spring has traits that give it an advantage in a particular environment, that organism will be more likely to survive and pass on those traits. As differences accumulate over generations, populations of organisms diverge from their ancestors.

Darwin's original hypothesis has undergone extensive modification and expansion, but the central concepts stand firm. Studies in genetics and molecular biology—fields unknown in Darwin's time—have explained the occurrence of the hereditary variations that are essential to natural selection. Genetic variations result from changes, or mutations, in the nucleotide sequence of DNA, the molecule that genes are made from. Such changes in DNA now can be detected and described with great precision.

Genetic mutations arise by chance. They may or may not equip the organism with better means for surviving in its environment. But if a gene variant improves adaptation to the environment (for example, by allowing an organism to make better use of an available nutrient, or to escape predators more effectively—such as through stronger legs or disguising coloration), the organisms carrying that gene are more likely to survive and reproduce than those without it. Over time, their descendants will tend to increase, changing the average characteristics of the population. Although the genetic variation on which natural selection works is based on random or chance elements, natural selection itself produces "adaptive" change—the very opposite of chance.

Scientists also have gained an understanding of the processes by which new species originate. A new species is one in which the individuals cannot mate and produce viable descendants with individuals of a preexisting species. The split of one species into two often starts because a group of individuals becomes geographically separated from the rest. This is particularly apparent in distant remote islands, such as the Galápagos and the Hawaiian archipelago, whose great distance from the Americas and Asia means that arriving colonizers will have little or no opportunity to mate with individuals remaining on those continents. Mountains, rivers, lakes, and other natural barriers also account for geographic separation between populations that once belonged to the same species.

Once isolated, geographically separated groups of individuals become genetically differentiated as a consequence of mutation and other processes, including natural selection. The origin of a species is often a gradual process, so that at first the reproductive isolation between separated groups of organisms is only partial, but it eventually becomes complete. Scientists pay special attention to these intermediate situations, because they help to reconstruct the details of the process and to identify particular genes or sets of genes that account for the reproductive isolation between species.

A particularly compelling example of speciation involves the 13 species of finches studied by Darwin on the Galápagos Islands, now known as Darwin's finches. The ancestors of these finches appear to have immigrated from the South American mainland to the Galápagos. Today the different species of finches on the island have distinct habitats, diets, and behaviors, but the mechanisms involved in speciation continue to operate. A research group led by Peter and Rosemary Grant of Princeton University has shown that a single year of drought on the islands can drive evolutionary changes in the finches. Drought diminishes supplies of easily



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