The Three Necessary and Sufficient Conditions of Natural Selection

Natural Selection is the key creative force in evolution. Natural selection, together with specific histories of populations (species) and adaptations, is responsible for the design of organisms. Most people have some idea of what Natural Selection is. However, it is easy to make conceptual errors when thinking about this important force of nature. One way to improve how we think about a concept like this is to carefully exam its formal definition.

[repost from gregladen.com]

In this post, we will do the following:

• Discuss historical and contextual aspects of the term "Natural Selection" in order to make clear exactly what it might mean (and not mean).
• Provide what I feel is the best exact set of terms to use for these "three conditions," because the words one uses are very important (there are probably some wrong ways to do it one would like to avoid).
• Discuss why the terms should be put in a certain order (for pedagogical reasons, mainly) and how they relate and don't related to each other.

When you are done reading this post you should be able to:

• Make erudite and opaque comments to creationists that will get you points with your web friends.
• Write really tricky Multiple Choice Exam Questions if you are a teacher.
• Evolve more efficiently towards your ultimate goal because you will be more in control of the Random Evolutionary Process (only kidding on this third one...)

Here are some definitions of Natural Selection I found on the web for your review:

• The differential survival and reproduction of organisms with genetic characteristics that enable them to better utilize environmental resources [source]
• Natural selection is the process in which some organisms live and reproduce and others die before reproducing. Some life forms survive and reproduce because they are better suited to environmental pressures, ensuring that their genes are perpetuated in the gene pool. [source]
• Process by which the genotypes in a population that are best adapted to the environment increase in frequency relative to less well-adapted genotypes over a number of generations. source
• The concept developed by Charles Darwin that genes which produce characteristics that are more favorable in a particular environment will be more abundant in the next generation. [source]
• the differential survival and/or reproduction of individuals within a population based on hereditary characteristics. [source]
• The process by which new species evolve when influenced by selective pressure (Martin et al, 2000). Natural selection occurs when the natural factors of environmental resistance tend to eliminate those members of a population that are least well adapted to cope and thus, in effect, select those best adapted for survival and reproduction (Nebel et al, 1998). [source]
• central thesis of the biologist Charles Darwin which suggests that within every population of living organisms there are random variations which have different survival value. Those which aid survival (or enhance reproductive capacity) are 'selected' by being genetically transmitted to succeeding generations. [source]

There are things I don't like about most of these definitions. A definition may focus on environmental conditions and thus ignore many very important other things such as developmental processes and mating. Definitions may focus on the individual's survival, etc., which works, but we may want to speak of traits as well as individuals. Some definitions use active verbs such as "ensuring that their genes are perpetuated in the gene pool." This may not be linguistically wrong but it incorporates teleological concepts, which don't need our help in creeping into our thinking (especially in the formal definition of a natural force!). There is often a direct link to Charles Darwin. This is good because it is true that this is his concept. However, a modern definition of Natural Selection needs to be Neo-Darwinian. So specifically referring in the definition to Darwin without more attention to the historical development is inadequate.

As a whole these definitions are not terrible, but they are mostly flawed for one reason or another. The definition I want to lay out here will have specific reference to the same process these definitions are about. However, there is also one very large problem with many of these definitions that is a bit more subtle than most but that is, to me, critical, that relates to the object of study. It is probably best to not assume a one-to-one correspondence between the process of Natural Selection that we are going to describe here and the concept of "adaptation." Ultimately, I would like to say that "adaptation" is the noun and "Natural Selection" is the verb in a key evolutionary process. But having said that, a useful and precise definition of Natural Selection may have to leave out processes that are nonetheless related to adaptations, both in terms of understanding the historical aspects of an adaptation and the functional aspects, but that do not fall under the process "Natural Selection" as it is best defined.

An adaptation may reach its particular form through the process of Natural Selection, but there are aspects to that form that have to do with, for instance, abiotic realities. You cannot have adaptations that involve swimming without bodies of water, for example. Our definition does not say "Oh, and there must be air, and water, and trees to climb in."

I actually want to provide TWO different (linguistically) but identical (functional) definitions of natural selection. The first is the coolest one, the simplest one, the one that makes you think. The second is a better pedagogical tool and serves better as the basis for adaptationist analysis of biological systems. The second also links better to certain historical aspects of the development of the concept.

The first definition is conceptually related to the following definition of evolution:

Change in allele frequency over time.

And in this context we can define Natural Selection as:

Nonrandom elimination of alleles.

I believe that this was suggested by Ernst Mayr.
http://en.wikipedia.org/wiki/Ernst_Mayr
This is a cool definition because it is short, sweet, and correct. Note the very important assymetry that this definition implies. There is no non-random generation of novel alleles. Only elimination. This jives with selection as a creative force, but neutral processes as providing the raw material. Neutral processes lay down the sediments that become the marble, Natural Selection is the sculptor. The adaptation is the sculpture.

The second definition, and the one I really want to get to, involves the so called "three necessary and sufficient conditions" and it goes something like this:

1. Variation in a trait
2. Heritability of the trait
3. Differential fitness conferred by the trait.

Just as important as these elements is the theoretical and logical framework in which they are placed. They are the THREE NECESSARY and SUFFICIENT conditions. Let's parse that out more.

Three ... That there are three is obviously because all important things happen in threes, sevens, or tens, for unknown cosmic reasons. Be that as it may, I want to point out that "three" implies "three different" things. If two of them could have been combined, then we would have only two. But there are three.

What this implies is the following: If there is a trait that varies, then it meets the first criterion. But "No," you say, "what about hair color? If I see a bunch of people with different color hair, and I KNOW they dyed their hair to get that way, this is not trait related to selection. So it does not meet the first criterion."

But you would be wrong. Remember, there are THREE DIFFERENT criteria. The first one is variation. If you see a bunch of dogs and they have different coats because they went to a very creative groomer, or a bunch of students standing around the cafeteria with blue, red, unnaturally black, and vivid yellow hair because they all went to Target and got dye and colored their hair, then in both cases you have met the first criterion because there is variation. By saying "these traits are not inherited" you have skipped ahead and cheated.

The second criterion is usually stated as "heritability" and that is a small problem, because the term "heritability" has a specific meaning in biostats. It is the measured variance in the genotype divided by the measured variance in the phenotype, squared. (Thus indicating something like the proportion of measured phenotypic variance that is accountable by genetic variation.) What is meant in our definition, however, is this: Is the variation in the trait conferred by genes? The dyed hair and the clipped poodle do not meat this criterion.

The third criterion is often stated as "Differential Reproductive Success" and that is simply wrong. The correct term is "Differential fitness" and it has to be differential fitness that is conferred by the trait. Why fitness instead of Reproductive Success (RS)? That is an important matter, and I will devote an entire post (at least) to that question. For now, let's just go with it.

OK, back to the theoretical context: Necessary.

Why are these three necessary? By necessary it is meant that ALL of them have to be true or it is not Natural Selection. This is fairly obvious. If you are missing any one of these three then what you are observing may be an interesting phenomenon but it is not Natural Selection. This also speaks to the need to be Neo-Darwinian. Darwin was aware of inheritance, but lacking an understanding of the mechanism, the necessary requirement of "heritability" (remember, shorthand for "the trait is passed on by genes"), any Darwinian definition (and I've avoided using his specific words here) is not good enough.

I had said at the beginning that the ordering of the three conditions is important. This is because they interact with each other in order. Here is how.

First, you need a trait that shows variation. Second you need to show that that trait is heritable, AND that the inheritance pattern relates to that variation. Third you need to show that the variation that is heritable maps on to differential fitness. So there is a strong logic to the order, that is embedded in the functional meaning of Natural Selection and any test criteria that are set up to investigate possible cases of it.

So the ordering works both for the understanding of the concept (pedagogy) and the investigation of the phenomenon.

Sufficient. That is a really important part of the context for this definition. If these three conditions are true, then Natural Selection IS happening. There is no alternative. The force of Natural Selection is activated when these three conditions are met, no matter what.

Does this mean that the selective force will have an effect? It depends. Natural Selection is a force, but there are other forces, including other instances of Natural Selection that may be operative in a particular organism. Gravity is a force but a fly can still walk on the ceiling. The gravity is still acting on the fly, but so is another force (adhesion) that keeps it up there.

This is an important point that bears emphasis. When the Three N&S Conditions of N.S. are working, Natural Selection happens. Period. The absence of an EFFECT is due to countervailing forces (including chance, because within it's operation there are still stochastic effect).

I believe it is incorrect and counterproductive to make "Sexual Selection" distinct from and parallel to "Natural Selection." Darwin was puzzled by apparent inconsistencies, especially along the lines of exaggerated traits mostly in males, and came up with Sexual Selection as a process to explain this. Fine. But I think it is best to think of both Sexual and Artificial Selection as subsets of Natural Selection.

The term "Selection" by itself is often used interchangeably with "Darwinian Selection," but often (usually?) as not really meaning the same thing as Natural Selection. Natural Selection works between generations on reproducing organisms and their genomes. Darwinian Selection, or selection in general can work on other things, like prospective students trying to get into law school, or neurons during culling, etc.