What kind of individual do you mean?

One of the discussions I find most interesting in the philosophy of science is about what exactly constitutes a biological individual (or organism). The discussion would be a lot less interesting if everything were a vertebrate. Vertebrates (nearly always) develop from a single fertilized egg, so the (mostly) genetically homogeneous and (usually) genetically unique unit is the same as the spatially bounded, contiguous and physiologically integrated unit (this doesn’t even cover all the proposed criteria; see Clarke 2010 for a fairly comprehensive list with citations). But when we look outside of the vertebrates, what we often find is that some biological units have some of these properties and either groups or parts of those units have others.

Aphid on dandelion by Amoceann. Public domain image from Wikimedia Commons.

Aphid on dandelion by Amoceann. Public domain image from Wikimedia Commons.

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Problems with major transitions: Maureen O’Malley & Russell Powell respond

The Great Oxidation Event by Adelle Schemm.

The Great Oxidation Event by Adelle Schemm.

In a recent series of posts, I reviewed Maureen O’Malley and Russell Powell’s paper in Biology and Philosophy, “Major Problems in Evolutionary Transitions: How a Metabolic Perspective Can Enrich our Understanding of Macroevolution.” Although they made several good points, I thought that some of their criticisms were off the mark and that their proposed solution to the real and perceived problems with the major transitions framework was unsatisfying.

Drs. O’Malley and Powell are both heavy hitters in the philosophy of biology, and as I usually do when I dig deeply into someone else’s paper, I invited them to respond to my criticisms. They kindly provided a thoughtful rebuttal and permitted me to post it here. I’ll have more to say later, but for now I’ll just say that they make some good points and (most importantly) fairly represent my arguments. As usual for guest posts, I have made no edits to the content of their response, only formatted and added links:

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Time for a revision? Maureen O’Malley and Russell Powell on Major Transitions, part 3

Maureen O’Malley and Russell Powell say that the major transitions framework is in need of repair. They have a point, or rather several good points. I have looked at their criticisms of three different versions (the original framework as laid out in the book by John Maynard Smith and Eörs Szathmáry, Rick Michod’s ‘evolutionary transitions in individuality‘ framework, and Szathmáry’s revised ‘Major Transitions 2.0‘). But what is their proposed fix, and will it have the intended effect?

Figure 4 from O'Malley and Powell 2016. Two major aeons of evolution (modified from Falkowski 2006). ‘Gya’ stands for ‘billion years ago’; the date for the origin of photosynthesis may need to be pushed back (see Crowe et al. 2013).

Figure 4 from O’Malley and Powell 2016. Two major aeons of evolution (modified from Falkowski 2006). ‘Gya’ stands for ‘billion years ago’; the date for the origin of photosynthesis may need to be pushed back (see Crowe et al. 2013).

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Time for a revision? Maureen O’Malley and Russell Powell on Major Transitions, part 2

One of the cool things about studying the so-called major transitions is that they are as interesting to philosophers of science as to biologists. So you really can’t help being exposed to the philosophy of science literature, and many (maybe most) biologists in the field cross the lines at least occasionally. I’ve been to both, and I’m here to tell you that philosophy conferences are more fun than biology conferences.

Last time, I briefly summarized the various forms of the major transitions framework and addressed one of O’Malley and Powell‘s criticisms, that the framework is progressivist. Now I’d like to look at their other two problems: lack of unity and missing events. By and large, I agree with these points, although there are some caveats I’d like to point out. Next time, I’ll consider their proposed solution, which I’m afraid I don’t find helpful.

Disunity is actually O’Malley and Powell’s first criticism, but it will be a bit more complicated than progressivism to address, and I was short on time on part 1. Essentially, they are arguing that the major transitions are not a natural kind, philosophese for groupings that belong together because of some fundamental commonality, as opposed to more arbitrary groupings whose members are only superficially similar. So what are the transitions? Here’s the list from the book:

Table 1.2 from Maynard Smith J, Szathmáry E (1995) The Major Transitions in Evolution. Oxford University Press, Oxford.

Table 1.2 from Maynard Smith J, Szathmáry E (1995) The Major Transitions in Evolution. Oxford University Press, Oxford.

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Levels of selection in biofilms: Ellen Clarke on individuality

Pseudomonas biofilm. From Spiers et al. 2013.

Pseudomonas biofilm. From Spiers et al. 2013.

The question of what constitutes a biological individual is intimately entangled with questions about levels of selection. Many authors implicitly or explicitly treat individuals as units of evolution or some variation on this theme. A recent appreciation for the complexity of bacterial biofilms has led to comparisons with multicellular organisms. A recent paper by Ellen Clarke bucks this trend by claiming that multispecies biofilms are not evolutionary individuals.

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Karen Kovaka on biological individuality

At the Philosophy of Science Association meeting in Chicago, I attended an interesting talk by Karen Kovaka, “Biological Individuality and Scientific Practice” (the abstract of her talk is here). Now the paper arising from that talk is out in the journal Philosophy of Science. It argues that biologists do not need to resolve the question of what constitutes an individual in order to do good empirical work, with which I agree. She contrasts two views of the relationship between individuality and scientific practice, the “quality dependence” account and the “content sensitivity” account:

Quality dependence: the quality of empirical work in biology depends in part on the resolution of the debate about biological individuality…

Content sensitivity: Biologists’ understanding of biological processes is sensitive to the individuals they take to be participants in those processes.

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Pathways to pluralism: Beckett Sterner on biological individuality, part 2

Aphids on dandelion

Aphids on dandelion. Public domain image from Wikimedia Commons.

Previously, I introduced Beckett Sterner’s new paper comparing and critically evaluating the views of Ellen Clarke and Peter Godfrey-Smith on biological individuality. For Clarke, individuality is recognized by the presence of ‘individuating mechanisms’: traits that increase the capacity for among-unit selection or decrease the capacity for within-unit selection. Godfrey-Smith recognizes different kinds of individuals, but at a minimum, populations of individuals must have Lewontin’s criteria of phenotypic variation, differential fitness, and heritability of fitness, i.e. be capable of adaptive change.

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Pathways to pluralism: Beckett Sterner on biological individuality, part 1

In grad school I wound up hanging around with John Pepper (yeah, Dr. Pepper) a good bit. I think I disagreed with him more than I agreed with him, sometimes to the point of exasperation, but conversations with him were never boring.

Dr.-Pepper

One of John’s most annoying refrains was “is it an organism?” I was studying (and still study) a group of algae for which this question can be genuinely confusing. Most people would say a Chlamydomonas cell is a single-celled organism, and most would agree that Volvox is a multicellular organism, but what about the four-celled species Tetrabaena? A four-celled organism or a collection of four single-celled organisms? What about an undifferentiated colony of 32 cells, such as Eudorina? Or Pleodorina, which is around the same size but with two cell types? Somewhere between a unicellular ancestor and Volvox, a new kind of individual emerged. Among the extant species*, where do we draw the line between organisms and groups of organisms, or can we (or should we) draw a line at all?

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Pierrick Bourrat responds

[I invited Pierrick Bourrat to respond to my two posts about his new paper and to comments to those posts. He kindly agreed, and he provided the following guest post, which I have edited only for formatting.]

First of all, I would like to thank Matthew Herron for his interest in my work and his invitation to respond to his posts. Also, I would like to thank Rick Michod and Deborah Shelton for their comments.

I will respond to several issues pointed out both in the posts and the comments.

About the usefulness of the export of fitness view of ETI: I agree that it is a useful way of thinking about it, as long as it is used as a heuristic. This means that I am not inclined to think that building models with the assumption that the fitness of a cell would have been 0 had it been in an environment with not social partners will be able to explain in some deep sense ETIs (and even more so the origin of fitness at some level). In his comment to Matthew’s first post, Rick Michod claims that I somehow confuse realized fitness from a more counterfactual notion of fitness.  Well, to be honest, I do not see how one could simulate (I do not mean ‘explain’) the evolution of a process if the variables in the model do not correspond to realized properties of the system. If I want to model a particular phenomenon, I ought to use variables and parameters that represent the target system and clearly, at least for me, this counterfactual notion of fitness does not represent any properties the cells have because they always have social partners. It is common to use expected rather than realized fitness in models, but this assumption is justified when we can assume that population are large and the environment is overall not fluctuating too much. With the counterfactual notion of fitness, aside from being useful for explaining the ETIs, I fail to see how it could be successfully integrated in models (by successfully, I mean how it could represent meaningfully the target system).

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Pierrick Bourrat on levels, time, and fitness, part 2: collective fitness

Last week, I posted some thoughts on Pierrick Bourrat’s new paper in Philosophy and Theory in Biology, focusing on his criticism of Rick Michod’s ‘export of fitness’ framework. This week, I’ll take a look at the second of Bourrat’s criticisms, regarding the transition from MLS1 to MLS2, as first defined by Damuth & Heisler, during a transition in individuality.
MLS1 and MLS2 refer to two different versions of MultiLevel Selection. As Bourrat describes it (and this is pretty much in line with other authors), fitness in MLS1 is defined in terms of the number of particles (or lower-level units, or cells) produced, while in MLS2 the fitnesses of the particles and collectives (or cells and multicellular organisms) are measured in different units. Cell-level fitness (for example) is defined in terms of the number of daughter cells, organism-level fitness is based on the number of daughter organisms. (As with last week’s post, I’ll generally stick to cells and organisms, though the principles apply equally to any two adjacent levels.

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