I'm currently reading "What Evolution Is" by Ernst Mayr, so far a great read, and I'm wondering about gradualism vs' punctuated equilibrium; specifically, if we have a set of fossils from one species to an ancestor such that the transitional fossils that exist within the set are such that it can be viewed as being virtually continuous, then would this not counter the claims of punctuated equilibrium?
I'm sure there is some subtle point I'm missing here because Stephen J. Gould was a rational fellow who devoted a life time of studying and investigating evolution and so is not likely to put forward an idea that seemingly runs counter to gradualism while having likely been aware of rather complete sets of fossils like that of the whale and its ancestor, horses, or from the therapsid reptiles to the mammals.
Thus, if someone could shed some light on this point I would much appreciate it.
>I'm currently reading "What Evolution Is" by Ernst Mayr, >so far a great read, and I'm wondering about gradualism >vs' punctuated equilibrium; specifically, if we have >a set of fossils from one species to an ancestor >such that the transitional fossils that exist within >the set are such that it can be viewed as being >virtually continuous, then would this not >counter the claims of punctuated equilibrium?
>I'm sure there is some subtle point I'm missing >here because Stephen J. Gould was a rational >fellow who devoted a life time of studying and >investigating evolution and so is not likely to >put forward an idea that seemingly runs counter >to gradualism while having likely been >aware of rather complete sets of fossils like >that of the whale and its ancestor, horses, or >from the therapsid reptiles to the mammals.
>Thus, if someone could shed some light on >this point I would much appreciate it.
Gould and, more properly, Eldredge, proposed punctuated equilibrium most specifically because of their familiarity with long stretches of fossil record of specific organisms. Wikipedia has a very nice essay on the subject http://en.wikipedia.org/wiki/Punctuated_equilibrium which carefully distinguishes between punc-eq and "saltational" evolution or macromutation where large jumps are made by single mutational events. That is not what punc-eq really refers to and it is not at all antagonistic to gradual change. It is just that a relatively large population of organisms is rather stable although at the geographic fringes of the range, gradually accumulated changes may result in populations rather different from the norm, new species in fact. Some type of ecological event may then cause the population to suddenly change with the formerly isolated group invading and taking over the habitat previously inhabited by the parent group. The fossil record taken from a single location would show a sudden change in phenotype even if the changes occurred gradually.
With the advent of evo-devo (the application of new data on developmental biology and gene regulation and expression to evolutionary biology) we now know that it is possible both for seemingly small mutational events to cause a rather large phenotypic effect and that it is also possible for the gradual accumulation of mutations to first produce no noticeable effect until the regulatory system becomes overwhelmed and suddenly reorganizes to produce a very large change. In this sense, saltations and macromutations are also quite possible mechanisms to produce the punc-eq effect.
It is also true that just how "smooth" vs. "step-wise" the fossil record appears depends strongly on how closely you look. The examples you mention, the evolution of the horse or of the whale involve such major and drastic changes that a series of small step-wise changes in each detail would look like gradual change when viewed over the long stretch.
conrad wrote: > I'm currently reading "What Evolution Is" by Ernst Mayr, > so far a great read, and I'm wondering about gradualism > vs' punctuated equilibrium; specifically, if we have > a set of fossils from one species to an ancestor > such that the transitional fossils that exist within > the set are such that it can be viewed as being > virtually continuous, then would this not > counter the claims of punctuated equilibrium?
No. But Gould never argued for saltation. In classical PE, transitions are gradual, with all the intermediates anyone would expect. In fact, the initial PE paper contained a claimed example of gradual change, in the evolution of Phacops trilobites. The claim is that such gradual changes are rare in the fossil record because they usually happen quickly in geological time, and in a restricted geographical area, such that we are unlikely to have a good series to show them.
But let's suppose you are referring to a series showing gradual change on a geological time-scale, i.e. a smooth transition over millions of years. Would this disprove PE? Sure, in that one case. But Gould never (hmmm, better make that "seldom", just in case) claimed that PE was the sole means of speciation, just the predominant one. You need to assemble a whole bunch of transitions and see how many of them match gradualism (however you want to define that) and how many match PE.
A bigger problem is that it's impossible to tell if one fossil species (really, a morphotype) is ancestral to another. An even bigger problem is that you can't really identify species in the fossil record, only morphotypes.
> I'm sure there is some subtle point I'm missing > here because Stephen J. Gould was a rational > fellow who devoted a life time of studying and > investigating evolution and so is not likely to > put forward an idea that seemingly runs counter > to gradualism while having likely been > aware of rather complete sets of fossils like > that of the whale and its ancestor, horses, or > from the therapsid reptiles to the mammals.
Gould would say that these series are not gradual transitions at the level of species. Instead they're a series of species showing a long-term trend. Nothing to do with PE vs. gradualism. Remember, PE is a theory about speciation.
> Thus, if someone could shed some light on > this point I would much appreciate it.
> conrad wrote: > > I'm currently reading "What Evolution Is" by Ernst Mayr, > > so far a great read, and I'm wondering about gradualism > > vs' punctuated equilibrium; specifically, if we have > > a set of fossils from one species to an ancestor > > such that the transitional fossils that exist within > > the set are such that it can be viewed as being > > virtually continuous, then would this not > > counter the claims of punctuated equilibrium?
> No. But Gould never argued for saltation. In classical PE, transitions > are gradual, with all the intermediates anyone would expect. In fact, > the initial PE paper contained a claimed example of gradual change, in > the evolution of Phacops trilobites. The claim is that such gradual > changes are rare in the fossil record because they usually happen > quickly in geological time, and in a restricted geographical area, such > that we are unlikely to have a good series to show them.
> But let's suppose you are referring to a series showing gradual change > on a geological time-scale, i.e. a smooth transition over millions of > years. Would this disprove PE? Sure, in that one case. But Gould never > (hmmm, better make that "seldom", just in case) claimed that PE was the > sole means of speciation, just the predominant one. You need to assemble > a whole bunch of transitions and see how many of them match gradualism > (however you want to define that) and how many match PE.
> A bigger problem is that it's impossible to tell if one fossil species > (really, a morphotype) is ancestral to another. An even bigger problem > is that you can't really identify species in the fossil record, only > morphotypes.
> > I'm sure there is some subtle point I'm missing > > here because Stephen J. Gould was a rational > > fellow who devoted a life time of studying and > > investigating evolution and so is not likely to > > put forward an idea that seemingly runs counter > > to gradualism while having likely been > > aware of rather complete sets of fossils like > > that of the whale and its ancestor, horses, or > > from the therapsid reptiles to the mammals.
> Gould would say that these series are not gradual transitions at the > level of species. Instead they're a series of species showing a > long-term trend. Nothing to do with PE vs. gradualism. Remember, PE is a > theory about speciation.
> > Thus, if someone could shed some light on > > this point I would much appreciate it.
> Sure.
I may be misremembering but having read the Blind Watchmaker, in retrospect, it would seem like Richard Dawkins misunderstood Gould's position then? I don't know if you have read this particular book, but from what I recall, Richard Dawkins took great pains in showing that PE was encapsulated by gradualism.
> >I'm currently reading "What Evolution Is" by Ernst Mayr, > >so far a great read, and I'm wondering about gradualism > >vs' punctuated equilibrium; specifically, if we have > >a set of fossils from one species to an ancestor > >such that the transitional fossils that exist within > >the set are such that it can be viewed as being > >virtually continuous, then would this not > >counter the claims of punctuated equilibrium?
> >I'm sure there is some subtle point I'm missing > >here because Stephen J. Gould was a rational > >fellow who devoted a life time of studying and > >investigating evolution and so is not likely to > >put forward an idea that seemingly runs counter > >to gradualism while having likely been > >aware of rather complete sets of fossils like > >that of the whale and its ancestor, horses, or > >from the therapsid reptiles to the mammals.
> >Thus, if someone could shed some light on > >this point I would much appreciate it.
> Gould and, more properly, Eldredge, proposed punctuated equilibrium > most specifically because of their familiarity with long stretches of > fossil record of specific organisms. Wikipedia has a very nice essay > on the subject > http://en.wikipedia.org/wiki/Punctuated_equilibrium > which carefully distinguishes between punc-eq and "saltational" > evolution or macromutation where large jumps are made by single > mutational events. That is not what punc-eq really refers to and it > is not at all antagonistic to gradual change. It is just that a > relatively large population of organisms is rather stable although at > the geographic fringes of the range, gradually accumulated changes may > result in populations rather different from the norm, new species in > fact. Some type of ecological event may then cause the population to > suddenly change with the formerly isolated group invading and taking > over the habitat previously inhabited by the parent group. The fossil > record taken from a single location would show a sudden change in > phenotype even if the changes occurred gradually.
> With the advent of evo-devo (the application of new data on > developmental biology and gene regulation and expression to > evolutionary biology) we now know that it is possible both for > seemingly small mutational events to cause a rather large phenotypic > effect and that it is also possible for the gradual accumulation of > mutations to first produce no noticeable effect until the regulatory > system becomes overwhelmed and suddenly reorganizes to produce a very > large change. In this sense, saltations and macromutations are also > quite possible mechanisms to produce the punc-eq effect.
Do you know of good books or research articles that discuss the cumulation of mutations not taking effect until a "regulatory system becomes overwhelmed"?
It sounds like great stuff, I'd like to read more on it.
> It is also true that just how "smooth" vs. "step-wise" the fossil > record appears depends strongly on how closely you look. The examples > you mention, the evolution of the horse or of the whale involve such > major and drastic changes that a series of small step-wise changes > in each detail would look like gradual change when viewed over the > long stretch.
conrad wrote: > On Jul 5, 10:15 pm, John Harshman <jharshman.diespam...@pacbell.net> > wrote: >> conrad wrote: >>> I'm currently reading "What Evolution Is" by Ernst Mayr, >>> so far a great read, and I'm wondering about gradualism >>> vs' punctuated equilibrium; specifically, if we have >>> a set of fossils from one species to an ancestor >>> such that the transitional fossils that exist within >>> the set are such that it can be viewed as being >>> virtually continuous, then would this not >>> counter the claims of punctuated equilibrium? >> No. But Gould never argued for saltation. In classical PE, transitions >> are gradual, with all the intermediates anyone would expect. In fact, >> the initial PE paper contained a claimed example of gradual change, in >> the evolution of Phacops trilobites. The claim is that such gradual >> changes are rare in the fossil record because they usually happen >> quickly in geological time, and in a restricted geographical area, such >> that we are unlikely to have a good series to show them.
>> But let's suppose you are referring to a series showing gradual change >> on a geological time-scale, i.e. a smooth transition over millions of >> years. Would this disprove PE? Sure, in that one case. But Gould never >> (hmmm, better make that "seldom", just in case) claimed that PE was the >> sole means of speciation, just the predominant one. You need to assemble >> a whole bunch of transitions and see how many of them match gradualism >> (however you want to define that) and how many match PE.
>> A bigger problem is that it's impossible to tell if one fossil species >> (really, a morphotype) is ancestral to another. An even bigger problem >> is that you can't really identify species in the fossil record, only >> morphotypes.
>>> I'm sure there is some subtle point I'm missing >>> here because Stephen J. Gould was a rational >>> fellow who devoted a life time of studying and >>> investigating evolution and so is not likely to >>> put forward an idea that seemingly runs counter >>> to gradualism while having likely been >>> aware of rather complete sets of fossils like >>> that of the whale and its ancestor, horses, or >>> from the therapsid reptiles to the mammals. >> Gould would say that these series are not gradual transitions at the >> level of species. Instead they're a series of species showing a >> long-term trend. Nothing to do with PE vs. gradualism. Remember, PE is a >> theory about speciation.
>>> Thus, if someone could shed some light on >>> this point I would much appreciate it. >> Sure.
> I may be misremembering but having read > the Blind Watchmaker, in retrospect, it > would seem like Richard Dawkins misunderstood > Gould's position then? I don't know if you have read > this particular book, but from what I recall, Richard > Dawkins took great pains in showing that PE was > encapsulated by gradualism.
This all depends on what you mean by "gradualism". Gould never claimed that PE was instantaneous; according to him, speciation would generally take somewhere around 10-100 thousand years. This would seldom be visible in the fossil record.
I don't remember what Dawkins said, but PE is really a theory about the relationship between speciation and morphological change. Under PE, species spend most of their time in stasis, and change only during speciation. Unfortunately, in the fossil record the only way to attempt to recognize speciation is by observing morphological change, and thus PE reduces to a theory that morphological change is coincident with morphological change.
On Jul 6, 3:15 am, conrad <con...@lawyer.com> wrote:
> I'm currently reading "What Evolution Is" by Ernst Mayr, > so far a great read, and I'm wondering about gradualism > vs' punctuated equilibrium; specifically, if we have > a set of fossils from one species to an ancestor > such that the transitional fossils that exist within > the set are such that it can be viewed as being > virtually continuous, then would this not > counter the claims of punctuated equilibrium?
It doesn't counter the claim that stasis is commonplace. -- __________ |im |yler http://timtyler.org/ t...@tt1lock.org Remove lock to reply.
>> On Sat, 5 Jul 2008 19:15:06 -0700 (PDT), conrad <con...@lawyer.com> >> With the advent of evo-devo (the application of new data on >> developmental biology and gene regulation and expression to >> evolutionary biology) we now know that it is possible both for >> seemingly small mutational events to cause a rather large phenotypic >> effect and that it is also possible for the gradual accumulation of >> mutations to first produce no noticeable effect until the regulatory >> system becomes overwhelmed and suddenly reorganizes to produce a very >> large change. In this sense, saltations and macromutations are also >> quite possible mechanisms to produce the punc-eq effect.
>Do you know of good books or research articles >that discuss the cumulation of mutations not >taking effect until a "regulatory system becomes >overwhelmed"?
>It sounds like great stuff, I'd like to read more >on it.
My comments were based mostly on my knowledge of complex systems theory and neurobiology where such things are commonplace. For example, a series of receptors with analog or graded behavior can act as a digital switch (on/off only) by interacting cooperatively or with positive feedback. Each component is analog, graded, or gradual but the ensemble acts digitally producing no effect until a threshold is reached and then suddenly switching to a new pattern of activity. The all-or-none action potential in neurons is a result of such positive feedback interaction. Biochemistry has many examples of such cooperativity: oxygen binding by hemoglobin is the classic example although the end product is not so extreme as to be all-or-none but rather shows a "sigmoidal" relationship instead of a linear one. The basic electronic digital storage unit, the flip-flop circuit, is another example.
If you want genetic types of examples, at the trivial level showing the effect of regulation there is the phenomenon of dominant alleles that produce the same effect whether one or two copies are present. So some flowers given either one or two copies of a functional gene that produces, say, red color, will always produce the same final amount of redness. Other species with different regulatory systems will vary the intensity of the color with gene dosage so that crossing red with white does not produce all red offspring but rather will produce an intermediate pink. That simply shows the phenomenon of gene regulation, not its "being overwhelmed" and "suddenly reorganizing".
To show a rather sudden change in phenotype with a gradation of genetic changes, consider Huntington's Disease. This is caused by mutations in the Huntingtin gene (yes, the spelling is different). This gene contains a repeating segment CAGCAGCAG... a so-called trinucleotide repeat that codes for a series of glutamine. If the series contains up to 36 glutamines in a row, there is no problem. However 36 to 39 produces a late onset and slow progression of symptoms of the disease and 40 or more produces severe disease. There are a number of diseases caused by variation in this trinucleotide repeat sequence. See http://en.wikipedia.org/wiki/Trinucleotide_repeat_disorders
As to true "genetic reorganization" on the basis of cumulative mutations, that was primarily a surmise on my part based, as I indicated earlier, on standard systems theory considerations, purely theoretical. I am not familiar enough with the molecular biological literature to give you better examples.
>>On Jul 5, 9:44 pm, r norman <r_s_norman@_comcast.net> wrote:
><snip>
>>> On Sat, 5 Jul 2008 19:15:06 -0700 (PDT), conrad <con...@lawyer.com> >>> With the advent of evo-devo (the application of new data on >>> developmental biology and gene regulation and expression to >>> evolutionary biology) we now know that it is possible both for >>> seemingly small mutational events to cause a rather large phenotypic >>> effect and that it is also possible for the gradual accumulation of >>> mutations to first produce no noticeable effect until the regulatory >>> system becomes overwhelmed and suddenly reorganizes to produce a very >>> large change. In this sense, saltations and macromutations are also >>> quite possible mechanisms to produce the punc-eq effect.
>>Do you know of good books or research articles >>that discuss the cumulation of mutations not >>taking effect until a "regulatory system becomes >>overwhelmed"?
>>It sounds like great stuff, I'd like to read more >>on it.
>My comments were based mostly on my knowledge of complex systems >theory and neurobiology where such things are commonplace. For >example, a series of receptors with analog or graded behavior can act >as a digital switch (on/off only) by interacting cooperatively or with >positive feedback. Each component is analog, graded, or gradual but >the ensemble acts digitally producing no effect until a threshold is >reached and then suddenly switching to a new pattern of activity. The >all-or-none action potential in neurons is a result of such positive >feedback interaction. Biochemistry has many examples of such >cooperativity: oxygen binding by hemoglobin is the classic example >although the end product is not so extreme as to be all-or-none but >rather shows a "sigmoidal" relationship instead of a linear one. The >basic electronic digital storage unit, the flip-flop circuit, is >another example.
>If you want genetic types of examples, at the trivial level showing >the effect of regulation there is the phenomenon of dominant alleles >that produce the same effect whether one or two copies are present. So >some flowers given either one or two copies of a functional gene that >produces, say, red color, will always produce the same final amount of >redness. Other species with different regulatory systems will vary >the intensity of the color with gene dosage so that crossing red with >white does not produce all red offspring but rather will produce an >intermediate pink. That simply shows the phenomenon of gene >regulation, not its "being overwhelmed" and "suddenly reorganizing".
>To show a rather sudden change in phenotype with a gradation of >genetic changes, consider Huntington's Disease. This is caused by >mutations in the Huntingtin gene (yes, the spelling is different). >This gene contains a repeating segment CAGCAGCAG... a so-called >trinucleotide repeat that codes for a series of glutamine. If the >series contains up to 36 glutamines in a row, there is no problem. >However 36 to 39 produces a late onset and slow progression of >symptoms of the disease and 40 or more produces severe disease. >There are a number of diseases caused by variation in this >trinucleotide repeat sequence. See > http://en.wikipedia.org/wiki/Trinucleotide_repeat_disorders
>As to true "genetic reorganization" on the basis of cumulative >mutations, that was primarily a surmise on my part based, as I >indicated earlier, on standard systems theory considerations, purely >theoretical. I am not familiar enough with the molecular biological >literature to give you better examples.
For starters, though, you could google "genetic regulatory reorganization".