[sci.bio] The Red Queen

werner@aecom.UUCP (Craig Werner) (12/17/86)

	It has often been argued that biological change (evolution, I daresay)
can occur even in the absence of environmental change.  To some this has
been used to discredit the entire evolutionary scenario.  However, it
was proposed (and I wish I know by whom) that interspecies competition may
play a more important role than previously assumed. They dubbed this
theory "The Red Queen Hypothesis", after the Red Queen's advice to Alice
in "Through the Looking Glass":  
	"Sometimes you have to run as fast as you possibly can just to 
stay in the same place."

	This (in my mind) has got to be the best-named theory in science.
And it also echoes the prefacery remarks in (again I don't remember which
text):  this book is not designed to be an introductory text.  The best
introductory text on this subject is Alice in Wonderland, but then again,
that goes for most subjects.

-- 
			      Craig Werner (MD/PhD '91)
				!philabs!aecom!werner
              (1935-14E Eastchester Rd., Bronx NY 10461, 212-931-2517)
		     "I tell you I'm a thief and you call me a liar?"

gallagher@husc4.harvard.edu (paul gallagher) (12/19/86)

In article <741@aecom.UUCP> werner@aecom.UUCP (Craig Werner) writes:
>
>	It has often been argued that biological change (evolution, I daresay)
>can occur even in the absence of environmental change.  To some this has
>been used to discredit the entire evolutionary scenario.  However, it
>was proposed (and I wish I know by whom) that interspecies competition may
>play a more important role than previously assumed. They dubbed this
>theory "The Red Queen Hypothesis", after the Red Queen's advice to Alice
>in "Through the Looking Glass":  
>	"Sometimes you have to run as fast as you possibly can just to 
>stay in the same place."
>

In the past (till the 1930's) many people believed in "internal" forces
directing evolution, with selection merely working to weed out the unfit.
These ideas are pretty much completely discredited.
Going to the opposite extreme,
the New Synthesis in the 1950's rather dogmatically asserted that almost all
evolutionary change was directed by adaptations to the environment.
Nowadays, people have begun to realize that evolution is controlled not
only by adaptations to the environment, but also by the constraints of
development, history, and body architecture - and also perhaps by chance.
Much allelic substitution occurs without controlling influence from
selection, and with no direct relaionship to adaptation.
An excellent source dealing with the profound changes occuring in evolutionary
theory - which far from discrediting it, improve its ability to explain the
history of life - is Stephen J. Gould's "Is a new and general theory of
evolution emerging" in Paleobiology, 6(1), 1980, pp. 119-130.

gallagher@husc4.harvard.edu (paul gallagher) (12/20/86)

In article <741@aecom.UUCP> werner@aecom.UUCP (Craig Werner) writes:
>	It has often been argued that biological change (evolution, I daresay)
>can occur even in the absence of environmental change.  To some this has
>been used to discredit the entire evolutionary scenario.  However, it
>was proposed (and I wish I know by whom) that interspecies competition may
>play a more important role than previously assumed. They dubbed this
>theory "The Red Queen Hypothesis", after the Red Queen's advice to Alice
>in "Through the Looking Glass":  
>	"Sometimes you have to run as fast as you possibly can just to 
>stay in the same place."

I think this refers to "character displacement".  When the ranges of two
similar species overlap, one may outcompete the other and cause its
elimination.  Alternatively, they can develop strategies to live together
in the same range.  One way is spatial segregation - one species lives in
one strictly localized habitat, the other lives in another so that they
avoid direct competition.   Another way is character displacement.  Either
one species comes to look very similar to the competing species, so that
selection acts upon all individuals as if they were just one species,
allowing the species to coexist, or the two species diverge in characters
so that the two species have slightly different habitats and thus again
can coexist.
For example, certain closely related agnostid trilobites seem to coexist
through size displacement.  It was predicted that size displacement would
have to be in the range of 1.28:1 in order to avoid competitive elimination.
The fossils in question were investigated and did indeed show ratios in
size of about 1.28:1.
Also, in the small area where the ranges of the trilobites Phacops iowensis
and Phacops rana overlap, P. iowensis diverged in morphology from P. rana,
while P. rana converged in some characters, diverged in others.
The source is Neil Eldredge, "Character displacement in evolutionary time"
American Zoologist 14 (1974), 1083-97.
However, Eldredge concludes that character displacement is not that important.


Another way a population might change without any change in its environment
is genetic drift.  Any very small population has only a limited, random 
sample of the genetic variability within a species, and whenever two
individuals reproduce,  some alleles will be lost, since only one chromosome
of each pair is passed down to the gametes.  Thus, in a small population,
there could be an evolutionary change in gene frequencies just through random
chance assortment.
Again, this process may not be very important.


Incidently, another way evolutionary change can occur far in excess of simple
responses to changes in the environment, is through changes in the timing
of development.
For example, neoteny: the occurence of sexual maturity in the juvenile animal.
It is believed that the early fish-like ancestors of vertebrates originated
when the free-swimming larvae of sea-squirts became able to reproduce, thus
eliminating the characters of the adult stage.
Similarly, the onset of maturity in the larvae of crustaceans may have given
rise to the small planktonic copepods.
Also, paedomorphosis: the retention of juvenile characters in the adult.
For example, in Cambrian trilobites the visual surface of the eye was surrounded
by an ocular suture - which caused the visual surface to be lossed during 
molting.  Probably through paedomorphosis, post-Cambrian trilobites abandoned
this system, replacing it with the juvenile form where the visual surface is
attached directly to the inner body.


In general, many people once thought that organisms were ideally adapted to
their environment.  So, in order to explain why they changed through time,
they had to say that their environment changed and that each change in a
characteristic corresponded to a perfection of its ability to function in
its environment.  Now, people realize that things are much more complex.
Stephen J. Gould uses a metaphor originated by Galton - the organism is not
a sphere which can be turned in any direction and any increment by changes
in the environment.  It is a polyhedron, each of whose faces is a point of
equilibrium.  Small environmental change will not turn the polyhedron to a
new face, and a big push which succeeds in changing the face of the polyhedron
may bring with it a whole set of changes not accounted for by natural
selection.
"Most of the changes in evolutionary viewpoint that I have advocated...fall
out of Galton's metaphor: punctuational change at all levels (the flip from
facet to facet, since homeostatic systems change by abrupt shifting to new
equilibria); essential non-adaptation, even in major parts of the phenotype
(change in an integrated organism often has effects that reverberate
throughout the system); channeling of direction by constraints of history
and developmental architecture.  Organisms are not billiard balls, struck
in deterministic fashion by the cue of natural selection, and rolling to
optimal position on life's table." - (Stephen J. Gould. "Is a new and
general theory of evolution emerging?" Paleobiology, 6(1), 1980, p. 129).


Paul G.

jc@cdx39.UUCP (John Chambers) (12/23/86)

> >It has often been argued that biological change (evolution, I daresay)
> >can occur even in the absence of environmental change.  To some this has
> >been used to discredit the entire evolutionary scenario. 

Anyone that argues that environmental changes are necessary to drive
evolution is simply ignorant of the low-level mechanisms.

Consider the scenario:  A stray alpha particle comes zipping along, 
smashes through the fringes of a DNA helix, and when the electrons
settle down again, a nucleotide or two have been changed.  About 3
times out of each billion such occurrences, this is in a cell that
produces a sperm or ovum, and the change is passed on to offspring.

Furthermore, there's no problem coming up with 'constant' features
of the environment that exert selective pressures.  One such is
called 'predators'.  Even if the predators were unchanging, they 
would still select for avoidance capabilities in a prey species. 

Alpha particles and predators are part of the environment of
all living creatures.

[OK, a flying alpha particle isn't a feature of a 'constant'
environment.  But in that sense, there is no evolution in a
truly constant environment, because everything is at absolute 
zero and there is no life.]

-- 
	John M Chambers			Phone: 617/364-2000x7304
Email: ...{adelie,bu-cs,harvax,inmet,mcsbos,mit-eddie,mot[bos]}!cdx39!{jc,news,root,usenet,uucp}
Smail: Codex Corporation; Mailstop C1-30; 20 Cabot Blvd; Mansfield MA 02048-1193
Clever-Saying: For job offers, call (617)484-6393 evenings and weekends.

michaelm@bcsaic.UUCP (Michael Maxwell) (01/05/87)

In article <927@husc6.UUCP> gallagher@husc4.UUCP (paul gallagher) writes:
>For example, certain closely related agnostid trilobites seem to coexist
>through size displacement.  It was predicted that size displacement would
>have to be in the range of 1.28:1 in order to avoid competitive elimination.
>The fossils in question were investigated and did indeed show ratios in
>size of about 1.28:1.

I suppose I'm being lazy (I could look up the reference), but I gather that
these species of trilobites were quite similar morphologically, and lived in
quite similar (or identical) environments.  Has anybody suggested that these
were actually individuals of the *same* species (possibly of different sexes)?
I know next to nothing about trilobites, but it certainly sounds suspicious...
I seem to recall that several living `distinct' species have turned out to be
members of the same species which underwent fairly radical changes during
ontogeny (Parrotfishes come to mind).  BTW, how easy is it to tell male and
female trilobites apart?

>Another way a population might change without any change in its environment
>is genetic drift.  Any very small population has only a limited, random 
>sample of the genetic variability within a species, and whenever two
>individuals reproduce,  some alleles will be lost, since only one chromosome
>of each pair is passed down to the gametes.  Thus, in a small population,
>there could be an evolutionary change in gene frequencies just through random
>chance assortment.
>Again, this process may not be very important.

I suggested this in answer to an exam question once, as the origin of blind
(eyeless) cave fish.  (The incidence of eyeless fish is quite high, but for
obvious reasons the eyeless ones seldom make it very far in life up here.)
The professor didn't like my answer...his point was that there had to be a
selective advantage to blindness in cave life.  I didn't believe so at the
time, and I'm still skeptical.  Anyone care to comment?
-- 
Mike Maxwell
Boeing Advanced Technology Center
	arpa: michaelm@boeing.com
	uucp: uw-beaver!uw-june!bcsaic!michaelm

dean@violet.berkeley.edu (Dean Pentcheff) (01/07/87)

In article <124@bcsaic.UUCP> michaelm@bcsaic.UUCP (Michael Maxwell) writes:
>I suggested this [genetic drift] in answer to an exam question once, as 
>the origin of blind
>(eyeless) cave fish.  (The incidence of eyeless fish is quite high, but for
>obvious reasons the eyeless ones seldom make it very far in life up here.)
>The professor didn't like my answer...his point was that there had to be a
>selective advantage to blindness in cave life.  I didn't believe so at the
>time, and I'm still skeptical.  Anyone care to comment?

I'm not sure, but I suspect that eyelessness in cave fish arose
independently from several groups of (eye-bearing) fish.  If this is
the case, it seems unlikely that cave fish stemming from different
ancestral groups would share the _same_ trait due to drift, given that
there are only a few characters that tend to be different in cave fish
(lack of eyes, lack of pigment).  If, on the other hand, the trait
conferred a selective advantage, then its appearance in several groups
is reasonable.  The advantage (presumably) has something to do with
energy savings for the eyeless fish (though this is a rampantly
"selectionist" argument).

-Dean	(dean@violet.berkeley.edu)

michaelm@bcsaic.UUCP (Michael Maxwell) (01/09/87)

In article <2103@jade.BERKELEY.EDU> dean@violet.berkeley.edu (Dean Pentcheff) writes:
>In article <124@bcsaic.UUCP> michaelm@bcsaic.UUCP (Michael Maxwell) writes:
>>I suggested this [genetic drift] in answer to an exam question once, as 
>>the origin of blind
>>(eyeless) cave fish.  (The incidence of eyeless fish is quite high, but for
>>obvious reasons the eyeless ones seldom make it very far in life up here.)
>>The professor didn't like my answer...his point was that there had to be a
>>selective advantage to blindness in cave life.  I didn't believe so at the
>>time, and I'm still skeptical.  Anyone care to comment?
>
>I'm not sure, but I suspect that eyelessness in cave fish arose
>independently from several groups of (eye-bearing) fish.  If this is
>the case, it seems unlikely that cave fish stemming from different
>ancestral groups would share the _same_ trait due to drift, given that
>there are only a few characters that tend to be different in cave fish
>(lack of eyes, lack of pigment).  If, on the other hand, the trait
>conferred a selective advantage, then its appearance in several groups
>is reasonable.  The advantage (presumably) has something to do with
>energy savings for the eyeless fish (though this is a rampantly
>"selectionist" argument).
>
>-Dean	(dean@violet.berkeley.edu)

This is what the prof said--there had to be a selective advantage.  My feeling
is that there simply couldn't be any selective DISadvantage--as there is in
fact no disadvantage to blindness or lack of pigment in caves.  There
WOULD be a disadvantage to almost any other simple change, hence the "only"
changes that show up in the distinct groups of cave fish are these two.
(Improved tactile perception would be an ADvantage--but probably much more
unlikely to arise in a short time frame than lack of eyes or pigment.)  Oh, one
other difference that has appeared in at least some groups of cave fish (I
doubt whether it has been extensively tested, it may appear in others) is
lack of circadian rhythms.  Again, it seems that if this were lost from a
couple individuals in a very small population, it might be perpetuated simply
by chance (i.e. due to chance fluctuations in the population which might kill
off all but their offspring).
-- 
Mike Maxwell
Boeing Advanced Technology Center
	arpa: michaelm@boeing.com
	uucp: uw-beaver!uw-june!bcsaic!michaelm

dbb@aicchi.UUCP (01/10/87)

In article <124@bcsaic.UUCP> michaelm@bcsaic.UUCP (Michael Maxwell) writes:
>
>I suggested this in answer to an exam question once, as the origin of blind
>(eyeless) cave fish.  (The incidence of eyeless fish is quite high, but for
>obvious reasons the eyeless ones seldom make it very far in life up here.)
>The professor didn't like my answer...his point was that there had to be a
>selective advantage to blindness in cave life.  I didn't believe so at the
>time, and I'm still skeptical.  Anyone care to comment?
>-- 

Well... Some thoghts on this;  First, it seems to me to be possible that traits arise that are simply no DISadvantage.  These traits may have disadvantageous 
effects in a different environment, but are OK in the special environment in
which they arise.  Blind fish would not survive well in a sunny pond.

After some number of the population is blind, we must THEN start to look for
some advantage to blindness.  Perhaps the production of dyes for use by the
retina saps the cave fish of amino acids that are not often found in his
world.  If there were a small famine due to unusual surface conditions, the
sighted fish might die faster than the blind ones.  Repeat this pressure many
times over thousands of years, and you have a totally blind fish population.

It is also possible that the sighted fish have a smaller range than the blind
ones.  They might choose to remain in the parts of the cave where light filters
in from the surface.  The blind fish, with no such tropism, would be free to
populate the nether regions...

Any constructive thoughts on these opinions?

-- 
-David B. (Ben) Burch
 Analysts International Corp.
 Chicago Branch (ihnp4!aicchi!dbb)

"Argue for your limitations, and they are yours"