Darwins Theory On his return to England, Darwin began to pull together a great deal of information from the voyage

and other sources, notably a reading of Thomas Robert Malthus book On Population. He was able to arrive at a mechanism for all the changes that he had observed. He proposed that evolution has taken place through natural selection. Summary of Darwins theory: 1. Species are mutable. Darwins observations on the Galapagos made it clear to him that species can alter over time, and can give rise to new species. This was part of Lamarcks theory too, but Darwin realized as Lamarck did not that it is the environment in which organisms find themselves that determines what kinds of organisms will survive there. 2. Natural selection can give rise to new adaptations and new species. The principle of Malthus (see graph below) applies to natural populations. Because unchecked reproduction leads to exhaustion of resources, not all individuals are able to survive and reproduce. The result is a struggle for existence, leading to an increase in the probability that the best-adapted individuals will survive and reproduce. It is these individuals that are more likely to pass their characteristics on to the next generation. This process is one of natural selection, and it operates on the individuals of each generation. Darwin was led to this idea by the writings of Thomas Robert Malthus (1766-1834), who pointed out that because populations increase exponentially while their resources increase at the most arithmetically, populations will always tend to outstrip their resources.

Resources (hectares of agricultural land)

3.5 106

3.5 106

3 106

3 106

Population

2.5 106

2.5 106

2 106

2 106

1.5 106

1.5 106

1 106 0 20 40 60 80 100

1 106 120

Time (years)
The contrast between the exponential growth of a human population (red) and the arithmetic growth of its resources (blue). 3. Natural and artificial selection are very similar but not identical. Darwin realized that one of the most convincing demonstrations of the power of selection comes from the artificial selection used by plant and animal breeders to produce rapid changes over short periods of time in a wide variety of domesticated breeds. He was particularly fascinated by pigeon breeding, and carried out experiments of his own that convinced him of the remarkable and swift effects of artificial selection on these birds.

Some of the many varieties (variants produced by artificial selection) of pigeons that have been produced by pigeon-breeders over a span of thousands of years, at least since the time of the ancient Mesopotamians (pigeons appear on coins dated from 4500 B.C.). All of these varieties are descended from an ancestral species still living in the wild, the blue rock dove Columbia livia (above).

Darwin recognized that the results of natural and artificial selection were different. The artificially selected pigeon varieties could not survive in the wild,

because they had been selected by their human owners for characteristics that have nothing to do with survival or indeed might actually decrease their ability to survive. Nonetheless, he emphasized that selection of any kind, natural or artificial, is capable of bringing about changes in any living species.

4. Variation must be heritable. For evolution to take place, the bestadapted individuals must be able to pass their characteristics on to the next generation a process that Darwin called descent with modification. The modifications, as we now know, arise through mutational change in the genetic material, but Darwin thought incorrectly that the inherited modifications were changes in an organisms characteristics that were acquired during its lifetime and passed on to its offspring. 5. Evolution is gradual. Evolutionary change happens gradually, in very small increments, and there are no large leaps or saltations. Darwin was aware that distinctly different new varieties of animals and plants could suddenly appear. In his time these were termed sports by plant and animal breeders, although we now call them mutants. But he thought that such mutants were not important to evolution, because they tended to be ill adapted to their environment. The question of whether mutations of large effect can sometimes be important in evolution remains open, however, as we will see. 6. Evolution has no goal. Evolution is not under the control of a guiding intelligence, nor is its direction dictated by innate properties of living organisms. It can go in any direction. Organs and structures can disappear as well as appear over time, and rudimentary organs and structures such as the flightless cormorant wing have become small because they no longer contribute to the survival of the individual. As a result, any changes that make them smaller are not selected against and may actually be selected for if for example the environment has changed and having wings capable of flight is not beneficial but is actually harmful. Any change in structure and function, which can be effected by small stages, is within the power of natural selection (Origin, Volume 2 p. 261).

Properties of natural selection. Public Issues. Social Darwinism. Much to Darwins distress, the philosopher Herbert Spencer (1820-1903) and others quickly applied his idea of natural selection to

contemporary human affairs. Spencer, who coined the phrase survival of the fittest in 1864 to describe natural selection, had begun to think about the effects of selection on human societies even before the publication of Darwins Origin. His adherents, who included the millionaire steel magnate Andrew Carnegie, tried to justify the exploitative capitalist system of the day as simply an example of social Darwinism. Eugenics. Darwins cousin, the pioneering statistician Francis Galton (1822-1911), went a step further that Spencer, proposing that some members of society were unfit to reproduce. He coined the term eugenics (good in birth) in 1883 and founded a Laboratory of Eugenics at the University of London in 1904. There have been many attempts during the twentieth century by various eugenics movements, including the Nazi-led movement in Germany, to improve our species. We will examine the history of this concept, and how it is being revived again, later in this course. The Conflict Between Evolution and Religion. From the moment of the publication of the Origin, religious groups registered strong objections to the assumption implicit in Darwins theory that there had been no special creation of living organisms. It is noteworthy that the Anglican Church, the first to object, has now accepted the principles of evolution, as have the Catholic Church and other major religions. Some religious fundamentalist groups (Creationists), believers in a literal interpretation of the bible, continue to raise objections to the theory, particularly in the United States.

A Victorian cartoon showing Darwin discovering that his ancestors were oysters.

Scientific Issues Human Origins. The first reaction of some in the general public and the religious community was horror at the idea that we should be descended from monkeys or perhaps even more primitive creatures. After his theory of evolution had become widely accepted, Darwin examined the question of human origins in an extremely important book, The Descent of Man and Selection in Relation to Sex. He concluded that natural selection, and especially sexual selection, had been responsible for the origin of our own species. Darwin stated clearly that our species had a natural rather than a supernatural origin. The Absence of Missing Links. Darwin envisioned his process of natural selection as acting on small differences between individuals, for reasons that will become apparent shortly. Such selection should result in gradual changes in populations of organisms over time This type of evolution would seem to imply that there ought to be missing links or a series of intermediate types of animals or plants that should fill in the many gaps in the fossil record.

Some of the gaps result from the fact that many organisms fossilize poorly.

The relative numbers of living and fossil species in different groups of animals. Note that animals that fossilize poorly, such as the insects, are underrepresented in the fossil record. Many missing links have, however, been found in the fossil record. Soon after the publication of the Origin, fossils of early dinosaur-like birds were discovered, and recently the fossils of many feathered dinosaurs have been found in China. Since Darwins time a clear picture of bird evolution has emerged, in which feathers appear to have begun as fuzzy down and decorative plumage on some dinosaur lineages, and were only later employed to aid gliding and flight.

Archaeopteryx, found in a fossil bed in Germany. Many other fossils of this animal have been found. Note the tail, which is clearly different from that of modern birds. How did birds evolve?

A single missing link is not enough to fill in an evolutionary story. How for example did feathers evolve? It seems unlikely that they evolved in such a way as to permit immediate flight, because flight feathers are large and complex and are unlikely to have appeared at once.

An artists reconstruction of the early Cretaceous, 125 million years ago, in what is now northeastern China. In the remarkable Liaoning formation, filled with superbly preserved fossils of both mammals and reptiles, a wide variety of feathered dinosaurs and early birds have been found. They include the large birdlike dinosaur Sinosauropteryx, with primitive downy feathers, the feathered dinosaur Protarchaeopteryx on the right, shown displaying its plumage, and the

primitive bird Confuciusornis on the branch above. In fossils found in these beds, all the stages of feather evolution have been found. It is now clear from both molecular and fossil evidence that feathers evolved in a series of stages, that they were brand-new structures and not simply modified reptilian scales, and that they must have had a variety of functions.

Two genes, sonic hedgehog and bone morphogenetic protein 2, play large roles in feather development, just as they do in other bone and skin structures. These genes produce positional signaling molecules that affect cell differentiation, resulting in a variety of different patterns according to how far the cells are from the source of the signaling. The pattern of expression of these genes in the follicle collar (the sonic hedghog pattern is shown in the figure) varies in collars that will give rise to different types of feather. Five stages of feather development can be distinguished, and fossils of these feathers have been found that represent all five stages. Perhaps most remarkably, the most advanced flight-type feather of stage 5 has been found in a reptile fossil from Liaoning, Microraptor gui, that was clearly dinosaur-like and not a bird, and that had in effect four wings! If it was a flyer, it must have been an extremely clumsy one. It seem more likely to have been able to glide for short distances. Thus, all the stages of feather evolution actually took place before the evolution of bird-like flight. What were feathers used for before

flight? Various non-exclusive hypotheses have been proposed insulation against the cold, sexual display, gliding and at the moment it seems likely that all may have played a role in the early stages of feather evolution.

A reconstruction of Microraptor gui.

Flight appears to have evolved through a process of exaptation, in which a pre-existing structure or behavior is coopted to take up a new function. A wide variety of elaborate feathers evolved for various reasons not related to flight, even before the evolution of birds and of feathered flight. As a result it was fairly easy for natural selection to produce alterations in feather number and shape, so that feathers were able to play a role in the evolution of new abilities for gliding and finally flight. Such exaptations have been common in evolution, and we will see a number of examples of them in this course. A combination of lucky fossil finds that have been carefully analyzed, along with advances in molecular biology, have allowed us to test a variety of hypotheses suggested by the data. We now have a picture of feather and bird evolution that is far more complete than the scanty information that was available to Darwin. (More on artificial selection: A recent paper in Science shows that an important determiner of the difference in size between small and large dogs is a single base change in an insulin-like growth factor gene IGF1. There are signs that a selective sweep has taken place in the region around this gene in small dogs, reducing the genetic variation in this region in small dogs vs large ones. Thus, small dogs have been subject to strong selection for this character. Several small genetic differences in or near the IGF1 gene separate large and small dogs. The authors suggest that selection for small dogs began when the first cities arose. Fifteen-thousand-year-old dog fossils from Russia are the size of Great Danes, and twelve-thousand-year-old dog fossils from the Middle East, at the time that towns and cities first began to appear, are the size of terriers. Look in your Readings for more information.)