EVOLUTION NOTES – PRINCIPLES OF BIOLOGY Modules 63,64,65,66, I. Definitions biological evolution – the change over time in the allele frequencies or genetic makeup of populations of organisms – Discuss – how long, genetics? , populations evolve !!!! Scientific theories – scientific explanations to complex phenomena that are supported by evidence. The theory usually remains the same but the evidence changes, so the theory becomes better understood. II. Natural Selection – a mechanism of evolution in which organisms possessing certain genotypic characteristics that make them better adjusted to an environment tend to survive, reproduce, increase in number or frequency, and therefore, are able to transmit and perpetuate their essential genotypic qualities to subsequent generations. Natural selection has the following aspects: i. Variation – individuals exhibit variation in a population, they have a unique set of traits. Some of these traits improve their chances of survival while others are less favorable. ii. Overproduction – populations produce too many young, many must die. iii. Struggle for existence – food, water and other resources are limited, organisms compete with one another for these resources. iv. Differential reproductive success – those individuals that have the most favorable characteristics in an environment, has higher chance of survival and reproduction. v. Descent with modification – different reproduction and survival can result in changes in the characteristics of organisms and the rise of new species. Charles Darwin and Alfred Wallace came up with the theory of evolution by natural selection. They based their theory on observations and many other evidence but did not know about molecular evidence. III. Evidence of Evolution -- data or information that is collected by scientists Direct observation (See examples of evolution) Molecular evidence Fossil evidence – radiometric dating, http://www.sciencefriday.com/segment/08/22/2014/tar-noir.html , only 1 % of all organisms ever lived left fossils in sedimentary rocks, tar pits, pitbugs Comparative anatomy – homology and analogy Vestigial organs Comparative embryology Biogeography IV. Examples of Evolution – specific evolutionary events that are/were observed Darwin’s finches Antibiotic resistance Herbicide and pesticide resistance V. Human birth weight evolution Industrial melanism Adaptations and natural selection Adaptation is any change in heritable traits that increase an organism’s evolutionary fitness Evolutionary fitness – shows the chance of an organism to survive and reproduce Must be able to give examples of various adaptations – See Module 68,69 Making of the fittest: http://media.hhmi.org/fittest/birth_death_genes.html -- Start at min. 2 1. What is the scientific discovery? 2. Why is it surprising? 3. What is the adaptation? 4. What is the evolutionary advantage? 5. Guide this story through the concept and aspects of natural selection. 6. What is the scientific explanation of the existence of antifreeze protein? Modules 68,69 VI. Mechanisms of Evolution – Natural processes that can change the allele frequency in a population and as a result cause evolution Natural Selection – According to the outcome of natural selection we can have the following types: i. Directional selection – Ex. Skin color gets darker in the tropics to protect against the damaging effects ii. Stabilizing selection – Ex. Human birth weight iii. Disruptive selection – Ex. Finch beak sizes change to very thin or thick depending on what type of seed they eat. Genetic Drift – fluctuations in the populations’ allele frequencies due to chance events (weather conditions, geological phenomena, etc.). Especially influential in smaller populations. i. Bottleneck effect ii. Founder effect Gene flow – mechanism of evolution in which alleles move between populations as a result of migration. Gene flow into the population (immigration) increases genetic variation, while gene flow out of the population (emigration) decreases genetic variation (Figure 8 in Module 67 is a good representation of all three mechanisms) Other mechanisms can include: Sexual Selection (Artificial Selection) Mutations VII. The Significance of Genetic Variation Genetic Variation – genetic differences among individuals within a population Genetic differences are frequently visible as phenotypic differences and may be passed on to the next generation Dutch study on epigenetics VIII. IX. Genetic variation drives evolution, because evolution can only work on existing genes. More variations of those genes give evolutionary processes more “options” Concerns about monocultures in our food supply Genetic variation is affected by geographic variation, because different environmental pressures are acting on the organisms of each population Ex. Human skin color, industrial melanism, mice fur coloration -http://www.youtube.com/watch?v=AMtT5_AQmLg Frequency-Dependent Selection The selection pressure on a phenotype is dependent on the frequency of that phenotype. Coevolution – result from beneficial or adverse interactions between species Ex. Pollinators and flower structure Lactose tolerance Sickle cell disease and malaria Lactose tolerance: http://media.hhmi.org/fittest/Got_Lactase.html EVOLUTION IN POPULATIONS – THE HARDY-WEINBERG THEOREM I. POPULATIONS THAT ARE NOT EVOLVING Because the allele frequency has to change for evolution, if the allele frequency remains the same, populations are not evolving. The allele frequency can be changed by any mechanism of evolution. In a non-evolving population none of the mechanisms of evolution are active. Non-evolving populations have the following five characteristics: i. Large populations – no genetic drift ii. Random mating – no sexual selection iii. No mutations iv. No natural selection v. No gene flow There are really no natural populations that would have all these conditions fulfilled. So we can conclude that evolution always takes place in natural populations. However, the pace at which it takes place can change. II. CALCULATING ALLELE FREQUENCIES A) Explanation of symbols used: p = the frequency of the dominant allele in a population q = the frequency of the recessive allele in a population p2 = the frequency of the homozygous dominant genotype or individuals in the population 2pq = the frequency of the heterozygous individuals or genotype in the population q2 = the frequency of the homozygous recessive individuals or genotype or recessive phenotype in the population p2 + 2pq = the frequency of the dominant phenotype in the population B) Hardy-Weinberg Equations: p+q = 1 P2 + 2pq + q2 = 1 These equations are used to determine the allele frequencies in various populations. Practice Problems: 1. In Drosophila the allele for normal-length wings is dominant over the allele for vestigial wings (vestigial wings are stubby little curls that cannot be used for flight). In a population of 1,000 individuals, 360 show the recessive phenotype. How many individuals would you expect to be homozygous dominant and heterozygous for this trait? 2. In the United States about 16% of the population is Rh negative blood typed. The allele for Rh negative is recessive to the allele for Rh positive. If the student population of Aurora High School is 1,078, how many students would you expect for each of the three possible genotypes? 3. In a certain population, the dominant phenotype of a certain trait occurs 91% of the time. What is the frequency of the dominant allele? The recessive allele? All three genotypes? THE BIOLOGICAL SPECIES CONCEPT AND SPECIATION I. WHAT IS A SPECIES? An interbreeding group of organisms that are similar to each other but different from other groups of organisms. Organisms of the same species are able to produce viable and fertile offspring in nature. However, there are many other species definitions exist. II. REPRODUCTIVE ISOLATION Reproductive isolation occurs when two organisms from different populations cannot produce viable and fertile offspring. If the organisms of two populations cannot reproduce with each other, they will eventually form two separate species. There are two major kinds of barriers that prevent reproduction: i. Prezygotic barriers – barriers that prevent the formation of a zygote 1. Gametic isolation 2. Mechanical isolation 3. Behavioral isolation 4. Temporal isolation 5. Habitat isolation ii. Postzygotic barriers – gametes can unite and form a viable zygote but any time after the formation of the zygote, failure can happen in the hybrid 1. Reduced hybrid viability 2. Reduced hybrid fertility 3. Hybrid breakdown WHEN DO POPULATIONS BECOME NEW SPECIES? New species form when they become genetically isolated from the ancestor species. Usually genetic isolation means that one or many beneficial mutations accumulate in one population. It may not be very clear when exactly a new species forms – See the case study SPECIATION AND GEOGRAPHIC BARRIERS Geography influences species formation in two ways: i. Allopatric speciation – occurs when two populations of the same species are separated from each other by some geographic barrier. As a result, these populations cannot reproduce with each other and over time become genetically so different from each other that they will not be able to reproduce with each other any more. ii. Sympatric speciation – occurs, when two populations are living in the same geographic area and have the potential to reproduce with each other but they do not reproduce. The reasons for the lack of reproduction are: 1. Polyploidy – Figure 4, Module 74 2. Sexual selection – Figure 9. Module 74 3. Habitat differentiation – Case study III. IV. V. THE PACE OF SPECIATION Species that exist today evolved from once lived ancestors that were very different from today’s species. This constant change of species is taking place today as well as millions of years ago. There are two different ideas about at what pace evolution is taking place at any given time: i. Gradualism – slow, incremental changes take place all the time ii. Punctuated equilibrium – species are mostly stable, but rare events apply selective pressures that will result in fairly rapid chanes. Evidence indicates that a combination of these two processes take place. What kinds of evidence would you use to prove this? The Origin and History of Life on Earth I. The Origin of Life and Evidence that Supports It There is no coherent, widely accepted scientific theory about the origin of life today. But we have several hypotheses that are supported by evidence. A. The Origin of Simple Organic Molecules: Review what an organic molecule is. The Earth, with the rest of the solar system formed 4.5 billion years ago. At this point, the Earth was hot, but slowly cooled down by about 4.2-3.9 billion years ago. The atmosphere at this point was mostly made up of CO2, N2, methane, ammonia, hydrogen sulfide and as the planet cooled, water condensed. o Evidence: air samples from rocks, radiometric dating of rocks, samples from other planets Haldane and Oparin Hypothesis: The composition of the early atmosphere, UV rays, energy from volcanic eruptions could create the simplest monomers of organic molecules, such as amino acids, hydrocarbons, simple sugars, simple fats and nucleotides. The Miller-Urey Experiment: Tried to replicate the conditions on the ancient earth under laboratory conditions and could form simple organic molecules such as amino acids. Today, we have hydrothermal vents on the ocean floor that still shows similar conditions to the ancient atmosphere. http://www.youtube.com/watch?v=BXGF3XS-yAI – hydrothermal vents Alternative theory: o Organic molecules arrived to the earth from outer space in meteorites. Scientists find meteorites today with organic molecules in them. Some asteroids also had ice in them which may be the source of water today on earth. B. The Origin of Macromolecules According to one theory, macromolecules could have formed when simple monomers attached to some substances with large surface areas. On these surfaces, the monomers could have reacted with each other to form long chains and rings. These surfaces are found on porous rocks and clay today. RNA world hypothesis – It is hypothesized that the very first macromolecules were used for multiple purposes. RNA was likely the first molecule that was responsible for being both the molecule of inheritance that could self-replicate and the catalyst of chemical reactions. Evidence of this hypothesis: i. Polynucleotide chains are able to self-replicate today under proper conditions ii. Ribozymes (enzymes that are made up of RNA not proteins) still exist today for example in the form of ribosomes iii. Some ribozymes can replicate with great accuracy Protocell formation: Membrane structures could develop from phospholipids that could also form on the surface of porous rocks. Once phospholipids form, they self-assemble into double layers in watery solutions. If the membranes become too large, they can split into two (divide). Once protocells could self-replicate, they could also evolve because their RNA or DNA could change over time. Evidence: i. Vague but it is possible to build protocells in ponds, lakes with varying temperatures if the monomers are found in the water. Figure 6 animation C. The Early Evolution of Life http://www.pbs.org/wgbh/evolution/library/03/3/l_033_01.html -- how was the age of the Earth determined. Radiometric dating – used to determine the age of fossils or rocks. Radioactive isotopes decompose at a steady rate. As their concentration decreases from the level that it was on during the formation of the rock or fossil, we can calculate the age of the rock or fossil. – Use Figure 6 in Module 65 By using radioactive isotopes, it was determined that the Earth is about 4.6 billion years old. Life first formed about 3.5 billion years ago. Evidence: The first fossil evidence comes from rocks called stromatolites that are about 3.5 billion years old. The first fossils were printed remains of groups of prokaryotic cells. Oxygen levels dramatically rose at about 2.5 billion years ago. Evidence: iron oxides increased in concentration in rocks. Significance: photosynthetic prokaryotes such as cyanobacteria appeared. This destroyed most life forms that existed previously, because all the previous life forms were anaerobic (poisoned by oxygen). Large areas became empty, so newly evolved aerobic life forms could reproduce without competition. First eukaryotes appeared 2.1 billion years ago. Evidence: i. The earliest eukaryotic fossils were found from that time ii. Available oxygen killed lots of organisms, so there were plenty of resources available for punctuated equilibrium type of evolution iii. The Endosymbiotic Theory – Mitochondria and chloroplasts evolved from free living bacteria, when they were engulfed by simple but larger cells. Evidence: 1. These organelles have circular DNA 2. They reproduce by binary fission like bacteria do 3. Some of their metabolic enzymes are similar to bacterial enzymes 4. They are about the same size as bacteria 5. They have double membranes, one of which was the original bacterial membrane, the other was the host’s cell membrane Watch the animation on Figure 5 – Module 77 Multicellular life evolved about 1.2 billion years ago. Evidence: i. Fossil record ii. DNA sequencing – scientists estimate that, based on the amount of change among the DNA sequences of multicellular eukaryotes, they had a common ancestor about 1.5 billion years ago. Major diversification of life – rapid speciation occurred about 500 million years ago. This is also when life started on dry land. – Figure 7 – Module 77 introduces phylogenetic trees Land organisms must have adaptations to avoid or survive drying out. A few of these: i. Thick skin, waxy outer covering ii. Inside transport system – blood vessels, vascular tubes iii. Root systems in plants iv. Unique respiratory systems v. Stronger support by having harder structures, bones, hard fibers D. MAJOR FACTORS THAT CONTRIBUTE TO THE DIVERSIFICATION OF SPECIES Mass extinctions – Catastrophic changes in the environment can destroy very large, 75% or more of species on the entire planet. These events are called mass extinctions. There were 5 of these in the history of life. These are very disadvantageous but can open up new environments for surviving species and with that speed up speciation again. Figure 9, Module 77 Adaptive radiation – When major ecological niches open up because of previous mass extinctions, surviving species can have major evolutionary changes that fill up all available niches. This happened with dinosaurs, mammals etc. Continental drift – a continuous process throughout the history of the Earth. Continental drift changes the climate and with that can radically change the environment. As a result, it speeds up speciation E. THE COMMON ORIGIN OF LIFE: All living organisms came from the same common ancestors and these evolved over the billions of years to the great diversity that we see today. Evidence: o Common genetic code o Common material of inheritance o Similar cellular processes o Same structure of the phospholipid bilayer o Same major types of macromolecules o Similar structure of amino acids PHYLOGENY I. II. III. EVOLUTION AND PHYLOGENY Phylogenies represent the history of relationships among evolving lineages of populations. Because anything that can reproduce and pass on their genes, over time will also evolve. Lineage is a series of organisms connected by genetic information passed from one generation to the next. At one point in their history of a lineage some members may become isolated from the original population. This isolation can be genetic or reproductive and can over time result in a split in the lineage – speciation. Phylogenetic trees can represent the history of a lineage in a simplified way. Nodes on these phylogenetic trees show the split between two or more branches. The node represents a common ancestor. These common ancestors are always historical, so they no longer exist. Diagram in Module 79 INTERPRETING PHYLOGENIES The closer to the leaves the branching for two species are on a phylogenetic tree the more recent common ancestors they have. Guidelines to read phylogenetic trees: o To understand relatedness, find the common ancestor o A phylogeny can be flipped around at any node without changing the meaning of the tree. o Trees can be drawn in many different formats but the guidelines for reading them don’t change o All currently living species should end on the same level of a phylogenetic tree – this rule is frequently not fulfilled, even in some AP materials, you will see an example CLADISTICS Sometimes homologous traits are used to determine common ancestry – cladistics, which groups species and other taxa into groups called clades. Clade includes one common ancestor and all of its potential descendants. Cladistic analysis can be complicated because it can include many different groups. o Monophyletic groups – include only one common ancestor and all of its descendants o Paraphyletic groups—include one common ancestor but not all of its descendants o Polyphyletic groups – include many common ancestors with many descendants Cladograms are charts that represent cladistics relationships Parsimony -- the rule of parsimony is used to determine the most likely evolutionary relationships in a cladogram. According to this rule, a tree with the fewest evolutionary events is the most likely to show the correct evolutionary relationship. PPT on how to build a cladogram is on Schoolpointe Figure 3 – Module 81 – Using sequencing data to determine molecular relationships – used in the lab