DEVELOPMENT OF EVOLUTIONARY THEORY (Rewritten from M J Farabee, 2001) The Ancient Greek philosopher Anaxiamander (611-547 B.C.) and the Roman philosopher Lucretius (99-55 B.C.) coined the concept that all living things were related and that they had change over time. The classical science of their time was observational rather than experimental. Aristotle (384-322 B.C.) developed his Scala Naturae, or Ladder of Life, to explain his concept of the advancement of living things from inanimate matter to plants, then animals and finally man. Post-Aristotlean “scientists” were constrained by the prevailing thought patterns of the Middle Ages – the literal creation of the earth in six days as laid out in the book of Genesis. According to Archbishop James Ussher of Ireland, the Earth was formed on October 22, 4004 B.C., and his ideas were readily accepted because they posed no threat to the social order of the times; comfortable ideas that would not upset the linked applecart’s of church and state. Prior to the sixteenth century dissection of human bodies was either illegal or severely limited so that most of the comparative anatomy was pure guesswork or was taken from the dissection of animals or during surgery, and it wasn’t until dissection laws had been relaxed and Andreas Vesalius (1514-1584) began to document his dissections culminating in his masterwork “The Seven Books on the Structure of the Human Body”. Through his work and that of many other anatomists, they gradually began to recognize humans as being one species among many, with a few unique traits but many others shared in common with other animals. Some 300 years later, Darwin used this vast stock of anatomical knowledge to build his theory of evolution. In the 1600’s the study of life changed forever. After relying on the authority of ancient writes like Aristotle, European naturalists began to look at life for themselves. Anatomists discovered that familiar organs, such as the heart, did not work the way that had previously been thought, and the English inventor, Robert Hooke, was able to look through a microscope to view previously unimaginable complexity hidden in tiny animals such as the flea. Some clergymen worried that this approach to life smacked of atheism but many naturalists, themselves religious, believed that they were on a religious mission to help demonstrate and appreciate God’s benevolent design. Natural theology dominated English thinking for nearly two centuries. The mid 1600’s heralded the birth of paleontology. Before this time fossils were thought to have fallen from the sky but an Italian, Nicholas Steno, made the leap and declared that “tongue stones” that had been previously found resembled shark teeth and did indeed come from the mouths of once-living sharks. Steno proposed that fossils belonged to living creatures and argued that fossils were snapshots of life at different moments in Earth’s history and that rock layers formed slowly over them over time. Fossils ultimately became some on the key evidence for how life evolved on Earth over the past four billion years. Homo sapiens, Tyrannosaurus rex, Escherichia coli – our English conversation is littered with pairs of Latin names for animals, plants, and microbes. Aristotle had begun to organize life in his ladder-like hierarchy with plants on the bottom, animals in the middle, and humans on the top. Swedish botanist Carl Linne (1707-1778; more popularly known as Linneus, after the common practice of the day which was to Latinize names of learned men), attempted to pigeon-hole all known species of the time into immutable categories. Many of these categories are still used in biology today. Georges-Louis Leclerc, Comte de Buffon (1707-1788) was the first to propose that species’ could change. This was a major break from earlier concepts that species were created by a perfect creator and therefore could not change because they were perfect. Buffon also provided evidence of “descent with modification” and speculated on various causative mechanisms. In his work, Buffon mentioned several factors could influence evolutionary change including: influences of the environment, migration, geographical isolation, overcrowding, and the struggle for existence. However, Buffon vacillated as to whether he believed or not in evolutionary descent, and professed to believe in special creation and the fixity of species. Erasmus Darwin (1731-1802; grandfather of Charles Darwin) a physician and poet, proposed that life had changed over time. His writings suggested the possibility of common descent based on changes undergone by animals during development, artificial selection by humans, and the presence of vestigial organs. However, he offered no mechanism to explain evolutionary descent. Georges Cuvier (1769-1832) was among the foremost proponents of catastrophism, the theory that the earth and geological events had formed suddenly, as a result of some great catastrophe (eg. Noah’s flood). This view was comfortable for the times and was widely accepted. Cuvier eventually proposed that there had been several creations that occurred after catastrophes. Louis Agassiz (1807-1873 – remember his name for the Africa unit) proposed 50-80 catastrophes and creations. Jean Baptiste de Lamarck (1744-1829) developed one of the first theories on how species changed. Lamarck, concluded that organisms of higher complexity had evolved from preexisting, less complex organisms. He was struck by the similarities of many of the animals he studied and was impressed with the ever-growing fossil record. It led him to argue that life was not fixed. When environments changed, organisms had to change their behavior to survive. If they began to use an organ more than they had in the past, it would increase in its lifetime. If a giraffe stretched its neck for leaves, for example, a “nervous fluid” would flow into its neck and make it longer. Its offspring would inherit the longer neck, and continued stretching would make it longer still over several generations. Meanwhile organs that organisms stopped using would shrink. Lamarck was proposing that life took on its current form through natural processes, not through miraculous interventions (ie. God, supernatural forces etc). His ideas were rejected and he was shunned and he died in poverty and obscurity. Despite all Lamarck got wrong, however, he can be credited with envisioning evolutionary change for the first time. Karl von Baer (1792-1876) was not a fan of evolution but his work on embryology provided Darwin with the most compelling evidence for evolution. Baer’s work on embryology showed that vertebrates all share an anatomy that invertebrates never acquire - humans bear a resemblance to fish in their early embryonic stages - and although over time organisms evolve further away from each other, natural selection modifies embryos in various ways, but some vestiges of common ancestry survive. (See embryo comparisons at the end of the handout) Evolution by Natural Selection The idea that species could change over time was not immediately acceptable to many: the lack of a mechanism hampered the acceptance of the idea as did its implications regarding the biblical views of creation. Charles Darwin (1809-1882) and Alfred Wallace (18231913) both worked independently of each other, traveled extensively, and eventually developed similar ideas about the change in life over time as well as a mechanism for that change: Natural Selection. Charles Darwin Darwin set sail on the Beagle, serving as the ship’s naturalist and spent much time ashore collecting plant, animal and fossil specimens, as well as making extensive geological observations. On his return to England in 1836, Darwin began to catalog his collections and ponder the seeming “fit” of organisms to their mode of existence. He eventually settled on four main points of the theory: 1. Adaptation: All organisms adapt to their environments. 2. Variation: all organisms are variable in their traits. 3. Over-reproduction: all organisms tend to reproduce beyond their environment’s capacity to support them. This is based on the work of Thomas Malthus (1766-1834) who studied how populations of organisms tended to grow exponentially until they encountered a limit on their population size). 4. Since not all organisms are equally well adapted to their environment, some will survive and reproduce better than others – this is known as natural selection. Sometimes this is also referred to as “survival of the fittest” (this phrase originated with Herbert Spencer after he finished reading Darwin’s Origin of Species). This merely deals with the reproductive success of the organisms, not solely their relative strength or speed. Alfred Wallace Wallace came from the lower class of England and struggled financially all of his life. He left England in 1854 to study the natural history of Indonesia where he contracted Malaria. During a fever Wallace managed to write down his ideas on natural selection. In 1858, Charles Darwin received a letter from Wallace, in which Darwin’s as-yet-unpublished theory of evolution and adaptation was precisely detailed. Darwin had begun formulating his theory of natural selection in the late 1830’s but he worked on it quietly for 20 years and was shocked to see that Wallace’s theory nearly replicated his own. Wallace’s paper published in 1858 (with the help of Darwin and several other colleagues) was the first to define the role of natural selection in species formation. As a result Darwin rushed to finish his book “On the Origin of Species by Means of Natural Selection” which remains one of the most influential books ever written. The Wallace-Darwin Theory 1. Individuals in a population have variable levels of agility, size, ability to obtain food, and different successes in reproducing. 2. Left unchecked, populations tend to expand exponentially, leading to a scarcity of resources. 3. In the struggle for existence, some individuals are more successful than others, allowing them to survive and reproduce. 4. Those organisms best able to survive and reproduce will leave more offspring than those unsuccessful individuals. 5. Over time there will be heritable changes in phenotype (and genotype) of a species, resulting in a transformation of the original species into a new species similar to, but distinct from, its parent species. Neither Darwin nor Wallace could explain how evolution occurred: how were these inheritable traits (variations) passed on to the next generation? Gregor Mendel had yet to publish his ideas about genetics. Gregor Mendel (1822-1884) is known as the “father of modern genetics” and working with pea plants he came up with his laws of inheritance. Although his work was criticized at the time it is now considered the foundation for modern genetics. In the twentieth century, several scientists were able to identify the processes of inheritance at a molecular level which greatly advanced the idea of evolution by descent with modification. These include: Hugo DeVries (Dutch botanist) – rediscovered Mendel’s work and in 1900 and proposed that gene mutations would create a new species in a single step. Thomas Hunt Morgan – in the 1920’s showed that gene mutation did not result in a new species but could alter a trait that could be passed down to the next generation. Theodosius Dobzhansky – In 1937 he published Genetics and the Origin of Species where he sketched out an explanation for how species’ actually came into existence. Mutations naturally crop up all the time. Some mutations are harmful in certain circumstances, but a surprising number have no effect one way of the other. These neutral changes appear in different populations and linger, creating variability that is far greater than anyone had previously imagined. It is this variability that serves as the raw material for making new species. Dobzhansky collaborated with many other scientists from other science disciplines (eg. geology, population genetics, paleontology, biology, and genetics) and they came up with one powerful explanation of evolution, showing how mutations and natural selection could produce large-scale evolutionary change. Their combined work is known as the “Modern Synthesis” and it became the foundation for future research. Ernst Mayr – redefined the definition of species and proposed various methods as to the creation of a new species (1942) including geographic isolation. Watson and Crick – 1953 proposed the double helix model for DNA. Evolutionary biology was revolutionized by the discovery of DNA. Scientists realized that a change in the “spelling” of a strand of DNA would be copied and those mutations might confer a selective advantage to an individual. The advantage may become more common over time, and ultimately these mutant genes may drive the older versions out of existence. As a result of the discovery of DNA and its structure scientist are now able to identify not just genes but individual bases. Embryo Comparisons