I am currently sitting in on a graduate philosophy of biology seminar and the theme of this semester’s seminar is evo-devo and we recently discussed the concept of modularity. I’m also sitting in on a history of biology course and we have talked a little about the early 19th century French scientist, Georges Cuvier. While attending the seminar, I was delighted to make a historical link between the two! (And oddly enough, one of the works we read in the seminar was a chapter from a book on modularity co-authored by Gunther Wagner which opens with the same link I had made.)
In my posts about the Spencer-Weismann debates, one of Spencer’s anti-Darwinian arguments I focused on can be called the “correlation of parts.” It was an idea articulated earlier by Cuvier, who said (as quoted by Wikipedia),
Today comparative anatomy has reached such a point of perfection that, after inspecting a single bone, one can often determine the class, and sometimes even the genus of the animal to which it belonged, above all if that bone belonged to the head or the limbs. … This is because the number, direction, and shape of the bones that compose each part of an animal’s body are always in a necessary relation to all the other parts, in such a way that – up to a point – one can infer the whole from any one of them and vice versa.
This article from the Academy of Natural Sciences quotes another passage from Cuvier:
Every organized being forms a whole, a unique and closed system, in which all the parts correspond mutually, and contribute to the same definitive action by a reciprocal reaction. None of its parts can change without the others changing too; and consequently each of them, taken separately, indicates and gives all the others.
The parts of an animal are so thoroughly integrated, that changing a single part – without proportionately modifying everything else involved as well – would produce a non-working animal. The correlation of parts prevented Cuvier from accepting Lamarckian evolution, or transmutation, in principle.
The correlation of parts later became an argument for Lamarckian evolution against Darwinian evolution,* as exemplified by Herbert Spencer. (I wonder if there has been any study on this shift in the principle’s use?) He believed, like Cuvier, that frequently a feature of an organism is tightly linked to a multitude of other parts and it would be very unlikely for all these parts to simultaneously vary to the appropriate magnitudes and directions required by neo-Darwinian evolution that prohibits the inheritance of acquired characteristics.
For example, an elk’s antlers are supported by thick skulls and strong back and neck muscles. If an elk were to gain an extra point to the rack through continued growth, increasing its weight, its muscles would be strained to keep the head up and the animal wouldn’t survive. However, in Lamarckian evolution, the muscles that the buck strengthens over its lifetime are passed on to its offspring. Over generations, the muscles would continue to strengthen to the point where extra growth in the rack could be supported. Thus the correlation of parts showed that pure Darwinian evolution couldn’t work in elk antlers, but Lamarckism could – according to Spencer, anyway.**
So, historically, according to some, evolution (or specifically, Darwinian evolution) couldn’t work because of the correlation of parts. I think the concept of modularity easily resolves this.
A module can be loosely defined as a semi-autonomous individual part or process. (I have written about modularity before in a previous post.) An example of modular part is the assortment of lobster appendages: they act and move mostly separate from the rest of their body. A modular process is exemplified by the development of the appendages themselves: each appendage develops indepently of the others, and these appendages are specified and created by modular genetic networks and pathways. Additionally, these modular structures, processes, and genetic networks can evolve freely from the rest of the organism-at-large.
Modularity answers Cuvier’s and Spencer’s objections to evolution. Yes, an organism may be well-integrated, but it is also built of semi-autonomous modules. A part or process of an organism can vary without the rest of the organism falling apart! Spencer argued that if Lamarckian inheritance, which sidesteps the correlation of parts, weren’t true, then evolution couldn’t happen, but perhaps he was half-right: modularity (which also sidesteps the correlation of parts), not Lamarckian inheritance, is what allows evolution to happen.
* I am obviously omitting Darwin’s views on the topic, but given what I read in Gould’s The Structure of Evolutionary Theory, his views seem a bit muddy and complicated. Perhaps I will investigate Darwin’s views on the correlation of parts at a later date, but here I am focusing on correlation of parts vs. modularity.
** I don’t understand why Spencer excludes the possibility of later fluctuations or mutations that could arise after the weight increase in the antlers. He also doesn’t mention how the skull would thicken as a result of Lamarckian processes. I don’t think Spencer’s argument works at all given his framework, but it still illustrates the use of the principle of correlation of parts as an argument against Darwinian evolution.