VIII.

Development of Evolutionary Theory

The origins of life on Earth have been a source of speculation among philosophers, religious thinkers, and scientists for thousands of years. Many human civilizations used rich and complex creation stories and myths to explain the presence of living organisms. Ancient Greek philosophers and scientists were among the earliest to apply the principles of modern science to the mysterious complexity and variety of life around them. During early Christian times, ancient Greek ideas gave way to Creationism, the view that a single God created the universe, the world, and all life on Earth. For the next 1,500 years, evolutionary science was at a standstill. The dawn of the Renaissance in the early 14th century brought a renewed interest in science and medicine. Advances in anatomy highlighted physical similarities in the features of widely different organisms. Fossils provided evidence that life on this planet was vastly different millions of years ago. With each new development came new ideas and theories about the nature of life.

A.

Ancient Views

The Greek philosopher Anaximander, who lived in the 500s bc, is generally credited as the earliest evolutionist. Anaximander believed that the Earth first existed in a liquid state. Further, he believed that humans evolved from fishlike aquatic beings who left the water once they had developed sufficiently to survive on land. Greek scientist Empedocles speculated in the 400s bc that plant life arose first on Earth, followed by animals. Empedocles proposed that humans and animals arose not as complete individuals but as various body parts that joined together randomly to form strange, fantastic creatures. Some of these creatures, being unable to reproduce, became extinct, while others thrived. Outlandish as his ideas seem today, Empedocles’ thinking anticipates the fundamental principles of natural selection.

The Greek philosopher and scientist Aristotle, who lived in the 300s bc, referred to a “ladder of nature”—a progression of life forms from lower to higher—but his ladder was a static hierarchy of levels of perfection, not an evolutionary concept. Each rung on this ladder was occupied by organisms of higher complexity than the rung before it, with humans occupying the top rung. Aristotle acknowledged that some organisms are incapable of meeting the challenges of nature and so cease to exist. As he saw it, successful creatures possessed a gift, or perfecting principle, that enabled them to rise to meet the demands of their world. Creatures without the perfecting principle died out. In Aristotle’s view it was this principle—not evolution—that accounted for the progression of forms in nature.

B.

Linnaeus and Scientific Classification

Many centuries later, the idea of a perfect and unchanging natural world—the product of divine creation—was predominant not only in religion and philosophy, but in science. Gradually, however, as knowledge accumulated from seemingly disparate areas, the beginnings of modern evolutionary theory began to take shape. A key figure in this regard was the Swedish naturalist Carolus Linnaeus, who became known as the father of modern taxonomy, the science of classifying organisms.

In his major work Systema Naturae (The System of Nature), first published in 1735, Linnaeus devised a system of classification of organisms that is still in use today. This system places living things within increasingly specific categories based on common attributes—from a general grouping (kingdom) down to the specific individual (species). Using this system, Linnaeus named nearly 10,000 plant and animal species in his lifetime. Not an evolutionist by any means, Linnaeus believed that each species was created by God and was incapable of change. Nevertheless, his orderly groupings of living things provided important insights for later theorists.

C.

19th-Century Foundations

Perhaps the most prominent of those who embraced the idea of progressive change in the living world was the early 19th-century French biologist Jean-Baptiste Lamarck. Lamarck’s theory, now known as Lamarckism and based in part on his study of the fossils of marine invertebrates, was that species do change over time. He believed, furthermore, that animals evolve because unfavorable conditions produce needs that animals try to satisfy. For example, short-necked ancestors of the modern giraffe voluntarily stretched their necks to reach leaves high in trees during times when food was scarce. Proponents of Lamarckism thought this voluntary use slightly changed the hereditary characteristics controlling neck growth; the giraffe then transmitted these alterations to its offspring as what Lamarck called acquired characteristics. Modern scientists know that adaptation and natural selection are far more complicated than Lamarck supposed, having nothing to do with an animal’s voluntary efforts. Nevertheless, the idea of acquired characteristics, with Lamarck as its most famous proponent, persisted for many years.

French naturalist and paleontologist Georges Cuvier feuded with Lamarck. Unearthing the fossils of mastodons and other vanished species, Cuvier produced proof of long-extinct life forms on Earth. Unlike Lamarck, however, Cuvier did not believe in evolution. Instead, Cuvier believed that floods and other cataclysms destroyed such ancient species. He suggested that after each cataclysmic event, God created a new set of organisms.

At around the same time that Cuvier and Lamarck were squabbling, British economist Thomas Robert Malthus proposed ideas extremely influential in evolutionary theory. In his 1798 work An Essay on the Principle of Population, Malthus theorized that the human population would increase at a much greater rate than its food sources. This theory introduced the key idea of competition for limited resources—that is, there is not enough food, water, and living space to go around, and organisms must somehow compete with each other to obtain resources necessary for survival. Another key idea came from Scottish geologist Charles Lyell, who supplied a deeper understanding of Earth’s history. In his book Principles of Geology (1830), Lyell set forth his case that the Earth was millions of years old rather than only a few thousand years old, as was maintained by those who accepted the biblical story of divine creation as fact.

D.

Darwin and Natural Selection

In 1831 Charles Darwin, who was intending to become a country minister, had an opportunity to sail as ship’s naturalist aboard the HMS Beagle on a five-year, round-the-world mapmaking voyage. During the journey, as the ship anchored off South America and other distant shores, Darwin had the opportunity to travel inland and make observations of the natural world. In the Galápagos Islands, he noted how species on the various islands were similar but distinct from one another. He also observed fossils and other geological evidence of the Earth’s great age. The observations Darwin made on that voyage seemed to suggest the evolution, rather than the creation, of the many local forms of life.

In 1837, shortly after returning to England, Darwin began a notebook of his observations and thoughts on evolution. Although Darwin had developed the major components of his theory of evolution by natural selection in an 1842 unpublished paper circulated among his friends, he was unwilling to publish the results until he could present as complete a case as possible. He labored for almost 20 additional years on his theory of evolution and on its primary mechanism, natural selection. In 1858 he received a letter from British naturalist Alfred Russel Wallace, a professional collector of wildlife specimens. Much to Darwin’s surprise, Wallace had independently hit upon the idea of natural selection to explain how species are modified by adapting to different conditions. Not wanting Darwin to be unfairly deprived of his share of the credit for the theory, some of Darwin’s scientific colleagues presented extracts of Darwin’s work along with Wallace’s paper at a meeting of the Linnean Society, a London-based science organization, in June 1858. Wallace’s paper stimulated Darwin to finish his work and get it into print. Darwin published On the Origin of Species by Means of Natural Selection on November 24, 1859. All 1,250 copies of the first printing were sold on that day.

Darwin’s book and the theory it popularized—evolution through natural selection—set off a storm of controversy. Some of the protest came from the clergy and other religious thinkers. Other objections came from scientists. Many scientists continued to believe in Lamarckism, the idea that living things could consciously strive to accumulate modifications during a lifetime and could pass these traits on to their offspring. Other scientists objected to the seemingly random quality of natural selection. If natural selection depended upon random combinations of traits and variations, critics asked, how could it account for such refined and complex structures as the human eye? Perhaps the most serious question—one for which Wallace and Darwin had no answer—concerned the inheritance of traits. How exactly were traits passed along to offspring?

E.

Mendel and Early Genetics

Darwin did not know it, but the answer was at hand—although it would not be acknowledged in his lifetime. In the Augustinian monastery at Brünn (now Brno in the Czech Republic), Austrian monk Gregor Mendel experimented with the breeding of garden peas, observing how their traits were passed down through generations. In crossbreeding pea plants to produce different combinations of traits—color, height, smoothness, and other characteristics—Mendel noted that although a given trait might not appear in every generation, the trait did not disappear. Mendel discovered that the expression of traits hinged on whether the traits were dominant or recessive, and on how these dominant and recessive traits combined. He learned that contrary to what most scientists believed at the time, the mixing of traits in sexual reproduction did not result in a random blending. Traits were passed along in discrete units. These units are now known as genes. Mendel performed hundreds of experiments and produced precise statistical models and principles of heredity, now known as Mendel’s Laws, showing how dominant and recessive traits are expressed over generations. However, no one appreciated the significance of Mendel’s work until after his death. But his work ultimately gave birth to the modern field of genetics.

In 1900, Dutch botanist Hugo Marie de Vries and others independently discovered Mendel’s laws. The following year, de Vries’s book The Mutation Theory challenged Darwin’s concept of gradual changes over long periods by proposing that evolution occurred in abrupt, radical steps. Having observed new varieties of the evening primrose plant coming into existence in a single generation, de Vries had subsequently determined that sudden change, or mutation, in the genetic material was responsible. As the debate over evolution continued in the early 20th century, some scientists came to believe that mutation, and not natural selection, was the driving force in evolution. In the face of these mutationists, Darwin’s central theory threatened to fall out of favor.

F.

Population Genetics and the Modern Synthesis

As the science of genetics advanced during the 1920s and 1930s, several key scientists forged a link between Mendel’s laws of inheritance and the theory of natural selection proposed by Darwin and Wallace. British mathematician Sir Ronald Fisher, British geneticist J.B.S. Haldane, and American geneticist Sewall Wright pioneered the field of population genetics. By mathematically analyzing the genetic variation in entire populations, these scientists demonstrated that natural selection, and not just mutation, could result in evolutionary change.

Further investigation into population genetics and such fields as paleontology, taxonomy, biogeography, and the biochemistry of genes eventually led to what is called the modern synthesis. This modern view of evolution integrated discoveries and ideas from many different disciplines. In so doing, this view reconciled the many disparate ideas about evolution into the all-encompassing evolutionary science studied today. The modern synthesis was advanced in such books as Genetics and the Origin of Species, published in 1937 by Russian-born American geneticist Theodosius Dobzhansky; Evolution: The Modern Synthesis (1942) by British biologist Sir Julian Huxley; and Systematics and the Origin of Species (1942) by German-born American evolutionary biologist Ernst Mayr. In 1942, American paleontologist George Gaylord Simpson demonstrated from the fossil record that rates and modes of evolution are correlated: New kinds of organisms arise when their ancestors invade a new niche, and evolve rapidly to best exploit the conditions in the new environment. In the late 1940s American botanist G. Ledyard Stebbins showed that plants display evolutionary patterns similar to those of animals, and especially that plant evolution has demonstrated diverse adaptive responses to environmental pressures and opportunities.

In addition, biologists reviewed a broad range of genetic, ecological, and anatomical evidence to show that observation and experimental evidence strongly supported the modern synthesis. The theory has formed the basis of evolutionary science since the 1950s. It has also led to an effort to classify organisms according to their evolutionary history, as well as their physical similarities. Modern scientists use the principles of genetics and molecular biology to study relationships first proposed by Carolus Linnaeus more than 200 years ago.

G.

New Techniques in Molecular Biology

In 1953, American biochemist James Watson and British biophysicist Francis Crick described the three-dimensional shape of DNA, the molecule that contains hereditary information in nearly all living organisms. In the following decade, geneticists developed techniques to rapidly compare DNA and proteins from different organisms. In one such procedure, electrophoresis, geneticists evaluate different specimens of DNA or proteins by observing how they behave in the presence of a slight electric charge. Such techniques opened up entirely new ways to study evolution. For the first time geneticists could quantitatively determine, for example, the genetic change that occurs during the formation of new species.

Electrophoresis and other biochemical techniques also demonstrated to geneticists that populations varied extensively at the molecular level. They learned that much of population variation at the molecular or biochemical level has no apparent benefit. In 1968 Japanese geneticist Motoo Kimura proposed that much of the variation at the molecular level results not from the forces of natural selection, but from chance mutations that do not affect an organism’s fitness. Not all scientists agree with the neutral gene theory.

H.

Sociobiology

In recent decades, another branch of evolutionary theory has appeared, as researchers have explored the possibility that not only physical traits, but behavior itself, might be inherited. Behavioral geneticists have studied how genes influence behavior, and more recently, the role of biology in social behavior has been explored. This field of investigation, known as sociobiology, was inaugurated in 1975 with the publication of the book Sociobiology: The New Synthesis by American evolutionary biologist Edward O. Wilson. In this book, Wilson proposed that genes influence much of animal and human behavior, and that these characteristics are also subject to natural selection.

Sociobiologists examine animal behaviors that are called altruistic—that is, unselfish, or demonstrating concern for the welfare of others. When birds feed on the ground, for example, one individual may notice a predator and sound an alarm. In so doing, the bird also calls the predator’s attention to itself. What can account for the behavior of such a sentry, who would seem to derive no evolutionary benefit from its unselfish behavior and so seem to defy the laws of natural selection?

Darwin was aware of altruistic social behavior in animals, and of how this phenomenon challenged his theory of natural selection. Among the different types of bees in a colony, for example, worker bees are responsible for collecting food, defending the colony, and caring for the nest and the young, but they are sterile and create no offspring. Only the queen bees reproduce. If natural selection rewards those who have the highest reproductive success, how could sterile worker bees come about by natural selection when worker bees devote themselves to others and do not reproduce?

Scientists now recognize that among social insects, such as bees, wasps, and ants, the sterile workers are actually more closely related genetically to one another and to their fertile sisters, the queens, than brothers and sisters are among other organisms. By helping to protect or nurture their sisters, the sterile worker bees preserve their own genes—more so than if they actually reproduced themselves. Thus, the altruistic behavior evolved by natural selection.

I.

Punctuated Equilibria

Evolutionary theory has undergone many further refinements in recent years. One such theory challenges the central idea that evolution proceeds by gradual change. In 1972 American paleontologists Stephen Jay Gould and Niles Eldredge proposed the theory of punctuated equilibria. According to this theory, trends in the fossil record cannot be attributed to gradual transformation within a lineage, but rather result from quick bursts of rapid evolutionary change. In Darwinian theory, new species arise by gradual, but not necessarily uniform, accumulation of many small genetic changes over long periods of geologic time. In the fossil record, however, new species generally appear suddenly after long periods of stasis—that is, no change. Gould and Eldredge recognized that speciation more likely occurs in small, isolated, peripheral populations than in the main population of the species, and that the unchanging nature of large populations contributes to the stasis of most fossil species over millions of years. Occasionally, when conditions are right, the equilibrium state becomes “punctuated” by one or more speciation events. While these events probably require thousands or tens of thousands of years to establish effective reproductive isolation and distinctive characteristics, this is but an instant in geologic time compared with an average life span of more than ten million years for most fossil species. Proponents of this theory envision a trend in evolutionary development to be more like climbing a flight of stairs (punctuations followed by stasis) than rolling up an inclined plane (Darwinian gradualism).

J.

Role of Extinction

In the last several decades, scientists have questioned the role of extinction in evolution. Of the millions of species that have existed on this planet, more than 99 percent are extinct. Historically, biologists regarded extinction as a natural outcome of competition between newly evolved, adaptively superior species and their older, more primitive ancestors. Recently, however, paleontologists have discovered that many different, unrelated species living in large ecosystems tend to become extinct at nearly the same time. The cause is always some sort of climate change or catastrophic event that produces conditions too severe for most organisms to endure. Moreover, new species evolve after the wave of extinction removes many of the species that previously occupied a region for millions of years. Thus extinction does not result from evolution, but actually causes it.

Scientists have identified several instances of mass extinction, when species apparently died out on a huge scale. The greatest of these episodes occurred during the end of the Permian Period, some 245 million years ago. At that time, according to estimates, more than 95 percent of species—nearly all life on the planet—died out. Another extensively studied extinction took place at the boundary of the Cretaceous Period and the Tertiary Period, roughly 65 million years ago, when the dinosaurs disappeared. In all, more than 20 global mass extinctions have been identified. Some scientists theorize that such events may even be cyclical, occurring at regular intervals.

In the view of many scientists, mass extinctions can be explained by changes in climate—episodes of global warming or cooling that destroy sensitive ecosystems, such as tropical or marine habitats. Other theories have centered on abrupt changes in the levels of the world’s oceans, for example, or on the effect of changing salinity on early sea life. Another theory blames catastrophic events for mass extinction. Strong evidence, for example, supports the theory that a meteorite some 10 km (6 mi) in diameter struck the Earth 65 million years ago. The dust cloud from the collision, according to this impact theory, shrouded Earth for months, blocking the sunlight that plants need to survive. Without plants to eat, the dinosaurs and many other species of land animals were wiped out.

Extinction as a cause of evolution rather than the result of it is perhaps best demonstrated in terms of our own ancestors—ancient mammals. During the time of the dinosaurs, mammals constituted only a small percentage of the animals that roamed the planet. The demise of dinosaurs provided an opportunity for mammals to expand their numbers and ultimately to become the dominant land animal. Without the catastrophe that took place 65 million years ago, mammals may have remained in the shadow of the dinosaurs.