Here’s a specific example, from my own work, of how the avoidance of mathematical modeling led to a fundamental discovery that eluded modelers and experimentalist for decades.
At least, that’s how I see it when I’m not feeling humble. It’s about resource allocation in ants, not the grand unified theory, after all.
For context, for those newer to the site, consider this post as a coda to an ongoing series (and discussion of sorts with Dynamic Ecology) about approaches to designing a research program. I have advocated that exploration by tinkering with unexplained curiosities within natural systems often leads to the best discoveries as well as the most consequential research programs. This post from a few weeks ago provides a good summary of that series. Another precursor to this post is a discussion about the relationship between mathematical modeling, hypothesis development, and how much math you need to become a scientist. That is also a precursor to this post, though it is a “long read,” for those averse to verbiage.
The subject of this post — the scientific discovery — came out in a paper last year (go read it if you wish), which I wrote with Sarah Diamond and Rob Dunn. In short, we discovered a fundamental pattern that could have been obvious to everyone, if anybody just looked in that direction. This pattern explains many unanswered ideas, going back to theories that E.O Wilson developed in the 1970s, along with George Oster.
Oster & Wilson set out to understand what regulates the varying levels of investment into the different members of ant colonies. Most inhabitants of ant colonies are functionally sterile, and in some species, there are multiple physical castes of sterile ants.
The genus Pheidole is the most species rich ant genus, and they’re found pretty much everywhere. All Pheidole (aside from a few exceptions) do something that isn’t found in many other lineages: they have two discrete sterile worker casts. They make big-headed soldiers and tinier minor workers, both of which do a variety of work for the colony. Some think that this dimorphic worker caste, and potentially the flexibility tied to its production, has enabled these ants to not only become ecologically successful but also to diversify.
Anyhow, Oster & Wilson made a number of predictions about the adapability of the ratio of soldiers to minor workers in Phediole colonies. One of their big testable predictions, or perhaps it could be seen as model to be falsified, is that the colonies actively adjust the ratio of soldiers to workers in response to environmental challenges.
It entirely makes sense. If a Pheidole colony is in an environment that requires more soliders, they would make more soldiers. Right? The problem is, despite a lot of looking carefully at Pheidole colonies, this wasn’t found. Finally in the mid ’90s it was found in the lab of Luc Passera, that P. pallidula colonies made more soldiers when they were exposed, without contact, to neighboring colonies. When I say it was found in the lab of Passera, I mean it happened physically in his lab. These were captive colonies.
A similar thing was found in the field in 2002, when I and Jeb Owen published a paper showing adaptive soldier production in another Pheidole species. (Also, my labmate Samantha Messier did the same thing before the Passera group, in a field experiment involving Nasutitermes termites and a machete.) Our studies were done in the field. In my experiment, when I put clumps of supplemental food in the field for months on end, the food was defended by soldiers, and in a short time colonies made more soldiers.
One thing I didn’t mention at the time, though, was that I didn’t find adaptive soldier production in a whole bunch of other species. However, I had less statistical power, and it was the most common species that showed this pattern. Maybe the less common ones did, but it was harder to detect.
If you were to ask around and dig into the literature, you’d see that it’s pretty clear that most species of Pheidole actually do not overtly shift their caste ratios when you mess around with their environment. Not every colony produces the same ratio, but a systemic environmental manipulation doesn’t cause an increase. Other than the two papers I just mentioned, I don’t think anybody else has found adaptive caste ratios in Pheidole. Others have looked, but it hasn’t emerged very clearly.
So, if most species just don’t ramp up and ramp down soldier production in response to the environment, what controls soldier production? For decades, there has been a consistent amount of work asking this question from behavioral, physiological and developmental angles. In the course of all of this excellent work (a lot of it being done by Diana Wheeler, Fred Nijhout, and their associates), we’ve made a lot of progress in understanding how colonies regulate their activity and how development is regulated through genetic, biochemical and physiological mechanisms.
One thing that I’ve always wondered about is, why do some species produce more soldiers than others? I’ve cracked open lots of twigs, and the numbers of soldiers are highly variable. And my experiments have shown that most species don’t obviously change their soldier production in response to environmental changes. There has been lots of great work to understand variation within a single species, but interspecific comparisons have been scant.
I can understand why there hasn’t been much comparative work. Measuring caste ratios of entire colonies can be hard. Find a Pheidole colony in the back yard and compare the number of soldiers and workers. See, not easy, huh? You’ve got to dig them up. Unless, of course, your backyard is a rainforest. In that case, you just pick up twigs. Over the years, I estimate that I and my students have picked up over 106 twigs over the years. Thousands of these have had Pheidole colonies inside. The rainforest is diverse, so I have data on many species. How do they compare?
Well, I learned that the caste ratios were different among species. Some species produced way more soldiers than others. Considering that we know so little about the natural history of these species, there wasn’t a great basis for comparing many of these species to one another. But one thing we could examine, quite easily, was body size. And, as it turned out, that was super-duper predictive of solider investment. Smaller species produced more soldiers than larger ones. When this pattern emerged on my laptop, it was one of those moments of elation that are very cool, but then you don’t have anybody with whom to share.
Then, I dug through the literature so see if the information that we had about caste ratios and body size shows the same pattern that I found in my rainforest. It turns out that the relationship is as identical as you can get. Our local scale pattern recapitulated Pheidole from around the world, and across the phylogeny.
Now, if you ask someone, what controls soldier production in Pheidole? You can say the answer is quite clearly body size. How and why does body size control this? There is some cool work that’s been done on this intraspecifically, that presumably is a mechanism that works more broadly.
How did my discovery of this generalized relationship come about from avoiding models? If you look at the work on soldier production, ever since Oster & Wilson published their monograph in the 1970s, there’s been a strong emphasis on modeling the mechanisms that trigger and regulate soldier production. Meanwhile, nobody before me bothered to step back to look at the big picture and ask, “how are species different and what is predictive of that?” If they did, then they would have found the caste ratio data in the literature as I had, and looked at the most obvious predictor: body size. Others were modeling solider production. I was merely trying to find a pattern.
I’m not claiming that the discovery of this pattern is earthshaking or that it explains mechanistically how colonies make more or fewer soldiers at the proximate level. The main take-home message from this paper is that many of the differences we find are driven by constraints rather than by adaptation, or that selection on body size is coupled with selection on soldier production. This leads to a lot of exciting thoughts about community structure, which we’re now working on.
This work by no means diminishes all of the careful experiments that others have done over the years on Pheidole. Though I’m not a developmental biologist nor as much of a behaviorist, I was able to find something that will be (or at least, I think should be) at the basis of future conversations about the evolution of caste ants.
This is why my choice is to keep asking “What is the pattern?” rather than attempting to model patterns.