Some stuff on Baldwin Effect
- From: "whitesickle@xxxxxxx" <whitesickle@xxxxxxx>
- Date: Wed, 4 Jan 2006 11:41:20 -0500 (EST)
Note: I read through some of the following below article "Organisms can
be proud to have been their own designers" and it stimulated a few
thoughts. First, I think the article exaggerates its case for adaptive
evolution with natural selection as an exception based on the Baldwin
Effect. It states, "Adaptive evolution may not require neither natural
selection nor the inheritance of acquired characteristics. An adaptive
evolutionary change in population without natural selection means that
an identical adaptive change in genetically different organisms of a
population can take place without a systematic difference in the
reproductive value between them, and these changes can also become
irreversible on the level of genome without the difference in the
reproductive value involved. The mechanisms which allow this are known
and sketched in this paper. Their description requires an approach on
the level of whole genome and a look to the organism as a
self-organizing and communicating system. Consequently, it is possible
to have a theory of adaptive evolution, for which the evolution with
natural selection is a special case."
While self-organization plays a role in adaptive evolution it is part
and parcel of natural selection. I'm omitting gene flow and genetic
drift. For example, the development of the human brain can be seen as a
product of emergence and related to self-organization but this can't be
separated from Darwinian evolution. I DO believe in the concept of the
Baldwin Effect of adaptive evolution not requiring natural selection
but don't see how this can occur without being guided or managed by an
outside source.
An online dictionary states, "Self-organization refers to a process in
which the internal organization of a system, normally an open system,
increases automatically without being guided or managed by an outside
source. Self-organizing systems typically (though not always) display
emergent properties. Is Darwinian evolution/natural selection an open
or closed system? The article is operating from the premise of adaptive
evolution without natural selection with the latter being a special
exception. I'm more interested in the idea of Darwinian evolution being
a closed or open system...whether our DNA represents a closed or open
system. For example, will future genetic engineering of *us* lead
largely to a genetic dead end or will it to profound changes consistent
with adaptive evolution and the Baldwin Effect? Or, under the current
definition of open system, can DNA be permeable to mass and energy and
yet efforts at genetic engineering still lead to a dead end?
The Baldwin Effect has often been associated with self-organization and
emergent properties without natural selection. Certainly I wouldn't
regard natural selection (assuming its an open system) as being guided
and managed totally by an "outside source". One could consider the
environment an "outside source" but that doesn't consider the
synergistic relationship between biota and its environment. How does
the Baldwin Effect take into account changing environments and biota
due to the slow effects of natural selection? Under Darwin, species
which DON'T adapt to their environment DIE. That is my understanding.
Am I wrong? Are we so arrogant and naïve to think the same doesn't
apply to ourselves?
We have not been our own designers in terms of biological evolution.
Nor are we today. Instead we are influenced by natural selection,
genetic drift, gene flow, etc. just as many other species are and we
share many of the same behaviors as other species.
Michael Ragland
[Published in: Cybernetics and Human Knowing vol 7(1), 2000, pp.
45-55.]
Organisms can be proud to have been their own designers
Kalevi Kull
Abstract
According to H. F. Osborn, one of the three authors of 'Baldwin
effect', adaptive evolution may not require neither natural selection
nor the inheritance of acquired characteristics. An adaptive
evolutionary change in population without natural selection means that
an identical adaptive change in genetically different organisms of a
population can take place without a systematic difference in the
reproductive value between them, and these changes can also become
irreversible on the level of genome without the difference in the
reproductive value involved. The mechanisms which allow this are known
and sketched in this paper. Their description requires an approach on
the level of whole genome and a look to the organism as a
self-organising and communicating system. Consequently, it is possible
to have a theory of adaptive evolution, for which the evolution with
natural selection is a special case.
Keywords: Self-organisation, autogenesis, Baldwin effect,
biosemiotics, post-Darwinism, individual adaptation, functional genome,
gene duplication, gene conversion, adaptive evolution
1. Introduction: Interpreting the 'Baldwin effect' as a
post-Darwinian mechanism of evolution
"The root-ideas of the conception of evolution are, first
differentiation, and secondly the interaction of the differentiated
products", wrote Lloyd Morgan (Morgan 1898: 487). This very general
statement can be interpreted, for instance, as a differential
reproduction, followed by competition and extinction of one of the
product. However, Lloyd Morgan's emphasis was evidently the
differentiation within an organism, or more precisely, the
differentiation into the non-self and self (Blitz 1992). That is the
problem of what is the role of distinctions made by organism in
evolution.
According to a principal statement of semiotic biology, organism as a
subject has activity and intentionality (cf. Uexküll 1940, Searle
1993), which are the main essentials of life. Also, if life has any way
to influence its evolution, these features of the subject should play
the role in this. However, it has not been easy to find an evolutionary
mechanism which may correspond to this view.
The role of self-organisation is generally accepted as evident for
ontogenesis. Also, a population can be seen as a self-organising system
in evolution. However, the question I would like to analyse here is
whether an organism can be seen as a self-organising system over the
generations, as a leader and designer of its own evolution.
The view according to which the self-organisation of an organism is
limited to its ontogenesis, comes from the well-known assumption that
the organism cannot influence in any way the internal determinants of
its offspring, i.e., its genome is inherited independently of the
behaviour during its lifetime. Still, for instance, F. J. Odling-Smee
(1994) has emphasised that organisms can influence the external
determinants of its offspring, via choosing or changing the environment
in which the offspring will develop. This is, however, not enough to
make any change irreversible, that is evolutionary. Thus, he requires
the natural selection in the second step to make the change
irreversible. My statement here is that this is not necessary. In order
to solve the problem we need to look how an organism is operating its
memory, and we also need to be precise in using and defining the notion
of natural selection.
One of the interesting alternative approaches to evolution was proposed
a century ago by paleontologist Henry Fairfield Osborn (1857-1953)
and psychologists James Mark Baldwin (1861-1934) and Conway Lloyd
Morgan (1852-1936), known as the concept of organic selection or
'Baldwin effect' (Baldwin 1896; Morgan 1986; Osborn 1896). The
Baldwin effect states that the ability of individuals to learn can
guide the evolutionary process. This effect was later analysed by G. G.
Simpson (1953) and C. H. Waddington (1953a, b). Waddington has used it
to develop his model of genetic assimilation. Recently the Baldwin
effect has gained attention again, which is expressed by the book of R.
K. Belew and M. Mitchell (1996), and several other publications
(Emmeche 1994; Deacon 1997; Jablonka et al. 1998; Kull 1998, 1999;
Ancel 1999; Robinson, Dukas 1999; cf. also Bowler 1992, Richards 1987).
With the work of Hinton and Nowlan (1987) the Baldwin effect came to
the attention of computer scientists, and since then it became a tool
in evolutionary computation (Turney et al. 1996, Turney 1996, Harvey
1996). Here, I propose a possible mechanism for the 'Baldwin
effect', which uses the concept of phenotype as an interpreter of
genotype, and develops the interpretation given by Simpson and
Waddington in a more radical way (e.g., as emphasised by Osborn (1896)
- adaptive evolution may not require neither natural selection nor
the inheritance of acquired characteristics, but may use natural
selection in some cases).
2. A mechanism of microevolution
The mechanism to which I would like to draw attention, can be briefly
described as consisting of the following statements. Most of these
steps are quite trivial, however, the consequences from the whole set
will not be so trivial.
(1) We need to notice that an organism can choose, which parts of its
genome to use. Accordingly, the genome includes a functional and
non-functional part. In most eukaryots, only a minor part of the genome
is in use. The unused part consists (a) from non-coding DNA, which,
still, may include some pseudogenes, and (b) from coding DNA, which is
not currently used, but can be used in some other circumstances (e.g.,
in other cells of the organism, other period of ontogenesis, or other
environmental conditions).
(2) An organism is able to change the used (functional) part of the
genome during its lifetime, as dependent, for instance, on the
conditions in which it lives.
(3) This change can be adaptive (due to self-organising nature of the
organism) even if the conditions are new, that means if the particular
combination of environmental factors has never been experienced before
by this organism during its phylogeny. This derives from the nature of
cell, as the cell is an adaptive system that changes its state
continuously according to the communicative activity of its functional
cycles.
(4) An organism as a self-organising system is able (due to its
mobility and an ability temporarily not to move; and also, due to its
ability to distinguish between different environmental situations) to
choose the environment for living.
(5) If a population of similar organisms will meet new environmental
conditions (when moving to a new place, or when the conditions change
by themselves, or when the change in conditions results from the
organisms local activity), then an adaptive change in the usage of
genome may take place almost simultaneously for most of the specimens
of the population.
(6) If the population keeps living in the new conditions (either due to
the organismal preference for it, or due to the constancy of the new
conditions), then the change in the usage of genome can be kept
(repeated) over a number of generations.
(7) In the case of biparentally reproducing organisms, the organisms of
a population keep to be similar. This is because if the genomes of the
mates are not similar enough (i.e., recognisable or complementary),
they cannot reproduce. Thus, the mortality due to the occasional big
genetic changes is not specific to a genotype, but depends on the
difference from the population mean.
(8) Due to the great similarity of individual genomes in a population,
the simultaneous adaptive change in the usage of genome (in gene
expression) may concern the same or similar areas of the genome in most
of the organisms of the population.
(9) Some stochastic mutations, which appear in the area of the genome
which is used (expressed) in the current conditions, lead to
inviability of a part of the offspring. This appears with a probability
which is proportional to the used part of the genome. Considering that
the size of the expressed part of the genome does not differ
significantly between the individuals of the population, it means that
also the probability for such mutations cannot differ significantly
between the individuals. I.e., the corresponding offspring mortality is
not specific to a particular parental genotype.
(10) The mutations which appear in that currently unused fraction of
the genome which has been in use in the previous environment, make the
return to the previous functioning of the genome impossible. There is a
great number of such mutations which may lead to this irreversibility.
Since they appear in non-functional part of the genome, they do not
influence the viability of organisms in the current conditions
specifically.
Natural selection is defined as differential reproduction of genotypes
(this being used as the standard definition of natural selection
according to Dobzhansky et al.). Differential reproduction means here
that there is a non-random (i.e., statistically significant) difference
in reproductive value between the groups in a population which
distinguish by some genetic marker. This type of difference was not
required to reach the statement (10), i.e., non of the statements
(1-10) assume any necessary difference in reproductive value between
specific genotypes.
Thus, the statements (1)-(10) described the way of adaptive
evolutionary change without a need for natural selection. This way of
evolution means that some genes may be dropped out from the expressible
genome during an adaptive specialisation of a population without any
differential reproduction of genotypes involved. It can be called an
evolution via forgetting of unused.
The mechanism described here is a Baldwin effect, since it assumes the
organisms' adjustment, or individual adaptation, which can be seen as a
form of individual learning. Accordingly, we have described a possible
particular mechanism, which allows individual learning to guide certain
type of evolution without natural selection being its requirement.
The importance of this mode of evolution concerns the possibility to
have an adaptive evolution without natural selection. There have been
described several ways of evolution which do not require natural
selection, the most well-known among them being the random walk, i.e.
neutral evolution. However, these cannot create adaptations. In this
sense the mechanism described here is exceptional.
3. Can a growth of functional genome be non-selectional?
If the forgetting of unused would be all of evolution, it would mean a
progressive narrowing of working genetic memory. Since this is not
generally the case in real evolution (one can find an evidence for this
probably only in some examples of specialisation), we need also to
analyse the mechanisms in which the genetic memory can grow.
It is very improbable that in many individuals of a population an
identical growth of their genome occurs simultaneously. Consequently,
the growth of functional genome without the work of natural selection
would be much less probable than its decrease. Still, the current
knowledge is not sufficient to state that the lower probability would
mean impossibility.
A phenomenon which appears very interesting in this context is the
ability of homologous chromosomes to stay extremely similar. This is
usually explained as a result of homolog recognition - in the case of
big differences meiosis does not go correctly and the sexual
reproduction will be inhibited. However, there are known few additional
exciting details. In the course of recombination also certain
recombination repair of DNA takes place. At least in some cases, it may
include a mismatch repair between the strands of DNA which are derived
from different parents. In this case, gene conversion can occur, where
the mismatch repair can convert one allele into the other. This
phenomenon can be detected through the offspring non-Mendelian ratios,
and is frequently observed, e.g., in fungal crosses (Stacey 1994).
Whether the gene conversion may account for distribution of gene
duplications in a population, we do not know. However, this seems to be
a reasonable hypothesis. Namely, assuming that gene conversions can
lead to an appearance of a gene duplication in the DNA strand derived
from the parent that did not possess it, and, at least sometimes, with
higher probability than the dropping out of the duplicated gene, then
this may have important consequences for the genome evolution. This
phenomenon - a rapid non-Mendelian distribution of a gene duplication
in population - if it occurs, should be quite rare. The speed of real
evolution is not great.
Considering the hypothesis above, we may sketch a mode of progressive
evolution without natural selection.
(1) The genome may grow, for instance, through duplication of genes.
Generally, there is no reason to assume that the reproductive values
differ significantly between the organisms which have a particular
duplication and which have not. The duplication can be not an exact
copy, but still a gene, which is simply not expressed and not needed
when it appears. A duplication, as well as any other mutation, can
spread over the population by random walk. In addition, it can be that
some recombination repair mechanisms enhance the distribution of
duplications.
(2) In the case of several copies of a functional gene, one may mask
the existence and slight differences of the other. The changes in the
masked copy are nevertheless restricted and kept similar within a
sexual population, in order to allow the sexual recognition (on the
level of pairing the homologous chromosomes) to take place.
(3) If most of the organisms of the population have obtained in this
way a masked and slightly different copy of some genes, these can be
used in a new way when the behaviour of the organisms will change in
the changed conditions of the population. This can be described as we
did through (1)-(8) in the previous chapter. The only difference is
that here a newly appeared gene will be taken into a use, i.e., the
functional genome may become larger, whereas in the previous case it
could only become smaller.
Thus, also here, we can see a mode of evolution which may occur without
differential reproduction (i.e., without significant differences in the
number of viable offspring between the different genotypes).
Still, a small nuance may be needed to explain in relation to the
notion of natural selection. If a gene is distributing in a population
in result of random walk, this is not natural selection, according to
its definition. But if the distribution is accompanied by gene
conversion, then the distribution can be shifted from random. However,
a non-equal copying itself is usually not assumed for being a
constituent of selection, it rather belongs to a special sort of
mutation. Therefore, to be more precise in defining natural selection,
we need to add that this is a significant difference in the viability
of offspring between the groups of parents with a particular genotypic
difference, except the difference which is resulted from non-identical
replication.
Accordingly, also in the case of the growth of functional genome, we
can see (at least hypothetical) possibility for a non-selectional
adaptive evolution.
4. Further comments on adaptation
Thus, we reach the conclusion that the adaptive evolution
(specialising, and may be also progressive) can occur without natural
selection necessarily involved. This does not mean, of course, that
natural selection is not working in evolution - there are many
examples documented where it does. However, this analysis demonstrates
that adaptive evolution is a more general process than the adaptive
evolution via natural selection. Which means that Baldwin effect
indicates to a possibility for a generalisation of theory of evolution.
It requires a special experimental work to find out in what extent the
adaptive evolution in the wild is non-selectional. Theoretically, it
allows much higher speed of adaptive evolutionary specialisation than
the evolution restricted by the mechanism of differential reproduction.
Consider the following scenario. A population of organisms will move
from one place to a new one, where the environment (e.g., food) will be
different from the initial one. As a result, the organisms of this
population will all need to use a part of their genomes which was not
used before (i.e., expressing other genes), and will not use another
part of the genome which was used before. If the individuals have an
ability for individual adaptation, which will include a change in gene
expression, then they can do it. Assume that the population will stay
in its new place, and thus the change in the usage of genome will be
preserved. Then, the stochastic mutations in the non-used region of the
genome fix this change on the genetic level and thus make it
irreversible, i.e., evolutionary.
Thus, the appearance of a new adaptation may occur during one
generation as a response of the organisms' self-organisation and
communication, and simultaneously for the whole population. The genetic
fixation of this change will take, of course, many generations
afterwards (or, in the case of gene duplications, it needed a number of
generations before it for the distribution of the duplication).
However, it is still enabling much quicker evolution than the case
where the new adaptation first appears in a single organism (a mutate),
and then via the competitive advantage distributes over the population.
This also indicates to a solution in the debate between punctualism and
gradualism. The data which led to the formulation of punctualism came
from the morphological studies of phylogeny, showing the periods of
stasis and change. The evidence from the molecular data, vice versa,
shows that the lineages change gradually. This is exactly what the
mechanism as described above predicts - the morphological change and
the genetic change may not be necessarily concurrent. E.g., the
morphological change may initially be an ontogenetic adaptation which
develops quickly, and it will be afterwards fixed in the genome
gradually.
The main statement from which the conclusions given here follow is that
for an organism, (a) there exist many different ways to behave and
build itself in the case of the exactly same genome, and (b) there
exist possibilities to behave in the same particular (constant) way for
a quite large variety of genomes. In other words, (a) an organism has
many ways to interpret its genome, and (b) there exist inheritance
mechanisms in addition to the genetic one (e.g., epigenetic, or just
the stability of the environment) which enable for an organism to keep
some features of its structure and behaviour unchanged even if some
changes in the genome take place.
Notably, the organism's phenotype and genotype are largely uncoupled
because (generally) an organism may not require its whole genome for
living, and there exist potentially more functionally expressible parts
in the genome than those which are currently in use. I.e., there are
many ways to live using the same genome, as, for instance, there are
many cell types in a multicellular organism which are using different
parts of the same genome. However, this freedom appears due to
behavioural activity (which includes perception and operation, i.e.,
the functional cycle, according to Uexküll (1928)), and particularly,
due to mobility, so changing and selecting the environment. Thus, the
uncoupling is a result of co-work of two levels of functional circles
which cells (or phenomes) possess - one of these acting toward the
genome, the other toward the environment.
A cell, having several ways to interpret its genotype (which means that
the cell can change the expression of particular genes and, together
with this, change the part of the genotype which is in use), also has
several ways to preserve a particular interpretation over a number of
generations (through epigenetic inheritance mechanisms, or due to a
permanent change of environmental conditions). This gives to a
stochastic genetic changes occurring in an unused part of the genotype
a time to accumulate and to fix the otherwise only phenotypic changes.
For instance, this means that if a particular cell type has not been
formed during many generations, this may not be possible to form it
later again, due to stochastic changes in the part of the genome used
by (required for) this type of cells.
It is important to admit that a similar interpretational shift can take
place simultaneously in many individuals of a population (e.g., as a
result of invasion of the population into a new environment, for
instance in the case of monophage or oligophage insects when they
inhabit a new host species). The phenetic shift which this implies may
be sufficient to decrease the efficiency of recognition of the source
population specimens (which is needed for mating) down to the level
which guarantees the sufficient isolation and provides time to the
mutation processes to fix this separation also on the level of genome
or cytoplasm incompatibility (Kull 1993, 1988, Paterson 1993).
Here, I would like to draw attention to an interesting paradox of
natural selection, which can be called the paradox of unique child.
That is, in the case of sexual reproduction, almost every descendant
has the genotype which has never been present before (e.g., in the
sense of a new combination of alleles of the whole genome), i.e., which
survival has never been checked by natural selection. Nevertheless, the
offspring usually includes (particularly in the species which have low
reproduction rate) a great percentage of individuals who stay alive.
A solution of this paradox states that every organism has many ways to
carry out the tasks they need to fulfil - the unique genetic memory
may not destroy this ability, and if some structures happen to be
corrupted, one can find several others in itself which still work, or
which may be put into work. Little quantitative changes in an
efficiency of one particular enzyme can often be compensated by small
changes in the production of other enzymes, without real influence to
the reproduction. Also, for the most part, it is not necessary for the
living organism to be digitally precise.
For a new character to appear in the phylogeny of species, it is not
necessary to assume that there had to be one specimen who gained this
character first due to mutation, and then this mutation had to be
spread over the species to all those whose grandparent that first
mutant individual is. This would be required only if the digital
preciseness in the determination of the character is assumed. As far as
this is not the case, many different genetic changes in many
individuals of the population may be simultaneously behind the same new
character. And the genetic fixations of the new character (in the sense
of making its appearance irreversible) could have been taken place
after the first appearance of this character in the paleontological
record.
Baldwin effect implies that evolution may take place without
differential reproduction of genotypes. Assuming that the mutations in
the expressed part of the genome cause the inviability of a certain
(generally - about equal) percentage of offspring in all individuals
of the population involved in the interpretational shift, we have a
mechanism of evolution which works without the differential
reproduction of genotypes. The neo-Darwinian mechanism is thus a
special case of this mechanism, since it requires an additional
assumption (e.g., that the percentage of inviable offspring is
systematically different in different individuals, and this difference
is correlated with a particular genetic character of parents).
The stochastic (entropic) changes in genotype preferentially lead to
the forgetting of "unused" and the keeping of "used". Due to
the large size of genomes in terms of the number of genes, there are
always many mutations which simultaneously distribute between the
individuals via sexual reproduction and thus enable the ontogenetic
change to become fixed for the whole population (making thus the
phenotypic change irreversible); theoretically, this is much quicker
than according to the classical mechanism which requires the
distribution of new mutations across the population through competitive
advantage.
The proposed 'Baldwin effect' mechanism is supported by recent
studies in epigenetic inheritance mechanisms (Jablonka, Lamb 1995), and
the stability of morphogenetic mechanisms (Webster, Goodwin 1996).
As a consequence of this mechanism, the activity of an organism as a
subject may play a role as an evolutionary factor. This is exactly what
the Baldwin effect originally claims. This also directs our attention
toward the functioning and evolution of the mechanisms of
interpretation (i.e., semiosis).
5. Conclusion: On the role of selection
It should be interesting to compare the behaviour of the described
model with the classical neo-Darwinian or synthetic theory of
evolution, and to investigate its applicability to the evolution of
linguistic systems.
According to the neo-Darwinian explanation (which is accepted in the
synthetic theory of evolution), the ability for learning (or
plasticity) as a characteristic of a phenotype may appear and develop
as a result of selective advantage of the corresponding genotype.
According to the semiotic view, and according to the explanation of
Baldwin effect as described above, the results of ontogenetic learning
themselves provide a factor and direction of evolution.
Jablonka et al. (1998) also see in the Baldwin effect an important
aspect of interpretation of evolution. They demonstrate the existence
of at least four different inheritance systems, all, according to them,
having "the properties necessary for Darwinian evolution" (Jablonka
et al. 1998: 209). However, they see it still in the framework of the
Darwinian mechanism of natural selection, as a Lamarckism within the
Darwinism.
In order to explain the relationship between the natural selection and
the mechanism described above, we need a very clear definition of
natural selection. Assuming that natural selection necessarily requires
a statistically significant difference in the survival between the
offspring of two genotypically distinguishable sets of organisms, we
may conclude that, according to the Baldwin mechanism (as described
above), the natural selection is not necessary for evolution (whereas
the notion of evolution is still kept in its traditional definition as
the irreversible change in the genetic structure of population).
Because if the offspring of two genotypically distinguishable sets of
organisms has the same percentage of non-viable organisms (e.g., if
these sets are indistinguishable according to their mortality, or some
other accepted measure of survival), there is no natural selection in
this case, according to the definition accepted.
The synthetic theory of evolution shows that evolution may occur
without natural selection, but in this case it is non-adaptive. We
tried to show here that this can also be adaptive.
Natural selection may certainly be effective in the case when the
selective factor (e.g., an antibiotic) is very strong, so that
population number decreases down to few individuals who only survive.
But if the population is permanently large, then natural selection
cannot be very effective; however, the Baldwinian mechanism may still
effectively work and consequently the evolution may go on.
Since the mechanism of Baldwin effect as described here is more general
than the Darwinian mechanism of natural selection (as differential
reproduction), and since the latter can be seen as a special case of
the former, it seems to be accurate to use the term post-Darwinism as a
name for this view (cf. Amundson 1998; Sermonti, Sibatani 1998; Kull
1999).
There certainly is selection in evolution. Most generally, the subject
of selection is organism (cf. Weingarten 1993).
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Notes
A longer version of this paper was presented at the Bennington
International Conference on Mind and Brain: An Integration of
Evolutionary Origins and Emerging Developmental Principles, in November
10, 1999, Bennington, Vermont, USA. Correspondence: K. Kull, Department
of Semiotics, University of Tartu, Tiigi Str. 78, Tartu, Estonia; or
Institute of Zoology and Botany, Riia Str. 181, Tartu, Estonia. E-mail
kalevi@xxxxxxx
I thank Mart Viikmaa, Sören Brier, and Terrence Deacon for helpful
comments to an earlier version of this paper.
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