Original study • DOI: 10.

2478/biocosmos-2025-0001 • BIOCOSMOS • 1 • 2025 • 12-17

BioCosmos

WHAT IS LIFE? AN OPERATIONAL DEFINITION

David S. Oderberg*

Department of Philosophy, University of Reading, Reading RG6 6EL, UK

Abstract

One way of defining life is via a real definition, which gives the essence of life. Another approach is an operational definition,
which shows how living things can be tested or measured in a way that is distinctive of the biological. Although I give a real
definition elsewhere, in this paper I provide an operational definition, echoing Canguilhem’s dictum that life is what is capable of
making mistakes. Biological mistakes are central to the behaviour of organisms, their parts and sub-systems, and the collections
to which they belong. I provide an informal definition of a biological mistake. I contrast mistakes with mere failures and malfunc-
tions. Although closely related phenomena, each is distinct. After giving some brief examples of mistake-making and how it can
be tested, I reply to some objections to the very idea of a biological mistake.

Keywords
life • mistakes • normativity • reductionism

There are two ways to approach that most difficult and well-being of the organism – whether or not the behaviour
fundamental of questions: what is life? One is to give a real be characterised by self-consciousness, language, free will,
definition – a statement of the essence of life (assuming it responsibility, knowledge of purpose, among other properties
has one). A real definition tells you what an object is. It of particular kinds of living thing. Immanent causation is to
doesn’t tell you primarily (or even at all!) about the meanings be distinguished from ordinary, common-or-garden efficient
of words, facts about grammar, or information about what is causation, which is, to co-opt the old saying, just ‘one damn
in someone’s head. The real definition of gold, for instance, thing after another’. With immanent causation, a living being
is that it is a metal whose atomic number is 79. The real acts not just on itself and does not merely do things to itself,
definition of a fish is that it is a water-dwelling vertebrate with as when a volcano erupts or a wave crashes on the shore.
gills at maturity. These definitions give you the very essence A living being acts on itself, and does things to itself, for itself.
of the thing being defined – what it is that separates it from To act for itself means that it acts to keep itself in a good
every other kind of thing in the universe. state, functioning well, alive, healthy, integrated, and so on.
No inorganic being, no matter how large or complex, does
The other way of approaching the question ‘what is life?’ is this. To the objection, ‘What about computers that maintain
to give an operational definition: one that, speaking loosely, their operating systems in good health?’ and so on, the
makes life measurable and testable – that characterises it in reply is that these are artefacts – man-made objects with
such a way as to give the experimental biologist something no intrinsic teleology of their own. Their ends and objectives
work on. These are not mutually exclusive approaches, of are programmed into them, so what they appear to do for
course, and they also interpenetrate at the more abstract themselves is really done for us. If there were no ‘us’, there
level, but they are methodologically quite different. would be no self-maintenance by a computer or any artefact,
because there would not even be artefacts – only inorganic
In previous work I have articulated what I contend to be things with no intrinsic purposive behaviour of their own.
the real definition of life: a living being is that which has the Their derivative ‘life’ – what they do based on what we direct
power of immanent causation – causation that originates them to do – would vanish without their users and creators.
with an agent and terminates in that agent for the sake of
its self-perfection (1, 2). It is not merely teleology that There is much that can and has been said about such a
defines life, but the kind of purposive behaviour directed at the definition, but here I focus on the fact that it is not immediately

*
Corresponding author e-mail: [email protected]
© 2025 Oderberg, This is an open access article distributed under the Creative Commons Attribution-NonCommercial-NoDerivs license
(http://creativecommons.org/licenses/by-nc-nd/3.0/).

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 Oderberg: What is life? an operational definition

obvious how to ‘cash out’ such a definition in experimental Mistake-making is then, as it were, a concrete entry on the
terms. What kinds of novel, testable hypotheses might be debit side of life’s accounting. It comes with the territory if
generated by such a high-level metaphysical definition? We something is alive. It indicates the fundamental normativity
can, however, operationalise this way of understanding life of life. Perhaps we could even give a definition of life halfway
by developing an insightful remark by the French philosopher between the highly abstract and the highly concrete: life is
and physician Georges Canguilhem (3): ‘life is what is capable that which is subject to norms. Note that we are, all the while,
of error’. Living things make mistakes; they get things wrong. talking about material life – the life of bodily substances,
A dog may forget where he buried his bone. A frog may be a whether or not – as in the human case, I believe – there is an
millisecond too late darting its tongue at an insect. A fish takes immaterial element as well. On an extended view, we could
the bait. Broody domestic hens are notorious for trying to hatch even say that purely spiritual beings – angels, for example –
golf balls and other vaguely egg-resembling objects. Animals are subject to norms of behaviour and are potential mistake-
are fooled by camouflage, traps, prey that hide. The living makers. God, by contrast, although purely spiritual is subject
world is full of tricks – deception, subterfuge, misjudgment, to no norms, does not act to perfect Himself since already
mistiming, miscommunication. It is astonishing how much an perfect, and cannot err.
organism – from the very large to the microscopic – has to get
right, how many mistakes it must avoid – just to approach the Normativity has very little place in current philosophy of biology.
end of its natural lifespan and propagate its own kind. That It is often equated, wrongly, with ‘values’, ‘oughts’, ‘shoulds’,
the environment is inherently dangerous and often unstable ‘prescriptions’, ‘commands’ (7). It is thought to involve the
– certainly quite unpredictable – renders mistake-making a ‘spooky metaphysics’ of teleology (8). It is considered to be
certainty and its avoidance a necessity. a matter of perspective, or context, or theoretical interest (9).
It is hard not to detect a certain cant (certainly not Kant!) in
Not only whole organisms, but also collectives of organisms such pronouncements, as though it were philosophically
and even parts of organisms make mistakes. Flocks of birds unproblematic literally to collapse the distinction between
fly into skyscrapers. Antibodies are fooled by pathogens – the health and disease, the normal and the pathological, welfare
classic example is antigenic mimicry, which is thought to be and what might be called ‘ilfare’. If anything in the natural world
at the root of many autoimmune diseases; it is not only the has a claim to intuitive obviousness, it is that there are ways in
self antigens that fool the antibodies in many cases but the which things go well or badly for an organism. We know it with
non-self antigens – such as viruses – themselves (4, 5). Blood certainty in our own case, yet why should our case be special?
platelets – essential for the formation of clots – are activated by
exposure to collagen released by endothelial injury. However, The point can be brought out vividly by the simple observation
in an individual with a certain genetic abnormality, they will that electrons do not make mistakes, neither do protons,
not respond correctly – as they are supposed to – and the or carbon atoms, or lumps of gold or grains of sand, or
individual will suffer potentially fatal blood loss (6). continents or rivers. The only mistakes in physics are those
made by physicists, found in textbooks or research articles.
It is arguably the case that if not actual mistake-making There are mistaken theories or hypotheses. Instruments may
then at least the potential for making mistakes is universal give erroneous readings. But the proper objects of physics
across biological systems. The connection with the real do not makes mistakes. No proton ever went the wrong way
definition of life offered above is clear enough: if it is definitive in a particle accelerator, except relative to some theoretical
of organisms (by which I mean, for convenience, whole perspective or expectation. By clear contrast, however, mice
organisms, pluralities, and also parts and sub-systems of do make mistakes – it’s the second mouse that gets the
organisms) that they act for their well-being, then we should cheese, so the proverb goes – and so do ants, grasshoppers,
expect them not always to get things right. Given the already- antelope, and we humans.
noted instability of nature – both external to the organism and
in its own internal constitution – we should expect mistakes to How should we define biological mistakes? Leaving aside
be found everywhere and at all times. Perhaps we can at least technical details (10), a biological mistake is a kind of
conceive of an organism in a hyper-stable environment, with behaviour that departs from a standard of correctness for
a simple life cycle, constitutively as simple as biology allows, the organism in its environment. The way it so departs is by
and thereby incapable of making a mistake – at least in its threatening the organism’s ability to ‘get on’, or ‘act effectively’,
actual world. Such an organism is likely not a denizen of our in its environment. There is no mathematical precision when
world, and if it were it would not take much imagination to evaluating what counts as a threat. We know that if a deer
posit a close possible world in which it could and did make gets caught in the headlights, then if the developing causal
mistakes. connection between the vehicle and the deer is not broken

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– say, by the deer’s coming to its senses and bounding over and biases is premised on such a fact about us humans (12).
the fence, or by the car’s braking hard – then the deer will Experimentally speaking, it would be a fascinating task to
end up injured or dead. We could say the same if it were map the limits of unavoidability in a given species – just one
fifty metres from the vehicle yet heading inexorably in that aspect of treating mistake theory as a way of operationalising
direction. There is no way to set a cut-off point beyond which organic teleology.
a connection is too remote to count, at least narrowly and
relatively insignificantly, as a mistake. More work should be The theory of biological mistakes is not supposed to supplant
done in this area, but for now I will say only that what this or compete with any other productive biological frameworks
teaches us is that mistakes shade into general risk. Life is full we already have. Rather, it is a different way of looking at
of risk: you risk your life, to some degree, whenever you get life with a view to developing novel and testable hypotheses.
out of bed in the morning. But it is not a mistake simply to live. Biologists investigate mistakes all the time: indeed the
It is, however, a mistake to play on your mobile phone while language of biological mistakes is far less problematic for
crossing a busy road. biologists, in my experience, than it is for philosophers who
are used to thinking of mistakes from the inside, as a human-
Mistakes are not the only marker of biological normativity. centred phenomenon. What mistake theory does is seek to
There are also malfunctions and what I call ‘mere failures’. organise the various concepts and phenomena involved in
Again, the ‘functions debate’ in philosophy of biology is a thinking about and observing mistakes, with a view to more
minefield of its own (11). For present purposes, a malfunction systematic investigation.
should be thought of as any systemic breakdown in an
organism that threatens welfare – sickness being the most Along with testing for unavoidable versus avoidable mistakes,
obvious. Sickness is a departure from correctness – being another avenue for research is testing for mistake prevention,
healthy. But it need not involve a mistake: being invaded by a minimisation, and correction mechanisms. For a given kind
parasite is not of itself a mistake, but drinking parasite-infested K, what capacities do Ks have for keeping themselves out
water is. Again, a mere failure also departs from correctness: of the way of certain kinds of mistake-making, or for keeping
it is what simply happens to an organism, not something mistakes to a minimum, or correcting mistakes that have been
the organism does to threaten its well-being. Being hit by made? Correction could involve getting back onto the right
lightning, or by a car, or rained on by radioactive particles, path there and then, or learning from the mistake in such a
are mere failures that need not involve doing anything wrong. way as to avoid the same one in future. It is highly likely that if
They might – say if you went outside during a thunderstorm Ks are mistake-prone but unable to prevent, avoid, correct or
against all advice – but mistake-making is not entailed either minimise those mistakes (at least some combination thereof),
by malfunctions or mere failures. they are already extinct or on the way to becoming so. Again,
one should expect a priori that no such kind does exist or ever
Interestingly, the converse is also true: mistake-making does has existed – otherwise how could it even have managed
not entail either malfunction or mere failure. The latter should more than a few seconds of existence in the first place?
be evident, but even the former is true. Our initial intuition
might be that making a mistake must involve a malfunction. As I have emphasised, mistake theory should enable the
But consider that a hen trying to hatch a golf ball, or a fish generation of novel, testable hypotheses. For example,
taking bait, are not malfunctioning (in the usual run of things): dopamine neurons have been proposed to perform an
they can be in the best shape possible for their kind, and yet evaluative function with respect to birdsong – in particular,
still get things wrong due to their intrinsic limitations. A hen that of zebra finches (13, 14). Dopamine spikes correlate
does not have the discriminatory capacity to distinguish an significantly with fluctuations in a combination of measures
egg from something looking quite a bit like one, nor a fish the such as song pitch, frequency, and Wiener entropy (a measure
ability to distinguish bait from real food. That’s just how it is of where the sound stands between a pure tone and white
with these kinds of organism. It would be a waste of a poultry noise), such that higher spikes are associated with closeness
farmer’s time to try to teach hens not to sit on golf balls; in fact of the song to the one it learned from its father and lower spikes
this mistake is exploited by poultry farmers to manage broody with greater fluctuations away from that template. Dopamine
hens. Most fish cannot be trained to avoid bait either, much seems to act like a mistake detection and correction system,
to the relief of fishermen the world over. These latter kinds of maintaining the song’s relative fidelity to what the finch
mistake I term unavoidable. They are an inevitable effect of learned. From this we can hypothesise that not only is the bird
the natural constitution of certain kinds of organism. It is likely, itself capable of mistake-making, but the dopamine neurons
a priori, that every species is liable to unavoidable mistakes, themselves might be capable of performing their evaluative
given their finiteness; much of the research into heuristics function mistakenly. This latter possibility will need further

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 Oderberg: What is life? an operational definition

investigation: does the dompaninergic system monitor or There are of course various objections that can be
regulate the song evaluation function of dopamine neurons, levelled at the very idea of a biological mistake (10).
and if so how? Further, how is the standard of correctness for I want to focus on two here. One is that since mistakes
song production represented by the zebra finch, especially presume the reality of biological normativity, and
since each one has a unique correct song that it is taught? biological normativity is reducible to physics and
Given that birdsong is a form of communication, mistake chemistry, there are no real biological mistakes. I have
theory focuses on whether and how a bird may get it wrong already pointed out that denying the reality of normativity
in, for instance, attracting a mate through song. Interestingly, (I will omit ‘biological’ from now on for convenience) means
the dopaminergic error signalling that operates when the bird denying evident distinctions between health and disease,
practises alone is turned off in the presence of a potential welfare and ilfare, and related phenomena. We can, however,
mate and retuned to feedback from the other bird (15). It say more (16). Consider a Laplacean all-knowing demon
would be informative to investigate whether this retuning was who comprehended all the physical and chemical facts in
itself mistake-prone. a given situation where one might postulate a biological
mistake. Suppose there to be three physico-chemical
Since relative fidelity is at stake, not mathematically exact pathways, as we might call them, traced by an organism
correspondence, we might hypothesise that the bird, via its – itself (ex hypothesi) nothing but a skinful of physics and
dopaminergic sub-system, evaluates the correctness of the chemistry, to put it metaphorically. The organism follows
song as correct rather than as containing a specific number of path A, which the mistake theorist points out, in agreement
pitches in a certain order, for example. But, one might object, with simple observation, is a mistake – say, mislocating
how could a bird evaluate something as correct without the a prey. It then backtracks and follows path B, which the
concept of correctness? In reply, we need to know what it mistake theorist, again conforming to observation, identifies
means for an organism to possess a concept: perhaps there as wandering around doing nothing in particular – neither
are kinds of practical mastery, for example, which constitute mistaken nor correct relative to the objective of finding prey.
a kind of concept possession but without the abilities It then backtracks and follows path C, which is identified
associated with reason, language, and self-awareness. If the as finding prey – and catching and eating it. From the
bird is able to keep its song faithful to the original, maybe normative perspective, these pathways are quite different
that is enough for us to assign to it a kind of mastery of the in their meaning – not just for us as observers, but for the
concept of a correct birdsong, for limited biological purposes. organism itself. One is right, one is wrong, one is neutral.
Still, we do not have to go even that far. It is doubtful a bird One promotes survival, one threatens it (a small amount,
has the concept of food any more than that it has the concept by delaying the hungry organism’s getting its lunch), and
of correctness. And yet birds are good at finding food and one is ‘neither here nor there’ – normatively indifferent, as
avoiding eating non-food; similarly, perhaps, for song: the we might say.
zebra finch can learn its song, can sing, and can keep its song
correct – for the most part. That requires an ability to assess The problem for the Laplacean demon – who is also a
the song as right or wrong, albeit instinctively. Perhaps the reductionist about normativity – is how he can make this
bird has some kind of primitive aesthetic standard; in other distinction. He has all the physico-chemical facts at his
words, the ability to tell whether the song sounds pleasant disposal, yet physico-chemically speaking pathways A, B,
(for mate-finding) or good enough (for an alarm call), and so and C are all on a par. None of them have any properties
on. One could test such a hypothesis, crudely speaking, by that mark them out as mistaken, correct, or indifferent. They
investigating whether the bird is counting or measuring units are, as we can call them, mere physico-chemical variations.
of song as opposed to evaluating the song as a whole pattern. It is, in principle, impossible for the demon to read off the
(Needless to say, such testing is extremely difficult to do, but normative properties from the physico-chemical ones. In
progress is being made all the time). If it can be established which case he is faced with a choice. He can deny normativity
that it evaluates a song for rightness or wrongness, then it altogether, in eliminativist fashion: there really is no distinction
is likely capable of making a bad judgment, e.g. evaluating between sickness and health, doing well in life and doing
a song as pleasant enough to attract a mate when in fact badly, flourishing and suffering, and so on. This, as suggested
it departs too far from the template to facilitate reproductive earlier, seems wholly implausible on its face. Or else, the
success. Evidence for a mistaken judgment would be found demon must concede that since normativity cannot be read off
in whether the song did successfully attract a mate; if not, the physics and chemistry on their own, it must be an addition
we could investigate whether and how quickly – perhaps too of being (to use a term from David Armstrong) – even if the
quickly for any calculations to be made – the mistake-prone normativity supervenes on the physical and the chemical.
bird was able to improve its song the next time around. (Or maybe it is emergent in some sense we cannot

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explore here.) This is a familiar non-reductionist line of obtaining, where the organism is directly or indirectly affected
argument but no less potent for that. by other things and/or capable of directly or indirectly affecting
other things. A sceptic might say: ‘All organisms carry on their
A second objection concerns the relation of mistakes to the life cycles in the universe as a whole, so this is the only real
environment. On mistake theory there is no such thing as environment in which mistakes can be made, especially given
a pure or absolute mistake. The job of every organism is to the “butterfly effect” and the causal connectedness of all things.’
get on well in its environment – to ‘make a living’, as Mark It is hard to know what to say that would convince the sceptic
Okrent (17) puts it. This applies to humans as well as to otherwise, nor is it clear that it is worth saying anything to such a
other organisms. To drink boiling water is a mistake – even sceptic. Given that mistake theory is about operationalising more
if, suppose fancifully, your stomach miraculously shrugs off abstract ideas about purpose and normativity, it is good enough
the assault and you are hailed by the crowds as a superman. to defer to biologists: they do not care about the boundaries of
We need to qualify, however. Assuming you had no idea this the universe when assessing whether an organism is doing well
would happen on drinking the boiling water, what you did or not. Nor do they consider the earth as a whole, or even the
was a mistake relative to the environment you were in – your vast bulk of it. They are interested only in whether an organism –
ordinary daily surroundings, let us suppose. You got off lucky, and its conspecifics, within a broader evolutionary context – is
to be sure, but you still acted against the norms of health and adapted to its niche, which is the specialised, relatively narrow
survival. But if you knew you had this miraculously strong spatio-temporal region in which its lives, maintains itself, and
stomach, then if, suppose further, you were demonstrating reproduces its kind. If that is good enough for biologists, it is
your prowess to an audience at a talent show, you would not good enough for mistake theorists.
have made a mistake in that environment. Note that we are
not considering morality or responsibility, merely what does The more stable a region is, the less mistake-making is possible.
or does not serve the person’s welfare. We could substitute With change comes greater potential to get things wrong. That
a bird and a mating display to make the same point. A bird is why organisms generally prefer stability and predictability, at
that shows off colourful plumage – with no mate in sight but a least when it comes to staying alive and healthy. The mistake
predator standing by and alerted to the bird’s presence – has theorist, from the experimental perspective, can choose
made a mistake, whether it knows there is a predator and no how broad a region to consider when evaluating behaviour
mate or not. Suppose – again somewhat fancifully – that the for actual or potential mistake-making. A mistake in one
bird is attacked mid-display by the predator but fights it off, environment may lead to success in a larger environment –
emerging battered but in generally good shape. A potential thus making it interesting to observe an organism’s strategies
mate then comes along, picks up cues as to the first bird’s over time for coping with different challenges coming from
health and fitness, and happy reproductive success ensues. novel causal interactions. There is a difference between,
Again, we should say that the bird made a mistake relative to say, a fish that accidentally swims upstream due to a sudden
the environment in which a predator was standing by, but not change in the current, thereby exhausting itself and dying of
relative to the predator-free environment in which a mate was fatigue, and a salmon swimming upstream so as to reproduce.
standing by. These are just facts about what the bird has done The first makes a mistake, the second doesn’t, and we make
– normatively laden facts about behaviour that does or does the distinction in terms of the different environments. They
not threaten effective action in a given environment. are two radically different, albeit superficially similar, kinds
of behaviour. But what if the salmon dies of exhaustion itself
We can, of course, add further facts into the mix in order to before managing to reproduce? The evaluation will depend on
make subtler evaluations; this is just what mistake theory is the detail. For example, if it was overwhelmed by a current
designed to encourage. If the bird knew it could fight off the stronger than it was capable of overcoming, it would have
nearby predator, exploiting its success in order to attract a been subject to a mere failure (see above). If it was diseased
mate, then we could plausibly assess the situation as one in and never capable of achieving its objective, it would have
which no mistake was made. We should say that there was been subject to a malfunction (again see above).
a single environment, containing both predator and potential
mate, in which the bird implemented a strategy for mate I submit that although there are genuinely important questions
attraction, with ensuing success. In other words, what counts about the relation of action to the environment, and hence
as an environment for mistake-theoretic purposes can involve about the individuation of mistakes, we should put aside more
what an organism knows, what it plans, and what its capacities factitious concerns about whether mistakes are possible at all
are. An environment is just a spatio-temporal region in which given the infinite divisibility of spatio-temporal regions, or the
an organism carries on its life cycle. The region is defined causal connectedness of the entire universe, or the potentially
at least partly by the biologically relevant causal relations huge variability in mistake-making from one period in an

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 Oderberg: What is life? an operational definition

organism’s life cycle to the next. What mistake theory enables Contemporary debates in philosophy of biology. Oxford: Wiley-
us to do is to organise many disparate ideas about action, Blackwell; 2010. p.72–85, at 83. (The authors do not use the
success and failure, welfare, avoidance, prevention, learning, word ‘spooky’).
among others, into a general framework for interrogating living 9. Searle JR. The rediscovery of the mind. Cambridge, MA: MIT
systems. In an age in which Big Data can overwhelm high- Press; 2002. p.238–239.
level theory, the investigation of mistakes enables biologists 10. Oderberg DS, Hill J, Bojak I, Gibbins JM, Austin C, Cinotti FM.
to stay close to the data while still entertaining the Big Ideas Biological mistakes: what they are and what they mean for the
of teleology and normativity. experimental biologist. British Journal for the Philosophy of
­Science. 2023. doi: 10.1086/724444
11. Garson JA. Critical overview of biological functions. Dordrecht:
References
Springer Cham; 2016.
12. Kahneman D, Slovic P, Tversky A. (eds.) Judgment under uncer-
1. Oderberg DS. Synthetic life and the bruteness of immanent tainty: heuristics and biases. Cambridge: Cambridge University
­causation. In: Feser E. (ed.) Aristotle on method and metaphys- Press; 1982.
ics. Basingstoke: Palgrave Macmillan; 2013. p.206–235. 13. Duffy A, Latimer KW, Goldberg JH, Fairhall AL, Gadagkar V.
2. Oderberg DS. Real essentialism. London: Routledge; 2007. Dopamine neurons evaluate natural fluctuations in ­performance
3. Canguilhem G. The normal and the pathological. New York: quality. Cell Reports. 2022;38: 110574. doi: 10.1016/j.­
Zone Books; 1991. p.22. celrep.2022.110574
4. Wildner G. Antigenic mimicry – the key to autoimmunity in 14. Gadagkar V, Puzerey PA, Chen R, Baird-Daniel E, Farhang AR,
­immune privileged organs. Journal of Autoimmunity. 2023;137: Goldberg JH. Dopamine neurons encode performance error in
102942. doi: 10.1016/j.jaut.2022.102942 singing birds. Science (New York, NY). 2016;354: 1278–1282.
5. Oldstone MBA. Molecular mimicry and immune-mediated doi: 10.1126/science.aah6837
­diseases. FASEB Journal. 1998;12: 1255–1265. doi: 10.1096/ 15. Roeser A, Gadagkar V, Das A, Puzerey PA, Kardon B, Goldberg
fasebj.12.13.1255 JH. Dopaminergic error signals retune to social feedback ­during
6. Arthur JF, Dunkley S, Andrews RK. Platelet glycoprotein VI-­ courtship. Nature. 2023;623: 375–380. doi: 10.1038/s41586-
related clinical defects. British Journal of Haematology. 2007;139: 023-06580-w
363–372. doi: 10.1111/j.1365-2141.2007.06799.x 16. Hill J, Oderberg DS, Bojak I, Gibbins JM, Austin C, Cinotti FM.
7. Garson J. What biological functions are and why they matter. Mistakes and correctness: a new look at the reductionism ­debate
Cambridge: Cambridge University Press; 2019. p.15. in biology. (forthcoming).
8. Cummins R, Roth M. Traits have not evolved to function the way 17. Okrent M. Nature and normativity: biology, teleology, and
they do because of a past advantage. In: Ayala FJ, Arp R. (eds.) ­meaning. London: Routledge; 2018.

17