Talk:Natural selection

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New version of Natural selection

Kim van der Linde and Gleng have developed a updated version of the Natural selection article at User:KimvdLinde/Natural selection. At the talk page there:

This page shows a proposed new version of the article on Natural Selection. This has been developed using parts of the existing version by Kim van der Linde and Gleng, following the discussions on the Natural selection talk page. We have tried to produce an article that will be both clear and rigorously accurate. If there is general agreement that this version is an improvement over the existing version, then we will replace it. Please put any comments and criticisms about this proposed new version on this Talk page, and in particular please indicate your support for, or opposition to, replacement of the existing version. Please do not edit this version except for any minor corrections at this stage.

Kim van der Linde ^{at venus} 05:16, 13 May 2006 (UTC)

• I like the new version. Let's go with it. Guettarda 23:05, 13 May 2006 (UTC)

what is born out of spite...

what is born out of spite...

"A well-known example of natural selection is the development of antibiotic resistance in microorganisms."

let's dissect the very first sentence above: "an example of selection is the development of antibiotic resistance..."

kim, i thought you agreed that selection is not evolution !? now the whole new article is full of such high-school-level imprecisions and sloppy formulations.

and, possibly more serious, the new article is mainly about evolution !

everybody should welcome additions and revisions to the existing article as long they are done orderly and carefully.

but one must correct any imprecisions, and one must refocus back to selection any additions that focus mainly on evolution.

note that i am an evolutionary biologist but i couldn't care less about selection for scientific reasons that are not simple.

however, the article about selection must be about selection, i.e. about the generation of fitness differences. Marcosantezana 04:23, 16 May 2006 (UTC)

See User_talk:KimvdLinde/Natural_selection#what_is_born_out_of_spite... for responses. Kim van der Linde ^{at venus} 20:46, 19 May 2006 (UTC)

Axel147 on new KimvdLinde version

I generally like it!! Here are some hopefully constructive comments...

1. I think the example is given too much prominence and is a bit scary for the layman.
2. On Sober's distinction the emphasis should be 'cause of selection' vs. 'free rider' (not trait vs. individual).
3. I think I object to the sentence 'For Darwin, fitness was equivalent to survival'. Darwin did not really use the term 'fitness' obviously undertood the importance of fecundity so I think this is misleading. ('Fittest' is really Spencers.) And is it survival of individuals or families of indiviuals etc.?
4. The article descibes natural selection in different places as a 'process', 'principle' and an 'idea'. Is this consistent?
5. The wider defintion is also a 'mechanistic' one: it is just that the mechanism of selection is combined with a mechanism for propagation (inheritance).
6. I hadn't read Petri Krohn trivia up until now! If only Darwin had used the term 'natural preservation' we wouldn't have had the mass discussions on this page! I'm not so sure this is trivial though as this confirms (if there was any doubt) that Darwin meant natural selection in the wide sense. However 'preservation' is not quite synonymous with 'evolution by natural selection'. New variation is required for (substantial) evolution but not for preservation. So let's get this part right!

— Axel147 19:55, 18 May 2006 (UTC)

I don't think KimvdLinde's point about 'result' quite makes sense: probable survival and reproduction of an individual is the result in KimvdLinde's 'phenotypic selection'. This is not fundamentally different from Darwin's probable preservation of favourable traits as the result of his 'natural preservation'.

(In any case it seems to me to be perfectly legitimate to define a process using a result: scoring a goal, boiling a liquid, lighting a match...)

— Axel147 13:00, 19 May 2006 (UTC)

The examples you give are perfect examples of a process, and that is exactly what the mechanistic version of NatSel does. As in scoring a goal, the ball is not part of the scoring, it is an object used to do the action with, resulting in a result (ball is in the goal). If you would include the ball in the mechanism, scoring a goal with for example a ice hockey puck should have its own name, just as scoring a goal with a discus. Kim van der Linde ^{at venus} 14:16, 19 May 2006 (UTC)

In your world the phenotype is the object used to do the action with, with a result (phenotype is selected). But you haven't answered my point: both defintions can be said to be mechanistic, and can be undertood in terms of a (probable) result. It is false to claim otherwise. — Axel147 15:22, 19 May 2006 (UTC)

You are right, the second has a mechanistic component in it. What would be your preferred choice of labeling those two definitions? Kim van der Linde ^{at venus} 15:30, 19 May 2006 (UTC)

It's difficult. Endler tries 'restricted' and 'general' but restricted sounds a bit pejorative. The only thing I can think of are more neutral words like 'phenotypic', 'wide', 'expanded', 'extended', 'general', 'broad','classic'. (Less sure about 'narrow', 'Darwin's') — Axel147 16:10, 19 May 2006 (UTC)

'Restricted' and 'general' contains a POV, so the words need to be neutral. The reason to use the current terms is that the inclusive version includes the result as wellas the mechanistic part. The mechanistic def describes what it is, just the mechanism. I think the terms should be clear about this. But I will leave it as it is untill there is a better set of neutral terms. Kim van der Linde ^{at venus} 17:22, 19 May 2006 (UTC)

What is it that causes a phenotype to be selected?

Natural selection we are told in the opening sentence is a 'process'. The beginning of the process, presumably, is the point of conception and the end, presumably, is the point at which an individual ceases to have any influence on its descendents (which could even be after death).

But it is imperative we define what it means to be a 'selected' phenotype. Is it defined in terms of capacity to survive and reproduce? Absolutely not — it is defined in terms of its capacity to propagate its characteristics. This is the only thing that matters. It is the only viable definition: and it demands a mechanism for propagation to be part of the definition of natural selection despite objections from KimvdLinde.

It is totally false to imply natural selection can happen without inheritance. If anyone can argue otherwise please do so. I think the article is a failure if we do not clearly define the criterion for selection.

I have since retracted the above!

— Axel147 04:23, 20 May 2006 (UTC)

It is totally false to imply natural selection can happen without inheritance. people try to artificially select for stuff that has zero heritability all the time. My understanding is that hip dysplasia has a heritability not significantly higher than zero in many dog breeds, but breeders still try to breed it out. The Alberta Eugenics Board sterilized citizens of my Province until the 1970s, they were trying to breed out traits with zero heritability. Both of these examples still constitute artificial selection despite there being no heritability. By analogy, traits with zero heritability may still determine which individuals get to breed, by Natural Selection. I've provided you with this argument before, I don't remember what your response was, feel free to just point me to the diff. Pete.Hurd 04:44, 20 May 2006 (UTC)

I am sorry Axel that you feel that this article uses the wrong definition. The only thing I can do is to suggest to you that you write an scientific article about it for a scientific journal and convince the scientific community. See for example Futuyma, D.J. 2005. Evolution. Sinauer Associates, Inc., Sunderland, Massachusettes, page 251. I quote: It (=natural selection) is a name for statistical differences in reproductive success among genes, organisms, or populations, and nothng more.. Or I can send you the article (pdf) of Lande, R. & Arnold, S.J. 1983. The Measurement of Selection on Correlated Characters. Evolution 37: 1210-1226, the key article concerning this topic. A screenshot of the crucial paragraph can be found here (Probably good as fair use for me as limited use, not for wikipedia). The article is cited 1277 times (!!!!!), primarily for this clear distinction. If you want to start a discussion with Russ (Lande), I have the e-mail address of him for you. Kim van der Linde ^{at venus} 04:50, 20 May 2006 (UTC)

I reverted the proposed changes by Axel, because the distinction had become a distinction between natural selection perse and evolution by means of natural selection.

Scientists use several, slightly different definitions of natural selection; the main difference is whether natural selection is viewed as the selection of phenotypes in a single generation or the selection of traits over several generations.

Selection continued over multiple generations is evolution, not Natural Selection. This is a key distinction between the two. Kim van der Linde ^{at venus} 04:59, 20 May 2006 (UTC)

I absolutely agree with Kim on this, and this is not a minor issue. To confuse a mechanism (natural selection) with its consequence (evolution) is to destroy the explanatory power of the mechanism; the "explanation" becomes tautological and vacuous, and the criticisms of creationists gain some validity. The power of the explanation lies in the simplicity, inevitability and "blindness" of natural selection as a mechanism. most importantly here, the mechanism (selection) has no ability to "tell" whether a trait is heritable or not. natural selection will act anyway. There will be consequences whether the trait is heritable or not, consequences that will affect evolution, but adaptive evolution of particular favored traits will only occur if the traits have a heritable component. Gleng 10:36, 20 May 2006 (UTC) In the opening sentence, "the process" could be replaced by "all the mechanisms by which", but I'm not sure that this would be an improvement in clarity.Gleng 10:39, 20 May 2006 (UTC)

In the end, Natural Selection is such a simple concept, that it becomes difficult to grasp. Because what it does is just giving the maladapted a hard time so that the better adapted produce more offspring. The rest is details on sub-mechanisms based on live stage, context, selection agent etc. Kim van der Linde ^{at venus} 15:13, 20 May 2006 (UTC)

Ok, I am forced to retract what I said yestersay as today I seem unable to defend it. I accept natural selection in the phenotypic sense can be both measured and defined 'without recourse to principles of heredity'. Of course selection always happens 'blindly' - I never meant to imply otherwise. In the Darwinian sense inheritance is only required to achieve preservation of variations.

The only case where 'capacity to survive and reproduce' differs from 'capacity to propagate characteristics' I think is with the green beard effect. A green beard may have good capacity to secure its characterisitcs in future generations despite low fitness. But even in this case I accept it is the fittest who are selected (and the descendents of those with the greatest inclusive fitness who survive). And I accept also that fitness can be defined without reference to genes.

Despite all this KimvdLinde's 2 extracts appear to contradict each other: how 'natural selection is statistical differences in reproductive success among genes' is compatible with 'natural selection acts on phenotypes, regardless of their genetic basis' I am not sure? I think the answer is that the second defintion goes out of its way to call itself 'phenotypic selection' and 'phenotypic natural selection' while the first leaves the door open to the extended definition.

The two are within the same scope. The key is difference is the Unit of selection. The first quote includes various levels, while the second was from an article that primarily dealt with whole individuals as the unit of selection, and for that reason used phenotypic. However, you can talk in the same way about it when discussing meiotic drive.Kim van der Linde ^{at venus} 00:48, 21 May 2006 (UTC)

(Continued)Anyway as a thought experiment I would like you to imagine for a moment that the principle of inheritance no longer exists: let's say that individuals still have a genetic basis, and that basis still influences their liklihood of reproducing. But here is the difference: in this world the children's genes have no relationship with those of the parent: we can imagine them to be chosen at random. The question is would natural selection still exist? Well in a Darwinian sense no, because there is no way of preserving particular variations. But in a phenotypic sense yes, and fitness would also continue to exist.

Exactly, it would still exist, the difference is whether it has an effect towards the next generation. Kim van der Linde ^{at venus} 00:48, 21 May 2006 (UTC)

(Continued)In answer to Gleng one of my main reasons for joining this discussion is with hope of adding to an article that would clearly express what Darwin intended, so as to counter creationist objections. I understood the argument and think the problem wouldn't exist if Darwin as he later wished had used the term Natural Preservation.

I think one of the manin faults of the scientists is that they get dragged into the discussion with creationists about Darwinism. We are 150 years ahead, and the current science is so much more sophisticated. It actually caters them because it is much easier to shoot holes into Darwinism than in the current thinking of Natural Selection. Furthermore, wikipedia is an encyclopedia, and it should reflect what we know. It is not a soapbox, nor has it an agenda. If an accurate description of an topic results in ammunition for a certain group, so be it. Kim van der Linde ^{at venus} 00:48, 21 May 2006 (UTC)

(Continued)In the current article we have an opening sentence 'Natural selection is the process by which individual organisms with favorable traits are more likely to survive and reproduce' . But if favourable traits are those which help an organism survive and reproduce it is this defintion which seems tautological. Darwin's 'preservation of variations' is much more substantial. Phenotypic natural selection therefore, not Darwinian natural selection is surely the one which is more easily reduced to tautology (see Endler). However the argument for evolution stays the same: all we are debating is whether inheritance falls inside natural selection of outside of it. The potential tautology just sits in a different place.

— Axel147 20:32, 20 May 2006 (UTC)

It may look like a tautology but those can be found by removing pieces of the sentence as they are redundant. So, if this sentence is tautological, we could remove the word: Natural selection is the process by which individual organisms with traits are more likely to survive and reproduce. This first version does bot mean anything anymore. The second version not even produces a coherent sentence. And you are correct, most of the discussion we had was about whether NatSel was evolution or not. Kim van der Linde ^{at venus} 00:48, 21 May 2006 (UTC)

To be consistent please correct sentence 'Natural selection need not apply only to biological organisms; in theory, it can be applied to any system in which entities 'reproduce' in a way that includes both inheritance and variation.' — Axel147 22:45, 20 May 2006 (UTC)

When I was at university I developed a genetic algorithm.

1. Generate a population of phenotypes ('best fit curves' for a quantum mechanical problem) from genes (input parameters).
2. Select phentotypes to reproduce using a fitness criterion
3. Reproduce (with inheritance by shuffling groups of genes of selected phenotypes)
4. Add variation (micro and macro mutations)

If step 2 is 'natural selection' I would say it is impossible that step alone can create complexity? — Axel147 09:55, 21 May 2006 (UTC)

Indeed, Natural selection on its own can not generate complexity. If the complexity is associated with higher fitness, those more complex individuals will be selected for. However, if the highest fitness is achieved by simplification, those will be selected for. You could have programmed that in to your genetic algoritm (I love them), and the result would be simplification, which for design purposes can be very usefull fitness criterion. The whole algoritm that you described is evolution by means of natural selection. Kim van der Linde ^{at venus} 14:38, 21 May 2006 (UTC)

My point was why did you put in the artice whether 'natural selection per se can generate complexity is strongly contested'. Surely selection (step 2) on its own cannot generate anything. Only in the extended sense (step 2 + 3) is there even a debate? — Axel147 17:04, 21 May 2006 (UTC)

Missed it, this section was copied from the old version,a nd although I have read over it, it has not endured the same scrutiny as the new sections that I wrote above. I will in the coming days look more into the isue, and I think I will rewrite it somewhat. The claim that NatSelEvol can not generate complexity is a creationist one, in biology, there is not an issue about that. Kim van der Linde ^{at venus} 17:20, 21 May 2006 (UTC)

In response to Gleng. I think we are on agreement with this (and apologies for giving a now retracted false argument for inheritance which appeared to show I was confused over the mechansism). Whhat I intended to say is that I think it is important to understand that the use of words has changed but the explanatory power has not. When Darwin says natural selection is [to paraphrase] 'the selection and preservation of variations which result from selection of the best adapted individuals' this does not undermine the explanatory role of phenotypic selection which is part of this process. (Nor is this equivalent to evolution as there is no mention of new variation.) There is no confusion over the mechanism (despite my strange contribution a couple of days ago!). It is just a question of semantics. — Axel147 09:55, 21 May 2006 (UTC)

I'm sure we are in agreement, and just struggling with the right words. I think though that it is easy to forget how important non heritable variations are for individual fitness. Most of the things that make individuals differ are differences of environment (whether involving interactions with genes or not), or are the results of chance and circumstance, or are acquired characteristics. These determine traits as we understand them much more than differences in genes, the effects of which will be more subtle and more widespread (each gene affects many traits, mostly in slight ways). It's still a bold idea that natural selection of individuals for these traits as a whole, but resulting in selective preservation and recombination of just the genetic components, can have such power.The argument is far from tautological, and the conclusions far from inevitable - the mechanism might be inevitable but the consequences still take a leap in vision to encompass the vast time, the large populations, and the effects of random change. Darwin grasped this vision, but he saw the big picture only - the devil in the details means that we are struggling with words and concepts whose meaning has shifted as the importance of the nuances has become clearer. Reading Darwin now, he is not wholly consistent and coherent in his usage; he was a thinker, not a preacher, and thinking is a process of continual change of meaning :) Gleng 12:40, 21 May 2006 (UTC)

In the Wikipedia evolution article I think we should clarify the sentence 'Natural selection is the idea that individual organisms which possess genetic variations giving them advantageous heritable traits are more likely to survive and reproduce and, in doing so, to increase the frequency of such traits in subsequent generations.' The is good description of Darwinian natural selection but we should more clearly distinguish this from phenotypic natural selection otherwise people will be confused. — Axel147 09:55, 21 May 2006 (UTC).

I had seen that sentence, and I am going to poke the editors there in some time. Currently I am focussing for a small time on less contentious articles (parrot species and systematics), but when I am in the mood, I will start talking there also. Kim van der Linde ^{at venus} 14:38, 21 May 2006 (UTC)

HIStory

Yo! Can we dump the history and sociology stuff from this article? It heavily retreads stuff that happens in evolution, and I think, as a secondary page, it doesn't really belong here. Brief mention of it and reference to better, secondary articles on the subject are in order, but not the 4-5 odd pages we've got here. I humbly submit. Anyone concur? Graft 23:24, 21 May 2006 (UTC)

Well dumping it completely no, but cutting out significant sections, fine with me. That are oieces that are left of the old article, I did not want to rewrite everything at once. Kim van der Linde ^{at venus} 01:16, 22 May 2006 (UTC)

Looking at the History and Impact sections here and in evolution I would say they are generally focussed on Natural selection here and on evolution there. There are two issues - the impact of the idea of Descent of species (covered in evolution - and the impact of the explanatory power of the mechanism, covered here. I don't know that an article on Natural selection can avoid either the history or discussing the very broad impact beyond biologyGleng 08:37, 22 May 2006 (UTC)

general comment

I want to thank all the people who have been active on this article over the past couple of weeks - I am commenting on the process, not the article itself - I think this has been a model of collaboration for writing Wikipedia articles. bravo! Slrubenstein | Talk 11:01, 22 May 2006 (UTC)

adaptation

"this can lead to adaptation" - I think this needs explanations. Most non-scientists think adaptation is an intentional act; I think in evolutionary biology it is an effect, not an act. "Adapt" is a verb and it is hard to think of it as anything other than an act, the act of the creature thad adapts (i.e. not the act of "nature"). Slrubenstein | Talk 11:04, 22 May 2006 (UTC)

I tweaked it a bit, and I think it is clearer now. Kim van der Linde ^{at venus} 11:21, 22 May 2006 (UTC)

Thanks. Now, please look at my latest edit to the article. My fear is that I am being redundant. My concern is to be crystal clear about adaptations being (1) inherited rather than learned and (2) effects rather than acts. I think the whole article is consistent with this view, but I also think these two distinctions are what trip up almost all non-scientists and even many scientists, so I think the two distinctions need to be highlighted and explained very clearly. Right now, they are explained in the context of a larger narrative about the development of the theory. There is nothing wrong with this narrative, I just fear non-scientists will fail to appreciate these subtle and counterintuitive distinctions. I invite Kim, Gleng, Graft, Axel, others to address this in the article. Slrubenstein | Talk 12:51, 22 May 2006 (UTC)

The way you have added it is actually causing more cinfusion, as this implies that adaptaitons are behaviour, which they are not perse. I will undo it for the moment, and add something to clarify it there as well later when I am back from workKim van der Linde ^{at venus} 13:02, 22 May 2006 (UTC)

That's fine - as long as you understand my point, then, if we agree, I am fine for you to figure out an effective way to communicate the point in the article, if you are willing, Slrubenstein | Talk 13:06, 22 May 2006 (UTC)

I think I will add a sentense making explicite what adaptation is in this context, more like a defining sentence. Or something like that. It is perfect clear what you want to say, and it is indeed one of those little nifty things that need to be addressed.-- Kim van der Linde ^{at venus} 13:09, 22 May 2006 (UTC)

I tweaked it, let me now if it is clear enough now. -- Kim van der Linde ^{at venus} 13:27, 22 May 2006 (UTC)

Kim, first, I re-indented this section - I think as long as it is just the two of us we should each be intented uniformly - otherwise, the text will get more and more cramped (and I think this indentation serves to make it clear who says what). I do not think your recent edit is enough ... but I am not impatient and can wait to see what others come up with. I want to stress that I do not think anything in the article is in any way "wrong." My concern comes only from my experience with non-scientists - undergraduate students mostly in the US but now in the UK, but also well-educated professionals, who really just don't get it. I think the language you use (including "passive") is correct and anyone with a good solid understanding of evolutionary theory will understand exactly what you mean and why it is important. I just think most lay-people will miss it. What is at stake here is not the accuracy of the article or even the clarity, as such, but who our audience is. If I asked an undergraduate student to read this article and prepare for a quiz, I am sure the student would read it carefully, memorize much of it, and be able to regurgitate it. But if I asked them to then talk about where new species come from, or ask them what makes Darwinian thought so radical, I'd discover that they just didn't get the distinctions I make above. Is this important? I think so, given how many students increasingly turn to Wikipedia, and given the popularity of creationism in the US which is tied to the abysmal standard of science education we have. But I do not expect you or other people contributing here necessarily to agree with me on this point - my point not about natural selection but about our target audience. So does the article really need more work in this regard? I think so but am happy to see what others think, if you or anyone else think it is worth the effort or see elegant ways to address this. Slrubenstein | Talk 13:49, 22 May 2006 (UTC)

If I am correct, the question you ask is about the scope of the article. I do not think that this article should detail speciation for example, there are a whole series of other articles about that on wikipedia. In the end, Natural selection perse is pretty dull. It is the embedding in the wider context that makes it so radical. Darwins idea was so radical because he explained evolution by means of natural selection. And maybe that is what we need, a seperate article about Evolution by means of Natural Selection (which might be actually already covered enough in Darwinism and evolution). What I question to a degree is how far we have to go in making each and every article creationist proof (it actually annoys me at times to see that some American controversies creep into every possible article that is even slightly related to it). Maybe we can do a littlebit here and there to clarify and point towards the appropriate articles, but in the end, I think we should keep the article as much as possible on the topic, and leave all the social controversies out of it. Just my 0.02 Euro cents. -- Kim van der Linde ^{at venus} 14:30, 22 May 2006 (UTC)

Fair enough! But just to be clear, I didn't mean that we should explicitly discuss creationism or any creationist objection (which we do in the evolution article) or go into detail on speciation - only highlight and explain the two distinctions I draw concerning adaptation. (My comments about creationists were only meant to explain why I feel strongly about being so clear about adaptation, not to spell out more turf the article should cover). Let's see what others think. Slrubenstein | Talk 14:58, 22 May 2006 (UTC)

I understand Slrubenstein's concern, but I think it's difficult to avoid using the word 'adapt' in some form or another. I can't think of anything better than Kim's wording at the moment. I think like Kim we could go too far in making the article creationist proof, but equally we could go too far in making it read like a science text. The audience for an encyclopedia no doubt includes many lay people particularly for an article with such great general interest as this. Ensuring the article accessible to non-scientists is surely a worthy goal if it can be done without compromising it. Hopefully that should also help in making it creationist proof. — Axel147 18:22, 22 May 2006 (UTC)

Did I get lucky with Endler?

Forgive me for presenting this topic once more but I do have a bit have a bit more evidence - so please bear with me. And this is related to the scope question, whether a separate 'evolution by natural selection' would still be useful and also the creationist proofing.

First let me say I welcome the paragraph on definition in the new version but just think it might go a little further. Kim's view that 'natural selection per se is pretty dull' and that it is the 'embedding in the wider context that makes it so radical' is of course perfectly coherent. The problem is that is true only when using a phenotypic definition while many scientists and creationists are using a wider one.

No, because Natural selection is a single generation thing regardless of definition, and what is interesting is the multigeneration thing where you get speciation by a passive process and do not need a God. -- Kim van der Linde ^{at venus} 14:41, 23 May 2006 (UTC)

Once we have included inheritance, we have the full mechanism, the full life cycle so that is enough for difference in trait frequency. — Axel147

Depends on your defintion. -- Kim van der Linde ^{at venus} 01:20, 24 May 2006 (UTC)

The crux of the difference on the definition issue is heritability and depending on one's taste (and wideness of definition) we know the role played by inheritance can be either:

1. A pre-requisite for or part of the mechanism of natural selection
2. Only required for natural selection to be 'effective' or to achieve 'results'

And that these equate to what I have been calling 'Darwinian natural selection' and 'phenotypic natural selection'. We also know there is no difference in understanding, in overall mechanism, merely in semantics, but it is still important to make this clear.

So are both definitions equally in play? Can we get rid of one? Well as a representative of the layman (and without easy access to scientific literature) I have to conclude the answer is 'no'. Let's for instance have a quick look at using the views of three eminent professors Sober, Futuyma and Dawkins.

Starting with Sober, he is clearly using the phenotypic definition. In 'The Nature of Selection' he tells us 'natural selection does not explain why I have an opposable thumb. This fact falls under the purview of mechanism of inheritance.' That is reasoanable enough. It implies, in the phenotypic sense, natural selection by itself cannot achieve or explain adaptation.

So when Futuyma says (in his interview linked to the article) ' Natural selection is the process by which species adapt to their environment. A mechanism of evolution ' he seems to be using a wider definition.

No, he is saying that NatSel is PART of evolution. Besides that, we do not distinguish every detail all the time, and speaking language is not nessecarily reflecting always the underlying details how we actually deal with it. -- Kim van der Linde ^{at venus} 14:41, 23 May 2006 (UTC)

It is difficult to square his 'a process by which species adapt' with your view that 'NatSel by itself is pretty dull' — Axel147 17:59, 23 May 2006 (UTC)

If you equate NatSel with adaptation, you are right. -- Kim van der Linde ^{at venus} 01:20, 24 May 2006 (UTC)

Dawkins on the other hand is unequivocal in his use of the wider Darwinian definition. He explicitly makes the point in this quote:

'Turning from the fact of evolution to the less secure theory of its mechanism, natural selection, the mechanism that Darwin and Wallace suggested, amounts to the nonrandom survival of randomly varying hereditary characteristics. ' [1]

• As explained within evolution context, not as a mechanism by itself.-- Kim van der Linde ^{at venus} 14:41, 23 May 2006 (UTC)

And you can hear him address the exact point on heredity here.

Just to show this position is very prevalent here are 10 serious links which explicitly make reference to heritability as part of the definition:

1. University of California, Berkeley
2. Mark Ridley author of 'Evolution', Oxford zoologist
3. Paper by Robert A. Skipper Jr., Roberta L. Millstein
4. Paper by Robert Quinlan, Assistant Professor Evolutionary & Cultural Anthropology, Washington State University
5. Article by Steven Rose, professor of biology and neurobiology at the Open University and University of London
6. Article by David H. A. Fitch, Assosciate Professor New York University
7. University of Michigan website
8. Howard Whiteman summary
9. Answers.com definition
10. Dictionary.com definition

I agree that some of these positions are difficult to separate from 'evolution by natural selection'. But I think the point is that evolution is the outcome and natural selection is the full mechanism including inheritance. (As I have mentioned before a possible distinction suggested by Endler is that evolution requires new variation whereas natural selection requires only initial variety.)

I think one of the problems here is that nearly all definitions of natural selection use the word 'reproduction'. And the mechanisms for reproduction and inheritance are equivalent, though the two are conceptually different. As a result if often unclear whether an author intends the role of inheritance to be read as 1 or 2. While other authors deliberately avoid the debate by never unbundling the phrase 'evolution by natural selection'.

I think here you go seriously wrong with your train of though. The mechanisms are not only conceptually different. Reproduction is a way of passing the inhertitable information on to the next generation. Dividing the cell, as bacteria do is another way of passing it on (basically a variant of reproduction, but without sex). Other ways of passing this on what we would call horizontal gene transfer, which happens a lot in plants and is caused by virusses transported by insects. Furthermore, if a trait is not heritable (See for example the article abiout DES and reproductive issues related to usage of the hormone during pregnacies DES), offspring is often seriously maladapted to reproduce. However, when they reproduce, they do not pass this on to he next generation because it is not heritable, but in the mean while, natural selection has eliminated part of the DES population because they are incapable to reproduce. In this example, non-heritable variation, NatSel and reproduction come together, and as you can see, all three have their own place in the story and are not equivalent at places. -- Kim van der Linde ^{at venus} 14:41, 23 May 2006 (UTC)

This is not the crux of the argument. I am just trying to give a bit of background as to the origin of the differences. But in any case I think you are missing the point here: reproduction always results in inheritance of some of the characteristics. (Even if some characteristics can transfer by another mechanism this does not invalidate the argument as fitness is defined with respect to reproduction) — Axel147 18:14, 23 May 2006 (UTC)

Sorry, reproduction does not result in inheritance. -- Kim van der Linde ^{at venus} 01:20, 24 May 2006 (UTC)

Oh yes it does. In case you are not aware inheritance is 'the process of genetic transmission of characteristics from parents to offspring'. It is not possible to have an offspring with no relationship to a parent. — Axel147 07:12, 24 May 2006 (UTC)

Another is that natural selection is often used as a kind of short or long hand. Darwin's 'preservation of favourable variations' is succinct enough but Darwin might be guilty of this himself in other parts of his text. In the evolutionary narrative, whether directly or indirectly, it is both individuals and characteristics that end up being selected.

Those who prefer the phenotypic view seem at pains to separate selection from its consequences: to emphasise its 'blindness'. Those who take wider view seem equally at pains to identify natural selection with the full idea: not just the existence of differential reproduction but the observation that this results in a change in trait frequencies. Many people do seem on the fence (either accidentally or purposefully). But the end point of 'evolution by natural selection' remains the same, so does the understanding and so does the mechanism. And in this sense this is an empty debate.

'We must not lose sight of the distinction between natural selection and evolution by natural selection. There may be selection for a trait without that trait's increasing frquency. That will happen if the trait has zero heritability.' Sober.

True, but nor should we lose sight of the difference in the uses of the term natural selection.

With respect to this article I think we can do a little more to make the distinction clearer. It could even be said that in preferring the phenotypic definition but giving us an example of 'evolution by natural selection' this article is adding to the confusion!

(Please add comments at the end) — Axel147 08:29, 23 May 2006 (UTC)

As I said, NatSel by itself is pretty dull; effectivly: you take a population, apply some mechanism of eliminating the maladapted and you have a selected population. That is all. Then there is a lot of mechanistic stuff we can talk about. The main problem is that NatSel is best understood by its consequences, and that is why the example includes the effect over time. I would be perfectly fine with eliminating that, but I think that it become less readable, and less understandable. I think it is not wise to spend to much time on digging in deeper on the definition issue. This is promarily an academic issue, an important one, but not the main thing of the article. I rather focus on the rst of the text and get that as clear as possible to explain the details of mechanisms of NatSel.

You have to keep in mind, that the way people talk about NatSel is in the general way, as in Evolution by NatSel. It is an easy shorthand. It is when you get to the details when it start to differentiate. For that reason, EvolNatSel will win the popularity contest with NatSel. Try to step back from your own preferred views, and look at the wider context. Look at what is actually the aim of Wikipedia, and loo at the other articles that are out here. The aim is to bring the state of the art, not what popularity polls suggest. Cut through the normal speak and see what is there underlying at the science basis. -- Kim van der Linde ^{at venus} 14:41, 23 May 2006 (UTC)

There is nothing particularly state-of-the-art about the phenotypic definition. Why don't you take a step back from your own preferred views. I have tried to present both positions in a balanced way. Both definitons give the same understanding. By saying 'NatSel' is this or that and persistnetly using the phenotypic selection which you have accepted as being different from Darwin, Dawkins, Endler etc. I think you're the one being dogmatic. — Axel147 17:41, 23 May 2006 (UTC)

I agree with Kim, here. Selection without heritability is not worth discussing. Yes, there's "selection" against "individuals who smoke", "individuals who ate lead paint", "individuals who got hit on the head with a really big rock". But, without heritability it's simply not interesting. And there's no way to discuss fitness without heritability, so I think we should ignore the issue entirely and stick to heritable traits.

Also, I note that we have no discussion of selection coefficients, which we ought to very early on, methinks. Graft 16:38, 23 May 2006 (UTC)

Well, I find the mechanisms themselves also interesting, but I agree with you, most of the realinteresting stuff is about the effects of natural selection on the underlying genetics. And maybe we have to split the current mechanism section into two, of which one is about the signature of NatSel on the genetics. I will play around with that a bit. I disagree with you about selection coefficient at the beginning, but I will play around with them and see where they make sense. -- Kim van der Linde ^{at venus} 01:20, 24 May 2006 (UTC)

I think you are more or less agreeing with me. But in case it's not clear in the wider definition 'selection' is not the same as 'natural selection'. Kim and I agree it's important to separate selection from its results. We also agree that if there was no inheritance whatsoever the whole thing would be a waste of time. And we also both agree there is more than one definition of natural selection. We just disagree on which should take the lead. — Axel147 17:41, 23 May 2006 (UTC)

Okay, now I'm not sure who I agree with, because this debate is kind of rambling. Can you concisely state what the issue is? I'm wholly unaware of an important distinction between 'selection' and 'natural selection' (excepting the fact that the former might also include artificial selection). Graft 19:51, 23 May 2006 (UTC)

The debate put simply is whether natural selection is this or selection due to fitness differences (differential reproduction). — Axel147 21:22, 23 May 2006 (UTC)

Well, I'm assuming you don't give a shit about variation, since that's merely a presupposition; selection is predicated on the existence of variation, so in its absence it obviously can't occur. So your beef is only with including heredity in there, i.e., does selection occur in the absence of heredity? I think this distinction amounts to, at best, a parenthetical, and I don't we shouldn't waste time with abstruse questions of usage. You're not going to achieve a consensus on this based on vernacular or even scientific usage, because I don't think there is any such consensus. Perhaps selection only represents differential reproduction (as in, attempting to select for traits with zero heritability), or perhaps it represents the outcome of differential reproduction. That is, selection has occured when something is selected for. I think you'll find this varies depending on circumstance. Does this really strike you as a major and important distinction to be made? And if so, why? Graft 22:41, 23 May 2006 (UTC)

Graft, what you see as Natural selection is only one of the definitions, and I would say that the one you use is the genetical theory of natural selection. There are good arguments for other definitions and they are also widely used (see for example Futuyma's book, or the ground breaking article of Lande and Arnold). What I have tried is to write it such that it avoids as much as possible those definitional questions.-- Kim van der Linde ^{at venus} 01:13, 24 May 2006 (UTC)