Steven Pinker Against Group Selection Or Against Evolution?

Sascha Vongehr

A new edge-essay by Steven Pinker is bound to lead to vehement reactions: The False Allure of Group Selection. It is worth a read – Pinker is a clear writer and so his position is easy to locate, however, I get the feeling that his position is to smooth talk whatever a certain establishment likes to hear being defended. The last time I listened, he told modern society that it is the most peaceful ever (an amazing feat of cherry picking data and re-interpretation). Now it seems he simply roots for the more well established guys in a heated turf battle: who may talk about evolution.

There is a lot I could nitpick, for example that he mentions Lee Smolin’s cosmological natural selection (CNS) of universes although one of Pinker’s most important criteria for natural selection is that success is strictly the number of copies in a finite population. There is no such finite number or even any good measure, that is precisely one of the two main problems with CNS. In other words: Don’t support your stuff with other stuff that you do not understand – it always backfires.

But let us leave the nitpicking and go straight to the core. This is basically his whole essay boiled down to fit into a nutshell:

Now, no one "owns" the concept of natural selection, nor can anyone police the use of the term. … But unless the traits arose from multiple iterations of copying of random errors in a finite pool of replicators, the theory of natural selection adds nothing to ordinary cause and effect.

He wants to police the use of “natural selection” to exclude anything that so called “multilevel selection” or similar more advanced concepts can draw on, so that in the end we are bound to yesterday's status quo of evolution: Naïve gene centrism. He fails via two main mistakes:

One is to think that modern design for example, he mentions touch-tone phones, is not largely very random. Note well, it is Pinker himself who defines ‘"random"in the sense that they do not anticipate their effects in the current environment’, which is a very good definition. Success, especially defined in the way he does, namely only counting how many of the same systems there happen to be, such success of technological advancements is mostly random, so is the design phase itself. Very few successful designs got successful the way the designers thought it should play out when designing.

The second and much graver mistake: He seems to be unaware of that already on the level of bacteria, evolution is not so much driven by random errors as by there being already evolved mechanisms in place that modify the genome in quite deliberate ways in targeted areas, mechanisms that we only just start to grasp. The mutation rate itself for example is adjusted as a response to stress. James Shapiro writes some interesting articles on that topic.

Stevens second error is deadly, as it not just inexcusable for a researcher who claims to be and stay informed about the cutting edge research in traditionally, narrowly defined evolution as biological evolution, but if we were to go along with what Steven Pinker writes about what is “natural selection” and especially what is not, almost the whole of biological evolution would no longer fall under the heading of natural selection!

Evolution is not just biological evolution, where stuff has to divide and die and all of that, and evolution is definitively not just the mere evolution of molecules, where nothing yet developed that tames the utter randomness of chemistry. Evolution is emergence via natural selection; something natural is there and it selects. Arguing what is natural, explicitly or implicitly, may defend your turf, but it is not providing any novel, useful distinction.

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Syntax. Sure. To help would be like creating another person, so I'm not offering. Most everything humans do is pretty weird to me, but I've got a good thing going so I'm not going to pretend I can help.

Here's the thing. Have you ever wondered why when people ~ professionals even ~ install a screen door, half of the time they install it with the swing the opposite of the main door? Somewhere, some when some tailless idiot thought that mismatching the hinges would be best, and what do you know, half of the cattle repeat the error, surprise surprise. Those mismatched door swings are a clear case when “traits arose from multiple iterations of copying of random errors in a finite pool of replicator.” It ain't good old cause and effect. It could be “already evolved mechanisms in place that modify the [doors wings] in quite deliberate ways in targeted areas.” Blame it on a union or the blind leading the blind. Its syntax is set in stone and there is no way you are going to be able to “help” except by watering it down and propagating your own syntax.

blue- green | 06/19/12 | 22:09 PM

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I couldn't get through the paper. Too much of it was the normal rubbish about gene replication without regard for the reality of biology. To his specific point, I am always disappointed when the insistence on treating group selection as merely a variation of individual selection.

Gerhard Adam | 06/19/12 | 22:08 PM

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I am always disappointed when the insistence on treating group selection as merely a variation of individual selection

Yes, this is the core issue once more in different words. It is like if I tell a biochemist to not use the term "replication" since it just a variation of quantum electro dynamics.

Sascha Vongehr | 06/19/12 | 22:36 PM

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From Pinker's article:

Granted, it's often convenient to speak about selection at the level of individuals, because it's the fate of individuals (and their kin) in the world of cause and effect which determines the fate of their genes. Nonetheless, it's the genes themselves that are replicated over generations and are thus the targets of selection and the ultimate beneficiaries of adaptations.

To illustrate the absurdity of this let's examine it from the perspective of a successful piece of music that was produced by a musician/artist.

Granted, it's often convenient to speak about selection at the level of songs, because it's the popularity of songs (and their performances) in the world of cause and effect which determines the fate of their notes. Nonetheless, it's the notes themselves that are replicated over generations and are thus the targets of selection and the ultimate beneficiaries of new recordings.

...and thus we have an explanation of why neither musicians or songs are important, but rather how notes are the "controllers" of all music and how they seek to replicate themselves in future songs.

... and thus is bullshit born.

Gerhard Adam | 06/20/12 | 20:45 PM

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To illustrate the absurdity of this

You might just as well say

'Why is a raven like a writing-desk?'
'Come, we shall have some fun now!' thought Alice. 'I'm glad they've begun asking riddles.—I believe I can guess that,' she added aloud.
'Do you mean that you think you can find out the answer to it?' said the March Hare.
'Exactly so,' said Alice.
'Then you should say what you mean,' the March Hare went on.
'I do,' Alice hastily replied; 'at least—at least I mean what I say—that's the same thing, you know.'
'Not the same thing a bit!' said the Hatter. 'You might just as well say that "I see what I eat" is the same thing as "I eat what I see"!'
'You might just as well say,' added the March Hare, 'that "I like what I get" is the same thing as "I get what I like"!'
'You might just as well say,' added the Dormouse, who seemed to be talking in his sleep, 'that "I breathe when I sleep" is the same thing as "I sleep when I breathe"!'
'It is the same thing with you,' said the Hatter, and here the conversation dropped, and the party sat silent for a minute, while Alice thought over all she could remember about ravens and writing-desks, which wasn't much.
'Have you guessed the riddle yet?' the Hatter said, turning to Alice again.
'No, I give it up,' Alice replied: 'what's the answer?'
'I haven't the slightest idea,' said the Hatter.
'Nor I,' said the March Hare.

You might just as well say

let's examine it from the perspective of a successful piece of music that was produced by a musician/artist.
Granted, it's often convenient to speak about selection at the level of songs, because it's the popularity of songs (and their performances) in the world of cause and effect which determines the fate of their notes. Nonetheless, it's the notes themselves that are replicated over generations and are thus the targets of selection and the ultimate beneficiaries of new recordings.
and thus we have an explanation of why neither musicians or songs are important, but rather how notes are the "controllers" of all music and how they seek to replicate themselves in future songs.

You might just as well say

... and thus is bullshit born.

derek_potter (not verified) | 06/29/12 | 16:04 PM

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Great comment reaper reminder, yellow card: This is a science column! Make sense or be deleted.

Sascha Vongehr | 06/29/12 | 20:42 PM

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Beautiful, Gerhard. So, gene-centrism is crap. You've summed it up nicely!

Steve Davis | 06/22/12 | 01:52 AM

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No - gene centrism is not "crap"! Gene centrism, much like Einstein-micro causality in physics or reductionism generally, is necessary for muddle heads like you to stay focussed on science! However, dogmatically refusing any emergent higher order description/terminology is the problem.

Sascha Vongehr | 06/29/12 | 20:50 PM

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Excellent!
Now justify it!

Steve Davis | 06/29/12 | 20:56 PM

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Can you clarify a bit more your position then? Many people seem to think that the theory itself is either completely flawed or never applies in the real world. My impression is that you think it has explanatory power in some situations but is over-represented to apply to everything, causing people to miss or deny higher order phenomena when they shouldn't.

Where do you stand on this, http://faculty.washington.edu/beecher/Nowak%20etal%20-%20evolution%20of%20eusociality%20-%20Nature%202010.pdf
and the response
http://whyevolutionistrue.wordpress.com/2010/08/30/a-misguided-attack-on-kin-selection/

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Thor Russell

Thor Russell | 06/30/12 | 22:44 PM

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Not sure how to clarify my position any further than I did in the comments and the article. If the physics analogy does not work for you, I do not know what will. Perhaps the following helps:

Always keep in mind that hype and fighting emerges from people trying to get ahead in the publish-or-perish game! For example the gradualism versus punctuated equilibrium issue, which was the previous big fight in bio-evolution, is just about two descriptions that use different time resolutions and thus slightly different, not even mutually exclusive, terminologies. Nobody would have bothered to defend one over the other (instead of just mentioning the time resolution under discussion) if it had not been about fame/money/academic positions/pride/ ...

Sascha Vongehr | 06/30/12 | 23:05 PM

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Your comments don't clarify that well because you appear to disagree with Steve, but not Gerhard when they both think gene centrism is completely wrong. Gerhard seems to think every biologist who gives even small credit to gene centrism is making a very basic mistake. However I think the gene centrism position has some value, do you?
It sure seems to me that the issue is ridiculously polarized, but I am not sure that you can blame pop science in this particular case given just as strong opinions exist in the general public who are not in the field. I think that politics may be involved and knee jerk reactions to the words "selfish gene". I wonder if people would have different opinions if it had been called "cooperative gene" instead.

I think gene centrism and group selection mechanisms are both correct, and useful in some situations although obviously gene centrism cannot explain everything and will give the wrong prediction in some situations. However I do not see why cooperation cannot potentially emerge as a result of different mechanisms, gene centrism, game theory etc included. Establishing what happened in a specific situation seems to be genuinely difficult and strong opinions for one or the other seem unjustified.

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Thor Russell

Thor Russell | 07/01/12 | 00:05 AM

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Gerhard seems to think every biologist who gives even small credit to gene centrism is making a very basic mistake.

I did not have time to follow the comment battle closely. To describe something centered on a certain concept (e.g. the gene as defined information theoretically) is good methodology as long as you stay aware of this being a concept etc.

However I think the gene centrism position has some value, do you?

Of course, just not to the effect that it is the last concept that may be suggested to be useful in the emergence of complex systems coevolving.

It sure seems to me that the issue is ridiculously polarized, but I am not sure that you can blame pop science in this particular case

I give you that in this case there is also the defending of good science against the religious that pushed the "selfish" gene down the track of being surounded by a trigger happy defense that will immediately bite into anything that slightly seems as if it could potentially be edging towards a slippery slope. Same for Einstein's relativity and so on - all surounded by "defenders" who misunderstand what they defend, and so use wrong arguments, ... . However, without POP, A) researchers would be much less selected to use such (any convenient) truncheons in order to stick it to their competitors, and B) they would not have so much pressure to hype their own stuff.

Sascha Vongehr | 07/01/12 | 03:14 AM

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These are good questions Thor.
The main point to make is the distinction between gene centrism and gene selection.
Gene selection may have some relevance but that will be limited because genes are invisible to selective pressures.
Gene centrism on the other hand, has no credibility because it is a world-view, an ideology that allows for no development of the concept as new evidence comes to hand.

Steve Davis | 07/01/12 | 03:17 AM

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"Gene centrism" as denoting an extreme world view rather than a deliberate "centering" on the gene concept (while being aware of this being in-effective methodology when describing higher order emergent biological structures) - I understand your point. New atheists may hold such necessary in the endfight against the religious. That may be why PZ Meyers supports Pinker.

Sascha Vongehr | 07/01/12 | 04:06 AM

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However I think the gene centrism position has some value...

What value do you suppose it has? I don't see discussions about "let's imagine selection from the gene's point of view". Instead the arguments are solely about how the genes influence outcomes [which they can't]. Selection operates directly on the phenotype and indirectly on the genotype.

If gene-centrism were true, then it would undercut all of evolution and natural selection, since the majority of genes are already conserved across species. Therefore there is no reason to suggest that a gene cares whether it is expressed in a bacteria or in a homo sapiens. Similarly for multi-celled organisms. The most reliable method of ensuring genetic integrity for replication is the single-celled asexual organism. Sexual reproduction already cuts the genetic odds for propagation in half.

Again, it produces all manner of other problems because it creates the illusion that genes are somehow "codes" that provide a direct result for selection to operate on. It completely ignores regulation and it has nothing to say for genes that never get expressed.

However, the ultimate negative aspect of it, is that when the conversation comes down to trying to explain cooperation and altruism because "genes are selfish". Then it illustrates just how silly and irrelevant that perspective has become. Then we see all the goofy definitions of what altruism is, by denying it if it doesn't actually incur a fitness cost but nevertheless using it as an example of altruism [i.e. bird giving warning]. In general, it's poorly thought out, because it is perpetually being fit in to the concept of selfishness as being the central governing element of "survival of the fittest".

Gerhard Adam | 07/01/12 | 08:04 AM

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I don't find your arguments convincing and I doubt either of us will learn anything from me trying to convince you it has value. Sascha also thinks it has so if you can understand why he does that would probably be more productive.

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Thor Russell

Thor Russell | 07/02/12 | 18:11 PM

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It seems that many of these articles are coming out in force to blast Edward O. Wilson's latest book and his continuing claim supporting a form of "group selection".

I always like how they make up some bogus example of group selection and then argue against it in a classic strawman argument. I have a simple example of group selection .... look in the mirror. When someone can explain how 10 trillion cells could assemble, with each "voluntarily" giving up its right to reproduce, so that the role could be relegated to the germ line after which only 50% of the genes will be passed on to future generations. Yet, because this is classified as a multi-cellular organism, the explanations are given as if this is a separate and distinct realm of biology.

Here's another example of some rather strange reasoning:

The core of natural selection is that when replicators arise and make copies of themselves, (1) their numbers will tend, under ideal conditions, to increase exponentially; (2) they will necessarily compete for finite resources; (3) some will undergo random copying errors ("random" in the sense that they do not anticipate their effects in the current environment); and (4) whichever copying errors happen to increase the rate of replication will accumulate in a lineage and predominate in the population.
http://edge.org/conversation/the-false-allure-of-group-selection

We already know #1 isn't true, although this is being hedged by the phrase "under ideal conditions". Number 2 isn't true, except within the sense that the earth is finite, although they MAY end up competing if their population growth is unconstrained. Number 3 doesn't even make sense, because while there are certainly errors, it would be hard to argue that something like Horizontal Gene Transfer (HGT) is not performed in "anticipation of effects" in the current or future environments. Number 4 is just silly, since it argues that the genes that will dominate are those that increase the rate of replication. Again, this is something that simply doesn't happen for the majority of animals.

Gerhard Adam | 06/22/12 | 03:20 AM

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I'm still pretty agnostic about the idea of group selection, since I have not dived in and read Wilson's book and don't know what his exact model is. I do find the idea of multiple levels of selection, as opposed to the purely gene-centered view, quite intriguing, though.

And unfortunately, my undergraduate education in evolutionary biology (mid 90s, Botany major) was a bit dogmatic, as I was told in no uncertain terms that group selection was "a myth", and that genetically encoded, non-kin altruistic behavior did not hold up from an evolutionary point of view, since the altruist was sacrificing its own fitness and therefore the predisposition to altruism would be selected against. That there had ever been a historical debate on the idea of levels of selection itself was not something I was introduced to, nor lesser known/newly discovered evolutionary mechanisms and theories like horizontal gene transfer, epigenetics, and evo-devo. I'm not sure if undergraduate education for biology majors has improved since the mid-90s, or if students are taught the straight neo-Darwinian model and not much else.

Iamcuriousblue (not verified) | 06/24/12 | 23:25 PM

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Yes, those are all part of the modern mythology. Of course, originally group selection was untenable, because it didn't make sense to consider that natural selection would [or could] determine what traits were beneficial from a group perspective. That would be even more problematic when the animals involved didn't necessarily live in groups.

However, what everyone failed to notice is that the gene-centric view is simply another type of group selection. Except that in this case, now we supposedly have the gene evolving [or being selected] for the good of the whole organism [i.e. 10 trillions cells in a human].

Again, this is just a mishmash if perspectives which is completely meaningless, since even the definition of a gene isn't consistent from Dawkins. In my view, most of biology is still suffering from physics-envy; the desire to have hard mathematical laws. As a result, most biologists immediately fall in love with equations, regardless of whether they represent anything in the real world. That isn't to say that there isn't some benefit from them, but anyone that has spent any time looking at life, would recognize that for every "law" there is likely, not just one, but multiple exceptions.

However, one of the main problems is in the area of fitness. There is this mistaken belief that animals just perpetually reproduce, which clearly isn't true. Interestingly enough, humans are one of the few animals that could reproduce more readily, and despite our huge population growth, we give birth very infrequently compared to the opportunities. As a result, it leaves gaping holes in concepts like Hamilton's rule, etc. Even Haldane's famous quote about sacrificing himself for two brothers or eight cousins, etc. is foolishness, since it depends on a nearly impossible set of probabilities to manifest itself.

I don't think there can be any doubt that the individual influences the group, while the group influences the individual, since essentially the group IS the "environment". So, like all environments, it will exert selection pressures on the individuals involved.

Gerhard Adam | 06/24/12 | 23:58 PM

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Granted, it's often convenient to speak about selection at the level of individuals, because it's the fate of individuals (and their kin) in the world of cause and effect which determines the fate of their genes.

I was surprised to learn that bacteria can undergo programmed cell death until I realised that this has a very favourable effect on survival. Bacteria are often attacked by bateriophages which can replicate very quickly. By instituting programmed cell death the single cells help prevent the spread of the pathogen because unlike necrotic cell death,which happens when the virus continues replicating, in apoptosis the cellular contents tend to be more contained within the cell wall and hence the viral particles are less likely to infect other bacteria in the colony. The individual genes sacrifice themselves because the colony benefits as a whole.

And of course, quorum sensing .... .

This observation suggests that PCD in bacteria serves to eliminate damaged cells, similar to apoptosis of defective cells in metazoa.

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Apoptosis-like elimination of defective cells in S. cerevisiae and protozoa suggests that all unicellular life forms evolved altruistic programmed death that serves a variety of useful functions.

John Hasenkam | 06/25/12 | 00:05 AM

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... and bacterial cannibalism
http://stke.sciencemag.org/cgi/content/abstract/sigtrans;2006/323/tw67

...and forming biofilms during periods of stress.

In my view groups are a natural consequence of organisms living in the same "geographic" area. It wouldn't make any sense to consider natural selection as operating only an individuals while ignoring those of the same species that surround you? This is even more obvious when an animal is dependent on others for finding mates.

There is no reason to believe that nature hasn't discovered one of the fundamental principles we employ in business; "economies of scale". It generally takes far less effort and energy to work with someone than to work against them. As a result, we tend to see groups forming in any situation where there are economies of scale. Of course, one of the primary exceptions would be in the case of large predators, and that's precisely what we see [i.e. a tendency to live isolated lives since there is no "energy" benefit from cooperation].

Gerhard Adam | 06/25/12 | 00:18 AM

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Gerhard -

The problem with your argument is that you seem to be confusing the evolution of cooperation, which can happen for all via kinds of perfectly neo-Darwinian routes, with group selection, which requires that individuals sacrifice individual fitness for group fitness and that this is transmitted in some way.

Iamcuriousblue (not verified) | 06/25/12 | 02:10 AM

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Not at all. Sorry if I wasn't more specific. The idea of suggesting that the "group" is the environment which creates a selection pressure makes that point.

For example, being a member of a wolf pack conveys advantages that a lone hunter wouldn't have. Therefore, it is advantageous to sacrifice the ability to reproduce for the benefit of surviving [with the idea being that you will survive long enough to reproduce].

From this we can reason that wolves that didn't take to the group, and became lone hunters likely (1) didn't do as well as groups and/or (2) had difficulty finding mates. These creates two strong selection pressures for wolves to become more socialized. Even if the inclination is to hunt alone, there is a pressure to associate with groups to find a mate. On the other hand, in grizzly bears we find that there is little "energy" benefit in being cooperative [for males or females]. So we find that these animals live a more isolated lifestyle. This creates a different kind of selection pressure, because bears that wanted to congregate together would have a more difficulty time finding sufficient resources for the group. Therefore, such groups would tend to be selected against. In fact, a good example of that is during salmon season in Alaska, where there is no difficulty in having grizzly bears congregate in a quasi-social fashion. With their "energy" needs satisfied, there is little impetus to engage in conflicts.

The point is that cooperation is a requirement for group behaviors, and that belonging to a group or being denied access to a group all create selection pressures that determine what the traits are of future generations. At some point, an animal may become so integrated into the group that it is literally impossible for it to survive individually. Such animals are unequivocally dependent on group selection. In the eusocial insects we see that the ability to reproduce is eliminated. In a eusocial mammal like humans, we still have the individual ability to reproduce, but it wholly dependent on the group. A human without group attachments has no biological future.

Gerhard Adam | 06/25/12 | 06:27 AM

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1) Pinker said that the mutations are random and citing the fact that the rate of mutation can be altered by the environement doesn't change that. The mutations are still random, not purposeful.
2) Evolution by natural selection is most definitely biological evolution. If you're referring to something else, then that is an entirely different topic. The group selectionists he is responding to were addressing biological evolution.
3) Your point about inventors not foreseeing what will come of their inventions has no affect on the argument it is addressing, and you are unwittingly (I hope) arguing that telephones and buildings ARE examples of something on which natural selection acts. If this is the case, then that is a fundamental misunderstanding of the basic idea of evolution by natural selection.
4) You still fail to show how group selection adds anything useful to the theory of natural selection. You've nitpicked a few phrases used without addressing the overarching thesis, which is a complete failure to rebut the article.

A Tyler (not verified) | 06/28/12 | 10:36 AM

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Pinker said that the mutations are random and citing the fact that the rate of mutation can be altered by the environment doesn't change that. The mutations are still random, not purposeful.

While this is often a point people like to quibble about, it is the latter part of this statement that I would also consider. For some reason, in discussing evolution there is some strange sense that "randomness" is important, despite the fact that it truly isn't random; it is only unpredictable [for the most part]. Similarly there's this quaint notion that everything depends on these "random" errors introduced either through cosmic rays or copying mistakes, etc.

While those are certainly elements that will affect heredity, it's also complete BULLSHIT! The majority of the "bush of life" consists of single-celled organisms, not the large multi-celled organisms that most of these ideas are formulated about. One of the primary traits that these single celled organisms have is the ability for Horizontal Gene Transfer (HGT).

Again, this is often mentioned and talked about as if it's some interesting little factoid about something insignificant like bacteria. However, since such life constitutes the majority of life on this planet, that it is fair to say that it represents the primary biological means by which genetic information is exchanged. Now, for the tricky bit.

HGT is intentional. There's nothing random about it. We're not talking about the exchange of a few molecules. We're talking about the exchange of genes. Completely functional units of hereditary materials. In other words ... while natural selection and evolution certainly occurs and is influenced by accidents, mistakes and the introduction of "random"/unpredictable changes, a significant majority of life operates with a much more directed approach to the exchange of genetic information.

In case anyone has missed the implications of this, it means that the largest component of life on this planet follows almost none of the "rules" we are fond of utilizing to explain it. Even the concept of a species doesn't really exist at this level.

Gerhard Adam | 06/28/12 | 16:35 PM

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"You still fail to show how group selection adds anything useful to the theory of natural selection."
No, the gene-centrics still fail to show that group selection does not occur.
Most people who have read The Selfish Gene think that Dawkins demolished group selection.
He did nothing of the sort.
If you read the passages on group selection carefully you will see that he admitted that he was merely presenting an alternative view to group selection.
It's only dogma that prevents acceptance of multi-level selection.

Steve Davis | 06/29/12 | 02:23 AM

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"No, the gene-centrics still fail to show that group selection does not occur."

You should probably avoid arguing about science until you become familiar with the burden of proof. That's a terrible, completely meaningless argument. It is for group selectionists to show why group selection has explanatory power that eludes the gene-centric explanation, which they have not done whatsoever.

J Taylor (not verified) | 06/29/12 | 09:58 AM

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There is no explanatory power in presuming that genes are the units of replication when:

(1) there is no specific definition of a gene that actually represents it as a "unit of selection".

(2) have a unit of selection that is powerless to ensure it is selected [i.e. expressed]. In other words, what is the basis by which a gene is "selected" for future generations? Invariably this gets muddled by looking at actual traits, however you can't ascribe specific traits to a gene alone because that isn't how they are produced. So it does no good to argue that the genes are "selfish" and looking to replicate themselves into the future, and then claim this is achieved by the genes acting as a group to do so.

(3) have a unit of selection that cannot consistently produce what it is supposed to be selected for [i.e. conservation of the same genes in different roles in different species]. The point here is that even genes that are identical cannot be predicted as to what trait they will specifically express without understanding their context. In other words, the "coding" is insufficient. Consider Darwin's finches again, in examining the beaks, we have different expressions all brought about by the same genes. This clearly indicates that it is the gene expression [i.e. trait] that is the unit of selection and not the gene itself. The gene merely carries the information that was necessary to produce the trait. This is where most of the confusion comes from, because there is obviously information necessary. However, by loose analogy, if a chef creates a new recipe and writes it down, which is the entity being selected? The food being produced [i.e. the trait], or the paper containing the recipe? Obviously the latter holds the information so that it can be reproduced, but it is the food itself that is being selected.

(4) Perhaps the most telling problem is that for sexually reproducing animals, the gene is insufficient to produce an organism [i.e. it must be coupled with those from another member of the species]. In the second place a fully expressed set of genes is still insufficient to produce a living organism [i.e. dependence on the microbiome]. This is probably one of the more significant demonstrates of group selection, where none of the organisms in question can actually survive on their own [although the microbiome probably has a better chance].

The second problem with that is that in the places where such reproduction is asexual, there is no meaningful definition to the concept of a species and we see indiscriminate exchange of genetic material promoting diversity. Again, the selection of gene transfer is occurring by the bacteria, not the gene [unless you wish to propose that the gene is instigating the transfers and acting as a kind of "command and control" center for bacterial and general cellular reproduction].
However, as stated previously, that would be a difficult argument to make since the gene can't even control it's own degree of expression, so attributing "command and control" seems unrealistic.

Gerhard Adam | 06/29/12 | 10:37 AM

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"However, since such life constitutes the majority of life on this planet, that it is fair to say that it represents the primary biological means by which genetic information is exchanged."

Group selection is not about single-celled organisms. Yes, that applies to single-celled life, but can not be used to support group selection, which is not about single-celled organisms, but about multicellular organisms that reproduce and evolve through copying errors. Bringing it up in this context, the only relevant implication could be that HGT happens in the organisms that group selection has been applied to, which is not the case.

If what you're saying is that HGT is a reason to dismiss the effect of randomness as important to vertical gene transfer, then this is invalid for the simple reason that randomness as an important component of vertical gene transfer is a scientifically supported observation. HGT is not observed in species to which group selection is applied by group selectionists.

And yes, the concept of species is definitely a fuzzy concept used for communication among humans and not a scientifically rigid or fundamental concept, not even when applied to multicellular life (there is necessary an unbroken continuum of "species" between every related branch of the tree of life).

J Taylor (not verified) | 06/29/12 | 10:12 AM

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Yes, that applies to single-celled life, but can not be used to support group selection, which is not about single-celled organisms, but about multicellular organisms that reproduce and evolve through copying errors.

So, you're arguing that "group selection" is only a minority explanation in biology? You then make "group selection" even more specific by focusing only on "copying errors".

HGT is not observed in species to which group selection is applied by group selectionists.

Based on what? Are you suggesting that a virus can never modify the genome?

It strikes me that the argument is simply a foolish one. We know that individuals get produced and that selection or loss at the individual level will affect the overall population. So this has foolishly been used to argue that only individual selection matters. However, that is like arguing that your body is depending on the success or failure only of single cell reproduction, and therefore the body doesn't matter, only the individual cells.

They are irrelevant and myopic views of what actually occurs.

There is a synergistic relationship between groups and individuals that has been ignored too long. Just like early "group selectionists" went too far in asserting that evolution occurred for the "good of the group", so have the gene-centrists gone too far the other way in claiming that groups play no role and that everything is based on selfish behavior.

If there is one serious failing in many biological discussions it is due to the ability for many biologists to engage in delusional behavior by insisting that the observed phenomenon doesn't exist in an effort to explain it in overly simplistic terms. Biology is complicated and messy, and every time someone provides a nicely wrapped up explanation, it can be pretty well assured that it is wildly inaccurate in most situations.

Gerhard Adam | 06/29/12 | 10:48 AM

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Group selection is not about single-celled organisms. Yes, that applies to single-celled life, but can not be used to support group selection, which is not about single-celled organisms, but about multicellular organisms that reproduce and evolve through copying errors.

So, who decided that HGT had nothing to do with groups? How was it decided that bacteria forming biofilms under stress has nothing to do with groups? How about quorum sensing? Perhaps bacterial cannibalism under stress?

I will say - straight up - that any biologist that doesn't consider single-celled organisms worthy of consideration in understanding groups is a fool.

Gerhard Adam | 06/29/12 | 15:34 PM

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"It's only dogma that prevents acceptance of multi-level selection."

It is the fact that multi-level selection fails to add any explanatory power to the theory of evolution. This is not dogma, it's a fundamental challenge that any scientific theory must aspire to meet. As Pinker pretty thoroughly demonstrated, none of the ideas put forth by group selectionists explain something that can't be explained by the gene-centric theory. All the explanations of group selection can be reduced to an argument based on genes, and there are always related problems that group selection can't explain, which gene-centric selection can explain (Pinker provided examples for each group selection explanation he addressed).

J Taylor (not verified) | 06/29/12 | 10:22 AM

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Gerhard Adam wrote:
"We already know #1 isn't true, although this is being hedged by the phrase 'under ideal conditions'."

No, it's completely accurate because of the phrase "under ideal conditions". That changes the meaning of the statement, and you have to take that into account before issuing a rebuttal to a strawman of your own making.

"Number 2 isn't true, except within the sense that the earth is finite, although they MAY end up competing if their population growth is unconstrained."

No, it is true. Resources are finite for any given population, end of story. You're mistaken.

"Number 3 doesn't even make sense, because while there are certainly errors, it would be hard to argue that something like Horizontal Gene Transfer (HGT) is not performed in 'anticipation of effects' in the current or future environments."

You read the statement wrong, and took a different meaning from what was said. You quoted "some will undergo random copying errors" but what you're responding to is "natural selection only works through random copying errors", which is a completely different statement--this is the definition of a strawman. It is completely accurate to say that some organisms undergo random copying errors; you're arguing with a basic, established fact in evolutionary science. This does not help group selection theory.

" Number 4 is just silly, since it argues that the genes that will dominate are those that increase the rate of replication. Again, this is something that simply doesn't happen for the majority of animals."

This is another basic element of natural selection. Those organisms that reproduce the most will become predominate. You're counter-argument is factually wrong. It does happen, and it's a basic element of evolutionary theory. It is at the core of the theory of natural selection, which I think you should be familiar with if you're going to respond to an actual scientist's writing. Pinker summarized the theory of natural selection as a preface to his arguments, and you unwittingly argued with the theory of natural selection instead of reading further and responding to his arguments. Very reactionary and not very productive.

Andrew TT (not verified) | 06/29/12 | 10:42 AM

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No, it is true. Resources are finite for any given population, end of story. You're mistaken.

So, you're arguing that populations never grow?

No, it's completely accurate because of the phrase "under ideal conditions". That changes the meaning of the statement, and you have to take that into account before issuing a rebuttal to a strawman of your own making.

Well, if you wish to be pedantic about it, but without defining the type of growth, it is a fundamentally meaningless statement other than to argue that growth will occur. However, this is contradicted by your second argument.

It is completely accurate to say that some organisms undergo random copying errors; you're arguing with a basic, established fact in evolutionary science.

Again, I have no problem with your pedantry.

Those organisms that reproduce the most will become predominate.

No, in this case you're wrong. That is only true between organisms [generally related] that compete for the same resources. There are countless examples of this occurring every time we see population blooms and die-offs. There is nothing that ensures dominance.

Gerhard Adam | 06/29/12 | 10:58 AM

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Gerhard, the fact that a single gene doesn't produce traits in isolation takes nothing away from gene-centric natural selection, and you haven't exaplained why you think it does. A gene is a sequence of DNA that is small enough to be reliably reproduced. The fact that some organisms reproduce sexually does not support group selection, and it is not incompatible with gene selection. You haven't explained how you're making that connection. You're stating facts and then stating that they disprove gene selection and support group selection, but not once have you explained how or why. Of course sexual reproduction is compatible with gene-centric natural selection; are you honestly sitting at your computer, thinking that the majority of evolutionary scientists are unaware of this? Only you and other group selectionists know that humans reproduce sexually? This makes no sense. You need to relate the facts you're stating to the theory you're disputing, and saying "this disproves it" is absolutely meaningless without making that connection. It's exactly as valid as me saying, "Some organisms reproduce sexually. This is very damning for group selection. Genes are not reliably copied. This disproved group selection." It's meaningless.

Andrew TT (not verified) | 06/29/12 | 10:54 AM

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A gene is a sequence of DNA that is small enough to be reliably reproduced.

That is a garbage definition and you know it.

...the fact that a single gene doesn't produce traits in isolation takes nothing away from gene-centric natural selection, and you haven't explained why you think it does.

If the gene is the "unit of selection", then the proposed arguments have always maintained that element of "selfishness" which is attributed to competition. This makes individual genes competitive with one another to be selected for greater representation in the future. However, it becomes useful to examine what "representation in the future" means, if the gene is powerless to actually achieve that objective. If it is then argued that multiple genes [in cooperation] do this by producing a trait which is selected [which is the only means by which genes CAN be selected since everything else would be a blind lottery], then the entire framework of "selfishness" or selection of genes individually becomes incorrect.

The problem here is that "selfish gene" attributed so much ability to genes to direct their own future, that organisms simply became lumbering robots destined solely to fulfill their objectives.

Therefore, every point I'm making is specifically illustrating that genes lack such a power and ability, and that the entire theory is premised on faulty analogies, and wishful thinking. Let me be quite specific.

Can you find a single example of where gene "behavior" has ever been selfish [not self-interested] that hasn't been detrimental to an organism, then there might be a basis for the theory. However, as stated, it is complete bullshit and has lead to more misunderstandings than helpful perspectives.

...are you honestly sitting at your computer, thinking that the majority of evolutionary scientists are unaware of this?

Oh, they're aware, and for decades haven't questioned the obvious, namely how does one propose "selfishness" when it is clear that reproduction depends on cooperation? Instead we got idiotic explanations of how selfishness is cooperation [depending on how one twists the perspectives]. So up became down just to satisfy a dumb metaphor.

It should also be intuitively obvious that "selfishness" is far too energy intensive of an activity to ever represent an actual trait in genes or anything else. However, when this is pointed out, then suddenly the "selfish gene" simply becomes a metaphor and wasn't really intended to be used in that way. So, the real problem is that the entire "selfish gene" theory has been a slippery concept that has never been properly applied and gene-centricism has attempted to attribute all manner of capabilities to genes, when a careful examination illustrates that they are merely the purveyors of information and have no intrinsic abilities to ensure that the information with be either (1) interpreted properly, (2) used at all, or (3) how it will be regulated. In short, they have NO abilities at all, any more than books possess abilities.

Gerhard Adam | 06/29/12 | 11:11 AM

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A gene is a sequence of DNA that is small enough to be reliably reproduced.

Much smaller parts of a gene---all the way to a nucleotide base---are reliably reproduced. Isn't a gene usually a part of DNA that codes for a protein?

Enrico Uva | 06/29/12 | 12:56 PM

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The gene is a union of genomic sequences encoding a coherent set of potentially overlapping functional products.
http://genome.cshlp.org/content/17/6/669.full

This is being suggested based on these assumptions:

There are three aspects to the definition that we will list below, before providing the succinct definition:

A gene is a genomic sequence (DNA or RNA) directly encoding functional product molecules, either RNA or protein.
In the case that there are several functional products sharing overlapping regions, one takes the union of all overlapping genomic sequences coding for them.
This union must be coherent—i.e., done separately for final protein and RNA products—but does not require that all products necessarily share a common subsequence.

Gerhard Adam | 06/29/12 | 13:09 PM

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"So, you're arguing that populations never grow?"

No, I'm not. You're really bad at strawman arguments. That does not follow from the statement you quoted.

"No, in this case you're wrong. That is only true between organisms [generally related] that compete for the same resources. "

Yes, we're talking about organisms that compete for the same resources, a given population of a single species. You're right of course, but again, the context in which you discuss natural selection should have made this pretty clear. If not, then add "a given population of a single species" and you understand.

Finally, a tendency to become predominate is not a claim that dominance is ensured. It only means that replicating the fastest will produce this tendency, all other factors being equal.

There is an understood use of shorthand in discussing evolution, so that every single sentence doesn't have to contain "a given population of a single species" and "all other factors being equal", but I apologize for not making these distinctions clear. Were I writing formally, I would have done so. I appreciate your enthusiasm and I'll try to return to this after work if you're still interested.

Andrew TT (not verified) | 06/29/12 | 11:30 AM

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That does not follow from the statement you quoted.

Of course it follows. If resources are finite, then you're arguing that population is limited by competition for those finite resources. However, this also creates the problem that population can't grow larger than those resources, otherwise the resources aren't actually finite [with respect to the population in question]. Certainly they are "finite" within the limitations of animal movement/migration and that they represent a bounded existence. If the set of resources is greater than the population dependent on those resources, then there is no fundamental competitive limit to the acquisition of those resources. In that sense, they are not "finite". [NOTE: the term "finite" is generally used to indicate a triggering point at which competition begins to have survival consequences].

The explicit requirement of population growth, reproduction, etc. is that there are always sufficient resources available to support the population [unless we're specifically addressing periods of stress]. In fact we specifically find seasonal variations that have produced migratory adaptations to deal with those situations.

So unless you're using the word "finite" in the trivial sense, it simply isn't true if the intent is to demonstrate competitive pressures. All one has to do is observe a herd of grazing animals to recognize that resources are "finite" for the entire group, not merely individuals. You will never find a grazing animal that is vigilantly guarding its grass supply to prevent others from accessing it. Therefore the resources are freely used by all members of the group, and consequently there is no opportunity for individuals to conserve against others. The group will survive or starve based on their collective actions, not merely those of individuals.

Pinker's actual statement is fundamentally meaningless. It is tautalogically true, but biologically irrelevant.

...whichever copying errors happen to increase the rate of replication will accumulate in a lineage and predominate in the population. After many generations of replication, the replicators will show the appearance of design for effective replication, while in reality they have just accumulated the copying errors that had successful replication as their effect.
http://edge.org/conversation/the-false-allure-of-group-selection

In short, his statement addresses nothing except genes that increase the rate of replication, being represented in future generations with the appearance of having been designed for higher replication rates.

The actual irony in the quote is the preceding sentence:

The core of natural selection is that when replicators arise and make copies of themselves...

Which, of course, is followed by the statement indicating that their success is governed specifically by their inability to accurately copy themselves. Again ... an overly simplistic representation filled with all manner of assumptions that may not hold true. Apparently resources aren't that "finite" with such replication.

Gerhard Adam | 06/29/12 | 11:49 AM

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Yes, we're talking about organisms that compete for the same resources, a given population of a single species. You're right of course, but again, the context in which you discuss natural selection should have made this pretty clear.

Why would you presume that natural selection requires such a context? I also didn't indicate a single species [since that is even more restrictive] but rather related species, such that one would tend to dominate and drive the other to extinction or into a different niche.

Natural selection still operates on unrelated species that are vying for the same resources, so it should not be presumed that the only context for this discussion is about "related species".

Gerhard Adam | 06/29/12 | 11:54 AM

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Actually Gerhard, I misspoke. The original quote from Pinker does give clear context with the phrase "in a lineage". This precludes the possibiltiy that we were talking about anything other than "a given populatio fo a single species", so your interpretation of what I said is invalid, and your criticism of Pinker's #4 is forcefully removing the context as well.

Andrew TT (not verified) | 06/29/12 | 11:36 AM

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Again, here's a perfect example against the notion of "selfish genes" and competition among genes as replicators.

When the researchers probed which sequences piRNAs tended to shut down, they found that if a cell has ever turned on a gene in the past, the piRNA system will recognize it as a "self" gene and allow it to be expressed. But if it hasn't been active in the organism before, the piRNA will set the silencing mechanism into action so it remains off.

If the piRNA doesn't silence a gene the first time it encounters it, it won't ever silence it. And if it silences it once, then every time that gene appears in the future, the system will turn it off.
http://www.sciencecodex.com/short_stretches_of_pirna_evaluate_cells_genetic_history-94244

This actually fits in quite well with my own personal theory of genes and the role of DNA. Of course, my view is purely speculative, but this does tend to support it.

Gerhard Adam | 06/29/12 | 12:17 PM

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Gerhard,
indeed it fits quite well, I'd say:

“This is really remarkable. It implies that an organism has a memory of all the previous gene sequences it’s ever expressed before.”
Craig C. Mello
http://www.hhmi.org/news/mello20120625.html

harry pinxteren (not verified) | 07/11/12 | 14:59 PM

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This suggests quite clearly that genes are incapable of "competing" within the genome and are subject to external "audit" and control.

Gerhard Adam | 07/11/12 | 19:44 PM

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"As Pinker pretty thoroughly demonstrated, none of the ideas put forth by group selectionists explain something that can't be explained by the gene-centric theory."
Except that every time Pinker looks at himself in the mirror he sees something that can't be explained by the gene-centric theory.
Gene-centrism is built around the myth of relentless competition.
If relentless competition was a fact, humans would cease to exist.

Steve Davis | 06/29/12 | 20:49 PM

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it would probably be better if you stuck to evolution topics. The world is a more peaceful place -- ask Genghis Khan.

Anonymous (not verified) | 08/12/12 | 22:08 PM

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Genghis Khan? Is he supposed to be a model for or against cooperation?

Gerhard Adam | 08/12/12 | 22:35 PM

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