[sci.bio] non-genetic evolution... not

colby@bu-bio.bu.edu (Chris Colby) (04/22/91)

In article <47570@ut-emx.uucp> bill@ut-emx.uucp (Bill Jefferys) writes:
>In article <79788@bu.edu.bu.edu> colby@bu-bio.UUCP (Chris Colby) writes:
>#In article <18637@csli.Stanford.EDU> cphoenix@csli.Stanford.EDU (Chris Phoenix) writes:

>#>Is this totally off base, or is non-genetic evolution a possibility?

>#	It's totally off base ;-) Non-genetic evolution is an
>#oxymoron.

>Actually, Chris P. is not so far off as one might think.
>_The Economist_ had a story this week (in the science section)
>about a certain species of wasp that reproduced parthenogenetically;
>all of the offspring are female. Yet, if the wasps are fed 
>antibiotics, they will produce males and females in 50%
>ratio. Why? It appears that a bacterium has co-evolved
>with the wasp. It is transmitted via the eggs (see Chris
>P's speculation above!!!), and has evolved the ability to
>supress the production of male offspring. Presumably this
>is because it provides an advantage to the bacterium!

	Yes, I read this when it came out in the literature.
There are also wasp species that can interbreed when fed
antibiotics, but not when they have the internal bacteria
present. Interesting stuff.

>Yet the wasp retains the ability to produce males, so it
>is the _conditions_ under which the embryo develops
>that determine the sex. A non-genetic "evolution" that
>"breeds true" as long as the bacterium is present.

	You are confusing the process of evolution with simple
change. If the gene pool of the wasps did not change (ie there
are still the same frequency of male and female determining 
genes), then the population did not evolve. This is true even
if there are more females than males compared to some time in
the past. Let me give an example to illustrate my point.

	Humans have been growing larger and heavier in recent
historical times. But this is not an evolutionary change; it
is a result of better nutrition and medicine. If the environment
returned to old conditions, humans would be shorter and thinner.
In other words the genes have stayed the same (roughly), but the mani-
festation of them has changed. The same is true in the wasps;
the gene pool has not changed (or at least we'll assume it
hasn't for the sake of argument) but the expression of the
genes has changed due to a change in environment (presence of an
internal endosymbiont). They have changed, but they haven't
evolved.

	Remember that evolution is _defined_ as a change in the
gene pool; therefore non-genetic evolution is by definition non-
sensical. Populations phenotypes (expression of genes) can change
without the gene pool changing. This can often be very interesting
(as in the wasp case), but it is not evolution.

>Bill Jefferys

Chris Colby
email: colby@bu-bio.bu.edu

bill@ut-emx.uucp (Bill Jefferys) (04/22/91)

In article <79798@bu.edu.bu.edu> colby@bu-bio.UUCP (Chris Colby) writes:
#In article <47570@ut-emx.uucp> bill@ut-emx.uucp (Bill Jefferys) writes:
#
#>Yet the wasp retains the ability to produce males, so it
#>is the _conditions_ under which the embryo develops
#>that determine the sex. A non-genetic "evolution" that
#>"breeds true" as long as the bacterium is present.
#
#	You are confusing the process of evolution with simple
#change. If the gene pool of the wasps did not change (ie there
#are still the same frequency of male and female determining 
#genes), then the population did not evolve. This is true even
#if there are more females than males compared to some time in
#the past.... 

Well, I _did_ put quotes around the word "evolution" in my
post, so I am not confused.

#	Remember that evolution is _defined_ as a change in the
#gene pool; therefore non-genetic evolution is by definition non-
#sensical. Populations phenotypes (expression of genes) can change
#without the gene pool changing. This can often be very interesting
#(as in the wasp case), but it is not evolution.

Well, Darwin would not have understood this definition of
evolution, as the concepts "gene" and "gene pool" postdate
him by a considerable time. For Darwin, evolution meant
"descent with modification." You may decide for yourself
whether this applies to the wasp example.

Bill Jefferys

-- 
If you meet the Buddha on the net, put him in your kill file
	--Robert Firth

cphoenix@csli.Stanford.EDU (Chris Phoenix) (04/25/91)

In article <79798@bu.edu.bu.edu> colby@bu-bio.UUCP (Chris Colby) writes:
>	You are confusing the process of evolution with simple
>change. If the gene pool of the wasps did not change (ie there
>are still the same frequency of male and female determining 
>genes), then the population did not evolve. This is true even
>if there are more females than males compared to some time in
>the past. 

This is getting away from my question, but what do you consider the
gene pool?  If the bacteria mutated and this caused some change in the
wasp, would you say the wasp had mutated?  If you say "no", I have
another question: would you say that we mutate if our mitochondria
mutate?  But the last theory I heard was that mitochondria used to be
bacteria that our cells took in, once upon a time.  (Bill, I wrote
this just before I got your mail.  I'll let it stand as a question to
others, if they're interested.)

Let me give an example to illustrate my point.
>	Humans have been growing larger and heavier in recent
>historical times. But this is not an evolutionary change; it
>is a result of better nutrition and medicine.

This is an extracellular change.  Obviously it is not a characteristic
of the individual cells (ie if you pulled a cell out of Plato and a
cell out of me and grew them in vitro you probably couldn't tell the
difference in their offspring.)  I am not interested in this kind of
change.  On the other hand, Discover did an article about the effects
of malnutrition in the womb, in wartime Poland I believe.  If the
first generation went hungry while the second generation was a
few-month-old fetus, then effects would be seen in the second *and*
third generation, even though the third had normal nutrition.

But I am interested in *intracellular* changes which do not involve
the cell's nucleic acid, and preferably do not involve any nucleic
acid.  Let me give you a hypothetical example that typefies what I am
looking for: Suppose a cell has a structure of tubules.  These tubules
serve as conveyors for various molecules, including some molecule
necessary to build the tubules.  So when the cell divides, the tubules
grow and then the structure splits.  If the structure were removed, it
would no longer be able to grow (and thus couldn't start growing)
since it was necessary for its own growth.  Assuming the cell was
still viable, its offspring would not have that structure.  (I've left
a lot of loose ends in this example, but I hope you see what I mean.)

>Populations phenotypes (expression of genes) can change
>without the gene pool changing. This can often be very interesting
>(as in the wasp case), but it is not evolution.

Should you distinguish between characteristics caused by expression of
the genes and characteristics caused by the environment?  Your human
size example is entirely dependent on the environment.  On a cellular
level, changes caused by hormones are part of the environment and not
part of the cell.  But if you take a cell and its descendent, wash
them carefully, and put them in petri dishes, and the offspring look
different, that is not caused by the environment but by a
transmissible change *inside the cell*.  If that change does not
involve nucleic acid, that is what I'm interested in.

Speaking of hormones, I've been following the discussion on hormonal
control of fetal growth with interest.  But I haven't seen anyone
address the question of where the hormones come from.  I think all of
them will be coded for by the DNA of the original fertilized egg.
Assume that if you take one dinosaur cell, and one chicken cell with
dinosaur DNA, and grow them in a dish, there is no difference in their
phenotype.  So a fertilized chicken-egg cell with dinosaur DNA will
begin by producing dinosaur hormones.  If these hormones spread
through the growing embryo in the same way (which is purely based on
the embryo's environment and physical structure) then the cells will
react to them just like dinosaur cells would.  So the embryo will keep
developing like a dinosaur, and will become a dinosaur.

Notice we've made several assumptions here.  1) The hybrid cells react
exactly like dinosaur cells to all stimuli.  2) The embryo starts the
same physically as a dinosaur embryo (eg. the original egg cell is the
same size).  3) The environment is the same (eg. the pH of dinosaur
yolk is the same as the pH of chicken yolk).  I am interested in case
1), in whether a two cells with the same DNA will have the same
phenotype--ie reaction to stimuli, internal structure, etc.

I hope this is still interesting for people...

gagen@bgsuvax.UUCP (kathleen gagen) (04/25/91)

From article <79798@bu.edu.bu.edu>, by colby@bu-bio.bu.edu (Chris Colby):
> 
> 	Remember that evolution is _defined_ as a change in the
> gene pool; therefore non-genetic evolution is by definition non-
> sensical. 

ORGANIC evolution is defined as a change in the gene pool.  There are many
other types of evolution.  EVOLUTION means change.  
ie:
the evolution of the solar system
the evolution of western religious thought
the evolution of Lake Erie.

Kathi Gagen

mroussel@alchemy.chem.utoronto.ca (Marc Roussel) (04/26/91)

In article <79798@bu.edu.bu.edu> colby@bu-bio.UUCP (Chris Colby) writes:
>	Remember that evolution is _defined_ as a change in the
>gene pool; therefore non-genetic evolution is by definition non-
>sensical.

     I'm not a biologist so I hope that what I'm about to write isn't total
nonsense.  If it is, reply by email and I will post a retraction with a
summary of the points people have made.  If not, we can debate things in
this forum.
     It seems to me that the definition presented above is useful in some
contexts, but overly restrictive in others.  The wasp example which
someone else posted is a good example.  Whether or not the wasps' gene
pool changed, there has been a change in the species which is persistent
against (one presumes) significant external perturbations.  It seems to
me that this qualifies as evolution in a less restricted sense than the
current dogma allows: a species can be said to have evolved a trait if
this trait is transmissible from generation to generation and stable
against reasonable environmental perturbations.  (I will let someone
else define "reasonable" and "external environment", if any of this makes sense.
Surely in the case of the wasps and their symbiotic partners, the
symbionts form one system and thus are not part of the external
environment.)  This definition of course excludes (quite rightly) trivial
cases like the "better nutrition = taller humans" effect which Chris mentioned.
     My definition has the advantage that it allows us to include in
evolutionary theory things which we may not have thought of yet.
Further, stability of evolved traits as the hallmark of evolution brings
the language of biology into line with modern thinking in other sciences.
(I am curious to hear comments on this last point.  Would people
think of this as an advantage, a disadvantage, or neither?)
     I look forward to your comments.

				Marc R. Roussel
                                mroussel@alchemy.chem.utoronto.ca

sarima@tdatirv.UUCP (Stanley Friesen) (04/28/91)

In article <1991Apr25.203311.20957@alchemy.chem.utoronto.ca> mroussel@alchemy.chem.utoronto.ca (Marc Roussel) writes:
>...  It seems to
>me that this qualifies as evolution in a less restricted sense than the
>current dogma allows: a species can be said to have evolved a trait if
>this trait is transmissible from generation to generation and stable
>against reasonable environmental perturbations.  (I will let someone
>else define "reasonable" and "external environment", if any of this
>makes sense.

I think it makes a certain amount of sense.  The main reason for the current
emphasis on the genetic component of change is because that is the one type
of change we *know* is transmissible, as opposed to externally imposed.

To Darwin, at least, the main point of evolution was the selection of
heritable (== transmissible) changes (or variation).

>Surely in the case of the wasps and their symbiotic partners, the
>symbionts form one system and thus are not part of the external
>environment.)  This definition of course excludes (quite rightly) trivial
>cases like the "better nutrition = taller humans" effect which Chris mentioned.

An interesting sidelight her is that there *is* a genetic component in this
system.  The *symbiotic* *microbe* does in contain genes coding for lack of
males in the wasp.  (Shades of night - a gene in a microbe coding for a
feature in its host?!?!).  And this gene is certainly beneficial to the
microbe, and is thus selected for.  What is happening here appears to be a
breakdown of the normal seperation between organisms.

It is certainly admitted that at some point a former symbiont becomes a
part of its host, at which point the distinction between 'genetic' and
'non-genetic' inheritance ceases to be meaningful.  In the case of
mitochondria and chloroplasts most of the genetic control has actually
been transfered to the eukaryotic host (which is the other direction than
the wasp example).  Nonetheless there are a *few* traits that are still
coded for in the organelle itself, leading to strange forms of 'cytoplasmic'
inheritence.  (Look at the 'Eve' studies in anthropoology).


So, it gets quite interesting, the borders of things are actually quite
fuzzy.  Which is very normal in biology.
-- 
---------------
uunet!tdatirv!sarima				(Stanley Friesen)

colby@bu-bio.bu.edu (Chris Colby) (04/29/91)

In article <18844@csli.Stanford.EDU> cphoenix@csli.Stanford.EDU (Chris Phoenix) writes:
>In article <79798@bu.edu.bu.edu> colby@bu-bio.UUCP (Chris Colby) writes:
>>	You are confusing the process of evolution with simple
>>change. If the gene pool of the wasps did not change (ie there
>>are still the same frequency of male and female determining 
>>genes), then the population did not evolve. This is true even
>>if there are more females than males compared to some time in
>>the past. 
>
>This is getting away from my question, but what do you consider the
>gene pool?
>If the bacteria mutated and this caused some change in the
>wasp, would you say the wasp had mutated?

	(Depending on what you mean by change) No, I would say the
environment (in this case an internal endosymbiont) caused a change
in the expression of the wasps genes.

>If you say "no", I have
>another question: would you say that we mutate if our mitochondria
>mutate? 

	Good analogy, in this case I would say yes. Why the difference?
Mitochondria and the rest of our cells are no longer autonomous; they
each depend on each other for survival. I would say that now the gene
pools count as one. In the wasp example, I don't know if the bacteria
could live without the wasp, but the converse is certainly true. For
that reason I would distinquish between the two gene pools.

>But the last theory I heard was that mitochondria used to be
>bacteria that our cells took in, once upon a time.  (Bill, I wrote
>this just before I got your mail.  I'll let it stand as a question to
>others, if they're interested.)

	Yes, this is the endosymbiotic theory. It is generally
accepted.

>Let me give an example to illustrate my point.
>>	Humans have been growing larger and heavier in recent
>>historical times. But this is not an evolutionary change; it
>>is a result of better nutrition and medicine.

>This is an extracellular change.  Obviously it is not a characteristic
>of the individual cells (ie if you pulled a cell out of Plato and a
>cell out of me and grew them in vitro you probably couldn't tell the
>difference in their offspring.)  I am not interested in this kind of
>change. 

>But I am interested in *intracellular* changes which do not involve
>the cell's nucleic acid, and preferably do not involve any nucleic
>acid.  Let me give you a hypothetical example that typefies what I am
>looking for: Suppose a cell has a structure of tubules.  These tubules
>serve as conveyors for various molecules, including some molecule
>necessary to build the tubules. 

	If you could get the tubules to grow in vitro supplying only
a nucleic acid free (or nucliec acid of random composition) cell
extract then I would say it counts as a genetic component. IE. it
contains the necessary info to replicate itself autonomously given
the appropriate environment. I would think this would be synonomous
with the mitochondrial example (mutual obligatory symbiosis).

>Should you distinguish between characteristics caused by expression of
>the genes and characteristics caused by the environment?  Your human
>size example is entirely dependent on the environment.

	Yes, one should distinguish between variation in a char-
actoristic due to environmental and genetic factors; this is key
to understanding evolution. In the human example, much of the variation
through time could be ascribed to environment, but not all. Two short
people are likely to have a short kid no matter how well you feed him/
her.

>Speaking of hormones, I've been following the discussion on hormonal
>control of fetal growth with interest.  But I haven't seen anyone
>address the question of where the hormones come from.  I think all of
>them will be coded for by the DNA of the original fertilized egg.

	Hormones are proteins coded for by genes.

>So a fertilized chicken-egg cell with dinosaur DNA will
>begin by producing dinosaur hormones.  If these hormones spread
>through the growing embryo in the same way (which is purely based on
>the embryo's environment and physical structure) then the cells will
>react to them just like dinosaur cells would.  So the embryo will keep
>developing like a dinosaur, and will become a dinosaur.

	The dinosaur DNA, however, would need the right cellular
environment to function (ie correct transcription factors to
activate the dinosaur genes, the appropriate equipment to pro-
cess dinosaur mRNA once transcribed (ribosomes and factors)). A
chicken cell may have diverged enough that it did not provide the
right environment. I don't know, experiment hasn't been done. 

>I hope this is still interesting for people...

	So do I.

Chris Colby
email: colby@bu-bio.bu.edu

colby@bu-bio.bu.edu (Chris Colby) (04/29/91)

In article <1991Apr25.203311.20957@alchemy.chem.utoronto.ca> mroussel@alchemy.chem.utoronto.ca (Marc Roussel) writes:
>In article <79798@bu.edu.bu.edu> colby@bu-bio.UUCP (Chris Colby) writes:
>>	Remember that evolution is _defined_ as a change in the
>>gene pool; therefore non-genetic evolution is by definition non-
>>sensical.

>     It seems to me that the definition presented above is useful in some
>contexts, but overly restrictive in others.  The wasp example which
>someone else posted is a good example.  Whether or not the wasps' gene
>pool changed, there has been a change in the species which is persistent
>against (one presumes) significant external perturbations.

	How does this differ (in the wasp case) from a persistent
change in a species induced by an external parasite (for ex lower
body weight due to the parasite tapping energy from the host)?
Would you consider that an evolutionary change? (Hint: it isn't)

	[some stuff deleted]

>    My definition has the advantage that it allows us to include in
>evolutionary theory things which we may not have thought of yet.

	So does the current definition. 

>Further, stability of evolved traits as the hallmark of evolution brings
>the language of biology into line with modern thinking in other
>sciences.

	You've been listening to creationists too long if you think
evolutionary biology is lagging behind the other sciences. (Hint:
go browse through recent copies of _Nature_ or _Science_ -big
league science journals- and see if you see any articles about
evolution. Hint: you will)

>(I am curious to hear comments on this last point.  Would people
>think of this as an advantage, a disadvantage, or neither?)
	
	Evolved traits (since they are by definition genetically
based) have been known to be stable since Mendel's work.

>				Marc R. Roussel

Chris Colby
email: colby@bu-bio.bu.edu