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Thinking Critically about Evolutionary Theory K. P. Mohanan Center for Integrative Studies Indian Institute for Science Education and Research (IISER), Pune April 2011 Note to the Reader This write up has two parts. Part I focuses on the creationism-evolution debate, and seeks to unpack evolution into its component propositions. It was originally meant as preparatory reading material for a class debate. Part II focuses on the task of constructing an explicit theory (as opposed to an operational definition) of species. Distributed to the class after the debate, it supplemented the debate, to clarify and consolidate some of the issues that emerged in class. The connection between the two parts can be outlined in terms of the following sequence of class activities: 1. Part I is distributed to students, with the instruction that they think about the tasks in it, first individually and then in their affinity groups, and come prepared for a class discussion. 2. Class session 1: We set up a debate between an actual evolutionary biologist (Priya Iyer) and a non-biologist who adopts the persona of a creationist (me), with a molecular biologist (Alok Srivastava) presiding. Students are encouraged to interrogate each of us. 3. Class session 2: In this round of debate, Dr Iyer and the students are the authors of a journal article supporting evolutionary theory. I quit the creationist persona and adopt that of a skeptical journal reviewer. Students have to convince me that each proposition of the evolutionary theory is correct. 4. In the course of the debate, the class discovers that the claims of evolutionary biology that we have been debating on cannot be critically evaluated unless the notion "species" is clearly defined. There are problems with the textbook definition (interbreeding with fertile offspring) but we decide to accept it tentatively. 5. In the course of the alleged evidence to support species divergence, we discover that the term "species" is used in such a way that two distinct species interbreed to produce a novel species. The concept of “species” implicit in this use logically contradicts the concept “species” in the definition. 6. Hence we search for a better definition, or a theory of species. Students don't have the background to pursue this task. Part II is meant to help them in this enterprise, while also summing up and consolidating the points that emerged in the class discussion. Part I: The Creation vs. Evolution Debate Creationism vs. Evolutionism There are those who support the idea of a divine creation of the world, and those who support biological evolution. Emotionally charged battles between these two groups have been going on ever since Darwin published his Origin of the Species. This controversy is salient in school education as well as in politics in many parts of the world. It is important, therefore, that serious students of science understand this issue properly, and be able to respond to it critically. One can gain a sense of this debate by looking at the Wikipedia entry at http://en.wikipedia.org/wiki/Creation%E2%80%93evolution_controversy, and J. R. Minkel’s Scientific American article, “Evolving Creationism in the Classroom.” (http://www.scientificamerican.com/article.cfm?id=evolving-creationism-in-the-classroom) Minkel reports the American Republican Vice-presidential nominee and Alaska Gov. Sarah Palin as saying that the Judeo-Christian version of creationism (the idea in the Bible that God created Earth and its life forms a few thousand years ago) should “get equal footing with evolution” in science classes. "Teach both," the article reports her as saying in a 2006 televised debate. "You know, don't be afraid of information." Presenting both “evolutionism” and “creationism” as “information” would be simply providing expositions of the two positions, where a textbook or teacher says, “Those who support creationism believe that X, Y, …Z are true, but those who support evolutionism believe that X’, Y’, …Z’ are true.” There is no harm in such expositions. After all, no teacher would find it difficult to tell her students, “Once upon a time, humans used to believe that the earth was flat, and there are still many of them around.” But if the teacher presents one of these positions as the Truth (whichever it may be) she would be requiring the students to believe what she believes to be true. That would be a form of indoctrination, something opposed to the very core of the value system that drives scientific inquiry. To take a closer-to-home example, it would be perfectly legitimate for a teacher to tell her students that Ptolemy believed that the earth is the unmoving centre of the universe, while Copernicus believed that the earth rotates on a tilted axis and revolves around the sun. If she is a teacher who values the students’ critical understanding of the issues, she may also provide an exposition of the evidence and argumentation in support of the two positions. What a teacher cannot do is dictate what the students should believe. After getting students to engage with the relevant evidence and argumentation, the teacher should leave the students to decide for themselves what they should believe. To decide whether they should believe the creationists or the evolutionists, students need: A) the critical thinking capacity to decide for themselves what to believe, what to reject as false, and what to keep on hold for further evaluation; and B) critical understanding of both positions, including the relevant evidence and argumentation that bears upon the choice between them. What follows is an attempt to help you develop a critical understanding of (the different versions of) creationism (including the version called intelligent design) and (the different versions of) the theory of evolution, such that you can decide for yourselves what position to take. The ultimate intention is to nudge you to pursue questions on biological evolution and engage in inquiry along novel paths. Thinking Critically about Creationism The website Allaboutcreation.org describes creationists as “…men and women who share a belief in God and a conviction that He created us and the world in which we live. Creation science, in its most general sense, is an effort to apply the scientific method to discover how God created the Heavens and the Earth.” Questions for you Q 1 What do you think they mean “the scientific method”? Q 2 What does it mean to say “God created the Heavens and the Earth?” Q 3 When they say “heavens”, are they referring to what you see when you look at the night sky, or to a place where you go when you die, which presupposes the assumption of a soul that continues to live after the body dies? 2 Q4 The formulation of the goal of creationism as “an effort to apply the scientific method to discover how God created the Heavens and the Earth” carries the presupposition that God did create the Heavens and the Earth. How scientific is it to accept that presupposition without critical scrutiny? Q5 Why do you think I have raised questions 1-3? Now take a look at their discussion of the issues in the creationism vs. evolutionism debate at http://www.allaboutcreation.org/evolution-vs-creation.htm, parts of which are given below: Evolution vs. Creation: The Great Debate The Evolution vs. Creation debate is often referred to as the "Great Debate." It's the emotion-packed question of "Origins" -- why, how, and where did everything come from? 20th century science has made the compelling discovery that, at some point, the universe began. Both sides of the Great Debate now agree that the universe has not existed eternally. However, this is where the agreement ends. As far as the "why" and "how" of the "origin event," this is where the division and contention begin. There are two basic theories in this Great Debate. The first is the historical default - the Creation Model of Origins. This theory maintains that the intricate design permeating all things implies a Designer. The second theory is the more recent, atheistic explanation - the Evolution Model of Origins. This theory postulates that the intricate design permeating all things is a product of random chance and excessive time. Evolution vs. Creation: The Contentions Evolution vs. Creation is indeed the Great Debate of our scientific times. In any scientific debate, the theories must be tested according to the evidence. We propose that the burden of evidence should be upon the Evolutionists, since Creation has been the historic and inherent default throughout virtually all cultures and religions until roughly the last 200 years. Of course, Evolutionists, who view themselves as the only "scientists" in the debate, insist that the burden of evidence be upon the Creationists. Evolutionists reason, we cannot see the Creator, we cannot hear the Creator, and we cannot touch, taste or smell the Creator. Therefore, we are unable to test for the Creator with any form of scientific equipment developed thus far. Creationists retort, we cannot see, hear, touch, taste, or smell the human mind. We cannot test for the human mind with any form of scientific equipment developed thus far. When we run an electroencephalogram, we are measuring salt flow and electrical activity within the human brain. We cannot so much as even locate the human mind. Yet we watch as human carcasses run about, making order of disorder, conscious decisions according to subconscious criteria. We see the design and complexity that result from the operation of the brain through the invisible realm known as the mind. Thus, we know with certainty that the human mind exists. Therefore, it's absolutely logical for Creationists to postulate the existence of a Creator based upon the same "evidence." The design we see all around us came from one, grand concept, and such a concept can only come from a complex Mind. Furthermore, the mathematical and physical laws inherent in all things (including, most dramatically, the Laws of Thermodynamics and the Law of Cause and Effect) effectively validate this evidentiary claim. Evolution vs. Creation: Origins In the Evolution vs. Creation conflict, Evolutionists do quite well in terms of theoretical science, but fail to find empirical evidence. Evolutionists theorize that the universe, with all that it contains (space, time, matter and energy), exploded from nothing. This is contrary to the First Law of Thermodynamics. Where did space, time, matter and energy come from in the first place? Thus, for Evolutionists, the ultimate question of Origins remains unsolved. To complicate the Evolutionary position, this original explosion of everything from nothing is unable to explain all of the complexity and fine-tuning in the universe, including cosmic "voids" and "clumps", retrograde motion of the galaxies, etc. Despite numerous problems, this explosion from nothing has been dubbed the "Big Bang" and is the accepted theory among the majority of Evolutionists. Evolution is a very unique "science." Typically, scientists observe evidentiary data and then formulate their conclusions. Evolutionists have formulated their conclusion, and now look for the missing data. Let us take a careful look at the charge the theory of evolution is not supported by evidence. 3 Null Hypothesis Consider the proposal that “… the burden of evidence should be upon the Evolutionists, since Creation has been the historic and inherent default throughout virtually all cultures and religions until roughly the last 200 years.” This in effect says that creationism is the null hypothesis. How would you react to this position? [Also, do all cultures and religions until roughly 200 years subscribe to creationism? e.g. What about Buddhism, Advaita, …] To respond to the question about the null hypothesis, you need to understand what the term means in scientific inquiry. A null hypothesis is a proposition that you take to be true if there is no evidence against it. Take the question, “Is there a correlation between the length of a person’s nose and her ability to do research in mathematics?” I doubt if there has been any research that investigates this question, and hence it is unlikely that there is any evidence that supports or refutes this hypothesis. In the absence of evidence for or against the claim, what should you accept as true? Should you conclude that there is a correlation or should you conclude that there is no correlation? We would all agree that the conclusion should be that there is no correlation. That is the null hypothesis, what we accept as true in the absence of evidence to the contrary. Not all choices among hypotheses involve null hypotheses. Suppose there is a correlation between two properties, say, the tendency to suffer from constant nausea and the tendency to eat cloves. Is one of them the cause of the other? We should consider the following possibilities. Eating cloves causes nausea. Nausea causes the desire to eat cloves. A certain gene causes both nausea and desire to eat cloves. In the absence of evidence, we would not be able to make a choice. That is to say, none of the above is the null hypothesis. A similar situation is found in mathematics. Suppose a mathematician, say, Ramanujam, proposes a mathematical conjecture, and no one has been above to prove it. No one has been able to prove that the conjecture is false either. Should we then take Ramanujam’s conjecture as the null hypothesis and conclude that, given that it has not been proved to be false, the conjecture is true? No. As far as mathematical conjectures are concerned there are no null hypotheses. If the conjecture has not been proved to be true or false, we say that it remains a conjecture, to be investigated further. In criminal law, the null hypothesis (called “presumption”) is that the accused is innocent. (“Innocent until proven guilty.”) As a result, the “burden of proof” is on the prosecution. All that the defense has to do is to pick holes in the prosecution’s argument. In civil law, in contrast, both parties in a controversy must present arguments in support of their respective positions, and the judge chooses the position supported by the strongest argument. This is called ‘probative weight’ in law. (The criteria of truth are different for criminal law and civil law.) What I am pointing to is the distinction between situations where the null hypothesis logic is legitimate and situations where the null hypothesis logic is illegitimate. In doing so, I am pointing to the need to listen to the creationist criticism, evaluate it critically, and decide for yourselves how much of the criticism you should accept. I would recommend that you spend some time thinking about it. An Alternative Version of Creationism Let us proceed. Consider the following proposition: 4 Creationism A. The universe we live in was created by a Divine Being outside the universe who set it in motion with the Big Bang singularity, and designed in such a way that it would evolve without her (= the Divine Being’s) intervention to be the way it is now, including the humans and other creatures on Earth. The Divinity in the above theory is non-interventionist. Once the universe is set in motion with the Big Bang, she does not intervene or control its evolution. Questions for you: Q 6 Is Creationism A in conflict with the scientific theories of the origin and evolution of the universe, the origin and evolution of the solar system and the planets, the physical history of the earth, the origin and evolution of life on earth, and the origin and evolution of the human species? If your answer is yes, identify and articulate the conflict. If your answer is no, modify the theory minimally such that it would result in a conflict. Q 7 Is Creationism A a scientific theory? If it is, say what makes it a scientific theory. If it is not, say why it is not a scientific theory. Q 8 Rev. Tom Honey is a vicar (a priest) in England. Watch his 18-minute TED talk on God and the Tsunami at http://www.ted.com/talks/lang/eng/tom_honey_on_god_and_the_tsunami.html. Is his theory of Divine Creation identical to Creationism A? If it is not, identify the differences. Are the two theories in conflict? If they are, say what the contradiction is. Q 9 Go to Khan Academy at http://www.khanacademy.org, and watch the 12 minute video on Intelligent Design and Natural Selection (in the section on Biology). Is Khan’s version of Intelligent Design the same as either Creationism A or the Creationism of Allaboutcreation.org? If they are not the same, are they in conflict? Thinking Critically about Evolutionary Theory Read the Wikipedia article “Evidence of Common Descent” at http://en.wikipedia.org/wiki/Evidence_of_common_descent and carefully examine the evidence and argumentation it outlines in support of shared ancestry in evolutionary theory. When you read it, it would be useful to bear in mind that the existence of analogical phenomena (e.g., same skeletal design of hands in humans and chimpanzees, same structure of six branches in snowflakes, belief in rebirth in ancient Greece and in ancient India…) can be explained in terms of the following strategies: Shared ancestry and divergent evolution ( = analogy as homology) Shared substrate at the micro-level and/or environmental pressures leading to convergent evolution at the macro level (= analogy as homplasy) The spreading of a given pattern from a given source to some other entity or domain. ( = analogy as borrowing) Pictorially, these three strategies can be represented as: Homology Ax Bx Cx figure 1 Homoplasy Spreading B’ C’ B’ C’ Bx Cx Bx Cx figure 2 figure 3 5 Take the emergence of writing in different cultures in human history. Take the x in the above figures as trait x. In figure 1, cultures B and C have a single ancestor, A, which had writing, and the writing was preserved (with or without modification) in the descendents. In figure 2, the ancestors of B and C are B’ and C’, and neither had writing: writing emerged independently in B and C, either because of (a) environmental pressures, or (b) the properties of the substrate (human brain-mind / human society), or a combination. In figure 3, ancestor B’ had writing, but C’ didn’t. The two cultures came into contact, and culture C borrowed it from B’ (or B). There are many examples of cultural analogies of this kind, analogous traits that appear in diverse cultures. They include incest taboos, ways of cooking, territorialism, weapons, speech, belief in demons, belief in ghosts, moral codes, institutional punishment for socially undesirable actions, music, dance, painting, mathematics, medicine, trade, money, religious institutions, family, tool use, … the list goes on. Anthropologists call them cultural universals. (See the University of California TV production “CARTA: Human Cultural Universals at http://www.youtube.com/watch?v=GdAILqOZ-4M (CARTA is Center for Academic Research and Training in Anthropogeny) Questions for you Q 10 What is the evidence to show that species undergo evolution, that is, changes of traits over successive generations? Q 11 Suppose it turns out that there is evidence for the claim that species undergo evolution. It could still be the case that while species evolve (= change their traits), those changes do not result in divergence (a split, such that a single ancestor species gives rise to two distinct species). What is the evidence to show that species undergo divergence? Q 12 Suppose it turns out that there is evidence for divergence. For instance, we might be able to show that humans, chimpanzees and orangutans descended from a single ancestor species, that all insects descended from a single ancestor species, and that both lions and tigers descended from a single ancestor species. However, that some species had a single ancestor species does not necessarily show that all species had a single ancestor species. What is the evidence to show that all multi-cellular and mono-cellular species existing today (and in the past) came from a single ancestor species? Q 13 We use each of the three strategies outlined above (in figures 1-3) for different subsets of analogical phenomena. Given this situation, for any explanation in terms of shared ancestry to be convincing, it should show that alternative explanations in terms of shared substrate or borrowing are not feasible. Given that the analogy of the camera eye of humans and octopuses had independent origins (homoplasy), why is it necessary to assume that the hand plan of humans and of chimpanzees had a shared ancestor hand plan? Why couldn’t it be that different mono-cellular species appeared on the earth independently of one another from the shared substrate of the large molecules, and these distinct cell types gave rise to different species groups of multi-cellular organisms? Q 14 The Wikipedia article cites comparison of the genetic sequence of diverse organisms as evidence for common ancestry. Do the analogies of gene sequencing necessarily demonstrate common ancestry? Q 15 The article also cites fossil evidence as evidence for common ancestry. It says, “The comparative study of the anatomy of groups of animals reveals structural features that are fundamentally similar or homologous, clearly demonstrating phylogenetic and ancestral relationships with other organism, most especially when compared with fossils of ancient extinct organisms.” Does “fundamentally similar” mean they are homologous? Does fundamental similarity “clearly demonstrate phylogenetic and ancestral relationships”? 6 Q 16 The article cites ‘Biogeography’ as further evidence for shared ancestry. Does it? (Now you do the interrogation.) When you address these questions, it might be useful to bear in mind the distinction between (a) shared ancestry and divergent evolution as an explanation for the observed patterns, (b) random mutation and selection as the mechanism for species divergence. The second part comes into play only for those cases where we have already accepted the first part. The existence of natural selection does not constitute an argument against homoplastic explanation. After having gone through the first round of reading-and-interrogating, it would be a good idea to read the attached article on Waddington (Johanan W. S. Slack “Conrad Hal Waddington: The Last Renaissance Biologist?”), as well as the attached article on Octopus by Simon Morris (Simon Conway Morris “Consider the Octopus.) And while you are at it, reflect on the following questions as well. Q 17 Why do you think I asked you to read Morris’s article, and the article on Waddington? Q 18 Consider how novel species emerge in the course of evolution. The standard model is that of diversification, one species splitting into two or more. But what about fusion, the possibility of a new species coming into existence because of a combination of two or more species? Is there any evidence to show that such a process cannot exist? If there is no such evidence, and if it turns out that fusion exists, what would the evolutionary tree look like, and what challenges would it bring to the theory of evolution? More on the CARTA talks on Cultural Universals: points to ponder 1. Donald Brown refers to homology (figure 1 above) as vertical transmission and spreading (figure 3 above) as horizontal transmission. Does horizontal transmission exist in biological evolution? Why is the term “transmission” inappropriate for homoplasy? 2. Consider the concepts of change, development, and evolution. Change is a difference in a given entity between two states of affairs at two different times. Development and evolution involve changes that are perceived to have direction or at least non-reversible changes. In biology, “development” and “evolution” refer to non-reversible changes within an organism and within a population across successive generations respectively. Linguists, like biologists, use the term development to non-reversible change within an individual (language development), but they use the term history when it comes to changes in a population across successive generation (historical linguistics), reserving the term “language evolution” to refer to the emergence of language in human history. Now consider the concepts of vertical and horizontal cultural transmission in the CARTA talks. How does this connect to biology and language? 3. In physics, change is studied under dynamics, investigated using the formal tool of differential equations. Does this suggest that the study of dynamical systems may be of value to researchers in biology, linguistics and anthropology? Should biologists, linguists and anthropologists start learning differential equations? 7 Part II: Towards a Theory of Species Preliminaries: Evolutionary and Non-Evolutionary Theories A traditional system of education expects you to understand what textbooks and classrooms assert as ‘knowledge’, accept what they tell you without questioning, and learn to apply what you have understood. The previous handout, Thinking Critically about Evolution (Round 1), was meant to make you doubt and question assertions in textbooks and classrooms, look for evidence and argumentation, and accept the assertions only if you are satisfied that they are rationally justified. These are habits of thought that education should help you develop to a point that they become part of your very way of thinking. This handout follows up with a more specific aim: it hopes to help you expand the range of concepts and modes of explanation when thinking about evolution and the systematic relationship across species. If you are not familiar with the relevant evidence for the theory of evolution, it is perfectly possible that you might imagine (and believe) that all the biological species found on this planet today have always existed, that they had no ‘origin’. It is equally possible that you might imagine that every species did have an origin — that they didn’t exist before a certain point in time. Let us dub these positions the no-genesis hypothesis and the genesis hypothesis respectively. The first chapter of the Old Testament (Genesis), and Darwin’s theory of evolution (The Origin of the Species), both subscribe to the genesis hypothesis, but provide different accounts of genesis. Those who adopt the genesis hypothesis have additional options to choose from: A. Cause: Did a supernatural agency cause the origin of life (create it), or did life emerge from natural causes, for instance, the self-organization of molecules? The Old Testament and its descendent called Intelligent Design subscribe to the position of supernatural causation, while evolutionary theory subscribes to natural causation. B. Simultaneity of specio-genesis: Did all the species on the planet emerge simultaneously, or did different species appear at different times? The Old Testament subscribes to simultaneous specio-genesis, while evolutionary theory subscribes to non-simultaneous specio-genesis. C. Divergence: If we adopt the non-simultaneous view, the next question is: Can a single ancestor species diverge into two or more descendent species in the course of time? Evolutionary theory holds that such divergent evolution of species is possible. D. Convergence: Independently of our position on divergence, another question is: Can two or more ancestor species converge into a single descendent species in the course of time? The standard evolutionary tree carries the implicit assumption that such convergence species does not occur. E. Explanation for analogies across species: There are at least four possible sources for the existence of analogies across species at the molecular and morphological levels: (1) shared ancestry (a pattern in an ancestor is inherited by the descendents); (2) horizontal transfer (i.e., borrowing: a pattern in one species spreads to another); (3) independent emergence (the same pattern emerges independently in different branches of the evolutionary tree, attributable either to environmental forces or to the self-organizational predispositions of the molecular substrate). Standard evolutionary theory employs (1) as its staple mode, occasionally allowing independent emergence stemming from environmental forces, but making no attempt to explain analogy through the properties of the substrate. 8 From a historical point of view, we find that human curiosity about the nature of reality, and about the origin of the physical universe, of life, and of humanity began with mythological story telling. The Old Testament and the creation stories of the other cultures are examples of using mythology to make sense of the world. At a later stage, we proceeded to rationally coherent speculations. The Rig Veda, the Upanishads, Plato, Heraclitus, and Sankara are examples of such metaphysical speculation. When conclusions based on systematically gathered evidence replace metaphysical speculation, it is then that science begins. Such evidence-based rationality employs two related strategies: hypothesis testing, and explanation-based conclusions. Darwin’s approach to evolution was that of hypothesis testing. Our discussions on evolution have been an attempt at nudging you to adopt the strategy of explanation-based conclusions. The theory of specio-genesis in the Old Testament, as we said earlier, involves a supernatural creator. This is inadmissible in scientific inquiry. Suppose we eliminate the creator from this theory; we can state the scientifically admissible part of that specio-genesis as follows: Non-evolutionary Genesis: All species came to exist simultaneously, independently of one another. Replacing the supernatural cause with a natural one, we may represent this view as follows: natural cause species S-1 S-2 S-3 S-4… S-n Figure 1: Non-Evolutionary Genesis In contrast, evolutionary theory advances the claim: “Each new species arises by descending, with modification, from an ancestral species.” (Darwin 1859) We may state this as a weak claim or a strong claim as follows: Evolutionary Genesis: Weak claim: At least some diverse species on the planet today evolved from a common ancestor. (Evolutionary Genesis Version 1) Strong claim: All the diverse species that exist on the planet today evolved from a single common ancestor. (Evolutionary Genesis Version 2) We may represent these two positions diagrammatically as follows: natural S-1 S-1a S-1ai S-1b cause S-2 S-1c S-1aii S-2a S-3 S-4 S-2b S-2bi S-4a S-2bii S-4ai ancestors S-4b S-4aii descendents … Figure 2: Evolutionary Genesis Version 1 natural cause ancestors SPECIES-0 S-1 S-1a S-1ai S-1b S-1aii S-2 S-1c S-2a S-3 S-4 S-2b S-2bi S-4a S-2bii S-4ai Figure 3: Evolutionary Genesis Version 2 9 descendents S-4b S-4aii … This way of unpacking the idea of evolution sheds light on the different positions that are possible to take, and the differences and similarities between them. We can derive the Old Testament theory from fig. 1 by changing the natural cause to a supernatural one. With similar modifications to fig. 2 and fig. 3, we can derive various versions of Intelligent Design theories. We could also have a combination of supernatural and natural causes, the supernatural cause generating the ancestor, and natural causes governing the rest of evolution. Common to both versions 1 and 2 is the idea that evolutionary divergence of traits within a species, in the course of time, can lead to evolutionary divergence of species themselves. Now, current evolutionary theories permit evolutionary convergence of traits, but not that of species: Divergence of traits of species YES YES Convergence of traits of species YES Figure 4 NO Figure 5 An evolutionary theory that allows both divergence and convergence of species (in addition to the already accepted divergence and convergence of traits) is a perfectly plausible conjecture that needs to be carefully investigated, not dismissed without consideration. A word of caution. It might be useful to distinguish the following kinds of species convergence: i. Convergence of two species into a single species through interbreeding, resulting in the reduction of diversity. ii. Convergence of two species into a single species through interbreeding, resulting in the formation of a new species and hence in an increase of diversity. iii. Independent evolution of two different ancestor species generating the same species, resulting in greater overall diversity (but less than what it might have been had the emergence resulted in distinct species.) The term "convergent evolution" in the literature refers to the convergence of traits, not to any of the convergences in (i)-(iii). This is what comes under homoplasy. The existence of (i) is and (ii) are acknowledged in the field, but, as mentioned above, this is logically inconsistent with the phylogenetic trees that carry the implicit assumption that such convergences do not exist. And furthermore, as we shall see, they are both inconsistent with the standard textbook definition of species. In the discussion above, we reviewed possible models for a theory of evolution. The review is a way to enlarge the space within which we can construct alternative evolutionary theories without taking the classical one for granted. There are many evolutionary theories of specio-genesis that contrast with the non-evolutionary one. One conjecture that they all share is that of divergent evolution of species. If we accept “evolution” as “changes resulting from successive cycles of reproduction,” the claim that unifies all evolutionary theories is: A single ancestor species diverges into more than one distinct species in the course of successive cycles of reproduction. To critically evaluate this claim, we need to answer an important question: Question 1: In the claim that species divergence happens in evolution (and species convergence does not happen), what does the term “species” mean? This question calls for either a definition or a theory of species, probably both. 10 Text Book Definition of Species I have not seen a single satisfactory definition of species. One textbook definition of species is ‘a population of organisms capable of interbreeding to produce fertile offspring’. This definition does not work for at least three reasons. First, it says nothing about species that engage in nonsexual reproduction. Second, it leads to a contradiction. There exist well-known instances of what are called “ring species”, in which: populations interbreed (and produce fertile offspring) √ A, B √ B, C √ C, D √ D, A But A, C X B, D X If the relation, “X belongs to the same species as Y,” is transitive, then populations A, B, C, and D must belong to the same species. Yet, by the definition of species, A and C cannot belong to the same species, nor can B and D. Thus, we end up with a logical contradiction. The third problem is the logical inconsistency of the definition. Though rare, we do come across instances of fertile interspecies hybrids; one such is that of dogs and wolves producing fertile offspring. A number of such cases are documented at http://www.ratbehavior.org/Hybridization.htm: “ If the parent species are extremely closely related, they may produce fertile offspring. These are the edge cases in which the biological species concept can become too rigid. The biological species concept states that animals belong to a separate species if they cannot interbreed. So, if animals supposedly from different species interbreed and produce fertile offspring, then according to the biological species concept they should be one species. However, we usually don't consider them the same species because they differ in other features, such as geographic location, appearance, behavior, and genetics. Matings between these species produce viable, fertile offspring: Canis species: Domestic dogs (Canis familaris), wolves (Canis lupus, Canis rufus) and coyotes (Canis latrans) can interbreed and produce fertile offspring: o Evidence for domestic dog DNA in wild Siberian wolves (Vila et al. 2003) o Evidence for domestic dog DNA in wild coyotes (Adams et al. 2003b) o Evidence for red wolf-coyote hybridization (Adams et al. 2003a); Northeastern coyotes may be the product of hybridization between Canadian wolves and Western coyotes. Cichlid species in Lake Victoria are extremely variable, displaying 500 color morphs. The species are isolated through mate choice, which is determined by coloration: mates choose each other by color pattern. However, in recent years, human activity has caused the water of Lake Victoria to become cloudy. In these areas, the cichlids can't differentiate between species. In these cloudy areas, bright color morphs have disappeared and the fish have become similar and dull in appearance through hybridization.” (Seehausen et al. 1997) These discussions indicate that biologists are using some criterion other than that of the textbook definition to categorize populations into species. Given this other criterion, we can reformulate the textbook definition as a biological law instead of as a definition: Prohibition of Interspecies Interbreeding: If two populations belong to two distinct species, they cannot interbreed to produce fertile offspring. This prohibition is stated as an absolute law (without exception). Given the examples cited above, this formulation cannot be right. If we state the law in probabilistic terms, however, it 11 appears to capture a true generalization. I would therefore suggest that we state the insight as three separate but related interbreeding laws: Interspecies Interbreeding Laws Law 1 a: If two populations belong to distinct species, the probability of their mating is low. Law 1 b: The probability of interspecies mating producing an offspring is low. Law 1 c: The probability of fertile offspring resulting from interspecies mating is low. As in the case of species divergence and species convergence, this discussion raises the following questions: Question 2: In the claims that: a) the probability of interspecies mating is low, b) the probability of interspecies mating producing an offspring is low, and c) the probability of fertile offspring resulting from interspecies mating is low, what does the term “species” mean? Other Problems There are other logical problems in the standard evolutionary theory that go beyond the wellknown flaws of the textbook definition of species. One of them arises from the application of the following universal law in development and evolution: Law 2: Like begets like. We all know that rabbits cannot give birth to mice and vice versa, and that the seed of a mango tree cannot grow into a cashew tree. Parents and offspring cannot belong to distinct species. This is the phenomenon of “Like begets like,” which is expressed as law 2. Notice that the law is about species, not about traits, or about varieties within a species. Thus, it does not say that parents and offspring cannot have different traits or belong to different varieties. Also, within an Evo-Devo theory, this law holds on reproduction, the -Devo part. When it comes to the Evo- part of the theory, we have a different story: Evolution takes place in the course of a chain of successive reproductions. Evolution can result in an ancestor and descendent belonging to distinct species. (fig. 4) If no single parent-offspring pair is allowed to be distinct in their species membership, how can a chain of such links result in an ancestor-descendent pair with distinct species memberships? What we see here is a different manifestation of the transitivity problem of the relation “x belongs to the same species as y” that we encountered in the ring species. species-j species-k Evolution: ancestor descendent Reproduction: …parent offspring-parent species-j species-j offspring-parent species-j offspring-parent species-j Figure 6 If no individual link in the chain allows a mismatch between the parent’s species and the offspring’s species, how can a chain of such links give rise to a new species? It is possible to eliminate the logical contradiction by constructing a theory of species that conforms to the law, “Like begets like,” in -Devo but not in Evo-. If so, our third question can be formulated as follows: 12 Question 3: In the claims that: a) parents and offspring cannot belong to distinct species, but b) ancestors and descendents need not belong the same species, what does the term “species” mean? Uniformity Principle One assumption that has always been at the center of astronomy and physics is the Uniformity Principle. The principle was extended to geology by James Hutton, and was subsequently popularized by John Lyles; it was available to Darwin, and was something he appealed to. The assumption that species diverge (fig. 4) needs to be tested now against this uniformity principle.A general version of the principle that applies to both space and time can be formulated as follows: Uniformity Principle: Unless there is evidence to the contrary, it is reasonable to conclude that a general law that holds on the present time and current location (here and now) also holds for a distant time and distant location (there and then, whether past or future.) The principle works as follows. Suppose someone tells us that there existed humans with wings a million years ago. We have not seen a single winged human; so, based on samples from the present, we formulate a law: "Humans have no wings." As the null hypothesis, this would apply to the past as well. Unless there is evidence to the contrary, we are justified in concluding that there were no humans with wings in the past either. Likewise, suppose someone asked you whether the gravitation of a particular planet in a far away galaxy that we have not observed obeys the inverse cube law, and not the inverse square law. We have no evidence to conclude either way, so we must go by the uniformity principle: the law that applies to our location (here) also applies to the far away location (there). By the same token, we must adopt the following null hypothesis for evolution: Null hypothesis: Like begets like in evolution. By this null hypothesis, ancestors and descendents cannot belong to distinct species. The apparent logical contradiction between evolution and development in fig. 6 lends support to the null hypothesis. In the absence of positive evidence to the contrary, we must assume that the null hypothesis holds, in which case the divergence-of-species hypothesis would be false. To establish the credibility of any version of evolutionary theory, in other words, we must show that divergence of species can be observed in the present, and that hence the null hypothesis is false. The kind of evidence that textbooks and popular literature commonly present for the existence of evolution in the present happens to be evidence for the divergence of traits within a species, not of species themselves. The story of the peppered moth, for instance, is often used to demonstrate observable evolution in the present. The population of this moth before the industrial revolution was predominantly light-coloured; but as industrial soot began to accumulate, dark-coloured moths began to appear, and soon became the most common form. But this does not point to the light-coloured and dark-coloured moths belonging to distinct species; it is only evidence for adaptation, which results in the divergence of traits, and then in divergent varieties within the same species. To give evidence for the divergence of species, we must show that the divergence in the traits of the two varieties result in the two populations bifurcating into two distinct species. In the research literature on the subject, unlike in popular literature and textbooks,, there is considerable experimental evidence for present day species divergence, summarized by Joseph Boxhorn in “Observed Instances of Speciation.” (http://www.talkorigins.org/faqs/faqspeciation.html) It is interesting that many of the examples involve hybridization of different 13 varieties or even different species resulting in the formation of a new species. Speciation through interspecies hybridization can be represented as follows: Species A Species B parent-i parent-j Species C Figure 7 Interestingly, this is an instance of the convergence of species (fig. 5) that is currently not part of the standard theory. Given this result, we must revise question 1 as follows: Question 1’: In the claim that both species divergence and species convergence happen in evolution, what does the term “species” mean? A First Step towards a Theory of Species I am not a biologist, so I will not attempt to construct a theory of species: I will leave that task to the experts in the field. As a trans-disciplinary theorist, however, I would like to sketch the bare outlines of the kind of theory we are looking for, and to offer a couple of suggestions. First, there is an important difference between an operational definition that we can use to mechanically decide on the grouping of an observed entity, and a theoretical definition of an abstract concept as an ingredient of an explanation. The definition of a woman as an adult human female is an operational definition: given any living or non-living observable entity, we can make a decision on its being a woman by asking “Is it a human being?” “Is it a female?” and “Is it an adult?” The definition of a female as a class of organisms with ovaries is also one that lends itself to a mechanical operation to decide on its femaleness. In contrast, a theoretical definition may not be directly connected to decisions on observables. The connection between the observables and the definition is made through the mediation of a set of theoretical propositions. The construct that is defined this way resides at nexus of the propositions. Unlike the distinction between male and female, for instance, the category of feminine is a theoretical concept. Likewise, unlike speed, for which we can provide an operational definition, no operational definition is possible for gravity. If we define gravity as the force that holds between any two bodies with mass, we still need to connect the definition to the observable entities through a number of other definitions (e.g., force, acceleration, velocity, mass, etc.), and the laws of gravitation and motion. The type of definition we are looking for in the concept of species is a theoretical one. Something like the following is a plausible candidate: Species (Def): Species-hood is an abstract property of a population of biological organisms that guides the reproduction and evolution of the population. This definition does not tell us whether or not we should group wolves and dogs into a single species or two distinct species, but it is adequate for the purposes of acting as a starting point for a theory. Next we must formulate a set of general laws in such a way that the concept of species appears at the nexus of these laws, and thereby acquires empirical substance. The following laws might be a reasonable starting point, even though they will have to be revised significantly: Law 3a: Resemblance: The aggregate of equivalences (analogies) in genes, proteins, anatomy, physiology, ontogenesis, and behaviour is greater within species than across species. Law 3b: Mating: The affinity for mating is greater within a species than across 14 species. Law 3c: Like Begets Like: Parents and offspring cannot belong to distinct species. (= law 2) Law 3d: Divergence: When two varieties of a species are prevented from mating by external conditions, they evolve into distinct species over time. Law 3e: Hybrid Offspring: Mating across species is unlikely to produce offspring. Law 3f: Fertile Offspring: Offspring resulting from mating across species is unlikely to be fertile. [Sidebar: Notice that Laws 3a, b, e, f are probabilistic laws. At some stage in the fleshing out of the theory of species, it should be possible to formulate these laws in quantitative terms. In particular, this is necessary for the law of resemblance, which cannot be critically evaluated without designing a way of numerically calculating the distance between populations in terms of their aggregate equivalences. We would also need a threshold of distance beyond which two populations would be treated as distinct species. These technical tasks are beyond the scope of the conceptual exploration we are currently pursuing.] What Kind of Categories are Species? In logico-mathematical terms, a category is a set. A set is a collection of entities. A collection consisting of an apple, a baby, the number six, and all the students in a university is a set; so is the collection of all the even numbers. Thus, the members of a set may or may not have anything in common. A category is a set of entities that have something in common. The set of even numbers forms a category, but the set consisting of an apple, a baby, the number six, and all the students in a university does not constitute a category. A category is a natural class, not an arbitrary class. In this sense, species is a category. But what formal concept of set must we use to model of the concept of species? To answer this question, we need to briefly review the available options. The classical notion of set is that of a crispy set with a clearly defined boundary, such that the definition of the set can tell us whether or not a given candidate belongs to that set. Such a set, by stipulating sufficient and necessary conditions in discrete terms, divides the universe into two, as in fig. 8, separating entities that are members of the set from those that are not. member non-member Figure 8: Classical Concept of Set The set of even numbers is such a crispy set. Discreteness is an essential characteristic of a classical set. The set of red objects, for instance, does not form a classical set, because redness is a gradient attribute, not a discrete attribute. The formalism that captures such categories is that of the fuzzy set. There are two properties that distinguish a fuzzy set from a crispy set. One is a center that expresses prototypical attributes, and the other is the absence of boundaries. At the center of the set of red objects for instance, is the prototype of red, receding to less protypical and marginal reds when moving away from the center, as in fig. 9: (the reader has to imagine the red receding gradually, without breaks). center 15 fuzzy margin Figure 9: Fuzzy set with center There are other ways of combining the discrete-gradient parameter with the parameter of the presence and absence of a center. For instance, we could have a fuzzy set without boundaries, but also without a center. Imagine, for instance, the rainbow spectrum where orange is gradually replaced by red on the one side and yellow on the other. Here, orange is a set and so is yellow, and in between orange and yellow are entities that could belong to either — orangish yellow or yellowish orange, as in fig. 10. Here, there is no center, and there are no boundaries. Likewise, we could have a series of inner and outer circles with crispy boundaries, as in fig. 11. Such a set has both boundaries and a center. Figure 10: Fuzzy set without center Figure 11: Crispy set with center Let us look at one last model of sets. Consider the following propositions: Condition 1: The minimum age for admission to the program is 20. Condition 2: A candidate will be admitted if (s)he has: (a) scored at least 90% in the DPLE test; or (b) completed a four-year undergraduate program in a North American or European university. Condition 1 is a necessary condition for admission. But it is not a sufficient condition. Conditions (2a) and (2b) by themselves are neither necessary nor sufficient. It is only when they combine that they jointly tell us what belongs to the set and what doesn’t. Now consider the following definition of scientific inquiry: Condition A: Only those forms of rational inquiry that justify their conclusions on the basis of independently corroborable observational grounds constitute scientific inquiry. Condition B: Academic inquiries that employ the following are forms of scientific inquiry: 1) quantitive observations 2) experimental observations or 3) instrumental observations Condition A is a necessary condition. Conditions B1-3 are sufficient conditions. A particular form of scientific inquiry may satisfies none, some, or all of these three conditions. Inquiry that satisfies all of them is a prototypical form of scientific inquiry. To the extent that it satisfies less conditions, it becomes less prototypical. non-typical typical prototypical Figure 12 16 What Fig. 12 shows is a prototype model of sets. Given such a range of formal concepts, the choice from among them will have important consequences for a theory of species. If we adopt the classical set, for instance, a population either belongs to a given species or doesn’t belong to it; there is nothing in between. In a fuzzy set with a center, on the other hand, membership is gradient: the attribute of belonging to a species can range from 0 to 1, instead of being either 0 or 1. This would mean that two varieties of a species can be located at two opposite margins, and can therefore be intermediate between belonging to the same species and belonging to two different species. This is precisely the consequence we want in cases that fit with fig. 7: interbreeding between two different species producing an offspring that then becomes the ancestor of a novel species. Likewise, consider the problem of the ring species. Suppose we take aggregate distance along the parameter of analogies as the criterion for species assignment. If the distance between two populations goes beyond a threshold, then they belong to different species. If so, fig. 10 allows for species assignments of the following kind: colours orange and yellow orange and red yellow and red belong to the same species the same species distinct species the distance between them below the threshold below the threshold above the threshold This is exactly the result we want in the ring species. In the classical set theory, if A and B belong to the same set along a given dimension, and B and C belong to the same set, then A and C must belong to the same set. This is not necessary in a fuzzy-prototype set. Finally, consider the logical contradiction we encountered in the application of the Like-begetslike law to development (where it should apply) and evolution (where it shouldn’t apply). Suppose “x and y belong to the same set” is determined in terms of aggregate analogy distance in a fuzzy-prototype set. Given this as the model for species, we would expect the distance between parent and offspring in any given parent-offspring pair to be very small, but we would also expect the distance to keep increasing over successive cycles of reproduction. If so, it follows that the law “Like begets like” would apply in development but not in evolution. The logical contradiction arising from the classical set disappears in the fuzzy-prototype set. In sum, replacing the classical set with a fuzzy-prototype set in modeling the idea of species allows us to escape all the problems of the classical species theory discussed in the previous sections. Issues to think about Adopting the fuzzy-prototype model requires us to rethink the species concept. Consider, for instance, the formulation of our laws of mating, hybrid offspring, and fertile offspring, repeated below for convenience: Law 3b: Mating: The affinity for mating is greater within a species than across species. Law 3e: Hybrid Offspring: Mating across species is unlikely to produce offspring. Law 3f: Fertile Offspring: Offspring resulting from mating across species is unlikely to be fertile. Let us compare this with the textbook definition of species: If two populations belong to different species, they cannot produce fertile offspring. The textbook definition, as we have seen, is inconsistent with the findings of biologists, and as formulated above, the law is false. Furthermore, it is contradictory to the idea of experiments of interbreeding to create new species to provide evidence for evolutionary theory. This falsity and 17 logical contradiction both disappear when the insight is formulated as three laws within the fuzzy-prototype model. The insight that the crossbreeding experiments show can be expressed as follows: When mating happens between two populations that are almost not members of the same species, their offspring is likely to become an ancestor of a novel species. This result should not be surprising. Creativity is the result of the coming together of two distinct strands that are sufficiently far apart but not in total opposition. One might also speculate (as one of the students in the class did), that the threat of extinction coming from near non-fertility leads to specio-genesis. If this account is along the right track, we might be pushed to a somewhat radical conclusion. In terms of the mathematics of graph theory, a tree is a rooted non-cyclic graph. The graph in fig. (13) is a tree, but those in (14) and (15) are not: Figure 13 Figure 14 Figure 15 If we allow convergent evolution of species, as in fig. 5 and fig. 7, we are forced to adopt fig. 15. And if we adopt fig. 15 and allow cyclic graphs, it naturally raises doubts about the singleroot requirement. What underlies fig. 2 is the idea expressed by fig. 14. Exploring the possibility of accepting the idea in fig. 14 for evolution would revolutionize the way we think about the phylogenetic tree. If the issues discussed above have aroused your interest, you could pursue further some of the related ideas in the following articles: (1) Doolittle, Ford W. (1999) “Phylogenetic Classification and the Universal Tree.” Science 284, 2124 (1999). [Argues against the traditional tree in figure 13.] (2) Pennisi, Elizabeth (2008) “Modernizing the Modern Synthesis.” Science 11 JULY 2008 Vol 321. [Reports on the Alternberg conference on Evolution that questions many of the assumptions of the mainstream evolutionary theory.] (3) Fuhrmann, G. Y (1991) “Note on the Integration of Prototype Theory and Fuzzy-Set Theory.” Synthese 86: 1-27, 1991. [Discusses precisely what the title says it does.] (4) Wake, David, Varvalee H Wake and Chelsea D Specht (2011) “Homoplasy: From Detecting Pattern to Determining Process and Mechanism of Evolution.” Science 331, 1032. [Has an excellent discussion of different categories of homoplasy.] (5) Slack, Jonathan M. (2002) “Conrad Hal Waddington: the last Renaissance biologist?” Nature Reviews Vol 3 November 2001 889. [Has an excellent discussion of Waddington’s revolutionary ideas, including that of the epigenetic landscape.] (6) Morris, Simon Convey (2011) “Consider the Octopus.” Embo reports Vol 12 / No3 182. [Discusses convergent evolution and homoplasy.] 18