Computational behavioural ecology…?

An image of a man wearing a t-shirt that reads "Deep down inside, we all love math".

I know I do.

I’ve written about the methods that we use in behavioural ecology, and the method that I use the most is definitely modeling.  To be more precise, I do a lot of computer simulation work on the evolution of behaviour (my focus is on evolutionary algorithms and individual-based models).  I do some formal mathematical modeling as well, primarily in game theory, but the bulk of my research is computational.  I admit it:  I’m a computer geek, and I always have been.  I love writing software, I love tinkering with code and hardware, and my natural approach to biological questions has always been to throw processor cycles at them.

Which leaves me wondering: what do we call computational studies of behavioural ecology?

The obvious answer, computational biology, is – I think – wrong.  At least, as it is currently defined, computational biology seems to be heavily focused on questions at the level of the cell or below.  If you look at the Wikipedia entry on computational biology, you’ll see that the examples given are all about cells, molecular biology, genomics, and so on.  Bioinformatics, computational genomics, “computational biomodeling” (not sure what that is, to be honest), systems biology, etc. are all examples of labels under the heading of computational biology, and none of them apply to the kind of work I do.  It’s natural that a lot of attention would be focused on this level of inquiry – people doing exciting work in genomics, cell biology, and proteomics are drowning in data and need computers to help them climb out of the well.  But I spend my time at the level of the individual and the evolution of their behavior, which doesn’t give me a lot to talk about with the computational biology people.

At the other end of the scale is the relatively new field of computational ecology.  If you forced me to chose right now, I would probably throw in with this camp, but it’s still a bit of an uneasy fit.  Computational ecology focuses on global population-level questions, and big ecosystems with many layers of complexity.  This is a fascinating area of work, but just like behavioural ecology differs from classical ethology / ecology in focusing on the individual, so too does the work I do focus on the evolution of mechanisms and behaviour at the level of the individual.  A typical question that I’m working on right now is the evolution of learning mechanisms for social foraging – how do animals learn to use the best strategy when foraging in a group, and what is the form of the mechanisms which allow them to do that?

And in the end, I’m left wondering where I fit.  There are others like me, of course;  for example, I’ve always admired the work of Dr. Graeme Ruxton, as well as the Laland group, both of which have done work in the same vein (this is by no means an exhaustive list, of either the people whose work I admire or who do work in the same area).  With the increasing specialization of scientists into subfields of subfields of major fields, I’m hesitant to invent a new term for myself and others like me, but maybe it’s time.

So:  computational behavioural ecology, anyone?

(Photo credit: Network Osaka)

How do we study behavioural ecology?

Yesterday, I wrote about my current field of study, behavioural ecology, attempting to explain what it is and what questions it tries to answer.  But I didn’t talk much about how it is studied.  How do we answer questions in behavioural ecology?

In general, there are three (overlapping) general techniques to be used.  Field studies are common, and have the flavour of the Wild West:  you pick your species, pack up your gear, and head into the wild blue yonder to collect data on your chosen animal.  You spend days, weeks, or months in the field, hoping that your data cooperate, and then come back to civilization to run some stats and publish your findings.  Field biology is romantic, and it draws a lot of outdoors types who like science and want to spend their time in the wilderness. I say this with a touch of envy;  I wish I had it in me to be a field biologist, but it just doesn’t work for me.

The second major method is the laboratory experiment.  Laboratory studies in behavioural ecology are important for their ability to control for confounding variables in a way you simply can’t in the wild, though this often comes at a corresponding cost to ecological validity (the match between the experimental setup and the real environment of the animal and its behaviour).  Labs tend to work on a few model species;  for example, one of the species that my lab does work with is the zebra finch (Taeniopygia guttata), which is one of the most widely-studied bird species.  Fish, like cichlids or guppies, are also common in lab work on behaviour.  Lab work is where you most often see the use of methods from other areas of biology, like molecular work or behavioural endocrinology, for obvious reasons of convenience.  I’ve done some lab work myself, but it is by no means where I spend the bulk of my time.

The third major method in the study of behaviour is modeling.  Biologists use mathematics, computer simulation, and statistics to create models of the behaviour and its evolution so that we can better understand it.  These models are used to drive new empirical work in the lab or the field, or explain results which have already been collected.  Because I’m primarily a modeler myself, you’ll see a lot about that on this blog, because one of the goals I have is to demystify the process and results of mathematical models of behaviour.

Now, I’ve talked about these three methods like they are distinct things, but it has to be stressed that everyone studying behavioural ecology has different mixtures of each tool in their tool box.  Some people are mainly field biologists, but even hardcore field people will do lab experiments to explore aspects of behaviour that they see in the wild in a more controlled experiment, and they will turn to models to help explain that behaviour.  Experimentalists love to see their work replicated in wild populations, and some will even head out into the field to do it themselves, while modelers have nothing to explain if there’s no data from experiments or field studies!  As a modeler myself, I never lose sight of the fact that the reason I’m trying to create these models of behaviour is to explain why real animals do the things that they do.

I know, I know;  this is a bit of a dry topic, reminiscent of the “methods” sections of your intro science courses in undergraduate studies.  But it gets better: in coming posts, I’m going to write about examples of each of these three methods from the best of both current and historical work, to showcase some of the more exciting examples of behavioural ecology past and present.  If you have any requests for topics to cover, or questions about how behavioural ecology is studied, please let me know by e-mail, Twitter, or leave a comment!

What is “Behavioural Ecology”?

The first and most obvious question to answer with this blog is what “behavioural ecology” (or “behavioral” ecology, for my American counterparts) is all about.  Historically, there has been some confusion on this matter, because of the overlap between various areas of study which deal with the question of animal behaviour. But I’m getting ahead of myself.

What is behavioural ecology?  In the introduction to an important textbook in the field, “An introduction to behavioural ecology“, John Krebs and Nick Davies write:

This book is about the survival value of behaviour. We call this subject “behavioural ecology” because the way in which behaviour contributes to survival and reproduction depends on ecology.  If, for example, we want to answer the question “How does living in a group contribute to an individual’s survival”‘, we have to start thinking in terms of the animal’s ecology; the kind of food it easts, its enemies, its nesting requirements and so on. These ecological pressures will determine whether grouping is favoured or penalized by selection. Behavioural ecology is not only concerned with the animal’s struggle to survive by exploiting resources and avoiding predators, but also with how behaviour contributes to reproductive success (p. 1).

The quote above shows that behavioural ecology is defined by the fields that intersect it.  These fields include ecology / ethology, evolutionary theory, and comparative psychology.  But the overriding focus of behavioural ecology is the study of behaviour through the lens of evolution, a laser focus that has set it apart from other fields in the past.  Behavioural ecology includes optimality theory and game theory (including evolutionary game theory) as part of its theoretical traditions, driven by the work of giants like William Hamilton and John Maynard Smith.  The kinds of questions that behavioural ecologists study include:

  • Information use by animals.  How do animals gather information about their environment, either personally or by observing others, and then use that information to improve their chances of survival or reproduction?
  • Behavioural syndromes / animal personality.  What drives the consistent differences in individual behaviour that we see in animals (like “aggressive” or “exploratory” behaviour) across different contexts?
  • Life history strategies – body size trade-offs, reproductive timing, etc.
  • Questions about foraging, whether as solitary foragers (optimal foraging theory) or foraging in groups (social foraging theory).  This is an area dear to my heart, because questions about social foraging are occupying the majority of my Ph.D.
  • The evolution of behaviour for mating and breeding;  mate selection, breeding habitats, and so on.
  • As above, the benefits and costs of group membership.  When should an animal join a group, and why?
  • The evolution of other social traits.
  • Learning mechanisms which drive animal behaviour.

And the list goes on.  The questions above come from a quick scan of chapter titles in a recent text on behavioural ecology (Behavioural Ecology, by Danchin, Giraldeau, and Cézilly), but that is not an exhaustive list by any means.

However, in our hurry to set ourselves apart from other fields, I hope that we don’t lose sight of what we owe them and what we can offer them.  I view behavioural ecologists as occupying a unique vantage point smack in the middle of those fields of study which occupy themselves with animal behaviour, working to both integrate the knowledge from those around them in an interdisciplinary way as well as extending that knowledge to give back.

But hey, I’m just a hopeless romantic.


Welcome to my new blog, A Bit of Behavioral Ecology.  I’m starting this blog because I recently realized that the field I’m currently knee-deep in – as I work on the last year of my Ph.D. – is not well-represented on the net.  There used to be a Behavioural Ecology blog written by Matt Macmanes, but that seems to have ground to a halt a couple of years ago with no signs of resurrection.  I hope that Matt won’t mind me picking up the torch, as it were, and attempting to carry on.

In any case, aside from Matt’s blog,  I haven’t found many resources for quality writing on behavioural ecology outside of journals. So I’m going to try and contribute something useful by using this forum to write about things that I think are cool in the world of behavioural ecology and science in general, and hopefully raise the profile of our field a little bit.   What kinds of things will this blog cover?  Well, I have pretty diverse interests but I do skew towards the modeling side of things, so I hope that potential readers can forgive me for that.  In general, though, you can expect to see material on recent papers in the field, “basic concepts” posts in areas that I’m familiar with, and whatever other random noise floats through my head on the topics of biology and science. And of course, I welcome contributions, comments, and ideas from you, the loyal reader.  (Of course, right now, I’m shouting into the void … but hopefully that will change!)  Leave a comment, send me an e-mail, reach out on Twitter, and hopefully I’ll see you around the blogosphere.