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Is Common Descent an Axiom of Biology?
[Editorial note: The following discussion paper was written
for the conference, The Darwinian Paradigm: Problems and Prospects,
held June 22-25, 1993, at the Pajaro Dunes beach community on Monterey Bay,
near Watsonville, California. The conference was organized by Phillip Johnson.
Attendees included Michael Behe, Walter Bradley, John Angus Campbell, William
Dembski, Dean Kenyon, Stephen Meyer, Paul Nelson, David Raup, Siegfried Scherer,
Jonathan Wells, and Kurt Wise.]
To:
Pajaro Dunes Conference Participants
From:
Paul Nelson and Jonathan Wells
Date:
15 June 1993
Re: Discussion
paper for Topic Area I (homology, etc.)
This paper is intended to highlight key points in the readings (being sent
to the participants by Phil Johnson), and to suggest some general issues for
discussion. The issues we raise do not, of course, exhaust the possibilities.
The readings are:
- Thomas Jukes and Syozo Osawa, "Recent Evidence for Evolution of the
Genetic Code," in Evolution of Life, eds. S. Osawa and T. Honojo.
New York: Springer-Verlag, 1991, pp. 79-95.
- V. Louise Roth, "The Biological Basis of Homology," in Ontogeny
and Systematics, ed. C.J. Humphries. New York: Columbia University Press,
1988, pp. 13-29.
- G.P. Wagner, "The Biological Homology Concept," Annual Review
of Ecology and Systematics 20 (1989): 51-69.
- E.B. Wilson, "The Embryological Criterion of Homology," Biological
Lectures Delivered at the Marine Biological Laboratory of Wood's Holl,
Summer Session of 1894. Boston: Ginn & Co., 1895
The Neo-Darwinian Explanation for Homology
We start by reviewing some familiar ground. The conventional neo-Darwinian
explanation for homology (call it the conventional explanation) is clearly outlined
by Romer and Parsons:
Homologous organs are those which are identical the same in
the series of forms studied. But what do we mean by "the same"?
One tends, unthinkingly, to believe that the same actual mass of material, the very same limb or leg or
bone, has been handed down, generation to generation, like an heirloom. This is quite
absurd, but such a concept has obviously influenced, unconsciously, the minds
of many workers.
In reality, of course, every organ is re-created anew in every generation, and
any identity between homologues is based upon the identity or similarity of
the developmental processes which produce them.
The development of the science of genetics has given us a firm base for the
interpretation of these processes. They are controlled by hereditary units,
the genes....If the genes remain unchanged from generation to generation,
the organ produced will remain unchanged (apart from environmental effects
upon an individual which may be obvious, but are not inherited by the next
or later generations), and the homology is absolute.
Changes, however, do occur in genes, as mutations; these mutations produces
changes in the structures to which the genes give rise....In a sense, a study
of organ homology is merely a study of phenomena produced by genes. If the
genetic constitution of all animal types were well known, the determination
of homology between structures might well rest upon the degree of identity
of the genes concerned in their production. What then are the best criteria
for the establishment of homology?...Best of all is similarity in developmental
history. Embryologic processes in vertebrates tend to be conservative, and
organs which are very different in the adult condition may reveal their homology
through similarity in embryonic stages.[1]
The conventional explanation can be represented
more schematically as follows:
A.
Main Principle: Descent with Modification (Material Continuity from a Common Ancestor)
B.
What Descends are Genes (Homologous Replicators)
C.
Genes Control Development (Homologous Developmental Processes and Patterns)
D.
Development Constructs Phenotypes (Homologous
Structures) (Q.E.D. )
And the conventional explanation is often embedded in the following story.
When Darwin arrived on the scene in the mid- nineteenth century, natural historians
were puzzled by the remarkable patterns of similarity they had discovered, such
as the pentadactyl limb or the vertebrate body plan. Darwin provided a material
causal explanation for the origin of these patterns, in the principle of descent
with modification from common ancestors (with all living things descending,
ultimately, from a single common ancestor). After the discovery of the rules
of inheritance in Mendelian genetics, the picture Darwin left unfinished was
completed, and "the problem of homology" was solved a major
scientific advance gained by the Darwinian revolution. As Ronald Brady observes,
We must remember that, for the minds of his contemporaries, much of the strength
of Darwin's hypothesis lay in its ability to account for the patterns of natural
history....Since that time, one may well agree with Dobzhansky (1973) that
"Nothing in biology makes sense except in the light of evolution,"
for without the theory of evolution we would stand again before the interlocking
patterns of common plan, homology, hierarchy, ontogenetic parallelism and
divergence, etc., without a causal hypothesis. The experience is now so distant
that it may be difficult to imagine.[2]
Some elements of the Darwinian interpretation of homology, such as the theological
suboptimality argument against the independent creation of homologous patterns,
are more properly the bailiwick of Topic Area 4. Still, we note the following.
"Nothing can be more hopeless," Darwin argues of the pentadactyl limb
pattern, "than to attempt to explain this similarity of pattern...by utility
or by the doctrine of final causes." Nowadays this has become, in Ayala's
hands, for instance, "an engineer could design better limbs in each case"[3], meaning, of course, a Really Competent
Divine Engineer. We expect that John Campbell will have illuminating things
to say about this aspect of Darwin's rhetorical program. And Paul Nelson will
be bringing copies of his paper on evolutionary theological arguments. As far
as Paul has been able to discover, the suboptimality of the pentadactyl limb
pattern has never been empirically demonstrated, but merely assumed. Darwin's
evidence for suboptimality, when the thread is wound to its end, turns out to
be only Richard Owen's unhappiness with the pentadactyl pattern. The pattern
squares...little," Owen opines, "with our idea of the simplest mode
of effecting the purpose."[4]
The fact that Richard Owen would have designed animals differently is helpful,
perhaps, for learning the content of Owen's mind but says nothing about
the optimality or suboptimality of actual tetrapod limbs.
Problems with the Conventional Explanation of Von Baer's
Laws,
I: The Evolutionary Interpretation
Now Brady implies that the patterns Darwin united by the theory of common descent
really do "interlock," and that after Darwin, there is a kind of evidential
inevitability in the matter, leaving reasonable investigators with only one
conclusion to draw. Such massive congruence of morphology, embryology, and genetics
can only be explained by material descent (whether the process was governed
or mediated by a Creator is a separate question). There really is a tree of
life, with a single root. But do the patterns interlock? That is, does
a consistent mapping obtain between levels B, C, and D of the conventional explanation
schema, in good accord with expectations generated by the theory of monophyletic
descent (level A)? Those who enter very far into the subject will find an astonishing
degree of confusion, and differences of opinion striking enough to
awaken the philosopher of science in anyone.
Take, for instance, early embryonic similarity in the vertebrates. Ayala sketches
the received view as follows:
Vertebrates, from fishes through lizards to humans, develop in ways that
are remarkably similar during early stages, but they become more and more
differentiated as the embryos approach maturity. The similarities persist
longer between organisms that are more closely related (man and monkey) than
between those less closely related (man and shark). Common developmental patterns
reflect evolutionary kinship.[5]
But this is simply not the case. The earliest developmental
patterns in vertebrates appear quite diverse:
Indeed, X. laevis [frog], G. domesticus [chicken], and M.
musculus [mouse] are radically different in such fundamental properties
as egg size, fertilization mechanisms, cleavage patterns, and morphogenetic
movements. This presents us with a conundrum: If early embryogenesis is conservative
[that is, functionally constrained; see below], how did such major changes
in the earliest events of embryogenesis occur?[6]
In fact, the most obvious structural characteristics of either the eggs or
the cleavage stages of a shark, a salmon, a frog, a bird, or a mammal are
unique each to its own class, not generally shared. We would not consider
them very much alike unless we had been taught so at a very early age....Each
class of vertebrates (in mammals we might almost say each particular order)
develops and then loses its own set of temporary structures like the parade
ground "formations of maneuver" during this period. The plain fact
is that evolutionary divergence has taken place at every stage in the life
history, the earliest no less than the latest.[7]
Empirically speaking these embryologists are not even in hailing distance of
Ayala. Nor is this unusual. The status of the evolutionary interpretation
of von Baer's laws of embryonic similarity[8]
(the received view; see Ayala, above) remains unresolved nearly a century after
the embryologist Adam Sedgwick urged that the interpretation "falls to
the ground."[9]
Stephen Jay Gould, for instance, writes of "von Baer's triumph,"
claiming that "his laws, in refurbished evolutionary dress, are now more
widely accepted than ever before."[10]
"These empirical laws," write Raff and Kaufman, "retain their
validity today and may be observed in operation in the development of any vertebrate."[11] Peter Medawar
argues that the laws (which he thinks are true) were important to the general
acceptance of descent:
There is an element of truth in the so-called law of recapitulation,
and it's embodied in Von Baer's law. This affirms that the embryos and young
of related animals resemble each other more closely than the adults into which
they develop. ...Similarities between mammalian embryos generally and fish
embryos in such things as the possession of yolk membranes by the human egg
were thought by Thomas Hunt Morgan, and also Thomas Henry Huxley, to be evidence
sufficient in itself to justify the acceptance of the evolutionary hypothesis.[12]
Properly restated, Løvtrup argues, von Baer's laws are not only true, but are
"the most parsimonious generalisation ever stated in biology."[13]
Others are unpersuaded. "We see the reality of von Baerian recapitulation,"
Wake and Roth demur, "as an open research question."[14] Michael Ghiselin is less politic.
"Von Baer's (1828) laws, which are false," he writes, "were replaced
by historical formulae of equally dubious status."[15]
Developmental biologist Lewis Wolpert waves von Baer aside, stating that, of
the laws, "both the first and third are just wrong: general characters,
as we have seen, do not, in early development, necessarily appear before special
characters, and neither does an animal depart more and more, during development
from the form of other animals."[16]
Reviewing what Løvtrup calls "von Baer's theorem during
their ontogenies the members of twin taxa follow the same course up to the stage
where they diverge into separate taxa"[17]
[note the agreement with Ayala's argument above] Dohle complains:
Everybody who is even slightly acquainted with ontogenetic facts knows that
there are hundreds of examples to which this theorem does not apply. In many
polychaete and prosobranch genera one species develops through a planktonic
larva, whereas another species has direct development. The telolecithal cephalopod
eggs cleave in a bilateral manner without any similarity to the spiral cleavage
of other related Mollusca. Triclad eggs have a blastomeric anarchy, whereas
the adults very closely resemble the polyclads which show spiral cleavage.
This list could easily be elongated.[18]
Problems with the Conventional Explanation,
II: The Ontogenetic Criterion of Homology
Or consider another persistent difficulty, closely related conceptually to
the question of von Baer's laws, namely, the ontogenetic criterion of homology.[19]
In the glossary (not written but approved by Darwin) appended to the sixth edition
of the Origin of Species, one reads, under "Homology":
That relation between parts which results from their development from corresponding
embryonic parts, either in different animals, as in the case of the arm of
a man, the foreleg of a quadruped, and the wing of a bird; or in the same
individual, as in the case of the fore and hind legs in quadrupeds, and the
segments or rings and their appendages of which the body of a worm, a centipede,
&c., is composed.[20]
The ontogenetic (or embryological) criterion
of homology widely accepted by the early Darwinians:
Huxley (1869, p. 137) considered the embryonic criteria to be the decisive
ones in the relationship of homology, for he defined it as follows: "Homology,
the relation between parts which are developed out of the same embryonic structures
......
Gegenbaur (1878) changed his early views in order to adopt the ancestral
criterion of homology while emphasizing the importance of embryological correspondences.
Thus he wrote...homology "is the name we give to the relations which
obtain between two organs which have had a common origin, and which accordingly
have also a common embryonic history."[21]
In his treatment of the "biogenetic law," Gareth Nelson argues as
well that "mode of development itself is the most important criterion of
homology."[22] Early in her research, V. Louise Roth (currently
one of the leading students of the concept of homology), went further: "A
necessary component of homology is the sharing of a common developmental
pathway."[23]
In 1895, however, E.B. Wilson (reading 4) warned that "similarity of embryological
origin," even when read with "great latitude," failed as a criterion:
It is a familiar fact that parts which agree closely in the adult, and are
undoubtedly homologous, often differ widely in larval or embryonic origin
either in mode of formation or in position, or in both. Innumerable cases
will suggest themselves to any embryologist of hollow organs that arise either
by invagination or delamination; of paired organs that arise from either single
or paired foundations, and vice versa. No one is disposed to question the
homology of the spinal cord of a teleost with that of a shark on the ground
that one arises as a solid cord, the other as an infolded tube. (pp. 107-8)
Citing this paper with approval several decades later, Nicholas Jardine writes,
"the Recapitulation Theory is now discredited, and with it the embryological
criterion of homology,"[24] a conclusion
reinforced in 1971 by Gavin de Beer. In a short monograph that has become something
of a minor classic, de Beer gives examples, such as the alimentary canal in
vertebrates, which violate the ontogenetic criterion.[25]
He then scolds Darwin, and, by implication, certain of his intellectual heirs:
It is therefore necessary to give the lie direct to the entry on 'Homology'
in the glossary by W.S. Dallas which Darwin most unfortunately appended to
the 6th edition of the Origin of Species. It defined homology as 'That
relation between parts which results from their development from corresponding
embryonic parts.' This is just what homology is not.
In his recent review of the homology concept
(reading 3) Gunther Wagner confronts the problem directly:
Homology is still the basic concept of comparative anatomy....However, there
are also quite problematic aspects of the current homology concept, which
has been in use since the time of Darwin. This is here called the historical
homology concept, since it is defined by historical continuity of descent
from a common ancestor....a large body of developmental data seems to contradict
certain implications of the current homology concept. (p. 51; second emphasis
added)
Roth (reading 2) points up these problems as
well. In 1988, having abandoned the embryological criterion of homology, she
argued:
Intuitively (and with some rational basis), biologists look to genetics and
developmental biology for the foundations of homology. Yet genetics and development
can provide inconsistent pictures. Homology becomes an elusive concept when
one attempts to tie it to specific biological processes or relationships or
mechanisms. (p. 4)
Both Roth and Wagner urge that the various
levels of homology in the conventional explanation be decoupled. Roth proposes
the notion of "genetic piracy" (p. 7), and Wagner, Epigenetic traps"
(p. 65), to account for the anomalies. Whatever the merits of these proposals
(and we might perhaps discuss them), we should note that neither biologist regards
the anomalies they have uncovered as casting any doubt on common descent.
That possibility never enters the conceptual picture.
So When is Common Descent at Risk Observationally?
This brings us to a significant problem. How is the theory of common descent
put at risk observationally?
One way of conceiving the empirical content of common descent (CD) is according
to the following schema:
(a)
CD
+ Independent auxiliary theory
Observational expectation
Here, common descent yields observational consequences via the "inferential
medium" of independently derived theories. In developmental biology, for
instance, we observe that disruptions of ontogeny are (generally speaking) increasingly
severe or deleterious the earlier they occur. The reason, as Van Valen argues,
is that development "ramifies out; later developmental decisions depend
on earlier ones which are much fewer and have consequences which interact."[26]
Earlier developmental decisions or events are thus "entrenched" relative
to what lies causally downstream from them, and carry a heavier "generative"
responsibility. As Leo Buss puts it,
It is axiomatic that a random alteration introduced early in ontogeny will
likely be manifested in a cascade of subsequent morphogenetic events, whereas
a modification introduced later in ontogeny can have relatively minor effects.
The validity of this interpretation can hardly be doubted. A random error
in the manufacture of the central processing unit of my computer would unquestionably
preclude any hope of my using it to write, while the various marketing decisions
reflected in the design of the exterior case have provided me with only minor
inconveniences.[27]
We might adopt Wimsatt's nicely evocative term,
generative entrenchment" (GE), to describe this theory.
What follows when generative entrenchment is coupled theoretically with common
descent? D.T. Anderson argues that
the highly integrated stepwise nature of animal development [GE] causes it
to be in many respects an extremely conservative process. Basic developmental
events established during the early evolution of a group are maintained repetitively
over hundreds of millions of years, since any change in them would spell extinction.[28]
This may be represented schematically as
(b)
CD
+ GE
Conservation of early development
But as we have just seen, early development in the vertebrates, for
example (not to mention throughout the Metazoa generally) looks pretty
diverse, not highly conserved. "Eggs, cleavage, gastrulation and germ layer
formation are very different in amphibians, bird and mammals" note Rudolf
Raff and his colleagues.[29] (Raff, as some of you may know, is a leading researcher
in this area.) Thus, there must be some way, Raff et al. conclude, of
escaping the functional constraints entailed by generative entrenchment
because, plainly, "early development does evolve, and sometimes dramatically."[30] Surveying the similar conclusions
of Keith Stewart Thomson, Van Valen concurs:
One heretical conclusion which Thomson comes to must, I think, be accepted.
This is that evolution occurs at all stages of development, often at early
stages of programs leading to the adult. His most conclusive argument for
this reminded me a bit of Descartes: Early development does often change;
therefore it can....At least such evolution can no longer be rationally dismissed
as Goldschmidt's folly.[31]
What has happened to the auxiliary theory, generative entrenchment?
It seems to have gone to the wall, blindfolded, for a last cigarette:
(c)
CD
+ GE
?
Non-conservation of early development
In this schema, which reflects the practice of evolutionary theorists, it is
generative entrenchment that is imperiled by the observations, not common descent.
The problem now however is that common descent's empirical content, vis-á-vis
the phenomena of development, is indeterminate. One really can't say what follows
observationally from the theory.
The grounds for the view that "early development does often change"
are, of course, almost wholly comparative. That is, given common descent,
the existence of radically differing ontogenetic patterns is prima facie evidence
that early development can indeed evolve. It then becomes a research problem
to learn by what mechanisms early development can be freed from its functional
entrenchment.
Experimental evidence that "early development does often change"
has not been readily forthcoming, however. As Jeffrey Levinton observes,
As a general rule, major developmental mutants give a picture of hopeless
monsters, rather than hopeful change. Epigenetic and genetic pleiotropy both
impart great burden to any major developmental perturbation....The cyclops
mutant of Artemia is lethal. The homeotic mutants of Drosophila
melanogaster suffer similar fates....But any geneticist interested in
major developmental mutants would be delighted to find viable hopeful monsters
in the laboratory, given the various tricks usually necessary to keep developmental
mutants in laboratory cultures. But, alas, major developmental mutants are
invariably sickly and show pervasive deformities. From both theoretical and
empirical points of view, hopeful monsters have led only to hopeless mooting.[32]
Under common descent, however, our responsibility for discovering the mechanisms
of macroevolution is not discharged by pointing out that earlier inquiries were
not successful. Dont forget, common descent is a fact! Shoulder to the
wheel!
The Universal Genetic Code Argument for Common Descent
Lest it be thought that this pattern of reasoning namely, sacrificing
the auxiliary theory to save common descent is an isolated example, we
offer another, perhaps more striking case.
Most of us are familiar with the universal genetic code argument for common
descent. The argument first appeared in the mid to late 1960s, after the structure
of the code was elucidated. It is now widespread.[33]
Here are two formulations:
If organisms had arisen independently they could perfectly well have used
different codes to connect the 64 trinucleotide codons to the 20 amino acids;
but if they arose by common descent, any alteration in the code would be lethal,
because it would change too many proteins at once. Hence the finding of the
same genetic code in microbes, plants and animals (except for minor variations
in intracellular organelles) spectacularly confirms a strong evolutionary
prediction.[34]
The universality of the code is easy to understand if every species is descended
from a common ancestor. Whatever code was used by the common ancestor would,
through evolution, be retained. It would be retained because any change in
it would be disastrous. A single change would cause all the proteins of the
body, perfected over millions of years, to be built wrongly; no such body
could live....Thus, we expect the genetic code to be universal if all species
have originated from a single ancestor.[35]
As in the developmental example above, we can conceive the empirical content
of common descent as following from its linkage with an independent auxiliary
theory. In the universal genetic code argument, the auxiliary theory is the
necessary functional invariance (FI) of the code:
This theory states that the code is universal because at the
present time any change would be lethal, or at least very strongly selected
against. This is because in all organisms...the code determines (by reading
the mRNA) the amino acid sequences of so many highly evolved protein molecules
that any change to these would be highly disadvantageous unless accompanied
by many simultaneous mutations to correct the "mistakes" produced
by altering the code.[36]
Consider what might happen if a mutation changed the genetic code. Such a
mutation might, for example, alter the sequence of the serine tRNA molecule
of the class that corresponds to UCU, causing them to recognize UUU sequences
instead. This would be a lethal mutation in haploid cells containing only
one gene directing the production of tRNAser, for serine would not be inserted
into many of its normal positions in proteins. Even if there were more
than one gene...this type of mutation would still be lethal, since it would
cause the simultaneous replacement of many phenylalanine residues by serine
in cell proteins.[37]
The universal genetic code argument may thus
be represented schematically as follows:
(d)
CD
+ FI
Universal genetic code
It should be clear that the theory of functional invariance
is what gives the deduction from common descent its empirical specificity.
That is, if the code were not functionally invariant, we might expect
that over time any number of different codes could have evolved. Functional
invariance, however, gives common descent considerable predictive strength.
It is, as it were, a strong lever or medium of inference, projecting the theoretical
and unobservable concept of common descent into the molecular phenomena.
Theoretical Consequences of the Discovery of the Non-Universality
of the Genetic Code
But life is never so simple. The genetic code now appears not to be
universal. (See Jukes and Osawa, reading 1.) The first variants, discovered
in 1979, occurred in the mitochondrial code, where "it was found that the
code in vertebrate mitochondria differed from the universal code by using codons
AUA for methionine and UGA for tryptophan."[38] As Fox argues, however, "mitochondria
could be thought of as exceptions that prove the rule: their genetic systems
produce only a very limited number of proteins and so might tolerate changes."[39]
Yet variants in the nuclear code discovered more recently, are, Fox argues,
of a different order: "Some 'real' (nuclear] exceptions have come to light
in both eukaryotic and prokaryotic free-living organisms, and the notion of
universality will have
to be discarded."[40] For instance, "in at least
four species of ciliated protozoa, the codons UAA and UAG [stop codons in the
universal code] occur in nuclear genes and are translated as Gln during cytoplasmic
protein synthesis."[41] In the bacterium Mycoplasma
capricolum, UGA encodes Trp, rather than termination (stop) as in the universal
code.[42] Other variants are given in the
Jukes and Osawa article.
Researchers now expect to encounter further variants. It seems obvious,"
Caron argues, "that the number of cases of deviations observed will increase
rapidly in the future."[43]
But what of the theory of functional invariance? Recognizing that, in the
face of variant codes, one cannot assume the truth both of common descent, and
functional invariance, nearly all researchers working on the problem have, either
explicitly or implicitly, rejected functional invariance:
Variations in codon assignments must arise as a result of mutations affecting
the codon specificities of tRNAs or the interactions between tRNAs and aminoacyl
tRNA synthetases. in either case the immediate result of such mutations in
a genetic system must usually be wholesale changes in the proteins produced
by that system, adversely affecting at least some and leading to a selective
disadvantage or inviability. Nevertheless, such variations have occurred during
evolution.[44]
Postulating that such fundamental variations occurred is, however, very far
from knowing how they occurred. "Direct replacements of one amino acid
by another throughout proteins," argue Osawa et al., "would
be disruptive in intact organisms and even in mitochondria."[45] That is, we should not think
that the body of molecular knowledge motivating functional invariance can be
jettisoned at will. (Yes, if common descent is true, and variant codes exist,
functional invariance has to go to the wall. Yet functional invariance still
seems to be true, or at least highly probable.) Rather, taking common descent
as given, we are now faced with another novel research problem: "How
could non-disruptive code changes occur?"[46] As Caron notes,
The scenarios have to answer the question: how, with our current knowledge
of molecular mechanisms, can we imagine a termination codon becoming a glutamine
codon or a leucine codon, a serine codon?[47]
Schema (e) illustrates the conceptual relation:
(e) CD
? FI
Non-universal genetic code
Note that, as with the earlier example concerning development, in this schema
the empirical content of common descent now, vis-á-vis the genetic code
is indeterminate. That is, the content of the theory awaits the outcome
of the new research program, to which the observational anomalies (the variant
codes) have been referred. But nearly all investigators (we know of only one
exception; see note 44) regard the finding of variant genetic codes as fully
consistent with common descent.
What Should We Make of All This?
So what is the empirical content of common descent, anyway? The theory is
certainly easy enough to state:
Evolution asserts that the pattern of similarity by which all known organisms
may be linked is the natural outcome of some process of genealogy. In other
words, all organisms are related.[48]
But is this not to put too fine a point
on it a testable proposition?
Or perhaps we should rather ask, do evolutionists (in practice) treat common
descent as a testable proposition? There are many indications that the answer
is "no, not really." On this view of the theory (which we offer for
discussion), common descent is actually something like an axiom or formal principle,
which is presupposed by evolutionary theory but is itself not at issue.
This view call it the axiom thesis is not as outlandish as it
may appear, and helps to make sense of the scientific practice of evolutionary
theorists, as in the examples given above. When reconciling a theoretical bundle
(common descent + independent auxiliary theory) with observation, evolutionists
act to conserve the truth of common descent. As philosopher of science Harold
I. Brown observes,
In science, not all propositions are treated as testable empirical hypotheses.
It is only because a large body of knowledge is taken as paradigmatic that
we can isolate individuals propositions for purposes of testing, and what
conclusions we draw from a particular test depends on what propositions we
take as paradigmatic.[49]
Kevin de Queiroz and Michael Donoghue argue that "the principle of common
descent" unifies the "patterns of living things in space, in time,
and in form under a single general theory."[50] But, as they also
argue,
The theory of common descent...is "evolutionary" only in the most
general sense, for it does not even refer to change. It certainly is not
tied to any particular model of the evolutionary process, nor is it at odds
with the results of systematic analysis.[51]
In other words, we need not worry that anything in our biological
experience will ever run afoul of the theory.
Suppose Darwin had it right, namely, that "all the organic beings which
have ever lived on this earth have descended from some one primordial form."[52] The existence of this "one
primordial form," the common ancestor, establishes a theoretical domain
that logically subsumes all biological and paleontological phenomena. That
is, even if life had multiple origins, we will be unable, having assumed the
truth of common descent, to provide any evidence for that possibility:
all observed organisms, whether recent or extinct, will necessarily lie within
what might be called the "common ancestor horizon."
If this seems counter-intuitive, try the following thought experiment. Assume
the truth of common descent, and then attempt to construct an empirical
argument against it. No imaginable evidence one might bring to bear, however
striking e.g., organisms for which no transitional stages seem possible,
multiple genetic codes will be able to overturn the theory. If there
really was a common ancestor, then all discontinuities between organisms are
only apparent, the artifacts of an incomplete history. An ideally fine-grained
history would reveal the begetting relations by which all organisms have descended
from the common ancestor.
If the axiom thesis is correct, then the theory of common descent will indeed
be refractory to the evidential challenges thrown up by biological experience.
One can see the point in Mayr's recent claim that common descent
has been gloriously confirmed by all researches since 1859. Everything we
have learned about the physiology and chemistry of organisms supports Darwin's
daring speculation that "all the organic beings which have ever lived
on this earth have descended from some one primordial form..."[53]
One wonders what we could have learned about
organisms, since 1859, that would not have confirmed common descent.
We offer the axiom thesis, not because we are persuaded of its truth, but to
provide a starting point or focus for discussion. How, really, do the patterns
of living things count for, or against, the notions of primary continuity (common
ancestry) or primary discontinuity (polyphyly)? If common descent cannot be
dislodged by the "evidence," then how should we go about evaluating
it?
[5] Ayala, "Theory of Evolution," p. 987.
[6] Richard P. Elinson, "Change in Developmental
Patterns: Embryos of Amphibians with Large Eggs," in Development as
an Evolutionary Process, eds. R. Raff and E.Raff (New York: Alan R. Liss,
1987), p. 3.
[8] As stated by von Baer, these are: (a) The general
features of a large group of animals appear earlier in the embryo than the
special features; (b) Less general characters are developed from the most
general, and so forth, until finally the most specialized appear; (c) Each
embryo of a given species [literally Thierform], instead of passing
through the stages of other animals, departs more and more from them, and
(d)
Fundamentally, therefore, the embryo of a higher animal is
never like [the adult of] a lower animal, but only like its embryo. Translation
from Entwicklungsgeschichte der Thiere [Borntrager, Konigsberg, 1828],
224; trans. S.J. Gould in Ontogeny and Phylogeny [Cambridge, Mass:
Harvard University Press, 19771, p. 56.)
[9] Adam Sedgwick, "On the Law of Development commonly
known as von Baer's Law; and on the Significance of Ancestral Rudiments in
Embryonic Development," Quarterly Journal of Microscopial Science
36 ([N.S.] 1894): p. 38.
[10] Gould, Ontogeny and Phylogeny,
p. 59
[11] Rudolf Raff and
Thomas Kaufman, Embryos, Genes, and Evolution (Bloomington, Indiana:
Indiana University Press, 1991), p. 9.
[12]
Peter Medawar, "The Evidences of Evolution," in Darwin's Legacy,
ed. C.L. Hamrum (San Francisco: Harper & Row, 1983), p. 50.
[13]
Søren Løvtrup, Darwinism, the Refutation of a Myth (London: Croom Helm,
1987), p. 378; italicized in original.
[14]
David Wake and Gerhard Roth, "The Linkage between Ontogeny and Phylogeny,"
in Complex Organismal Functions, eds. D.B. Wake and G. Roth (New York:
John Wiley, 1989), p. 363.
[15]
Michael Ghiselin, "The origin of molluscs in the light of molecular evidence,"
Oxford Surveys in Evolutionary Biology, Volume 5, eds. P. Harvey and
L. Partridge (Oxford: Oxford University Press, 1988), p. 84.
[16]
Lewis Wolpert, "The evolution of development," Biological Journal
of the Linnean Society 39 (1990): p. 120.
[17]
Løvtrup, Darwinism, p. 378.
[18]
Wolfgang Dohle, review of Darwinism: The Refutation of a Myth, by Søren
Løvtrup. Journal of Evolutionary Biology 1 (1988): p. 285.
[19]
"Homology" should here be understood phylogenetically: "The
definition of homology most commonly used by biologists...uses phylogeny as
a defining criterion, and can be expressed as follows: Homologous features
(or conditions of the features) in two or more organisms are those that stem
phylogenetically from the same feature (or condition) in the immediate common
ancestor of these organisms" (Walter Bock, "The homology concept:
its philosophical foundation and practical methodology," Zoologische
Beitrage Neue Folge 32 [1989]: 331).
[20]
Charles Darwin, The Origin of Species by Charles Darwin: A Variorum Text,
ed. Morse Peckham (Philadelphia: University of Pennsylvania Press, 1959),
p. 765; emphasis added.
[22] Gareth Nelson,
"Ontogeny, Phylogeny, Paleontology, and the Biogenetic Law," Systematic
Zoology 27 (1978): p. 335.
[23] V. Louise Roth, "On homology,"
Biological Journal of the Linnean Society 22 (1984): p. 17; emphasis
in original.
[24] Nicholas Jardine, "The Concept
of Homology in Biology," British Journal for the Philosophy of Science
18 (1967): p. 127.
[25] "Structures as obviously homologous
as the alimentary canal in all vertebrates can be formed from the roof of
the embryonic gut cavity (sharks), floor (lampreys, newts), roof and floor
(frogs), or from the lower layer of the embryonic disc, the blastoderm, that
floats on top of heavily yolked eggs (reptiles, birds)" (Gavin de Beer,
Homology, An Unsolved Problem, Oxford Biology Readers No. 11, eds.
J.J. Head and O.E. Lowenstein [Oxford: Oxford University Press, 1971] p. 13.)
[27] Leo Buss, The Evolution of Individuality
(Princeton: Princeton University Press, 1987), p. 33.
[29] Rudolf Raff, Gregory Wray, and Jonathan
Henry, "Implications of Radical Evolutionary Change in Early Development
for Concepts of Developmental Constraint," in New Perspectives on
Evolution, eds. L. Warren and H. Koprowski (New York: Wiley-Liss, 1991),
v. 189.
[30] Raff et al., "Implications,"
p. 145.
[31] Leigh Van Valen, "A morphogenetic
basis for macroevolution, Evolutionary Theory 9 (1988): p. 329.
[32] Jeffrey Levinton, Genetics, Paleontology,
and Macoevolution (Cambridge: Cambridge University Press, 1988), pp. 252,
254.
[33] See A.G. Cairns-Smith, The Life
Puzzle (Edinburgh: Oliver & Boyd, 1971) 148; Theodosius Dobzhansky,
"Nothing in biology makes sense except in the light of evolution,"
American Biology Teacher 35 (1973):125-129; John Maynard Smith, The
Theory of Evolution (New York: Penguin, 1975), 82; Theodosius Dobzhansky,
Francisco Ayala, G. Ledyard Stebbins, and James Valentine, Evolution
(San Francisco: W.H. Freeman, 1977), 28; Douglas Futuyma, Evolutionary
Biology (Sunderland, Mass.: Sinauer, 1979), 38; Ernst Mavr, "Darwin,
intellectual revolutionary," in Evolution from Molecules to Men,
ed. D.S. Bendall (Cambridge: Cambridge University Press, 1983), 30-31; David
Raup and James Valentine, "Multiple origins of life," Proceedings
of the National Academy of Sciences USA 80 (1983): 2981; Bernard Davis,
"Molecular Genetics and the Foundations of Evolution," Perspectives
in Biology and Medicine 28 (1985); 256; Richard Dawkins, The Blind
Watchmaker (New York: W.W. Norton, 1986), 270; Mark Ridley, Evolution
and Classification (London: Longman, 1986), 119-20; Colin Patterson, "The
impact of evolutionary theory on systematics," in Prospects in Systematics,
ed. D.L. Hawksworth (Oxford: Clarendon Press, 1988), 61; Elliot Sober, Reconstructing
the Past (Cambridge, Mass.: MIT Press, 1988), 9; Antoni Hoffman, Arguments
on Evolution (Oxford: Oxford University Press, 1989), 8-9; Ernst Mayr,
One Long Argument (Cambridge, Mass.: Harvard University Press, 1991),
23.
[34]
Davis, "Molecular Genetics," p. 256, emphasis added.
[35]
Mark Ridley, The Problems of Evolution (Oxford: Oxford University Press,
1985), pp. 10-11; emphasis added.
[36]
Francis Crick, "The Origin of the Genetic Code," Journal of Molecular
Biology 38 (1968): p. 369; emphasis added.
[37]
James Watson et al., Molecular Biology of the Gene, 4th ed.
(Menlo Park, California: Benjamin/Cummings, 1987), p. 453; emphasis
added.
[39]
Thomas Fox, "Diverged genetic codes in protozoans and a bacterium,"
Nature 314 (1985): p. 132.
[41]
Thomas Fox, "Natural Variation in the Genetic Code," Annual Review
of Genetics 21 (1987): p. 77.
[43]
F. Caron, "Eucaryotic codes," Experientia 46 (1990):
p. 1111.
[44]
Fox, "Natural Variations in the Genetic Code," p. 84; emphasis added.
As far as we know, only Hubert Yockey has suggested that the discovery of
multiple codes may imply multiple origins of life. "The pervasiveness
of the standard genetic code," he writes, "has often been cited
as evidence of a single
codes is
evidence of a number of independent origin of life events" (Information
theory and molecular biology [Cambridge: Cambridge University Press, 1992],
pp. 202-203).
[45] Osawa, Muto, Jukes, and Ohama, "Evolutionary
changes," p. 21.
[48] Niles Eldredge and Joel Cracraft,
Phylogenetic Patterns and the Evolutionary Process (New York: Columbia
University Press, 1980), p. 2.
[49] Harold I. Brown, Perception, Theory
and Commitment (Chicago: University of Chicago Press, 1979), p. 106.
[50] Kevin de Queiroz and Michael Donoghue,
"Phylogenetic Systematics or Nelson's Version of Cladistics," Cladistics
6 (1990): p. 62.
[52] Charles Darwin, On the Origin
of Species, 1st ed., p. 484.
[53] Ernst Mayr, One Long Argument
(Cambridge: Harvard University Press, 1991), p. 163.
Copyright 1993, 2001 Paul Nelson and Jonathan Wells. All rights
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