 261–267.

ADDITIONAL KEYWORDS: evo-devo – evolution – phenotypic plasticity – philosophy of science –

supervenience.

‘Emergence’ is a controversial concept with a convo- and similar ideas as obscurantist and leading only to
luted history, in both science and philosophy. It is never ending and fruitless debates – think, say, of
therefore not surprising that it has been misused and Richard Dawkins’ (1976) dismissal of anything smell-
vilified, as well as more often than not misunderstood. ing like group selection, or his famous quip that living
Typically, the idea of emergence is brought up by organisms are nothing but ‘lumbering robots’ control-
researchers who are – for one reason or another – led by their selfish genes.
unhappy with an ultra-reductionist scientific pro- It is undoubtedly the case that methodological
gramme, preferring instead some kind of holism or reductionism has an enviable track record both in
interactionism in the way they approach their biology specifically and in science more generally. By
research questions (think of the always current the onset of the 20th century chemistry had already
debates on gene–environment interactions: Lewontin, for all effective purposes been reduced to physics (Le
1974a). Just as surely, biologists who are embedded in Poidevin, 2005; although this is by no means an
reductionist programmes are skeptical of emergence accepted conclusion in philosophy of chemistry:
Hendry & Needham, 2007), and hopes were high
that biology would soon follow suit. Indeed, the
*E-mail: [email protected] demise of vitalism and the contemporary surge of

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262 M. PIGLIUCCI

model-based genetics at the beginning of the 20th One of the earliest articulations of the concept of
century seemed to nail the argument in favour of emergence in biology is due, perhaps surprisingly, to
reductionists. In 1931 J. B. S. Haldane could confi- John Stuart Mill (1843: III(6)1): ‘To whatever degree
dently declare that ‘Biologists have almost unani- we might imagine our knowledge of the properties of
mously abandoned vitalism as an acknowledged the several ingredients of a living body to be extended
belief ’, although he was at the same time uncon- and perfected, it is certain that no mere summing up
vinced by exclusively mechanistic explanations of of the separate actions of those elements will ever
biological phenomena: ‘We must find a different theo- amount to the action of the living body itself.’ The
retical basis of biology, based on the observation that idea being expressed here is familiar in both biology
all the phenomena concerned tend towards being so and statistics nowadays: we know of plenty of exam-

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coordinated that they express what is normal for an ples, from chemistry and biology, where the whole –
adult organism’ (cited in Bedau & Cleland, 2010: 95). as the saying goes – is more than (or, to be more
The Modern Synthesis (MS) of the 1930s and 1940s accurate, different from) the sum of its parts. Mill
was, in some important respects, another manifesta- himself contrasted an example from physics and one
tion of the same debate. On the one hand, population from chemistry to make his point. In Newtonian
geneticists like Dobzhansky and Fisher provided the physics (which was, of course, the dominant paradigm
experimental and theoretical bases for a reduction of at the time), mechanical forces can be combined in an
organismal biology to genetics; on the other hand, the additive fashion by means of vector analysis. So, for
infamous exclusion of developmental biology from the instance, the total force being applied to a given
MS, as well as Dobzhansky’s (1981) own research on object from two different sources is simply the sum of
reaction norms and gene–environment interactions in the individual forces thus applied. In chemistry,
Drosophila pseudoobscura pointed to the limitations however, the result of a chemical reaction is pretty
of straightforward reductionist approaches in biology. much never a simple sum of the reactants: we can
As is well known, a literature parallel to the MS thus combine, say, an acid and a base to obtain a salt (and
originated (e.g. with the early work of Schmalhausen, water). The properties of the salt are in no meaning-
1949, and Waddington, 1942) and eventually led – ful sense simply the sum of the properties of the
among other factors – to the onset of ‘evo-devo’ as a pre-reaction acid and base.
separate field of investigation (Love, 2009), as well as Arguably the next big step in the development of
to the resurgence and widespread appreciation of emergentism in philosophy of science was due to C. D.
research on phenotypic plasticity (Pigliucci, 2001). Broad (1925), who framed the issue in terms of the
More recently, the genomic revolution led initially independence (or lack thereof) of the ‘special’ sciences
to wildly optimistic claims of what would be learned (i.e. of anything that is not fundamental physics).
after the human genome had been sequenced This is very much still a topic of discussion today,
(Gannett, 2008). Yet, it soon became clear that having usually pitting physicists on the side of one or another
the sequence on a CD would not straightforwardly ‘theory of everything’ (Weinberg, 1992) and (some)
lead to the cure of diseases or to a deep understand- philosophers who defend the notion of a fundamental
ing of how to make a human being. Moreover, com- disunity of science (Dupré, 1995). Broad aligned
parative genomics has quickly led to a shift in focus himself with the disunity camp, on the basis that he
from what individual genes do or don’t do, to emer- thought there are some emergent ‘trans-ordinal’ laws
gent complexities and systemic properties of genetic that are compatible with, and yet irreducible to, more
networks (Vidal, Cusick & Barabási, 2011). fundamental principles of physics. Moreover, he did
Given this convoluted and controversial history, it not think these laws presented any particular reason
may pay to get a bit clearer about where the concept for the mystically oriented to rejoice: ‘There is
of emergence came from and how it evolved qua nothing, so far as I can see, mysterious or unscientific
concept (O’Connor, 2006). Perhaps the first thing to about a trans-ordinal law or about the notion of
appreciate is that emergentism is usually put forth ultimate characteristics of a given order. A trans-
as a third way between mechanism-reductionism ordinal law is as good a law as any other; and, once
and vitalism-holism. Emergentists – particularly in it has been discovered, it can be used like any other
biology – reject quasi-mystical appeals to vital forces, to suggest experiments, to make predictions, and to
but at the same time argue for the appearance of give us practical control over external objects’ (Broad,
genuinely novel phenomena at various levels of com- 1925: 79).
plexity of matter. Of course, much hinges on what It is worth sketching out the approach of two of the
‘novel’ means in this context, as well as on the idea major modern critics of the standard reductionist
that there are meaningful ‘levels’ of complexity in programme in science, a programme that seldom
nature. I will return to two (not mutually exclusive) appears to be questioned by many practising scien-
interpretations of both these ideas shortly. tists. John Dupré (1995) has advanced the idea that

© 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112, 261–267
 PRIMER ON EMERGENCE 263

the sciences are fundamentally dis-unified, which can think of emergent properties from either an onto-
means that any talk of a ‘theory of everything’ is logical or an epistemological stance, although the two
nonsense (and, in his view, even socially dangerous, are not mutually exclusive. Ontology, of course, has to
because it gives priority to a particular, mechanistic, do with the nature of things, while epistemology has
view of science and knowledge). Dupré’s argument to do with how we (think we) come to have knowledge
begins with the (uncontroversial, among philoso- of the nature of things. Ontological claims are inher-
phers) observation that science itself depends on ently metaphysical, while epistemological claims are
metaphysical assumptions that are not justifiable not (they can be purely pragmatic, or derived from
empirically, and proceeds to question three related principles of logic). To complicate things further,
theses underlying the project of unification: determin- several philosophers (though by no means all!) from

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ism, essentialism, and reductionism. The first and the mid-20th century on began to agree that meta-
last metaphysical positions are challenged by the idea physical statements ought to be evaluated in terms of
of emergence. This is not the place for an in-depth our epistemic access to the world, meaning that what
analysis of Dupré’s views (which do have their critics, we can know empirically should constrain how we
e.g. Mitchell, 2003), my point simply being to note think metaphysically (e.g. Ladyman & Ross, 2009).
that there is a vibrant ongoing discussion in philoso- In terms of emergence and reductionism, my sense
phy of science about issues that many scientists take of the literature is that most philosophers nowadays
for granted without further examination. are in agreement with most scientists: they reject
A similar example is provided by Nancy ontological emergence and accept ontological reduc-
Cartwright’s (1983) criticism of the unity project, this tionism. What this means is that the standard meta-
time hinging on her analysis of the concept of natural physical position is that there are no true emergent
laws (her positions too, of course, have been critically phenomena, only phenomena that cannot currently
evaluated, e.g. McArthur, 2006). Cartwright sees laws (or even ever) be described or understood in terms of
as empirical generalizations with more or less broad fundamental physics, and yet are, in fact, only
(but, crucially, not universal) domains of application. complex manifestations of the microscopic world as
Indeed, she maintains that the empirical evidence understood by fundamental physics. A simple way to
itself strongly suggests that laws cannot be both make sense of this idea is to deploy the concept of
universal and true: the only circumstances when we supervenience: in philosophy a property A is super-
can verify a law of nature (say, Newtonian mechanics) venient on another one, B, just in case A cannot
to a precise extent is when we create artificial worlds change unless there is a change in B. For instance, if
characterized by highly controlled conditions. In the the total amount of money in my pocket is $20, this
real world, by contrast, laws only apply given a more fact cannot change unless the number of coins and/or
or less large number of ceteris paribus conditions. And notes that make up that amount somehow diminishes
this, of course, is the case for physics, where usually or increases inside said pocket (as opposed to, say,
scientists take the very idea of a law of nature to be simply exchanging a dollar bill for four 25 cent coins).
uncontroversial. In biology there are still plenty of Analogously, higher-order phenomena in physics or
debates – among both philosophers and biologists – biology supervene on micro-physical phenomena just
about whether we can even sensibly talk about laws in case the only way to change the former is to change
(as opposed, again, to mere empirical generalizations) the latter (i.e. there are no genuinely emergent
in the first place (Mikkleson, 2003; Lange, 2005; phenomena).
Carroll, 2006; Elgin, 2006). While at first sight the I will not comment much further on the issue of
relationship between Cartwright’s views on natural ontological emergence versus reductionism because it
laws and emergence is a bit less obvious than in is of hardly any concern to the practising biologist. I
Dupré’s case, consider that one of the metaphysical will note, however, that the position I just described is
ideas that Cartwright needs to challenge surely is rather odd, because it actually contradicts the prima
reductionism, and that emergence arguably repre- facie empirical evidence: as Jerry Fodor (1974)
sents the major challenge for reductionist explana- famously put it, an ‘immortal economist’ would in
tions of the world. vain attempt to derive the principles of his discipline
from knowledge of fundamental physics. It simply
cannot be done. But if our epistemology tells us that
TWO TYPES OF EMERGENCE
the universe behaves as if it contained genuine emer-
Having briefly examined the conceptual history of gent properties (say, the properties of economic
emergence, it is time to make a crucial distinction systems, which do not seem to have much to do with
between two ways of thinking about it (or about the properties of quarks), then is it not the case that
reductionism, for that matter) that have been lurking rejection of ontological emergence is a flagrant viola-
unaddressed throughout the discussion so far. One tion of the principle that epistemology should inform

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264 M. PIGLIUCCI

metaphysics? All in all, I think the most reasonable actual complex genetic networks in living organisms.
course of action is actually to take a neutral, agnostic, Robustness measures the tendency of genetic net-
stance on the matter and to proceed to where we are works to withstand internal disruptions (e.g. muta-
going next: epistemological emergence. tions) while maintaining functionality, and is related
O’Connor (2006) helpfully describes two types of the to the broader concept of evolvability (Pigliucci, 2008).
latter, which he labels predictive and irreducible- While the mathematics of Romero & Zertuche
pattern. Predictive emergence is the idea that in (2007) is not for the faint of heart, their results are
practice it is not possible to predict the features of a relatively straightforward: robustness emerges from
complex system in terms of its constituent parts, even the statistical properties of a genotype–phenotype
if one were to know all the laws governing the behav- map modelled as an NK Kauffman-type network,

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ior of said parts. Irreducible-pattern emergentists although there are restrictions on the range of
maintain that the problem is conceptual in nature, values of both N and K that yield robust mapping.
i.e. that the lower-level laws simply do not have the Interestingly, these restrictions are within the
tools to deal with higher-level phenomena – as in the empirical NK ranges that are derived from studies of
already mentioned case of Fodor’s unfortunate econo- organisms as disparate as yeast and our own
mist with a misguided penchant for fundamental species. Here, then, emergence is the appearance of
physics. a biological property (robustness) as a result of a
As O’Connor himself acknowledges, the distinction particular type of non-linear interaction among
between predictive and irreducible-pattern views of lower-level entities (the genes in the network).
epistemic emergence is not sharp, but it does draw Clearly, Romero & Zertuche are not making any type
attention to the fact that emergent phenomena of ontological statement here (indeed, there is no
present both pragmatic and conceptual issues for the reason they should even be concerned with ontology),
practising scientist and aspiring reductionist. It is not but are rather deploying something like O’Connor’s
just, for instance, that it would be too computation- predictive concept of epistemic emergence. It is an
ally cumbersome to develop a quantum mechanical open question whether research such as that of
theory of economics (the predictive issue), it is that Romero & Zertuche may lead to the stronger claim
one would not know where to start with the task of of irreducible-pattern-type emergence. In this case,
deploying the tools of quantum mechanics (indeter- my hunch is that this may not be the case, as the
minacy principle, non-locality, etc.) to somehow levels of analysis – individual genes and gene net-
account for the phenomena studied by economists works – with which Romero & Zertuche (2007) are
(relation between supply and demand, boom-and-bust concerned are sufficiently close to each other to be
cycles, etc.). So, again, one does not need to be an described in terms of the same conceptual arsenal.
ontological emergentist to firmly reject a greedy But, again, this is an open question.
reductionist programme in biology or the social My second example is a classic one, in reference to
sciences. which ‘emergence’ has often been brought to bear – or
disparaged, depending on a researcher’s metaphysical
preferences: the study of gene–environment interac-
tions (Lewontin, 1974a; Pigliucci, 2001). The debate
TWO EXAMPLES OF EMERGENT
about nature and nurture has been going on at least
PROPERTIES: NK NETWORKS AND
since Plato’s idea of learning as recollection in the
G-BY-E INTERACTIONS
Phaedo, and later John Locke’s (opposite) contention
It will be instructive to anchor the somewhat esoteric that the human mind is a tabula rasa on which
discussion we have engaged in so far with a couple of experience writes out our character. In modern times,
examples from the actual biological literature, to similar discussions have pitted social scientists who
focus our ideas about what emergence may sensibly are inclined toward a Lockean position (think of B. F.
mean in the context of biological research. Skinner’s emphasis on operant conditioning) versus
The first such example comes from a paper by those more taken by a genetic perspective (as in The
Romero & Zertuche (2007) on NK networks, also Belle Curve book by Herrnstein & Murray, 1994) (see
known as Kauffman-type networks. Stuart Kauffman Keller, 2010, for an analysis of the conceptual confu-
(1969) proposed these models – which are a type of sions underlying the modern version of the debate).
cellular automaton – as an early attempt at exploring A better tool for thinking about gene–environment
the properties of genetic networks (characterized, spe- interactions has been available since the beginning
cifically, by N elements each with K input connections of the 20th century in the form of the idea of a
and one output). Romero & Zertuche were interested norm of reaction: a genotypic- and environment-
in the relationship between the properties of NK specific function that displays the range of pheno-
cellular automata and the robustness displayed by types produced by a given genotype within a given set

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 PRIMER ON EMERGENCE 265

of environments. As Lewontin (1974b) elegantly house-building metaphor, another one is to cash out
showed in reference to the specific case of the herit- on the promise of evo-devo in order to understand the
ability of human IQ, grasping the concept of a reac- emergence of phenotypes in biological organisms.
tion norm allows one to understand seemingly Regardless, the point remains that this – as well as
paradoxical ideas such as, for example, that a change the previous case of robustness – seems to represent
in environmental variance may affect estimates of a genuine case of emergence, at least at the epistemic
heritability (as it has been empirically demonstrated level (as I mentioned above, ontological emergence is
several times since: Pigliucci, 2001: ch. 4), even a metaphysical notion that is likely not to be settled
though the ‘genetic’ component of phenotypic variance empirically, and about which the best course of action
remains unaltered. is to maintain philosophical neutrality).

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But what does it mean to think of gene–
environment interactions as ‘emergent properties’?
WHY EMERGENCE?
The phrase can be given at least two distinct inter-
pretations, one statistical and pretty straightfor- A good number of scientists are understandably wary
ward, the other one a bit more vague but of the notion of emergence, for the simple reason that
particularly relevant to evolutionary developmental it sounds a bit too mystical and wool-eyed. Of course,
biology. Consider the statistical meaning first. In a if emergence turns out to be an ontological reality,
typical reaction norm diagram one can disentangle then these scientists would simply be mistaken and
the average effect of the environment on a given would have to accept a new metaphysics. However,
trait – measured by the mean slope of the measured even if emergence is only an epistemic phenomenon,
reaction norms – from the average effect of genotype, there are good reasons to take it seriously, for
measured by the mean height of the reaction norms instance because it points toward current methodo-
sampled. These are both additive effects, respectively logical or theoretical deficiencies that make straight-
quantifiable by the so-called Environmental and forward reductionist accounts unfeasible in practice,
Genetic (E, G) variances in a standard analysis of if not in principle.
variance. In many cases, however, the individual (i.e. Still, in order for more scientists to take emergence
genotype-specific, as opposed to average) reaction seriously we need a coherent account of why we see
norms are characterized by different heights in the emergent phenomena to begin with. One such account
Environment versus Phenotype space, and they also has been provided recently by Brian Johnson (2010),
have different slopes (indicating genetic variation for and it is worth considering briefly. I am not suggest-
phenotypic plasticity). This so-called G-by-E interac- ing that Johnson is necessarily correct, or that his
tion variance is the result of (statistically) non- explanation is the only one on the table. But it rep-
additive effects that cannot simply be reduced to a resents a good example of the contribution that phi-
sum of genetic and environmental effects. A popula- losophy of science (in this case, actually done by a
tion with a significant G-by-E variance, therefore, scientist) can give to the way in which scientists
exhibits a quantifiable ‘emergent’ (at the statistical, themselves think of a given issue. Besides, Johnson
population-level) property. may very well turn out to be exactly right.
There is a less straightforward, but more interest- Johnson’s basic idea is simple: (at least some kinds
ing, sense in which G-by-E represents a case of emer- of) emergent properties are the result of a large
gence in biology. As again Lewontin pointed out, if we number of interactions among parts of a complex
think in terms of genetic and environmental effects as system, all going on simultaneously in time and
distinct causes shaping phenotypes in a more or less space. In order to be able to grasp emergent outcomes
additive-linear fashion, we put ourselves in the naive our brains should be able to think in parallel at the
position of trying to understand how a house is built conscious level (parallel unconscious thinking does
by simply weighing the total amount of bricks and occur, but it leads to an ‘intuitive’, not rational, grasp
lime that goes into it. Clearly, the key to building the of phenomena). As the human brain is not capable of
house lies in the specific alternating pattern in which parallel conscious processing of information, we are
bricks and lime interact to yield the final construc- faced with the impossibility of reasoning our way
tion. Similarly, genes and environments continuously through the mechanics of emergence.
interact to build phenotypes throughout the process How do we know that the human brain cannot do
we call development. And this is a major reason why parallel processing consciously? There are several
one simply cannot understand evolution without reasons to think so, but Johnson provides a simple
development (and vice versa), an idea that has lurked little exercise in figure 1 of his paper (Johnson, 2010)
around for many decades before finally flourishing which is worth trying out to see how difficult that sort
into a distinct field of evo-devo studies (Love, 2009). of thinking actually is, and how unsuitable we are at
Of course, one thing is to appreciate Lewontin’s carrying it out. (The exercise involves summing up

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266 M. PIGLIUCCI

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