11/02/2005

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In this lesson, we wish to ask:

• How did observations in nature lead to the formulation of the theory of evolution?
• What are the main points of Darwin's theory of evolution?
• How does the process of natural selection work?
• What evidence do we have for local adaptation?
• How can natural selection affect the frequency of traits over successive generations?

The (R)Evolution of Theory

In other words, all of the similarities and dissimilarities among groups of organisms that are the result of the branching process creating the great tree of life (see Figure 1), were viewed by early 19th century philosophers and scientists as a consequence of omnipotent design.

Figure 1: A phylogenetic "tree of life" constructed by computer analysis of cyochrome c molecules in the organisms shown; there are as many different trees of life as there are methods of analysis for constructing them.

However, by the 19th Century, a number of natural historians were beginning to think of evolutionary change as an explanation for patterns observed in nature. The following ideas were part of the intellectual climate of Darwin's time.

• No one knew how old the earth was, but geologists were beginning to make estimates that the earth was considerably older than explained by biblical creation. Geologists were learning more about strata, or layers formed by successive periods of the deposition of sediments. This suggested a time sequence, with younger strata overlying older strata.
• A concept called uniformitarianism, due largely to the influential geologist Charles Lyell, undertook to decipher earth history under the working hypothesis that present conditions and processes are the key to the past, by investigating ongoing, observable processes such as erosion and the deposition of sediments.
• Discoveries of fossils were accumulating during the 18th and 19th centuries. At first naturalists thought they were finding remains of unknown but still living species. As fossil finds continued, however, it became apparent that nothing like giant dinosaurs was known from anywhere on the planet. Furthermore, as early as 1800, Cuvier pointed out that the deeper the strata, the less similar fossils were to existing species.
• Similarities among groups of organisms were considered evidence of relatedness, which in turn suggested evolutionary change. Darwin's intellectual predecessors accepted the idea of evolutionary relationships among organisms, but they could not provide a satisfactory explanation for how evolution occurred. 
• Lamarck is the most famous of these. In 1801, he proposed organic evolution as the explanation for the physical similarity among groups of organisms, and proposed a mechanism for adaptive change based on the inheritance of acquired characteristics. He wrote of the giraffe:

"We know that this animal, the tallest of mammals, dwells in the interior of Africa, in places where the soil, almost always arid and without herbage, obliges it to browse on trees and to strain itself continuously to reach them. This habit sustained for long, has had the result in all members of its race that the forelegs have grown longer than the hind legs and that its neck has become so stretched, that the giraffe, without standing on its hind legs, lifts its head to a height of six meters."

In essence, this says that the necks of Giraffes became long as a result of continually stretching to reach high foliage. Larmarck was incorrect in the hypothesized mechanism, of course, but his example makes clear that naturalists were thinking about the possibility of evolutionary change in the early 1800's.

• Darwin was influenced by observations made during his youthful voyage as naturalist on the survey ship Beagle. On the Galapagos Islands he noticed the slight variations that made tortoises from different islands recognizably distinct. He also observed a whole array of unique finches, the famous "Darwin's finches," that exhibited slight differences from island to island. In addition, they all appeared to resemble, but differ from, the common finch on the mainland of Ecuador, 600 miles to the east. Patterns in the distribution and similarity of organisms had an important influence of Darwin's thinking. The picture at the top of this page is of Darwin's own sketches of finches in his Journal of Researches.

• In 1858, Darwin published his famous On the Origin of Species by Means of Natural Selection, a tome of over 500 pages that marshalled extensive evidence for his theory. Publication of the book caused a furor - every copy of the book was sold the day that it was released. Members of the religious community, as well as some scientific peers, were outraged by Darwin's ideas and protested. Most scientists, however, recognized the power of Darwin's arguments. Today, school boards still debate the validity and suitability of Darwin's theory in science curricula, and a whole body of debate has grown up around the controversy (see the WWW site Talk.Origins for an ongoing dialogue). We do not have time to cover all of Darwin's evidence and arguments, but we can examine the core ideas. What does this theory of evolution say?

 

Darwin's Theory

Darwin's theory of evolution has four main parts:
1. Organisms have changed over time, and the ones living today are different from those that lived in the past. Furthermore, many organisms that once lived are now extinct. The world is not constant, but changing. The fossil record provided ample evidence for this view.
2. All organisms are derived from common ancestors by a process of branching. Over time, populations split into different species, which are related because they are descended from a common ancestor. Thus, if one goes far enough back in time, any pair of organisms has a common ancestor. This explained the similarities of organisms that were classified together -- they were similar because of shared traits inherited from their common ancestor. It also explained why similar species tended to occur in the same geographic region.
3. Change is gradual and slow, taking place over a long time. This was supported by the fossil record, and was consistent with the fact that no naturalist had observed the sudden appearance of a new species. [This is now contested by a view of episodes of rapid change and long periods of stasis, known as punctuated equilibrium].
4. The mechanism of evolutionary change was natural selection. This was the most important and revolutionary part of Darwin's theory, and it deserves to be considered in greater detail.

The Process of Natural Selection

Natural selection is a process that occurs over successive generations. The following is a summary of Darwin's line of reasoning for how it works (see Figure 2).
• If all the offspring that organisms can produce were to survive and reproduce, they would soon overrun the earth. Darwin illustrated this point by a calculation using elephants. He wrote:

 
"The elephant is reckoned the slowest breeder of all known animals, and I have taken some pains to estimate its probable minimum rate of natural increase; it will be safest to assume that it begins breeding when 30 years old and goes on breeding until 90 years old; if this be so, after a period from 740 to 750 years there would be nearly 19 million elephants descended from this first pair."
•  

Figure 2: The Process of Natural Selection

• This unbounded population growth resembles a simple geometric series (2-4-8-16-32-64..) and quickly reaches infinity.
• As a consequence, there is a "struggle" (metaphorically) to survive and reproduce, in which only a few individuals succeed in leaving progeny.
• Organisms show variation in characters that influence their success in this struggle for existence. Individuals within a population vary from one another in many traits. (Animal behavioralists making long-term studies of chimps or elephants soon recognize every individual by its size, coloration, and distinctive markings.)
• Offspring tend to resemble parents, including in characters that influence success in the struggle to survive and reproduce.
• Parents possessing certain traits that enable them to survive and reproduce will contribute disproportionately to the offspring that make up the next generation.

To the extent that offspring resemble their parents, the population in the next generation will consist of a higher proportion of individuals that possess whatever adaptation enabled their parents to survive and reproduce.

The well-known example of camouflage coloration in an insect makes for a very powerful, logical argument for adaptation by natural selection. Development of such coloration, which differs according to the insect's environment, requires variation. The variation must influence survival and reproduction (fitness), and it must be inherited.

During the early and middle 20th Century, genetics became incorporated into evolution, allowing us to define natural selection this way:
 
 

Natural Selection is the differential reproduction of genotypes.

Natural Selection Requires...

For natural selection to occur, two requirements are essential:
1. There must be heritable variation for some trait. Examples: beak size, color pattern, thickness of skin, fleetness.
2. There must be differential survival and reproduction associated with the possession of that trait.
Unless both these requirements are met, adaptation by natural selection cannot occur.

Some examples:

• If some plants grow taller than others and so are better able to avoid shading by others, they will produce more offspring. However, if the reason they grow tall is because of the soil in which their seeds happened to land, and not because they have the genes to grow tall, than no evolution will occur.
• If some individuals are fleeter than others because of differences in their genes, but the predator is so much faster that it does not matter, then no evolution will occur (e.g. if cheetahs ate snails).
In addition, natural selection can only choose among existing varieties in a population. It might be very useful for polar bears to have white noses, and then they wouldn't have to cover their noses with their paws when they stalk their prey. The panda could have a much nicer thumb than the clumsy device that it does have.

When we incorporate genetics into our story, it becomes more obvious why the generation of new variations is a chance process. Variants do not arise because they are needed. They arise by random processes governed by the laws of genetics. For today, the central point is the chance occurrence of variation, some of which is adaptive, and the weeding out by natural selection of the best adapted varieties.
 

Evidence of Natural Selection

Let's look at an example to help make natural selection clear.

Industrial melanism is a phenomenon that affected over 70 species of moths in England. It has been best studied in the peppered moth, Biston betularia. Prior to 1800, the typical moth of the species had a light pattern (see Figure 3). Dark colored or melanic moths were rare and were therefore collectors' items.

Figure 3. Image of Peppered Moth

During the Industrial Revolution, soot and other industrial wastes darkened tree trunks and killed off lichens. The light-colored morph of the moth became rare and the dark morph became abundant. In 1819, the first melanic morph was seen; by 1886, it was far more common -- illustrating rapid evolutionary change. 

Eventually light morphs were common in only a few locales, far from industrial areas. The cause of this change was thought to be selective predation by birds, which favored camouflage coloration in the moth.

In the 1950's, the biologist Kettlewell did release-recapture experiments using both morphs. A brief summary of his results are shown below. By observing bird predation from blinds, he could confirm that conspicuousness of moth greatly influenced the chance it would be eaten.

Recapture Success

	light moth	dark moth
non-industrial woods	14.6 %	4.7 %
industrial woods	13 %	27.5 %

Local Adaptation - More Examples

So far in today's lecture we have emphasized that natural selection is the cornerstone of evolutionary theory. It provides the mechanism for adaptive change. Any change in the environment (such as a change in the background color of the tree trunk that you roost on) is likely to lead to local adaptation. Any widespread population is likely to experience different environmental conditions in different parts of its range. As a consequence it will soon consist of a number of sub-populations that differ slightly, or even considerably.

The following are examples that illustrate the adaptation of populations to local conditions.

• The rat snake, Elaphe obsoleta, has recognizably different populations in different locales of eastern North America (see Figure 4). Whether these should be called geographic "races" or subspecies is debatable. These populations all comprise one species, because mating can occur between adjacent populations, causing the species to share a common gene pool (see the previous lecture on speciation).
 



Figure 4: Subspecies of the rat snake Elaphe obsoleta, which interbreed where their ranges meet.

• Galapagos finches are the famous example from Darwin's voyage. Each island of the Galapagos that Darwin visited had its own kind of finch (14 in all), found nowhere else in the world. Some had beaks adapted for eating large seeds, others for small seeds, some had parrot-like beaks for feeding on buds and fruits, and some had slender beaks for feeding on small insects (see Figure 5). One used a thorn to probe for insect larvae in wood, like some woodpeckers do. (Six were ground-dwellers, and eight were tree finches.) (This diversification into different ecological roles, or niches, is thought to be necessary to permit the coexistence of multiple species, a topic we will examined in a later lecture.) To Darwin, it appeared that each was slightly modified from an original colonist, probably the finch on the mainland of South America, some 600 miles to the east. It is probable that adaptive radiation led to the formation of so many species because other birds were few or absent, leaving empty niches to fill; and because the numerous islands of the Galapagos provided ample opportunity for geographic isolation.



Figure 5

Stabilizing, Directional, and Diversifying Selection

Finally, we will look at a statistical way of thinking about selection. Suppose that each population can be portrayed as a frequency distribution for some trait -- beak size, for instance. Note again that variation in a trait is the critical raw material for evolution to occur.

What will the frequency distribution look like in the next generation?

Figures 6a-c

First, the proportion of individuals with each value of the trait (size of beak, or body weight) might be exactly the same. Second, there may be directional change in just one direction. Third (and with such rarity that its existence is debatable), there might be simultaneous change in both directions (e.g. both larger and smaller beaks are favored, at the expense of those of intermediate size). Figures 6a-c capture these three major categories of natural selection.

Figure 7

Under stabilizing selection, extreme varieties from both ends of the frequency distribution are eliminated. The frequency distribution looks exactly as it did in the generation before (see Figure 6a). Probably this is the most common form of natural selection, and we often mistake it for no selection. A real-life example is that of birth weight of human babies (see Figure 7).

Under directional selection, individuals at one end of the distribution of beak sizes do especially well, and so the frequency distribution of the trait in the subsequent generation is shifted from where it was in the parental generation (see Figure 6b). This is what we usually think of as natural selection. Industrial melanism was such an example.

Figure 8

The fossil lineage of the horse provides a remarkable demonstration of directional succession. The full lineage is quite complicated and is not just a simple line from the tiny dawn horse Hyracotherium of the early Eocene, to today's familiar Equus. Overall, though, the horse has evolved from a small-bodied ancestor built for moving through woodlands and thickets to its long- legged descendent built for speed on the open grassland. This evolution has involved well- documented changes in teeth, leg length, and toe structure (see Figure 8).

Under diversifying (disruptive) selection, both extremes are favored at the expense of intermediate varieties (see Figure 6c). This is uncommon, but of theoretical interest because it suggests a mechanism for species formation without geographic isolation (see the previous lecture on speciation).

 

 

Summary

Darwin's theory of evolution fundamentally changed the direction of future scientific thought, though it was built on a growing body of thought that began to question prior ideas about the natural world.

The core of Darwin's theory is natural selection, a process that occurs over successive generations and is defined as the differential reproduction of genotypes.

Natural selection requires heritable variation in a given trait, and differential survival and reproduction associated with possession of that trait.

Examples of natural selection are well-documented, both by observation and through the fossil record.

Selection acts on the frequency of traits, and can take the form of stabilizing, directional, or diversifying selection.
  

Suggested Readings

• Darwin, C. 1958. On the Origin of Species by Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life. London: J. Murray.

• Futuyma, D.J. 1986. Evolutionary Biology. Sunderland, Mass: Sinauer Associates, Inc.

• Dawkins, R. 1989. The Selfish Gene. Oxford: Oxford University Press.

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