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	<title>Comments on: For Darwin Day: On the Origin of Genetic Information</title>
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	<link>http://metamodern.com/2009/02/12/for-darwin-day-on-the-origin-of-genetic-information/</link>
	<description>The Trajectory of Technology</description>
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		<title>By: Darwin portraits on sale, ₤10</title>
		<link>http://metamodern.com/2009/02/12/for-darwin-day-on-the-origin-of-genetic-information/comment-page-1/#comment-4567</link>
		<dc:creator>Darwin portraits on sale, ₤10</dc:creator>
		<pubDate>Mon, 26 Dec 2011 13:42:32 +0000</pubDate>
		<guid isPermaLink="false">http://metamodern.com/?p=1635#comment-4567</guid>
		<description>[...] For Darwin Day: On the Origin of Genetic Information [...]</description>
		<content:encoded><![CDATA[<p>[...] For Darwin Day: On the Origin of Genetic Information [...]</p>
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		<title>By: Evolution: The concept and how we talk about it</title>
		<link>http://metamodern.com/2009/02/12/for-darwin-day-on-the-origin-of-genetic-information/comment-page-1/#comment-2490</link>
		<dc:creator>Evolution: The concept and how we talk about it</dc:creator>
		<pubDate>Sun, 03 Jan 2010 09:32:56 +0000</pubDate>
		<guid isPermaLink="false">http://metamodern.com/?p=1635#comment-2490</guid>
		<description>[...] For Darwin Day: On the Origin of Genetic Information [...]</description>
		<content:encoded><![CDATA[<p>[...] For Darwin Day: On the Origin of Genetic Information [...]</p>
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		<title>By: Great Science, Great Scientists, and Icons</title>
		<link>http://metamodern.com/2009/02/12/for-darwin-day-on-the-origin-of-genetic-information/comment-page-1/#comment-2184</link>
		<dc:creator>Great Science, Great Scientists, and Icons</dc:creator>
		<pubDate>Fri, 27 Nov 2009 10:33:25 +0000</pubDate>
		<guid isPermaLink="false">http://metamodern.com/?p=1635#comment-2184</guid>
		<description>[...] For Darwin Day: On the Origin of Genetic Information [...]</description>
		<content:encoded><![CDATA[<p>[...] For Darwin Day: On the Origin of Genetic Information [...]</p>
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		<title>By: Will Ware</title>
		<link>http://metamodern.com/2009/02/12/for-darwin-day-on-the-origin-of-genetic-information/comment-page-1/#comment-569</link>
		<dc:creator>Will Ware</dc:creator>
		<pubDate>Tue, 24 Feb 2009 15:30:32 +0000</pubDate>
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		<description>Thanks all for the reference to the MacKay book. It looks very interesting and I am amazoning myself a copy this week.</description>
		<content:encoded><![CDATA[<p>Thanks all for the reference to the MacKay book. It looks very interesting and I am amazoning myself a copy this week.</p>
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		<title>By: Eric Drexler</title>
		<link>http://metamodern.com/2009/02/12/for-darwin-day-on-the-origin-of-genetic-information/comment-page-1/#comment-543</link>
		<dc:creator>Eric Drexler</dc:creator>
		<pubDate>Fri, 20 Feb 2009 23:08:38 +0000</pubDate>
		<guid isPermaLink="false">http://metamodern.com/?p=1635#comment-543</guid>
		<description>@ Bruce Smith --
You make good criticisms of my slapdash argument, which needs some modification.

First I should say that I assume that analysis made by David MacKay is correct, given its assumptions. Any conflict is only with hasty conclusions that might be drawn from it.

For background, I’m not concerned with biological realism here, but about constraints that might be incorrectly be thought to apply to all evolutionary systems, including computational systems &lt;em&gt;designed&lt;/em&gt; to evolve their information quickly.

For simply dividing and re-merging a population, let’s substitute ongoing partial division: We have one big, gene-mixing population when viewed on a long time scale, but not with the fastest and most uniform possible mixing.

Now consider “epistasis”, that is, non-additive gene effects. It is both reasonable and permissible for present purposes to postulate that many mutations produce a small benefit that becomes large only when combined with a second mutation which  (for example) fixes a problem created by the first, and would be disadvantageous in itself. The second mutation can’t spread until the first becomes common, and in a small population, this will happen in fewer generations. The story from there involves mixing from one small population to another, or perhaps favorable linkage disequilibrium when mixing into a larger population. This example is enough to show that conclusions drawn from a mathematically simple, idealized population model need not hold in general. (Note that GAs are often coded with structured populations of the sort I described.)

There is also the question of  what the question is. If (as tends to happen) “rate of information gain” is tacitly regarded as meaning “rate of evolution”, there is another problem, because conclusions about the former don’t necessarily tell us much about the latter. (“Rate of evolution” is clearly a meaningful concept, even if it lacks a unique metric).

Here, the argument is simple: It is permissible for present purposes to postulate a mapping from genotype to phenotype such that very rare mutations are very important; if so, then for a given mutation rate, a large population will find them faster, simply because there are more mutations in total. In this model, for a given rate of information gain, a larger population would achieve a faster increase in information value.

Here again, I am not saying that this is (or isn’t) a good description of biology, merely that it’s unwise to leap quickly from a theorem about information gain to a conclusion about the rate of significant evolutionary change.</description>
		<content:encoded><![CDATA[<p>@ Bruce Smith &#8211;<br />
You make good criticisms of my slapdash argument, which needs some modification.</p>
<p>First I should say that I assume that analysis made by David MacKay is correct, given its assumptions. Any conflict is only with hasty conclusions that might be drawn from it.</p>
<p>For background, I’m not concerned with biological realism here, but about constraints that might be incorrectly be thought to apply to all evolutionary systems, including computational systems <em>designed</em> to evolve their information quickly.</p>
<p>For simply dividing and re-merging a population, let’s substitute ongoing partial division: We have one big, gene-mixing population when viewed on a long time scale, but not with the fastest and most uniform possible mixing.</p>
<p>Now consider “epistasis”, that is, non-additive gene effects. It is both reasonable and permissible for present purposes to postulate that many mutations produce a small benefit that becomes large only when combined with a second mutation which  (for example) fixes a problem created by the first, and would be disadvantageous in itself. The second mutation can’t spread until the first becomes common, and in a small population, this will happen in fewer generations. The story from there involves mixing from one small population to another, or perhaps favorable linkage disequilibrium when mixing into a larger population. This example is enough to show that conclusions drawn from a mathematically simple, idealized population model need not hold in general. (Note that GAs are often coded with structured populations of the sort I described.)</p>
<p>There is also the question of  what the question is. If (as tends to happen) “rate of information gain” is tacitly regarded as meaning “rate of evolution”, there is another problem, because conclusions about the former don’t necessarily tell us much about the latter. (“Rate of evolution” is clearly a meaningful concept, even if it lacks a unique metric).</p>
<p>Here, the argument is simple: It is permissible for present purposes to postulate a mapping from genotype to phenotype such that very rare mutations are very important; if so, then for a given mutation rate, a large population will find them faster, simply because there are more mutations in total. In this model, for a given rate of information gain, a larger population would achieve a faster increase in information value.</p>
<p>Here again, I am not saying that this is (or isn’t) a good description of biology, merely that it’s unwise to leap quickly from a theorem about information gain to a conclusion about the rate of significant evolutionary change.</p>
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		<title>By: Eliezer Yudkowsky</title>
		<link>http://metamodern.com/2009/02/12/for-darwin-day-on-the-origin-of-genetic-information/comment-page-1/#comment-529</link>
		<dc:creator>Eliezer Yudkowsky</dc:creator>
		<pubDate>Wed, 18 Feb 2009 10:24:03 +0000</pubDate>
		<guid isPermaLink="false">http://metamodern.com/?p=1635#comment-529</guid>
		<description>MacKay is talking about gaining bits like bits on a hard drive - flipping zeroes in a string to ones; this is not the same as gaining bits in an information-theoretic sense.

Although I originally thought that an error rate of 10^-8 would imply a maximum genome size of around 10^8, a simple Python simulation failed to validate this.  Although it takes one death to remove one mutation from the gene pool, more than one mutation can be removed by the same death.  And this indeed gets you an info bound that goes as the square root of genome size, not just selection pressure.

The simulation was written for perfect selection on an entire gene pool (the exact bottom half being eliminated, and such) and it&#039;s not clear to me what happens when you start introducing more realistic assumptions like stochastic selection on small bands or families.

It&#039;s at least &lt;i&gt;worth noting&lt;/i&gt; that in real life the genome seems to be mostly junk, and even if it&#039;s not, it still contains vastly less info than it &quot;could&quot; according to MacKay&#039;s bound and the age of life on Earth.

Merging subpopulations is not a trustworthy argument against various attempted speed limits, because the combined population doesn&#039;t instantly get the best of every subpopulation.  If you split up into a million subpopulations, then, after merging, every beneficial adaptation promoted in them goes to a frequency of 1 in a million.  That&#039;s going to take some time to rise to universality, and any collisions will lose information, and any complex adaptations will be fragmented.

See &lt;a href=&quot;http://www.overcomingbias.com/2007/11/natural-selecti.html&quot; rel=&quot;nofollow&quot;&gt;this discussion on Overcoming Bias&lt;/a&gt; for much more.</description>
		<content:encoded><![CDATA[<p>MacKay is talking about gaining bits like bits on a hard drive &#8211; flipping zeroes in a string to ones; this is not the same as gaining bits in an information-theoretic sense.</p>
<p>Although I originally thought that an error rate of 10^-8 would imply a maximum genome size of around 10^8, a simple Python simulation failed to validate this.  Although it takes one death to remove one mutation from the gene pool, more than one mutation can be removed by the same death.  And this indeed gets you an info bound that goes as the square root of genome size, not just selection pressure.</p>
<p>The simulation was written for perfect selection on an entire gene pool (the exact bottom half being eliminated, and such) and it&#8217;s not clear to me what happens when you start introducing more realistic assumptions like stochastic selection on small bands or families.</p>
<p>It&#8217;s at least <i>worth noting</i> that in real life the genome seems to be mostly junk, and even if it&#8217;s not, it still contains vastly less info than it &#8220;could&#8221; according to MacKay&#8217;s bound and the age of life on Earth.</p>
<p>Merging subpopulations is not a trustworthy argument against various attempted speed limits, because the combined population doesn&#8217;t instantly get the best of every subpopulation.  If you split up into a million subpopulations, then, after merging, every beneficial adaptation promoted in them goes to a frequency of 1 in a million.  That&#8217;s going to take some time to rise to universality, and any collisions will lose information, and any complex adaptations will be fragmented.</p>
<p>See <a href="http://www.overcomingbias.com/2007/11/natural-selecti.html" rel="nofollow">this discussion on Overcoming Bias</a> for much more.</p>
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		<title>By: Bruce Smith</title>
		<link>http://metamodern.com/2009/02/12/for-darwin-day-on-the-origin-of-genetic-information/comment-page-1/#comment-524</link>
		<dc:creator>Bruce Smith</dc:creator>
		<pubDate>Tue, 17 Feb 2009 07:53:46 +0000</pubDate>
		<guid isPermaLink="false">http://metamodern.com/?p=1635#comment-524</guid>
		<description>Clarification: by &quot;information retained in a population&quot; above, I meant &quot;new information (derived from selection) retained in a population, per generation&quot;.</description>
		<content:encoded><![CDATA[<p>Clarification: by &#8220;information retained in a population&#8221; above, I meant &#8220;new information (derived from selection) retained in a population, per generation&#8221;.</p>
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		<title>By: Bruce Smith</title>
		<link>http://metamodern.com/2009/02/12/for-darwin-day-on-the-origin-of-genetic-information/comment-page-1/#comment-523</link>
		<dc:creator>Bruce Smith</dc:creator>
		<pubDate>Tue, 17 Feb 2009 07:51:16 +0000</pubDate>
		<guid isPermaLink="false">http://metamodern.com/?p=1635#comment-523</guid>
		<description>You wrote:
&lt;cite&gt;
If I divide the population into a million separate populations, the supposed limit now would allow useful genetic information to increase a million times faster. After a few generations, I merge the populations, and could have achieved advances that would supposedly have required ten million generations. A principle with this consequence obviously makes no sense.

Not so fast -- when you mentally re-merge the populations, i.e. admit that they were really one population all along, &lt;i&gt;with a single gene pool&lt;/i&gt;, the whole-population gene pool is forced to average out what the subpopulation gene pools &quot;learned&quot;, which reduces the amount of information it can retain from the &quot;raw input&quot; of offspring-counts per individual.

It&#039;s the mixing of genomes within a natural population which justifies thinking of it as a unit (a single &quot;gene pool&quot; from which new genomes are randomly drawn), and which also necessarily reduces its retained information from the raw input (as well as increasing its accuracy as a measurement of the environment, to the extent the environment is uniform). If this mixing breaks down, more information might be retained; but also, the organisms might speciate (perhaps not a coincidence?).

My conclusion from this: your thought experiment isn&#039;t a good reason to doubt the plausibility of a population-size-independent upper limit on information retained in a population, if by a &quot;population&quot; we mean something justifiably thought of as having &quot;a single gene pool&quot;.

(I&#039;m not saying I&#039;ve given evidence for such a limit -- ale&#039;s reply sounds like it points to something more sophisticated, which does that -- only for your thought experiment not ruling one out.)

Of course there can be ambiguity in what counts as &quot;one population&quot;, just like &quot;one mountain&quot; (e.g. it might have subpopulations with limited but nonzero gene exchange between them; one mountain with two peaks might be thought of as two nearby mountains), but that doesn&#039;t make the concept of a population arbitrary or unnatural -- it&#039;s based on what genome mixing is actually occurring.</description>
		<content:encoded><![CDATA[<p>You wrote:<br />
<cite><br />
If I divide the population into a million separate populations, the supposed limit now would allow useful genetic information to increase a million times faster. After a few generations, I merge the populations, and could have achieved advances that would supposedly have required ten million generations. A principle with this consequence obviously makes no sense.</p>
<p>Not so fast &#8212; when you mentally re-merge the populations, i.e. admit that they were really one population all along, <i>with a single gene pool</i>, the whole-population gene pool is forced to average out what the subpopulation gene pools &#8220;learned&#8221;, which reduces the amount of information it can retain from the &#8220;raw input&#8221; of offspring-counts per individual.</p>
<p>It&#8217;s the mixing of genomes within a natural population which justifies thinking of it as a unit (a single &#8220;gene pool&#8221; from which new genomes are randomly drawn), and which also necessarily reduces its retained information from the raw input (as well as increasing its accuracy as a measurement of the environment, to the extent the environment is uniform). If this mixing breaks down, more information might be retained; but also, the organisms might speciate (perhaps not a coincidence?).</p>
<p>My conclusion from this: your thought experiment isn&#8217;t a good reason to doubt the plausibility of a population-size-independent upper limit on information retained in a population, if by a &#8220;population&#8221; we mean something justifiably thought of as having &#8220;a single gene pool&#8221;.</p>
<p>(I&#8217;m not saying I&#8217;ve given evidence for such a limit &#8212; ale&#8217;s reply sounds like it points to something more sophisticated, which does that &#8212; only for your thought experiment not ruling one out.)</p>
<p>Of course there can be ambiguity in what counts as &#8220;one population&#8221;, just like &#8220;one mountain&#8221; (e.g. it might have subpopulations with limited but nonzero gene exchange between them; one mountain with two peaks might be thought of as two nearby mountains), but that doesn&#8217;t make the concept of a population arbitrary or unnatural &#8212; it&#8217;s based on what genome mixing is actually occurring.</cite></p>
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