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Honors Biology Lesson Notes 1 Unit 11 - Biological Evolution Lesson Objectives: You will: 1. describe the atmosphere and energy conditions thought to exist early in Earth's history. 2. explain the results of experiments designed to test whether simple organic compounds might have formed in the early atmosphere. 3. explain the steps that might have led to the evolution of the first cells according to the heterotroph hypothesis. I. Formation of the Universe and Solar System A. Big Bang Theory - all matter was originally concentrated in one mass that blew apart about 18 billion years ago. B. The smaller atoms, hydrogen and helium, which make up about 99% of all matter, fused to produce the heavier elements found in our solar system. C. Stars were borne out of huge masses of interstellar gases several light years across. D. Our sun was formed about 6 billion years ago. E. About 4.6 billion years ago the planets of our solar system formed by the condensing of peripheral gases and matter around our sun. II. Formation of Organic Compounds A. Fossils of photosynthetic bacteria have been found that date back 3.5 billion years, but how did simple life arise in the first place? B. Frankly, there are many theories and we really don't know. However, there is some direct and indirect evidence that gives us an idea of how life began. C. We know that the young earth was a violent place. UV radiation and lighting bombarded the surface. The early atmosphere consisted mainly of gases released by volcanic activity hydrogen, nitrogen gas, carbon monoxide and carbon dioxide. Hydrogen sulfide, methane and ammonia also appeared to be present in small quantities. Water vapor was also likely abundant in the atmosphere due to much higher temperatures and some have suggested that it rained for millions of years as the earth cooled. D. Oxygen which now makes up 21% of our atmosphere was not present. This means that the atmosphere was reducing (favored the buildup of more complex organic compounds). It is doubtful that the processes which led to the formation of the earliest organic molecules could take place in our current oxidizing atmosphere (favors the breakdown of more complex organic compounds). E. In the 1920's Oparin and Haldane postulated that the reducing atmosphere coupled with vast amounts of free energy (i.e. useable energy) from volcanic activity, lightning, radioactive minerals, and tremendous amounts of energy from the sun (there was no ozone layer back then) enhanced reactions which joined simple molecules together to form the first organic molecules. F. This hypothesis was tested in 1950's by Miller and Urey who duplicated these early earth conditions in the lab. They constructed an artificial system which contained an "atmosphere" and "ocean". They introduced hydrogen, methane, ammonia and water into the system and turned on an electric spark as a supply of energy. G. In less than one week amino acids and other small organic molecules formed. H. Other scientists repeated their work, utilizing updated primeval atmospheric models, eventually producing: other amino acids, ATP, glucose and other sugars, lipids and the bases which form RNA and DNA, and adenine the key component of ATP and NAD. I. Over the course of hundreds of millions of years these compounds would be washed by rains into the oceans where they were built up in large concentrations forming a "primordial soup". C. Pace III. Origin of Life A. Abiotic synthesis (not from life) of organic monomers (simple organic molecules) - many experiments demonstrated that it is possible to get many different types of organic monomers by starting with a primitive atmospheric model and adding energy. 1. This includes the work of Stanley Miller in 1953. 2. You may have read about abiotic synthesis of living organisms is also known as spontaneous generation which was once widely believed. Work done by Redi, van Leeuwanhoek, Spallanzani, Pasteur, and Tyndall from the late 17th century to the mid-ninetenth century essentially disproved this concept. However, they were working under modern atmospheric conditions. B. Synthesis of polymers - recall that polymers are chains of monomers (i.e. monosaccharides, amino acids, and nucleotides) synthesized by condensation or dehydration synthesis reactions (i.e. eliminating a water molecule. 1. Experiments have demonstrated that polymers can be made by dripping organic monomers onto hot sand, rocks or clay. Clays and other minerals attract amino acids and repeated wetting and drying out promotes condensation reactions. The mineral pyrite may have played a major role in abiotic polymerization because it provides a charged surface and electrons freed during its formation could support bonding between abiotically synthesized organic molecules. 2. Sidney Fox found that heating a dry mixture of amino acids to about 60oC produced proteinoids, protein-like molecules of roughly 100 amino acids. 3. Molecules such as chlorophyll and the cytochromes involved in cellular respiration may have arisen from formaldehyde, a component of cosmic clouds. C. The Formation of Protobionts 1. Protobionts - Aggregates of abiotically produced molecules able to maintain an internal environment different from their surroundings and exhibiting some life properties such as metabolism, excitability, and self replication (yet not able to precisely reproduce). Probable antecedents of first true cells. Sometimes called proto-cells or pre-cells. 2. Proteinoids placed in water form aggregates called proteinoid microspheres which exhibit some cell-like attributes. They have a protein bilayer that forms a boundary between the interior and external environment. This boundary layer is differentially permeable and allows primitive nucleotide-like molecules to enter. They can contain lipid-like areas, watery areas, and boundarylayer areas that provide distinct places for specific chemical reactions. When they grow to unstable size, they split to form daughter microspheres. 3. Liposomes can form spontaneously when phospholipids form a bilayered membrane similar to those of living cells. 4. Coacervates (colloidal drops of polypeptides, nucleic acids, and polysaccharides) self-assemble under the right conditions. D. Chemical Evolution (Selection) 1. Chemical selection is similar to natural selection, but acts on nonliving systems. 2. Early protobionts probably enjoyed an abundance of sustaining organic monomers and polymers in their environment. But, as raw materials became scarce, protobionts with more stable combinations were favored. 3. Stable protobionts would likely have catalysts that would allow for primitive metabolic activities. Some of these catalysts may have been inorganic, while others may have been organic. Any protobiont that developed a useful catalytic pathway would have a selective advantage over other competing protobionts. 4. It is thought that complex, multistep metabolic pathways evolved backwards from useful, rapidly diminishing end products to less directly useful raw materials. C. Pace E. Origin of self replication - it appears that one of the first polymers was RNA and like now it can selfreplicate in the presence of nucleotides. 1. Simple self-replicating systems of RNA, enzymes and coenzymes have been created in the laboratory. Applying heat to nucleotides causes them to assemble into short strands of RNA. Adding zinc as a catalyst (Zinc is part of the RNA Polymerase enzyme) allows the formation of longer strands of RNA and for the formation of RNA complimentary to existing strands of RNA, much like DNA. 2. However, RNA is a relatively unstable molecule and the strands are too fragile to become very long. Perhaps the more stable, double stranded DNA eventually replaced RNA as the carrier of the genetic information. 3. Whether proteins or RNA came first is an intriguing question. Some feel that they coevolved eventually forming a relatively complex feedback cycle involving many primitive “genes” and “enzymes”. F. Formation of plasma membranes - cells are separated from their environment by a plasma membrane. Recall that the plamsa membrane is a lipid bilayer and that such layers will spontaneously form when lipids are placed in water. 1. Proto-cells, may have developed plasma membranes similar to those of living cells early in the chemical evolutionary process. Needless to say, plasma membranes became extremely important structures as proto-cells evolved into the first versions of true-cells. G. What do most scientist's agree on? A summary follows. 1. There was production of simple organic compounds. 2. Larger, more complex molecules were synthesized from simpler ones. 3. Concentrations of many complex molecules became surrounded by membranes. 4. A means of obtaining energy for life functions developed. 5. A reliable means of reproduction evolved. IV. Evidence of Early Life A. The earliest evidence of life occurs as 3.8 billion year old organic carbon deposits in Greenland. B. Stromatolites, banded domes of sediment formed by bacteria, have been dated at about 3.5 billion years old. C. Also, 3.5 billion year old spherical and filamentous prokaryotic fossils have been found in western Australia and south Africa. D. At first energy requirements were met by living off the abundance of organic matter in the seas. 1. This is called the Heterotroph Hypothesis. The idea that heterotrophs came first because their was an abundance of raw materials available in the environment to support primitive living systems. E. However, over millions of years available nutrients would become scarce and competition for food would become intense. Those most likely to survive, primitive autotrophs, could make their own food. F. The most successful organisms were those that developed ability to make direct use of the suns energy, i.e. photosynthetic. An early form of photosynthesis (cyclic photophosphorylation) appeared between 3.5 - 3.2 billion years ago. G. By about 2.5 billion years ago the most common form of photosynthesis, which included noncyclic photophosphorylation, evolved. 1. This was a major milestone in the evolution of the earth and its organisms because the byproduct, oxygen, began to accumulate in the atmosphere. 2. Evolution of photosynthesis is well documented in the geological record. Iron deposits began to oxidize (rust). H. Recall the early atmosphere lacked oxygen. This oxygen was important for two reasons: 1. Some of the oxygen was converted to ozone (O3) which filters out harmful solar radiation. C. Pace I. J. K. L. M. 2. Oxygen build up allowed for the evolution of aerobic respiration. Recall how much more energy aerobic respiration produces compared to anaerobic. This allowed for larger and more complex organism to evolve. It took at least a billion years for atmospheric oxygen levels to reach the current 21% and atmospheric oxygen levels appear to fluctuate over time. For example, we know that oxygen levels in the atmosphere has been high enough (greater than 21%) in the past to support gigantic forms of arthropods, especially the insects. The first Eukaryotes appeared about 1.4 billion years ago. It seems certain that the eukaryotic cell arose via endosymbiosis. 1. Recall the endosymbiont theory - prokaryotic organisms began symbiotic relationships in which they lived together within a common cell. 2. Recall that chloroplasts and mitochondria have two membranes, their own DNA and ribosomes, and they can self-replicate. 3. The nucleus probably arose by an infolding of the plasma membrane to protect the genetic material. By about 900 million years ago there are abundant fossils of various types of marine algae. By 700 million years ago simple multicellular marine organisms had evolved all over the earth. The invasion of land by plants took place about 450 million years ago. This probably took place in intertidal zones. Once land plants evolved they provided the food necessary for higher animals to evolve. V. Plate Tectonics A. Plate tectonics (continental drift) has had a profound effect on the evolution of life. 1. According to plate tectonic theory, the plates of the earth’s thin crust are constantly moving apart at ridges and colliding at trenches. 2. Basically, the land masses on earth have been drifting around almost since they were first formed 4 billion years ago. B. At one time all the continents formed one large land mass called Pangea. C. Later there were two major land masses: 1. Gondwanaland - the current southern hemisphere continents plus India (which was part of eastern Africa). 2. Laurasia - the current northern hemisphere continents. VI. Biological Evidence for Continental Drift A. Fossil Glossopteris (seed ferns) date back about 250 million years ago. Their distribution was: South America, Antarctica, South Africa, Australia and India. This is a classic Gondwana distribution. B. There are three extant genera of lungfishes, one each in Australia, Africa and South America. This is also a classic Gondwana distribution. C. Lack of native placental mammals in Australia - Australia has no native placental mammals. 1. They are replaced by marsupials - opossums, kangaroos, koalas and other mammals that complete their development in a marsupium (pouch). 2. Marsupials arose several million years before placental mammals. Later on placental mammals arose in Laurasia and spread southward into Gondwanaland. However, Australia had already drifted away from Gondwanaland before placental mammals got there. C. Pace These notes are revised from lecture notes written in 1997 by Chuck Pace for his A.P. Biology students that were themselves based on lecture notes originally published on the World Wide Web at <http://arnica.csustan.edu/biol1010/Evolution/evolution.htm> by Dr. Steven J. Wolf of California State University Stanislaus. C. Pace