n the mid-1960s laboratory techniques such as electrophoresis and selective assay of enzymes became available for the rapid and inexpensive study of differences among enzymes and other proteins. The application of these techniques to evolutionary problems made possible the pursuit of issues that earlier could not be investigated—for example, exploring the extent of genetic variation in natural populations (which sets bounds on their evolutionary potential) and determining the amount of genetic change that occurs during the formation of new species.
Comparisons of the amino acid sequences of corresponding proteins in different species provided quantitatively precise measures of the divergence among species evolved from common ancestors, a considerable improvement over the typically qualitative evaluations obtained by comparative anatomy and other evolutionary subdisciplines. In 1968 the Japanese geneticist Motoo Kimura proposed the neutrality theory of molecular evolution, which assumes that, at the level of the sequences of nucleotides in DNA and of amino acids in proteins, many changes are adaptively neutral; they have little or no effect on the molecule's function and thus on an organism's fitness within its environment. If the neutrality theory is correct, there should be a “molecular clock” of evolution; that is, the degree to which amino acid or nucleotide sequences diverge between species should provide a reliable estimate of the time since the species diverged. This would make it possible to reconstruct an evolutionary history that would reveal the order of branching of different lineages, such as those leading to humans, chimpanzees, and orangutans, as well as the time in the past when the lineages split from one another. During the 1970s and '80s it gradually became clear that the molecular clock is not exact; nevertheless, into the early 21st century it continued to provide the most reliable evidence for reconstructing evolutionary history. (See below The molecular clock of evolution and The neutrality theory of molecular evolution.)
The laboratory techniques of DNA cloning and sequencing have provided a new and powerful means of investigating evolution at the molecular level. The fruits of this technology began to accumulate during the 1980s following the development of automated DNA-sequencing machines and the invention of the polymerase chain reaction (PCR), a simple and inexpensive technique that obtains, in a few hours, billions or trillions of copies of a specific DNA sequence or gene. Major research efforts such as the Human Genome Project further improved the technology for obtaining long DNA sequences rapidly and inexpensively. By the first few years of the 21st century, the full DNA sequence—i.e., the full genetic complement, or genome—had been obtained for more than 20 higher organisms, including human beings, the house mouse (Mus musculus), the rat Rattus norvegicus, the vinegar fly Drosophila melanogaster, the mosquito Anopheles gambiae, the nematode worm Caenorhabditis elegans, the malaria parasite Plasmodium falciparum, the mustard weed Arabidopsis thaliana, and the yeast Saccharomyces cerevisiae, as well as for numerous microorganisms.
The Earth sciences also experienced, in the second half of the 20th century, a conceptual revolution with considerable consequence to the study of evolution. The theory of plate tectonics, which was formulated in the late 1960s, revealed that the configuration and position of the continents and oceans are dynamic, rather than static, features of Earth. Oceans grow and shrink, while continents break into fragments or coalesce into larger masses. The continents move across Earth's surface at rates of a few centimetres a year, and over millions of years of geologic history this movement profoundly alters the face of the planet, causing major climatic changes along the way. These previously unsuspected massive modifications of Earth's past environments are, of necessity, reflected in the evolutionary history of life. Biogeography, the evolutionary study of plant and animal distribution, has been revolutionized by the knowledge, for example, that Africa and South America were part of a single landmass some 200 million years ago and that the Indian subcontinent was not connected with Asia until geologically recent times.
Ecology, the study of the interactions of organisms with their environments, has evolved from descriptive studies—“natural history”—into a vigorous biological discipline with a strong mathematical component, both in the development of theoretical models and in the collection and analysis of quantitative data. Evolutionary ecology (see community ecology) is an active field of evolutionary biology; another is evolutionary ethology, the study of the evolution of animal behaviour. Sociobiology, the evolutionary study of social behaviour, is perhaps the most active subfield of ethology. It is also the most controversial, because of its extension to human societies.
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