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Basis of Darwinism 1. Organisms vary 2. Some of that variation is inherited 3. All organisms produce more offspring than can survive 4. On the average, those that survive will be the ones better adapted or suited to local environments (natural selection) N.B. As environments change, different characteristics will become advantageous How did Darwin develop his principles? Were they an incredible burst of creativity? Answer: Not entirely. The first two (organisms vary, some of that variation is inherited) certainly weren’t original with Darwin, but based on his own work. Darwin studied artificial selection in various species. The fact selection works says that some of the variation is inherited. There is incredible variety in the pigeons coming from human selection among the varying characters of pigeons. (N.B. natural variation in pigeons) Selection produced bald-headed pigeons, pigeons with fan tails, and hundreds of others. Darwin wrote a whole book about artificial selection. Statement 3 - all organisms produce more offspring than can survive Darwin only understood this after reading Malthus’ An Essay on the Principle of Population. In it, based on data from the American colonies, Malthus said that populations increase exponentially, but that resources can increase only in a linear fashion. Thus, inevitably, a population outgrows the resources it needs. For the human population, Malthus, a religious man, felt that overpopulation would lead to “misery and vice”. Darwin turned this into the 4th principle - Statement 4 - On the average, those that survive will be the ones better adapted or suited to local environments (natural selection). This has been the most difficult of the statements to verify experimentally. However, there have been some recent successes. How can experiments test the tenets of evolution? First, the general pattern of the hypotheco-deductive (“the scientific) method: Steps in the scientific method: a) make an observation that is interesting/ unexpected b) propose an explanation for the observation this is called a hypothesis. It should make testable predictions, i.e. the hypothesis should be falsifiable. c) test the hypothesis experimentally. If results fit the hypothesis, don’t stop - do other experiments. d) If many experiments that could potentially falsify the hypothesis don’t, then accept the hypothesis. Otherwise modify or reject it. The hypothesis of evolution by natural selection is testable: → to more specific deductions DEDUCTION 1. Natural selection can operate if more offspring are born than can survive DEDUCTION 2a. If the hypothesis is true, species of the remote past must be different from those alive now. - fossils in sedimentary rocks -- few of the species alive now occur as fossils in the remote past DEDUCTION 2b. The older the sedimentary strata, the less chance of finding contemporary species as fossils - this issue was first tested by Sir Charles Lyell (1797-1875) who had collected fossil shells from the Tertiary period - some species are remarkably persistent, e.g. the horseshoe crab current fossil EPOCH Fossil species Alive today % of fossil species still alive 15 MYA (recent Pliocene) 226 216 96 older Pliocene 569 238 42 Miocene 40-50 MYA (Eocene) 1021 1238 176 42 17 3 * if all species had been created in 4 days, you’d expect all rocks to have the same array of species DEDUCTION 3. - The age of the earth must be very great, possibly many millions of years (contrary to biblical dating) - first estimates based on the thickness of strata, assuming rate of deposition was constant and layers had steadily built up over time (but depostion varies with rainfall, slope, material, etc.) - 1940s: age of rock estimated in terms of the rate of radioactive decay of materials in the rock - C12:C14 If we know the ratio of C12 : C14 when the organism was alive, and we know how much C14 is in it now, and we know the rate of decay of C14 (its ‘half-life’=5730 years), then we can calculate how many years since the organism died. - Carbon dating is especially useful for dating organisms K/Ar is used for older rocks Aging rocks permits scientists to view earth’s history as a sequence of layers, as is evident if you were to travel down the Grand Canyon: DEDUCTION 4. - If the members of a taxonomic unit, e.g. phylum Chordata, share a common ancestry. That should be reflected in their structure (“Homology” = similarity of structure based on common descent. C.f. ‘Analogy’) - e.g., the anterior paired appendages of vertebrates: - pectoral fins of fish, whales, dolphins - wings of birds, bats, pterodactyls - hoofed forelegs of horses and cattle - tool-using arm of humans DEDUCTION 5. If the members of a taxonomic unit share a common ancestry, that should be reflected in their embryonic development. Ernst Haeckel (1834-1919) said “Ontogeny recapitulates Phylogeny” e.g. in the basic chordate body plan 1. Possession of a notochord 2. Gill slits or pouches in the pharynx 3. Dorsal hollow nerve tube - human adult has only one of these (the dorsal nerve tube which becomes brain and spinal cord) - AS EMBYOS we have all three! DEDUCTION 6. If there is unity of life based upon evolution and common ancestry, that should be reflected in the molecular processes of organisms. There is considerable evidence of common metabolic heritage -1. Amino acids. When produced in the lab, there are two stereo-isomers D-form (right handed) L-form (left handed) Life is all left-handed amino acids (with the exception of cell walls of some bacteria) 2. There are only 23 amino acids in biological systems 3. There are very similar sequences of amino acids in proteins of very different organisms - e.g., of 104 amino acid positions in the sequence for cytochrome c, yeast and horse are identical for 64 - e.g., insulin in pigs and humans is very similar 4. Genetic programs are common: there are only 2 nucleic acids in all life forms: DNA (or in some viruses RNA) provides the basic code. - each have only 4 bases (ATGC in DNA and AUGC in RNA) 5. ATP is the energy currency in all organisms e.g. microcomplement fixation method - compare antibodies produced by test organisms (rabbit/rat) versus antigens from different taxa (using specific proteins in blood serum) - ‘immunologic distance’ data show conservatism, e.g. primates: gibbon siamang orangutan human chimp gorilla Old world monkeys (mandrills & macacs) Does the evidence suggest a slow, steady pace for evolutionary change? Not always. The fossil record seems to show “jumps” at many points. Sometimes called “macroevolution”, a misnomer, it seems that the fossil record shows long periods of stasis in the morphology of a species, then rapid change as a new species appears. There is an explanation, and it isn’t that the pace of selection changes. Until recently studies have been observational, rather than experimental. An example of a modern experiment documenting evolution: Studies of guppy life histories in ponds in Central America Observations: Pike-cichlids predate larger, mature guppies in ponds where the pike cichlids occur. Killifish predate juvenile guppies (but less intensely) in ponds where they are found. Have the different predators driven evolution differently in ponds where the differing predators occur? Do the guppies differ? The life histories of guppies differ in ponds with pike-cichlids and in ponds with killifish. Guppies in ponds with pike-cichlids mature earlier, are smaller at maturity, and produce larger broods than those living with killifish. Good start! But is it the predators that drive the difference in life history? Hypothesis: Feeding preferences of predators select for the size and age of maturity in guppies. Test: Transplant guppies from an area with pikecichlids to one with killifish (but no guppies prior to transplantation). This is the experimental group. Remember: we need to use a number of ponds to ensure sufficient replication. We also need to keep track of a number of ponds with guppies and pike-cichlids both present, using those guppies as controls. Prediction: Transplanted guppies should (after generations of selection) mature later, at larger size, and produce smaller broods Results: Compared to controls (guppies not transplanted) over 11 years (or 30-60 generations) maturation of transplanted guppies occurred significantly later, at significantly larger size, and they produced significantly smaller broods. The life histories of the transplanted guppies had “evolved” exactly as the hypothesis had predicted. It is very difficult to design and execute experiments that can test the hypothesis of evolution. This experiment, done by David Reznick and John Endler, is a very good test. It supports evolution as Darwin presented it. Does it “prove” evolution? Of course not. There have been other tests, in sufficient numbers that we have elevated evolution to the status of a theory, and recognize it as the cornerstone of modern biology.