Sunday, December 6, 2015

An Evolutionary Biologist's Apology

A few months ago, I was attending the joint lab meeting of Rosemary Gillespie and George Roderick at UC Berkeley, where I am on sabbatical. At the start of the meeting, Philip Spieth showed us a review in Bioscience about a book called A Mathematician’s Apology that was, this year, celebrating its 75th anniversary. What made the book very interesting to evolutionary biologists was that it was written by George Hardy, a British mathematician most of us know as the (co)originator of the Hardy-Weinberg equilibrium. If you do any work in population genetics or evolution or indeed in many other aspects of biology, you will know about this equation. “Hmmm,” I thought at the time “that might be a cool read,” and so the same day I ordered it from Amazon. The slim book, of which about a third was an excellent introduction by C.P. Snow, arrived a few days later and it became my reading material for the next few nights.

In his book, Hardy used the term apology “in the sense of a formal justification or defence (as in Plato's Apology of Socrates), not in the sense of a plea for forgiveness” (from Wikipedia). In particular, Hardy mounted a defense of his brand of “pure” or “real” mathematics, in contrast to applied mathematics, which he described in terms such as “trivial,” “ugly,” and “dull.” Now, I am all for defending a science, or any endeavor really, in the sense of intrinsic interest or beauty – the pure delight of discovery. And this is what Hardy logically did for his real mathematics; yet the bizarre additional facet of his apology was that real mathematics could be justified because of its very uselessness. That is, if an endeavor can’t be of any use, then it can’t be of any misuse either. Indeed, the applied math that Hardy discussed was often done so in the context of its use in war. Thus, because real mathematics had no use, it couldn’t be used for horrible things, just providing a further justification for its existence. Thus, Hardy’s justification boiled down to beautiful and useless.

I was recently caused to reflect (again) on how these sorts of justifications relate to my own avocation – evolutionary biology. I have been back in Montreal this past week for a meeting of our long-standing bioGENESIS core project, which was originally a part of DIVERSITAS and is now transitioning to Future Earth. Within DIVERSITAS, an NGO focused on specifically biodiversity, our role was to bring evolutionary perspectives, in both deep-time (e.g., phylogenetics) and contemporary time (e.g., genetic variation within species), to biodiversity science. The value of this role would initially seem straightforward given that all past, current, and future biodiversity is the product of evolution; yet we sometimes found ourselves having to “apologize” for our existence within the context of a growing emphasis not on biodiversity per se but on ecosystem services. In this context, we wrote a number of papers about the importance of evolutionary thinking not only for biodiversity but also for ecosystem services. Most directly, we pointed out that EVOsystem services were the foundation of all current and future ecosystem services, as well as many other useful and non-useful aspects of biodiversity. And, over the years, a few of these papers and the debates surrounding the ideas made it into some of my blog musings.

DIVERSITAS has now ended and its core projects, including bioGENESIS, are being folded into Future Earth, along with a number of other global change NGOs. Future Earth is a much bigger and more encompassing enterprise than was DIVERSITAS: its focus goes beyond biodiversity to immediate human concerns, such as health and wellbeing, alternative energy sources, sustainable development, social structures, and so on. Thus, in the context of transitioning to Future Earth, bioGENESIS again needs justify its continuance. (I am not being pejorative here because, quite reasonably, all core projects transitioning into Future Earth need to do the same thing.) As a result, we spent several days writing a “transition document” that describes how we will fit into Future Earth, how we will address its core concerns and questions, and how we will interface with other core projects, as well as with likely stakeholders. In essence, one can think of the transition document as having elements of an “apology” in the sense of Hardy and Socrates, which made me wonder: What would an evolutionary biologist’s apology actually look like? (This apology is my own and does not necessary reflect the views of bioGENESIS or Future Earth.)

Following Hardy, and countless other commentators, we might divide any scientific discipline into “basic” (Hardy’s “real”) and “applied” contexts. Basic evolutionary biology is interesting, fascinating, inspiring, and enjoyable but, at the same time, often useless. We might here consider paleontology. Just think of how much richer our understanding and appreciation of the world has become simply because of all those cool dinosaurs that have been described. But this discovery and knowledge is useless, right? Well, perhaps not in the sense that such discoveries increase revenues at museums and lead to Hollywood blockbusters. But what about more specific discoveries, such as the fact that dinosaurs had feathers. This finding is super cool but surely the information is truly useless. Jurassic Park would not have been any scarier, and perhaps less so, if the velociraptors had feathers (see the video below). Thus, paleontology is really about wonder and beauty that we appreciate in the sense of great art or music while being useless in applied context. So it seems to me that this branch of evolutionary biology is pretty close to justifiable on the grounds by which Hardy justified “real” math.

Of course, most evolutionary biologists applying for grants do not say their work is useless. Instead, they often say precisely the opposite. That is, they find ways to make their work sound applied and relevant even if it isn’t, really. Sometimes they even write grants for applied work not because they want to address the applied question but rather because they think it is more likely to attract money, which will then allow them to piggy-back “real” science onto the “trivial”, “ugly”, and “dull” applied science that the grant outlines. So, in reality, many evolutionary biologists spend time justifying their existence in precisely the opposite way to Hardy – that their work is useful. Examples abound and our transition document for Future Earth makes four main cases (which I here rephrase in my own words and meanings).


Evolutionary history is relevant to many human endeavors. As just one example, knowing the evolutionary tree of life means that we can be sure to preserve particularly distinctive branches of life that might harbor properties that are useful for us in one way or another (remember, we are here justifying evolutionary biology in relation to its usefulness to humans). This perspective is often discussed in the context of conserving many diverse forms of life as “optional values” for the future.

Contemporary (sometimes called “rapid”) evolution is essential for projecting and shaping future, including how changing environments will shape populations, communities, and ecosystems. For instance, evolutionary potential – and natural selection acting on it – unquestionably determines whether or not populations can persist in the face of climate change, invasive species, pollution, habitat loss, harvesting, and so on. Evolutionary potential thus also shapes all of the community and ecosystem properties (including “services”) that stem from organisms.

Evolutionary thinking has direct benefits in many directly applied contexts. As one key example, the pervasiveness of evolutionary thinking in medicine has allowed us to make incredible advances in the control of infectious diseases and cancer by slowing the evolution of resistance in pathogens and cancerous cell lines. As another example, evolutionary thinking has been very effective in agriculture in slowing the evolution of resistance to pesticides, herbicides, and fungicides. And, of course, we have biological control and the diversification/domestication/improvement of crops and so on.

Evolutionary tools can be applied to many other contexts. For instance, evolutionary thinking (we should seek a polymerase that can function at high temperatures in test tubes by finding hot spring bacteria that are naturally adapted to high temperatures) is what led to efficient PCR methods, which has completely revolutionized genetic analyses and therefore medicine and agriculture and much else. In addition, the basic ingredients of evolution (variation, selection, inheritance) provide an algorithm that has been useful in many engineering and design contexts (e.g., the use of “genetic algorithms” in many optimization procedures).


Clearly, evolutionary biology as a general field is critically important, indeed essential, for pretty much any human endeavor. “But wait,” I hear you saying “this suggests that, beyond the gee-whiz dinosaur argument, we should not give any more money to basic evolutionary biology.” I too would – at this juncture of the apology – start to be concerned on the same account, not the least because much of my research is focused simply on understanding the way that various aspects of the world works. How fast do salmon evolve? What forces drove the evolution of Darwin’s finches? How do natural selection and gene flow oppose each other in threespine stickleback? How do different predation environments cause reproductive isolation to evolve between guppy populations? None of these – and countless other – evolutionary studies have any obvious immediate use. Yet these studies are important, perhaps more so than any of the other angles described above, for several reasons. First, the results of such studies are interesting, beautiful, amazing, inspiring, and just damn cool. Thus, they are justifiable in the same way as is paleontology and real math. Second, basic evolutionary biology elucidates patterns and mechanisms, the understanding of which can subsequently be adopted and used in applied evolutionary questions. For instance, the basic studies showing that evolution in natural populations can be rapid has subsequently had profound influences on conservation biology, natural resource management (fisheries!), agriculture, medicine, and so on. Thus, basic evolutionary biology studies of natural populations are justifiable on all levels: they have artistic appeal, like Hardy’s real math, and they have potential future utility, like Hardy’s applied math.

Hardy was concerned that if something was useful it could also be misused and thus cause harm: trigonometry helps design buildings but it also helps aim artillery shells. This same concern could well be leveled at evolutionary biology. Indeed, eugenics, which in extreme forms attempted to justify and promote the unfair treatment of various categories of humanity, took at least some twisted inspiration from applications of Darwin’s initially innocent ideas regarding “survival of the fittest.” Thus, evolution – like any other science or, for that matter, art – can be used for both good and bad. Here is where societal values and controls need to come into play. That is, science itself is neither good nor bad – these judgments must be rendered only to our uses of it. I am reminded of the "I have not come for what you hoped to do. I've come for what you did." scene in V for Vendetta. Nuclear physics can provide energy but it can also destroy the world. Fortunately, modern societies seem pretty good at – at least with time – sorting the good from the bad of any new scientific advance.

All species are the product of evolution and will evolve in the immediate and long-term future. Thus, all services and disservices that species provide have been, are being, and continue to be shaped by evolution. Without applying evolutionary thinking to species and their biological communities, we will have a drastically reduced ability to respond to ecological and societal challenges. But we also need basic evolutionary biology because it is first fine (at least some of it is) art in that we appreciate and enjoy its discoveries, including literally in the form of museums and nature documentaries and also in an enhanced appreciation for how the world works as we walk or swim through it. Moreover, the fundamental truths revealed by basic evolutionary biology will often have applications that we can’t even envision. I am optimistic that these applications will be all (or nearly all) to the good in the years to come. But it is up to us and to you.

Dobzhansky’s famously overused phrase “Nothing in biology makes sense except in the light of evolution” is clearly a vast understatement. In fact, nothing in the world makes sense except in the light of evolution. Of course, much of the world still doesn’t make sense and so only the widespread application of evolutionary thinking will bring the necessary illumination.


Of course, I am not the first to consider justifications for the study of evolutionary biology, with a good previous example being:

Futuyma, D. J. 1995. The uses of evolutionary biology. Science 267:41-42.


  1. "Fortunately, modern societies seem pretty good at – at least with time – sorting the good from the bad of any new scientific advance.” Wow, that’s one of the more staggeringly optimistic statements I’ve ever heard. I would say instead that modern societies seem pretty good at – at least with time – finding military applications for any new scientific advance. It would be nice to think that evolutionary biology is relatively immune to this, but sadly I'm not sure that's true. Evolutionary algorithms are being used to refine the designs of drones and killer robots, to hone machine learning and artificial intelligence, and so forth. As you say, knowledge is neither good nor evil in itself; it depends on how we choose to use it. When I look at the state of the world today, it does not seem to me that we are choosing to use it in particularly positive ways.

  2. From Michel Loreau, former head of DIVERSITAS.

    Dear Andrew,

    I did enjoy reading your post, which asks an old and important question about the social values of science.

    I personally have a different view on the « beautiful » vs « useful » dichotomy, though. I believe almost everything is both useful and beautiful, and what can be viewed as useless in one context can be very useful in another context. Evolutionary biology is no more beautiful or less useful than any other science. In a collective enterprise such as Future Earth, curiosity-driven basic science can be very useful for other parts of the enterprise. Of course, if you plan to focus bioGENESIS on the feathers of Velociraptor, that might not be the most straightforward argument to contribute to Future Earth...

    All the best,

  3. From Doug Futuyma.

    Dear Andrew,

    Thank you for connecting me to your excellent essay on a subject I increasingly care about. Four days ago, I had the final meeting of my undergraduate evolution course. I ended it by reading the beginning of the last section of my textbook, which starts:

    "All art," said Oscar Wilde, "is perfectly useless." He meant that as high praise: art is a human creation that needs no utilitarian justification, a creation that is justified by being an expression -- indeed one of the defining characteristics-- of humanity.

    Much of what is most meaningful to us is "perfectly useless": music...gardening, spiritual inspiration-- and understanding. Whether the subject be mathematics, the natural world, philosophy, or human nature, attempting to understand is rewarding in itself...

    The passage continues in that vein -- this at the end of a chapter that relates evolutionary biology to social concerns, and describes a range of practical applications. That is, it's much the same dualistic apology as yours. I'm strongly, emotionally committed to the importance of the perfectly useless, of truth and beauty threatened by the crassly economic,acquisitive, materialistic.

    Thanks again,


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