Serial Endosymbiosis and Intelligent Design

AUTHOR: Allen MacNeill

SOURCE: Original essay

COMMENTARY: That's up to you...

It's very gratifying to see Lynn Margulis finally getting the recognition that she deserves. As the most effective exponent of the serial endosymbiosis theory (SET) for the origin of eukaryotes, Lynn's work provides an excellent example of how ID should (but currently doesn't) proceed. During the late 1960s, Lynn published a series of revolutionary papers on the evolution of eukaryotic cells, culminating in her landmark book Symbiosis and Cell Evolution, in which she carefully laid out the empirical evidence supporting the theory that mitochondria, choloroplasts, and undulapodia (eukaryotic cilia and flagella) were once free living bacteria (purple sulfur bacteria, cyanobacteria, and spirochaetes, respectively).

Her theory was greeted with contempt and scorn by almost all evolutionary biologists (sound familiar?), who believed at the time that all eukaryotic cellular organelles evolved by gradual elaboration of invaginations of the plasma membrane. But Lynn didn't give up, or continue to simply restate her original theory (sound familiar?). Instead, she continued to do extensive field and laboratory research, publishing hundreds of papers and dozens of books in which she presented the accumulating empirical evidence supporting her theory. With time, other researchers (encouraged by the success of her field and lab research) began to test her hypotheses themselves, and discovered yet more empirical evidence supporting her theory.

And so today, Lynn Margulis's SET has become the dominant theory explaining not only the origin of eukaryotes, but also the origin of evolutionary novelty at dozens of different levels in biology (see her book, Acquiring Genomes for a comprehensive review). So well accepted has her work become by evolutionary biologists that finally, after almost four decades, creationists and ID supporters have begun to attack her theories. As she said at our Darwin Day celebration at Cornell this past February, no greater affirmation of one's "having arrived" as a major theorist in evolutionary biology could be imagined.

The point here is that, if ID wants to become accepted as part of evolutionary biology in the same way that Lynn Margulis's SET has become accepted, then ID supporters have to do the same thing she did: get out in the field and get your hands dirty, and get into the lab and do the same thing. Her ideas were just as unorthodox and unacceptable in 1969 as ID is now. However, she didn't put all of her effort into public relations and political propaganda. No "Symbiosis Institute" dumped millions into the production of deliberately distorted press kits and one-sided propaganda films. Legions of self-appointed experts whose only exposure to biology was in high school classes or what they read on Answers in Genesis or Uncommon Descent bloviated on SET and declared themselves experts after a week of superficial study of articles on Wikipedia.

No, Lynn and her colleagues did the hard work of finding the empirical evidence that eventually carried the day and established her SET as one of the bedrock foundations now worthy enough of respect as to earn the ire of the creationists and IDers. Her ideas are still radical, and still raise the blood pressure of many evolutionary biologists. Her dismissal of the "modern evolutionary synthesis" in particular is not popular among many evolutionary biologists, who are largely still mired in paradigms that are at least four decades of out of date. She has said some things about the "modern synthesis" that have brought smiles to the faces of the creationist quote-miners. The difference between her and them is that they can't even begin to claim any credibility in science; their "work" is entirely parasitic on hers, and deserves nothing but contempt.

When the history of evolutionary biology in the 20th century is written (I hope to contribute to it myself, if I live long enough), the work of Lynn Margulis will rank right up there with the work of Fisher, Haldane, Wright, Dobzhansky, Mayr, Simpson, Stebbins, Gould, Lewontin, Kimura, Williams, Hamilton, Trivers, and the two Wilsons. And unless and until IDers decide that it's finally time to stop doing agitprop and start doing science, they and the creationists will at best be a trivial footnote.

Comments, criticisms, and suggestions are warmly welcomed!

--Allen

Scientists Say Darwin's 'Tree of Life' Not The Theory Of Everything

ARTICLE: physorg.com

AUTHOR: Lisa Zyga

COMMENTARY: Allen MacNeill

First the news item, followed by some commentary:

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There is only one figure in On the Origin of Species, and that is a tree diagram. As Darwin’s model for the theory of evolution, he used the Tree of Life (TOL) to clearly and visually explain the interrelatedness of all living things, implying that from one common ancestor (the root) sprung branches, which produced smaller offshoots as genetic progeny, etc.

In this model, similarities between species reveal a common ancestor, and differences result from (and explain) Darwin’s main catalysts: competition and natural selection, which generate improvement in future generations. As a simile, the TOL served a vital purpose for introducing the theory of evolution to the community in an understandable way. Although there is no external evidence to support the idea that evolution is inclusively hierarchical, many evolutionists believe the TOL provides an accurate general representation of the history of life, which could potentially be completely reconstructed by knowing the relevant data.

In recent times, however, a minority of biologists and evolutionists have questioned the accuracy of the TOL hypothesis, including W. Ford Doolittle and Eric Bapteste. In a recent paper in the Proceedings of the National Academy of Sciences, “Pattern Pluralism and the Tree of Life Hypothesis,” the scientists investigate the shortcomings of the TOL, as well as propose alternative models that would better explain how to classify the history of evolving life forms.

Much of the initial concern over TOL was provoked by biologists studying the complex relationships among prokaryotes, the most primitive life forms that include bacteria and archaea. Prokaryotes have a much simpler DNA structure than eukaryotes (all other life forms). Because of this, prokaryotes often transfer their DNA via processes such as lateral gene transfer as opposed to vertical gene transfer (direct transmission form parent to progeny) which is the basis for the “phylogenetic” (evolutionary relatedness) TOL scheme.

“Surely a tree is the right model for most multi-cellular animals and plants,” Doolittle explained to PhysOrg.com. “Thus the TOL is great for fossils and museums and dinosaurs and most of visible life, over the last billion years. But unicellular eukaryotes and prokaryotes represent the bulk of the biomass and diversity of life on earth, as well as the first two-thirds of its history.”

In their paper, Doolittle and Bapteste highlight research that shows other causes of genetic modification, suggesting that evolutionary history is more complex than described by the TOL. For example, recombination, gene loss, duplication, and gene creation are a few of the processes whereby genes can be transferred within and between species, causing variation that’s not due to vertical transfer. These transfer methods give results that don’t fit on the TOL, including species that cannot be traced to a common ancestor.

While such diverse methods might appear to obviously point to a more complex nonhierarchical evolutionary scheme, Doolittle and Bapteste explain that the TOL thinking persists due to confusion between the roles of “process” and “pattern.” The above methods are processes and are widely accepted by modern evolutionists, whereas the TOL is a pattern that, as Doolittle and Bapteste explain, has been ingrained in biologists’ minds from early education as a single, unifying model. As the researchers explain of the current biology scene, “We may be process pluralists, but we remain pattern monists.”

If this combination of thinking seems to clash, Doolittle and Bapteste suggest that the Western philosophical tradition of thinking in universal patterns has caused biologists to cling to classification without realizing it. The authors point out that many algorithms used to study evolutionary hierarchies impose or extract the TOL structure due to their intrinsic design. TOL is a paradigm that has stuck. But Doolittle sees ways to alter this mentality.

“Sure we can [re-train Western thinking]. That's what ‘postmodernism’ is about,” he said. “I would agree that the need to classify might be built in, but the coupling of this practice to a specific theory about what classifications are ‘natural’ is surely not.

As an alternative to the TOL, the scientists suggest that relationships among life forms may be represented by whatever model fits for a certain purpose, a certain taxonomic group, or a certain scale. In contrast to pattern monism, they call this belief “pattern pluralism.” While parts of evolution certainly are tree-like, other parts may be nets or webs or other complex models. Most importantly, however, there seems to be no “theory of everything” in evolution, no metanarrative to unify all life forms.

“In 2006, our understanding of evolution at the molecular, population genetic, and ecological levels is rich and pluralistic in character,” the scientists conclude, “and does not require (or justify) a monistic view of the phylogenetic pattern.”

As for any blow to Darwin’s ego, the scientists point out that he never wrote about reconstructing the tree in an attempt to relate every living thing, but rather used the model as a general guide.

“I'd like to think he would adjust,” Doolittle said about Darwin. “After all, his theory was developed before there was any understanding of genetics and when bacteria were still believed to be spontaneously generated.”

REFERENCES CITED:
Doolittle, W. Ford, and Bapteste, Eric. “Pattern pluralism and the Tree of Life hypothesis.” PNAS, February 13, 2007, vol. 104, no. 7, 2043-2049.

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COMMENTARY:

Ford Doolittle has been saying this for many years, and has been joined by Lynn Margulis, who has argued that the concept of "species" does not apply to prokaryotes.

But, that's not what I want to talk about. What I want to talk about is how "intelligent design theorists" quote-mine and otherwise distort reality to the point of outright lying. Why do I say this? Because former lawyer and professional propagandist, Casey Luskin, has a postat Evolution News, propaganda outlet for the Discovery Institute in which he says

"By invoking insufficient data, horizontal gene swapping, rapid evolution, and other ad hoc explanations, Darwinists reveal that neo-Darwinism is trying explaining away the data; it is not explaining the data. Perhaps the inability to construct robust phylogenetic trees using molecular data stems from the fact that common descent is simply wrong."

And the alternative? Why, magic of course. All of the living forms on Earth today were created, all at once, on Tuesday 25 October 4004 BC at 9 AM...or some other time in the past, exact date unspecified. Because, of course, that is the only alternative to common descent. Either you and your siblings are the offspring of your parents (i.e. common descent) or you aren't, and if the latter is true, then you must all have been created separately, not by your parents, but rather by God...excuse me, the Grand Omnipotent Designer. And with your memories of your childhood already inserted into your minds, so that the jarring discontinuity of your separate creations would not disturb you too much...would lead you, in fact, into the damning conclusion that you were, in fact, the offspring of your parents, and thus consign yourself to everlasting torment in the lake of fire.

Or not. You see, this is what the kind of egregious propagandizing of the folks of the Disco Institute leads to. Not discussion of the science of biology on its merits, on the basis of reason and evidence, but pure assertions without any alternatives at all (not that they want you to read in public, anyway).

I'm tired (our new baby is keeping Leah and me awake at night), and so I'll sign off now. But damn, people like Luskin just make me want to scream sometimes...

"Against stupidity, the gods themselves contend in vain."
- Friedrisch Schiller

--Allen

A Third Way: James Shapiro and the Post-Modern Synthesis

AUTHOR: James A. Shapiro

SOURCE: The Boston Review

COMMENTARY: Allen MacNeill (following the article)

The recent reviews in your columns of books by Dennett, Dawkins, and Behe are testimony to the unflagging interest in controversies about evolution. Although such purists as Dennett and Dawkins repeatedly assert that the scientific issues surrounding evolution are basically solved by conventional neo-Darwinism, the ongoing public fascination reveals a deeper wisdom. There are far more unresolved questions than answers about evolutionary processes, and contemporary science continues to provide us with new conceptual possibilities.

Unfortunately, readers of Boston Review may remain unaware of this intellectual ferment because the debate about evolution continues to assume the quality of an abstract and philosophical "dialogue of the deaf" between Creationists and Darwinists. Although our knowledge of the molecular details of biological organization is undergoing a revolutionary expansion, open-minded discussions of the impact of these discoveries are all too rare. The possibility of a non-Darwinian, scientific theory of evolution is virtually never considered. In my comments, then, I propose to sketch some developments in contemporary life science that suggest shortcomings in orthodox evolutionary theory and open the door to very different ways of formulating questions about the evolutionary process. After a discussion of technical advances in our views about genome organization and the mechanisms of genetic change, I will focus on a growing convergence between biology and information science which offers the potential for scientific investigation of possible intelligent cellular action in evolution.

The past five decades of research in genetics and molecular biology have brought us revolutionary discoveries. Upsetting the oversimplified views of cellular organization and function held at mid-century, the molecular revolution has revealed an unanticipated realm of complexity and interaction more consistent with computer technology than with the mechanical viewpoint which dominated the field when the neo-Darwinian Modern Synthesis was formulated. The conceptual changes in biology are comparable in magnitude to the transition from classical physics to relativistic and quantum physics.

Four categories of molecular discoveries are especially important in opening up exciting new ways of thinking about the biological processes that underlie evolutionary change.

(1) Genome Organization. Our current ideas of genome organization are completely different from the "beads on a string" view that dominated genetics in the 1940s and 1950s. At that time genes were "units" which corresponded to individual organismal traits, and the "one gene-one enzyme" hypothesis told us that the essential business of each gene was to encode a specific protein molecule linked to a particular phenotype. We have now deconstructed each genetic locus into a modular assembly of regulatory and coding motifs. Most of these motifs are shared among many loci, suggesting that genomes are assembled Lego-like from a repertoire of more basic sequence elements, many of which do not encode proteins but determine other important functions (transcription, translation, RNA processing, DNA replication, chromatin condensation, etc.). As we analyze genome expression during cellular proliferation and multicellular development, we have learned that diverse genetic loci are organized hierarchically into interconnected genome-wide networks which function dynamically. Not confined to a single pathway, many genetic loci are active at different times, participating in the expression of more than one phenotypic trait. Comparisons of genomes in different organisms have revealed unexpected patterns of evolutionary conservation across large taxonomic distances, while closely-related genomes frequently differ significantly in the arrangement of repetitive DNA elements which do not encode proteins.

How all of this modularity, complexity, and integration arose and changed during the history of life on earth is a central evolutionary question. Localized random mutation, selection operating "one gene at a time" (John Maynard Smith's formulation), and gradual modification of individual functions are unable to provide satisfactory explanations for the molecular data, no matter how much time for change is assumed. There are simply too many potential degrees of freedom for random variability and too many interconnections to account for.

Studies of the molecular sources of genetic variability have taught us two major lessons about how cells take care of their genomes--one about self-protection, the other about self-reorganization.

(2) Cellular Repair Capabilities. First, then, all cells from bacteria to man possess a truly astonishing array of repair systems which serve to remove accidental and stochastic sources of mutation. Multiple levels of proofreading mechanisms recognize and remove errors that inevitably occur during DNA replication. These proofreading systems are capable of distinguishing between newly synthesized and parental strands of the DNA double helix, so they operate efficiently to rectify rather than fix the results of accidental misincorporations of the wrong nucleotide. Other systems scan non-replicating DNA for chemical changes that could lead to miscoding and remove modified nucleotides, while additional functions monitor the pools of precursors and remove potentially mutagenic contaminants. In anticipation of chemical and physical insults to the genome, such as alkylating agents and ultraviolet radiation, additional repair systems are encoded in the genome and can be induced to correct damage when it occurs.

It has been a surprise to learn how thoroughly cells protect themselves against precisely the kinds of accidental genetic change that, according to conventional theory, are the sources of evolutionary variability. By virtue of their proofreading and repair systems, living cells are not passive victims of the random forces of chemistry and physics. They devote large resources to suppressing random genetic variation and have the capacity to set the level of background localized mutability by adjusting the activity of their repair systems.

(3) Mobile Genetic Elements and Natural Genetic Engineering. The second major lesson of molecular studies into the origins of genetic change is that all cells possess multiple biochemical agents for natural genetic engineering--processes that include the cutting and splicing of DNA molecules into new sequence arrangements. Most frequently, natural genetic engineering capabilities reveal themselves through the activities of mobile genetic elements--DNA structures found in all genomes that can move from one position to another. Mobile genetic elements are the most fluid components of the genome and also the most taxonomically specific. In human cells, mobile elements include retrotransposons, like the half-million or more Alu sequences dispersed over all our chromosomes, as well as the inherited gene fragments which our lymphocytes assemble daily to form active genetic loci encoding the key antigen recognition molecules of our immune system. The biochemical agents of DNA restructuring include the enzymes used in our own genetic engineering for research and biotechnology (nucleases, ligases, reverse transcriptases and polymerases) as well as other proteins that combine to form molecular machines capable of mobilizing different genomic components.

The existence of cellular biochemical activities capable of rearranging DNA molecules means that genetic change can be specific (these activities can recognize particular sequence motifs) and need not be limited to one genetic locus (the same activity can operate at multiple sites in the genome). In other words, genetic change can be massive and non-random. Some organisms, such as the ciliated protozooan Oxytricha, completely reorganize their genetic apparatus within a single cell generation, fragmenting the germ-line chromosomes into thousands of pieces and then reassembling a particular subset of them into a distinct kind of functional genome. Furthermore, natural genetic engineering systems can operate premeiotically during the somatic development of tissues that will ultimately produce gametes. This means that major chromosome reorganizations can be present in multiple gametes. Consequently, the appearance of new genome architectures during evolution is not necessarily limited to isolated individuals.

The discovery that genome reorganization is largely a biological process traces back to Barbara McClintock's pioneering studies of mutation and chromosome rearrangement in maize from the 1940s through the 1960s. She linked these genetic events to changes in the regulation of gene expression programs during plant development. We can now appreciate her tremendous wisdom and foresight by seeing how the Lego-like patterns of integrated genome organization mentioned above could be created by the activity of cellular natural genetic engineering systems. Because, like all cellular functions, natural genetic engineering systems are subject to control circuits, they can be held in abeyance for long periods and then called into action at certain key times. Sometimes these activations can be regularly programmed, as in the development of our immune systems, and sometimes activations can occur in response to crisis, as McClintock documented in maize.

The point of this discussion is that our current knowledge of genetic change is fundamentally at variance with neo-Darwinist postulates. We have progressed from the Constant Genome, subject only to random, localized changes at a more or less constant mutation rate, to the Fluid Genome, subject to episodic, massive and non-random reorganizations capable of producing new functional architectures. Inevitably, such a profound advance in awareness of genetic capabilities will dramatically alter our understanding of the evolutionary process. Nonetheless, neo-Darwinist writers like Dawkins continue to ignore or trivialize the new knowledge and insist on gradualism as the only path for evolutionary change.

(4) Cellular Information Processing. While it is easy to see how advances in our understanding of genome organization and genetic change will impact theories of evolutionary processes, another development in contemporary biology is of less obvious but even more basic relevance. This is the growing realization that cells have molecular computing networks which process information about internal operations and about the external environment to make decisions controlling growth, movement, and differentiation. This realization has come, in large measure, from detailed genetic analysis of cellular processes and multicellular development. The inducible repair systems mentioned above provide a relatively simple, well-studied example. Bacterial and yeast cells have molecules that monitor the status of the genome and activate cellular responses when damaged DNA accumulates. The surveillance molecules do this by modifying transcription factors so that appropriate repair functions are synthesized. These inducible DNA damage response systems are sophisticated and include so-called "checkpoint" functions that act to arrest cell division until the repair process has been completed. When the checkpoints do not function, cell division proceeds before repair is completed, and the damaged cells die or produce inviable progeny. One can characterize this surveillance/inducible repair/checkpoint system as a molecular computation network demonstrating biologically useful properties of self-awareness and decision-making.

There are many other cellular systems that display comparable information-processing capabilities. Fro example, it is now common among molecular biologists who study the cell cycle to speak of various checkpoints (Is DNA replication complete? Are the chromosomes properly condensed and aligned on the metaphase plate?) and decision points (e.g., when to initiate chromosome movement and cytokinesis).

A recent special issue of Scientific American [1] describes beautifully how cancer is now seen as a disease of the molecular information processing routines that ensure orderly cell growth and behavior in the healthy organism. Aberrant tumor cell growth appears to result from at least two kinds of malfunction: the loss of checkpoint controls, or the failure of decision-making routines that dictate programmed cell death (apoptosis) for cells in inappropriate surroundings. During embryonic development, cells make decisions about differentiation based on multiple molecular signals picked up from their environment and from their neighbors by means of surface receptors. These receptors are linked to intercellular molecular cascades called "signal transduction pathways" which integrate the inputs from the receptors to generate appropriate patterns of differential gene expression and morphogenesis of specialized cell structures.

Signal transduction is not limited to multicellular development. We are learning that virtually every aspect of cellular function is influenced by chemical messages detected, transmitted, and interpreted by molecular relays. To a remarkable extent, therefore, contemporary biology has become a science of sensitivity, inter- and intra-cellular communication, and control. Given the enormous complexity of living cells and the need to coordinate literally millions of biochemical events, it would be surprising if powerful cellular capacities for information processing did not manifest themselves. In an important way, then, biology has returned to questions debated during the mechanism-vitalism controversy earlier this century. This time around, however, the discussion is informed by two new factors. One is that the techniques of molecular and cell biology allow us to examine the detailed operation of the hardware responsible for cellular responsiveness and decision-making. The second is the existence of computers and information networks, physical entities endowed with computational and decision-making capabilities. Their existence means that discussing the potential for similar activities by living organisms is neither vague nor mystical.

What significance does an emerging interface between biology and information science hold for thinking about evolution? It opens up the possibility of addressing scientifically rather than ideologically the central issue so hotly contested by fundamentalists on both sides of the Creationist-Darwinist debate: Is there any guiding intelligence at work in the origin of species displaying exquisite adaptations that range from lambda prophage repression and the Krebs cycle through the mitotic apparatus and the eye to the immune system, mimicry, and social organization? Borrowing concepts from information science, new schools of evolutionists can begin to rephrase virtually intractable global questions in terms amenable to computer modelling and experimentation. We can speculate what some of these more manageable questions might be: How can molecular control circuits be combined to direct the expression of novel traits? Do genomes display characteristic system architectures that allow us to predict phenotypic consequences when we rearrange DNA sequence components? Do signal transduction networks contribute functional information as they regulate the action of natural genetic engineering hardware?

Questions like those above will certainly prove to be naive because we are just on the threshold of a new way of thinking about living organisms and their variations. Nonetheless, these questions serve to illustrate the potential for addressing the deep issues of evolution from a radically different scientific perspective. Novel ways of looking at longstanding problems have historically been the chief motors of scientific progress. However, the potential for new science is hard to find in the Creationist-Darwinist debate. Both sides appear to have a common interest in presenting a static view of the scientific enterprise. This is to be expected from the Creationists, who naturally refuse to recognize science's remarkable record of making more and more seemingly miraculous aspects of our world comprehensible to our understanding and accessible to our technology. But the neo-Darwinian advocates claim to be scientists, and we can legitimately expect of them a more open spirit of inquiry. Instead, they assume a defensive posture of outraged orthodoxy and assert an unassailable claim to truth, which only serves to validate the Creationists' criticism that Darwinism has become more of a faith than a science.

A sounder perspective on the history of science would be very helpful to all concerned. For example, a parallel has been drawn by Allen Orr and others between criticisms of Darwinian orthodoxy and assaults on the Law of Gravity, presenting them as equally deplorable examples of anti-science obscurantism. Yet, if truth be told, gravity is far from a settled matter. The relativistic Law of Gravity at the end of the 20th century is not the same as the classical Law of Gravity at the end of the 19th century, and discovering how the continuous descriptions of general relativity can be integrated into a single theory with the discrete accounts of quantum physics is still an active field of research. From a scientific point of view, then, the Law of Gravity has quite properly been under continuous challenge. Dogmas and taboos may be suitable for religion, but they have no place in science. No theory or viewpoint should ever become sacrosanct because experience tells us that even the most elegant Laws of Nature ultimately succumb to the inexorable progress of scientific thinking and technological innovation. The present debate over Darwinism will be more productive if it takes place in recognition of the fact that scientific advances are made not by canonizing our predecessors but by creating intellectual and technical opportunities for our successors.

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NOTES:

[1] Robert Weinberg, "How Cancer Arises," Scientific American 275, no. 3 (September 1996), pp. 62-70.

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COMMENTARY:

In general, I agree with Shapiro: the paradigms established during the "modern evolutionary synthesis" of the 1930s-50s are by and large defunct. In particular, the idea that evolutionary change is necessarily gradual has been under fire since at least 1972, and the evidence available now is leaning toward what Ernst Mayr once called "evolution by jerks" - that is, punctuated equilibrium. And new work in the field of evolutionary developmental biology ("evo-devo") has provided a new paradigm for the production and evolution of variation at speeds and in degrees that are largely incompatible with the "modern synthesis."

But this doesn't mean, by any stretch of the imagination, that current evolutionary theory now needs to have "intelligent design theory" incorporatd into any of its newly emerging paradigms. On the contrary, the new empirical studies in evo-devo, paleontology, and especially evolutionary genetics have provided us with new explanations for the sources of variation in natural and laboratory populations, and new explanations for the mechanisms by which that variation can be fixed and modified in evolving populations. Nowhere in this exciting ferment of laboratory and field research has anyone found any need to invoke any supernatural "intelligent designer" to explain where this variation comes from, or where it is going.

"Intelligent design theory" isn't a forward-looking science at all; it's a rear-guard action in a "long twilight struggle" against the startling and overwhelming successes that evolutionary biologists have had recently in explaining the origin and evolution of life on Earth. We biologists have indeed begun to give up our tired old "modern" paradigms, but unless the supporters of "intelligent design theory" are ready to do the same, they will almost certainly fade into a footnote in the history of science.

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ORIGINAL PUBLICATION REFERENCE:

Location Online:
The Boston Review
URL: http://www.bostonreview.net/br22.1/shapiro.html

Original posting/publication date timestamp:
February 1997

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