How Evolution Works? An introduction The “dead-leaf moth” (Oxytenis modestia) has evolved camouflage that resembles a dead leaf. Observations on the diversity of life • When you look around in nature you see a great diversity of life in different forms. • We can call these forms species • A species is a group of living organisms consisting of similar individuals capable of interbreeding. That is to exchange genes. • It is estimated there are 8.7 million eukaryotic species on earth. • Why are there so many diverse species and why do many of them look the same? Observations about the diversity of life • An example of this diversity is the Dead leaf moth (Oxytenis modestia). This species is one of over 120,000 members of the Lepidoptera (butterflies and moths). • Found in the Peruvian rain forest it has evolved a shape and colouring that makes it look like a dead leaf. • The caterpillar of this species has a head shaped like that of a snake that it waves from side to side to warn off predators. • These strategies are examples of the natural selection of traits that increase survival of the fittest in a particular environment. • Can you spot this camouflaged moth Observations on the diversity of life • The Lepidoptera (butterflies and moths) with over 120,000 members are classified on the basis of common characteristics. • A juvenile stage characterised by a welldeveloped head and many chewing mouthparts. • The adults share other features; • three pairs of legs. • two pairs of wings covered with small scales. • But the lepidopterans also differ from one another. • How did there come to be so many different moths and butterflies, and what causes their similarities and differences? Observations on the diversity of life The similarities between this moth and its 120,000 relatives illustrates three key observations about life. 1. The striking ways in which organisms are suited for life in their environments (the term environment refers not only to the physical surroundings but to the presence of other organisms as well). 2. The many shared characteristics (unity) of life. 3. The rich diversity of life. This is the result of evolution. Definitions of Evolution •Darwin defined evolution as “descent with modification”. •Meaning that present day species are descendants of ancestral species that were different. •It can also be defined as a change in the genetic composition of a population from generation to generation. Definitions of Evolution • Evolution can be seen in two related but different ways: (1) as a pattern and (2) as a process. 1. The pattern of evolutionary change is revealed by data from many scientific disciplines, including biology, geology, physics, and chemistry. These data are facts—they are observations about the natural world—and these observations show that life has evolved over time. 2. The process of evolution consists of the mechanisms that cause the observed pattern of change. These mechanisms represent natural causes of the natural phenomena we observe. Definitions of Evolution •Indeed, the power of evolution as a unifying theory is its ability to explain and connect a vast array of observations about the living world. •As with all theories, Scientists constantly challenge evolutionary theory to advance our understanding of the processes involved. Evolution of evolution theory • Before Darwin many people had considered how life had formed. • Empedocles (c. 494–434 B. C. E.), proposed that the origin of life had taken place in a manner that suggested evolution. • Aristotle (384–322 B. C. E.) viewed the universe and species as unchanging. This view dominated for over 2000 years as it agreed with religious teachings that species were fixed (unchanging) since the act of creation. • Al-Jahiz (776-868) (Basra) described three mechanisms of evolution: Struggle for Existence, Transformation of Species into each Other, and Environmental Factors. • Erasmus Darwin (Darwin’s grandfather) published “Zoonomia” in 1794 proposing that species changed to adapt and survive. • Others had similar ideas. But none had facts to support them. Evolution of evolution theory • Carolus Linnaeus (1707–1778) a Swedish physician and botanist developed a system to classify life’s diversity. • He developed the two-part, binomial, format for naming species (such as Homo sapiens - the wise Man for humans) that is still used today. • His system classified plants based on the number of male and female reproductive parts of flowers (sexual system). • He used a nested classification system, grouping similar species into increasingly general categories. • For example, similar species are grouped in the same genus, similar genera (plural of genus) are grouped in the same family, and so on (more in a later lecture). • Linnaeus considered the similarity among species was due to the pattern of their creation. • We now know his system had faults. But at the time this was a great advance in scientific understanding. • See videos below. • His system revolutionized the classification of species. • See how his system is still used today. Linnaeus Evolution of evolution theory The concept of geological time • In the 1700’s the study of fossils (Paleontology) led to new theories on the age of the earth. 1 Rivers carry sediment into aquatic habitats such as seas and swamps. Over time, sedimentary rock layers (strata) form under water. Some strata contain fossils. • Previously scholars (e.g. Sir Isaac Newton) had estimated the earth to have formed in approximately 4000 B. C. E. • Many fossils are found in sedimentary rocks formed from the sand and mud that settle to the bottom of seas, lakes, and swamps. As new layers cover older ones they compress them into layers of rock called strata (singular, stratum). • The fossils in a particular stratum provide a view of some of the organisms that populated Earth at the time that layer formed. • Later, erosion carves through upper (younger) strata, revealing deeper (older) strata that had been buried. 2 As water levels change and geological activity pushes layers of rock upward, the strata and their fossils are exposed. Younger stratum with more recent fossils Older stratum with older fossils Evolution of evolution theory The concept of geological time • Georges Cuvier (1769–1832): This French scientist developed the science of palaeontology. He noted that fossils in older strata of rock were different from current life forms. And that in each strata new species appeared while others disappeared. • He proposed that catastrophic events lead to extinction of local species followed by colonisation by new species migrating from outside. • He opposed the idea of evolution. Was I an immigrant? Image: Palais_de_la_Decouverte_Tyrannosaurus_rex Evolution of evolution theory • • • • • The concept of geological time In contrast to Cuvier’s emphasis on sudden events, others suggested that profound change took place through the cumulative effect of slow but continuous processes. James Hutton (1726–1797): This Scottish geologist proposed that Earth’s geologic features could be explained by gradual mechanisms, such as valleys being formed by rivers. Charles Lyell (1797–1875), the leading geologist of Darwin’s time incorporated Hutton’s thinking into his proposal that the same geological processes are operating today as in the past, and at the same rate. And that these changes take place over enormously long periods of time. These ideas would come to strongly influence Darwin’s thinking regarding the time required for evolution to occur. Evolution of evolution theory Lamarckism - The first modern theory of evolution • Jean-Baptiste de Lamarck (1744-1829) proposed the first testable mechanism for how life evolves over time in response to a changing environment. • Published in 1809, the year of Darwin’s birth, this was revolutionary as it recognised that evolutionary changes explained patterns of fossils and the match of organisms to their environments. • By comparing living species with fossils he identified several lines of descent from older to younger fossils leading to a living species. Jean-Baptiste de Lamarck Evolution of evolution theory Lamarckism - The first modern theory of evolution He based his theory on two principles. 1. Use and disuse, the idea that parts of the body that are used extensively become larger and stronger, while those that are not used deteriorate. 2. Inheritance of acquired characteristics, stated that an organism could pass these modifications to its offspring. Evolution of evolution theory Lamarckism - The first modern theory of evolution • An example used was that giraffes necks grew longer by stretching for leaves in high branches. This trait was then passed on with each generation. Evolution of evolution theory Lamarckism - The first modern theory of evolution • Lamarck thought that evolution happens because organisms have an innate drive to become more complex. • Darwin rejected this idea but did think that variation was introduced in to the evolutionary process by the inheritance of acquired characteristics. • Lamarck’s theory eventually failed as it breaks the rules of genetic inheritance proposed by Gregor Mendel. • It is important to remember that: • He recognized the fact that organisms are well suited for life in their environments and this can be explained by gradual evolutionary change. • It was only in the early 1900’s, after the deaths of Darwin and Mendel, that the significance of Mendel’s identification of genetic inheritance was realised. Evolution of evolution theory • • • • • • • • • Darwin’s theory of evolution; a theory that changed the course of history The world has seen many scientific breakthroughs. But none as revolutionary in the way that Darwin’s theory of evolution has been. This theory was REVOLUTIONARY, overturning over 2000 years of scientific and cultural thinking. The impact of this theory was felt throughout society. Scientific Darwinism has lead the advancement of science leading in to the age of molecular genetics. Social Darwinism used to justify imperialism, racism, eugenics and social inequality at various times over the past century and a half. Political Darwinism was used drive political movements such as Communism, and Fascism resulting in World War 2. Despite the overwhelming evidence for evolution its impact is still controversial today. This is how it began. Charles Darwin (1809 – 1884) • Darwin studied botany at Cambridge University and had a life long interest in nature. • On graduating his supervisor recommended him to Captain Robert Fitzroy the commander of the Royal Navy survey ship HMS Beagle. • The Beagle was to survey the coast line of south America then sail round the world. • Darwin was to be a companion for Capt. Fitzroy and the ships unpaid biologist. Voyage of HMS Beagle (1831 – 1836) Voyage of HMS Beagle • When ever the ship stopped to survey the coast Darwin was taken ashore where he observed and collected 1000’s of plant and animal specimens. • He described the features that allowed them to survive in such diverse environments as the humid forests of Brazil, grasslands of Argentina, and the Andes mountains. • Also noting that plant and animals in temperate regions of South America more closely resembled those in tropical regions of S. America than those in temperate regions of Europe. Voyage of HMS Beagle • He read Lyell’s Principles of Geology during the voyage. • Experience geological change first hand during an earthquake in Chile. Observing that the land had been thrust up several metres. • And found fossils of ocean species high in the Andes mountains. • This led him to conclude that Lyell was correct in that the physical evidence indicating the earth was very old. • This contradicted the view held then that the earth had been created around 4000 B. C. E. • This would be a critical point when he developed his theory of evolution by natural selection. Voyage of HMS Beagle The Galapagos Islands • His interest in species diversity was further stimulated in the Galapagos islands, near the equator, 900 km west of S. America. • He observed several types of mockingbirds. That looked similar but seemed to be different species. Some unique to a single island some to more than one. • Similar observations were made for Finches and Tortoises. Darwin’s Finches Galapagos Islands Tortoises Variation among tortoises: Hood island tortoise has a long neck and a curved shell open round the neck and legs, allowing it to reach the sparse vegetation found on the island. The tortoise from Isabela Island has a dome-shaped shell and a short neck. On this island vegetation is more abundant and close to the ground. The Tortoise from Pinta Island has a shell intermediate between the two forms. Voyage of HMS Beagle Darwin’s finches • There are more than a dozen species of closely related finches on the islands. Some found on only one island. • We now know their beaks are adapted to specific diets Voyage of HMS Beagle The Galapagos islands • Darwin saw that animals on the Galapagos islands were similar to those on the S. American mainland but not found any where else in the world. • His observation of the diversity of amongst species such as finches, mocking birds and tortoises led him to a revolutionary idea. THE BIG IDEA • That the islands had been colonised by South American species carried across the Pacific ocean. These species had then adapted so diversifying in to different species on the various islands. What is adaptation • During his voyage, Darwin saw many examples of adaptations. • These are inherited characteristics that improve an organisms chances of survival and reproduction in a specific environment. • Later, he realised that adaptation to the environment and the origin of new species were closely related processes. • He thought, and we now know, that new species could arise from an ancestral form by the gradual accumulation of adaptations to a different environment. • He realized that explaining such adaptations was essential to understanding evolution. His explanation of how adaptations arise centred on natural selection. This is a process in which individuals that have certain inherited traits tend to survive and reproduce at higher rates than do other individuals because of those traits. To publish or not? • By the early 1840s, Darwin had worked out the major features of his hypothesis. • In 1844, he wrote a long essay on descent with modification and its underlying mechanism, natural selection. • He was reluctant to publish his ideas, because of the controversy it would generate. • He continued to collect evidence. • By the 1850’s he had described his ideas to Lyell, and few others. • So controversial were his ideas that he only told those he trusted. • Lyell, although not yet convinced by evolution encourage him to publish before someone else had the same idea. Lyell’s prophecy comes true • In 1858, Darwin received a manuscript from Alfred Wallace a British naturalist working in the islands of Malaysia. • Wallace had developed a hypothesis of natural selection almost identical to Darwin’s. • Darwin wrote to Lyell “Your words have come true with a vengeance. . . . I never saw a more striking coincidence . . . so all my originality, whatever it may amount to, will be smashed.” • In July that year Darwin and Wallace’s papers were presented at the Linnean Society of London. Alfred Russel Wallace (1823 – 1913) On the Origin of Species by Means of Natural Selection • Darwin quickly finished his book and published it in 1859. • Publication caused protests from all levels of society and the establishment. • However, within 10 years most scientists of the time were convinced that life’s diversity is the result of evolution. • Darwin succeeded by presenting a plausible scientific mechanism argued with simple logic and supported with overwhelming evidence. Ideas from “The Origin of Species” • Darwin attributed the unity of life to the descent of all organisms from an ancestor that lived in the remote past. • He also thought that as the descendants of that ancestral organism lived in various habitats, they gradually accumulated diverse modifications, or adaptations, that fitted them to specific ways of life. • He reasoned that over a long period of time, descent with modification eventually led to the rich diversity of life we see today. Ideas from “The Origin of Species” • Darwin viewed the history of life as a tree. • In this tree each fork in a branch indicated a common ancestor of all the lines arising from that branch. • Those labelled A-D represented living species those unlabelled were extinct. • He thought this branching, along with extinction could explain morphological gaps that sometimes exist between related groups of organisms. • He formulated this idea in 1837 soon after returning from his voyage on the Beagle. • In the corner he writes “I think”. Artificial and natural selection, and adaptation • Darwin used artificial selection that is used to domesticate plants and animals to support his idea of natural selection. • Arguing that domesticated crops and animals bear little resemblance to their wild ancestors. • He then argued a similar process occurred in natural selection. He based his argument on two observations. Artificial and natural selection, and adaptation These two observations were as follows: Observation #1: Members of a population often vary in their inherited traits. Observation #2: All species can produce more offspring than their environment can support, and many of these offspring fail to survive and reproduce. Inference #1: Individuals whose inherited traits give them a higher probability of surviving and reproducing in a given environment will leave more offspring than do other individuals. Inference #2: This unequal ability of individuals to survive and reproduce will lead to the accumulation of favourable traits in the population over generations. Artificial and natural selection, and adaptation • • • • • Evolutionary pressure Darwin had read a famous essay by the economist Thomas Malthus, who stated that much of human suffering—disease, famine, and war—resulted from the human population’s potential to increase faster than food supplies and other resources. This led to competition for resources (energy, food, land, water). Similarly, Darwin realized that the capacity to over reproduce was a characteristic of all species. This introduces competition for resources – evolutionary pressure. In an environment with such pressure adaptation can happen. Artificial and natural selection, and adaptation Key Features of Natural Selection • Natural selection is a process in which individuals that have certain heritable traits survive and reproduce at a higher rate than do other individuals because of those traits. • Over time, natural selection can increase the frequency of adaptations that are favourable in a given environment. • If an environment changes, or if individuals move to a new environment, natural selection may result in adaptation to these new conditions, sometimes giving rise to new species. • Heritable traits that give an advantage in one environment may not do in another. Artificial and natural selection, and adaptation Remember the following • Only populations evolve not individuals. • Natural selection only amplifies or reduces heritable traits that differ within the population. If a population is genetically identical for a heritable trait, evolution by natural selection cannot happen. • Environmental factors vary from place to place and over time. A trait that is favourable in one place or time may be useless—or even detrimental—in other places or times. • Natural selection is always operating, but which traits are favoured depends on the environment in which a species lives and mates. The power of adaptation • Through natural selection adaptation can be both subtle or extreme. • This depends on the selection pressure and how favourable is the outcome. • Examples below are for camouflage. • Different environments shaped the adaptation of form and colour in these species of Preying Mantis. South African flower-eyed mantis Common form of Mantis Malaysian orchid mantis The power of adaptation • • • • • • Natural selection for antibiotic resistance Penicillin, the first antibiotic was a major break through in the treatment of bacterial diseases. Large numbers of antibiotics are now in use. 1943, Penicillin was first widely used. By 1945, 20% of Staphylococcus aureus bacterial strains in hospitals were resistant. They had an enzyme penicillinase that destroyed penicillin. Other S. aureus strains quickly developed resistance to other antibiotics. Finally in 1959, Methicillin was used. Methicillin deactivates an enzyme the bacteria uses to synthesize its cell walls. However some used a different enzyme so resistance soon developed. Leading to the designation MRSA (Methicillin resistant S. aureus) Now some S. aureus strains are resistant to multiple antibiotics. Many other bacteria have also developed multi-drug resistance e. g. gonorrhoea and tuberculosis. Natural selection in action • The following video shows E.coli bacteria developing antibiotic resistance in real time. • This is an example of Darwin's evolution by natural selection and his “tree of life”. The Evolution of Bacteria on a “Mega-Plate” Petri Dish (Kishony Lab) https://www.youtube.com/watch?v=plVk4NVIUh8 • The generation time of E.coli is 20 minutes. • This experiment took 11 days that is 792 E. coli generations. • If the generation time for humans is 20 years it is the equivalent of 15,840 years. • This is longer than recorded human history. NEXT LECTURE: PHYLOGENETICS Phylogenetics http://tolweb.org/tree/ Investigating the tree of life Definition Phylogenetics is defined as the study of evolutionary history that reveals relationships among biological entities - often species, individuals or genes (which may be referred to as taxa). Nature can be deceptive. • This looks like a snake but it is a legless lizard, the European glass lizard (Ophisaurus apodus). • What tells us it is not a snake? • It does not have the classic traits of a snake such as; • a highly mobile jaw. • a large number of vertebrae • a short tail located behind the anus • These three traits are shared by all snakes. Phylogenetics Phylogenetic tree • Snakes and lizards are part of the continuum of life extending from their common ancestor to the most recent species. • We will now look at hypotheses regarding how diversity evolved from the point of the view of the pattern of evolution – observations of evolutions products over time. Rather than the evolutionary process (mechanisms). Phylogenies show evolutionary relationships How are organisms named? • Common names - such as monkey and finch are useful but confusing as there are many species of monkeys and finches. • Others do not accurately describe the organism they refer to. An example are the three “fishes”: jellyfish (a cnidarian), crayfish (a small, lobster like crustacean), and silverfish (an insect). • And organisms have different names in different languages: Frog in English becomes kurbağa in Turkish. Phylogenies show evolutionary relationships • • • • • • How are organisms named? Binomial nomenclature The two-part format of the scientific name, commonly called a binomial, was developed in the 18th century by the botanist Carolus Linnaeus. The first part is the genus (plural, genera) to which the species belongs. The second part is unique for each species within a genus. Example Panthera pardus, the scientific named for a Leopard. The first letter is always capitalised and the whole binomial is italicized. Phylogenies show evolutionary relationships • • • • • • • Hierarchical classification of organisms This taxonomic system was developed by Linnaeus and is called the Linnaean system. Closely related species are grouped into the same genus. Example: The leopard (Panthera pardus), is in the genus containing the African lion (Panthera leo), the tiger (Panthera tigris), and the jaguar (Panthera onca). In addition taxonomists use progressively more comprehensive categories for classification. This system places related genera in the same family, families into orders, orders into classes, classes into phyla (singular, phylum), phyla into kingdoms, and, more recently, kingdoms into domains. Each level is a taxon. Panthera is a taxon at the genus level and Mamalia is a taxon at the class level. Taxon above the genus level are not italicized. Phylogenies show evolutionary relationships Hierarchical classification of organisms • This hierarchical system allows us to group similar species together and group them into increasingly broader categories. • This allows us to catalogue/classify the world of diversity. • Example: Pine and Fir trees look similar (remember the legless lizard and the snake) but different enough to be different species (genera). But are similar enough to be in the same family, the Pinaceae. • As categories become broader the characters used to classify one group of organisms will be of no use in another. • For example the level of morphological and genetic diversity between an Order of snails and an Order of mammals is not the same. • The placement of species within this hierarchy does not always fit evolutionary history. • Watch these videos they explain phylogenetic trees. • Example of how to make a tree https://www.youtube.com/watch?v=6_XMKmFQ_w8 • Phylogenetics https://www.youtube.com/watch?v=fQwI90bkJl4 Phylogenies show evolutionary relationships • • • • • Linking Classification and Phylogeny Evolutionary history can be shown as phylogenetic tree like Darwin’s original tree of life. This was originally based on species having key morphological features in common (taxonomy). The Linnaean system distinguishes groups (amphibians, mammals, reptiles etc.). But it tells us nothing about their evolutionary relationship. An alternative system is cladistics where organisms are classified based on evolutionary relationships. With species assigned to groups including a common ancestor. In this system taxon's are called clades. This tree shows the evolutionary relationships within the Carnivora. Evolutionary relationship between members of the Carnivora Phylogenies show evolutionary relationships • • • • • • Phylogenetic trees Phylogenetic trees represent hypothetical evolutionary relationships. On the left is the root, the common ancestor. The relationships are represented by two-way branch points. Each branch point represents a common ancestor of the two evolutionary lineages diverging (arising) from it. The order that Taxa appear on the tree does not indicate the sequence of evolution. They show patterns of descent not phenotypic similarity. Phylogenies show evolutionary relationships Phylogenies show evolutionary relationships Practical applications of phylogenetic trees • Maize originated in the Americas, and is now a global crop. • A tree based on DNA sequence data identified two wild grass species that may be maize’s closest living relatives. • These may be useful “reservoirs” of beneficial genes that can be introduced to cultivated maize by cross breeding. Wild Maize Darraq et al 2010 DOI: 10.1186/1471-2164-11-233 Phylogenies are inferred from morphological and molecular data • Phenotypic and genetic similarities due to shared ancestry are called homologies. • Morphological homology: The similarity in the number and arrangement of bones in the forelimbs of mammals is due to descent from a common ancestor. • Genetic homology: The similarity of genes or other DNA sequences if descent is from sequences carried by a common ancestor. • Generally organisms with similar morphologies or DNA sequences are likely to be closely related. Phylogenies are inferred from morphological and molecular data • However, morphological divergence between related species can be greater than the genetic diversity. The opposite is also true. • Example: Hawaiian silversword plants, some species are tall trees. Others are dense ground-hugging shrubs. Despite the morphological differences the genetic differences are very small. • The molecular divergence indicates the group began diverging 5 million years ago. This is called a molecular clock. Dubautia scabra Dubautia linearis Phylogenies are inferred from morphological and molecular data Convergent evolution • This is when similar environmental pressures and natural selection produce similar adaptations in organism from different lineages. • Example: the two mole-like animals shown. • They look similar. But their internal anatomy, physiology and reproductive systems are different. Differences in the genetic and fossil evidence indicates their common ancestor lived 140 million years ago. • Phylogenetically, they are far apart. An organism’s evolutionary history is documented in its genome • Comparison of nucleic acid and other molecules can be used to deduce relationships. • Molecular data can uncover evolutionary relationships between groups that have little common ground for morphological comparison, such fungi which are more closely related to animals than plants. • Molecular methods allow reconstruction of phylogenies among groups of living organisms for which there is little or no fossil record. An organism’s evolutionary history is documented in its genome • • • • • • Evolution leaves a footprint in nucleic acids Different genes evolve at different rates even in the same evolutionary lineage. Molecular trees can represent short or long periods of time depending on which genes are used. Slow change: DNA coding for ribosomal RNA (rRNA) changes slowly. Comparing DNA sequences in these genes is used to investigate relationships between taxa that diverged 100’s millions of year ago. Example: Such studies show that fungi are more closely related to animals than to plants. Rapid change: Mitochondrial DNA (mtDNA) evolves rapidly and is used to investigate recent evolutionary events. Example: This method was used to trace relationships among Native American groups. It confirmed other evidence that the Pico (Arizona), May (Mexico) and Yanomami (Venezuela) are closely related. And probably descended from the first of three waves migrants that crossed the Bering land bridge form Asia to the Americas 15,000 years ago. Image source: https://dna-explained.com/2013/09/18/nativeamerican-mitochondrial-haplogroups/ An organism’s evolutionary history is documented in its genome Gene duplication and gene families • Gene duplication plays an important part in evolution as it increases the number of genes in the genome. This increases the opportunity for further evolutionary changes. • These molecular phylogenies must account for repeated duplications that result in gene families. These are groups of related genes within an organisms genome. • By accounting for duplication we can distinguish two types of homologous genes. Orthologous and paralogous genes. An organism’s evolutionary history is documented in its genome • • • • Gene duplication and gene families A) Orthologous genes (from the Greek orthos, exact): Homology results from speciation (one species becoming two or more species) and occurs in genes found in different species. Example: Genes coding for cytochrome c in humans and dogs are orthologous. B) Paralogous genes (from the Greek para, in parallel): Homology results from gene duplication; multiple copies have diverged from one another within a species. Example: Olfactory receptor genes have under gone many duplications in vertebrates. Humans have 380 and mice have 1200 functional copies of these paralogous genes. An organism’s evolutionary history is documented in its genome • • • • Gene duplication and gene families Orthologous genes diverge only after speciation; that is after the genes are found in different species. Example: Although cytochrome c serves the same function in humans and dogs, the gene sequence in humans has diverged from that of dogs since they last shared a common ancestor. Paralogous genes can diverge within a species as there is more than one copy in the genome. Example: Human olfactory receptor gene family members have diverged from each other during our long evolutionary history. They now specify proteins conferring sensitivity to varied molecules, ranging from food odours to sex pheromones. An organism’s evolutionary history is documented in its genome • • • • • Genome evolution We can now compare the entire genomes of different organisms and two patterns have emerged. Pattern 1: Lineages that diverged long ago often share many orthologous genes. Example: Although the human and mouse lineages diverged about 65 million years ago, 99% of human and mice genes are orthologous. As are 50% of human and yeast genes despite one BILLION years of divergent evolution. This explains why many divergent organisms have the same biochemical and developmental pathways. Because of these shared pathways, the function of genes linked to human diseases can be studied in yeast and other distantly related organisms. An organism’s evolutionary history is documented in its genome • • • • • Genome evolution Pattern 2: The number of genes a species has does not seem to increase through duplication at the same rate as phenotypic complexity. Humans have about four times as many genes as yeast, a single-celled eukaryote, even though—unlike yeast—we have a large, complex brain and a body with more than 200 different types of tissues. There is evidence that humans genes are more versatile than those in yeast. A single human gene can encode multiple proteins that perform different tasks in different tissues. The mechanism underlying this genomic versatility and phenotypic variation are an ongoing challenge. Molecular clocks help track evolutionary time Introduction • Evolutionary biology aims to understand the relationships between organisms. To do this it is important to know when lineages diverged from each other. • Earlier we saw that the ancestral Hawaiian silversword diverged 5 million years ago. This estimate relied on the concept of a molecular clock. Molecular clocks help track evolutionary time How molecular clocks work • Molecular clocks are based on the observation that some genes and other regions of the genome appear to evolve at constant rates. • An assumption underlying the molecular clock is that the number of nucleotide substitutions in orthologous genes is proportional to the time that has elapsed since the genes branched from their common ancestor. • In the case of paralogous genes, the number of substitutions is proportional to the time since the ancestral gene was duplicated. Molecular clocks help track evolutionary time How molecular clocks work • We can calibrate the molecular clock of a gene that has a reliable average rate of evolution by graphing the number of genetic differences—for example, nucleotide, codon, or amino acid differences—against the dates of evolutionary branch points that are known from the fossil record. • The average rates of genetic change inferred from such graphs can then be used to estimate the dates of events that cannot be discerned from the fossil record, such as the origin of the silverswords discussed earlier. Molecular clocks help track evolutionary time The mammalian molecular clock The number of accumulated mutations in seven proteins has increased over time in a consistent manner for most mammal species. The three green data points represent primate species, whose proteins appear to have evolved more slowly than those of other mammals. The divergence time for each data point was based on fossil evidence. By averaging the number of mutations in more than one gene or protein variation in the mutation rate is smoothed out. Molecular clocks help track evolutionary time Differences in clock speed • Some mutations are selectively neutral –they are not beneficial or detrimental. • Others are harmful, and are quickly removed from the gene pool by natural selection. • In genes were mutations are neutral or have little or no effect of fitness, then the rate of mutation will be regular like a clock. • The less important the gene is for fitness the faster its clock. • The more important the gene is for fitness the slower its clock will tick. Molecular clocks help track evolutionary time • • • • • • Applying a molecular clock: Dating the origin of HIV Phylogenetic analysis shows that HIV, the virus that causes AIDS, is descended from viruses that infect chimpanzees and other primates. When did it jump the species barrier in to humans? HIV’s genetic material is RNA, and like other RNA viruses, it evolves quickly. Using HIV-1M, the most common HIV strain in humans, samples were compared that had been collected at different times. The earliest sample was from 1959. Sequence comparison showed the virus had evolved in a clock like fashion. It indicated the jump to humans of HIV-1M happened around 1930. Later work using a more advanced molecular clock indicated a date of 1910. Molecular clocks help track evolutionary time Applying a molecular clock: Dating the origin of HIV • Sequence comparison showed the virus had evolved in a clock like fashion. • It indicated the jump to humans of HIV-1M happened around 1930. Later work using a more advanced molecular clock indicated a date of 1910. • Black dots based DNA sequences of an HIV gene in patients blood. The tree of life continues to change based on new data • As phylogenetic data has accumulated we have gone from two Kingdoms (plants and animals) to five Kingdoms (Monera (prokaryotes), Protista (a diverse kingdom consisting mostly of unicellular organisms), Plantae, Fungi, and Animalia). • This system highlighted the two fundamentally different types of cells, prokaryotic and eukaryotic. Prokaryotes were separated from all eukaryotes by placing them in their own kingdom, Monera. • But genetic phylogenies showed that some Prokaryotes were more different from each other than they were from Eukaryotes. • This resulted in the domain system (Bacteria, Archaea, and Eukarya) a taxonomic level higher than Kingdoms. The tree of life continues to change based on new data • In the new Domain system. • The domain Bacteria contains most of the prokaryotes. • The domain Archaea are diverse prokaryotes inhabiting a wide variety of environments. • The domain Eukarya contains all organisms containing true nuclei. This includes single celled organisms and multicellular plants, fungi, and animals. • This system reflect the fact that most of evolutionary history has been about single celled organisms. 180 years of progress in phylogenetics First tree of life. Darwin 1838 A speculatively rooted tree for rRNA genes, showing the three life domains: bacteria, archaea, and eukaryota. The black trunk at the bottom of the tree links the three branches of living organisms to the last universal common ancestor. (https://en.wikipedia.org/wiki/Phylogenetic_tree) NEXT: HOW EVOLUTION WORKS -microevolution How Evolution Works? MICROEVOLUTION Terminology review • Population – group of individuals belonging to same species in a defined area. • Allele – alternate form of a given gene sequence. • Genotype – allele combination of a specific set of genes. • Phenotype – physical apperance of an individual. • Gene pool – all allele forms of a given population. Microevolution • How does it work? • Microevolution is driven by changes in the genetic make up of individuals and ultimately populations. Exactly how do populations evolve? • There are many different alleles in a population but a frog can only have two. • These two frogs have different alleles for coloration. GENE POOL : All possible allele combinations present in a population of a species GENETIC EQUILIBRIUM • States that allele frequencies (genetic variation) in a population will remain constant from generation to generation if there are no factors causing those frequencies to change. • This situation in which allele frequencies remain constant is called genetic equilibrium. • Populations that are not evolving are said to be in Hardy-Weinberg equilibrium. • At Hardy-Weinberg equilibrium, allele frequencies don’t change. The sum of allele frequencies is equal to 1 • Genotypes frequencies: • Genotype AA Aa aa • Frequency p2 2pq q2 Conditions required for HardyWeinberg equilibrium to hold true: – Very large population – No gene flow into or out of the population – No mutations – Random mating – No natural selection • Departure from these conditions results in a change in allele frequencies in the population. – Evolution has occurred! In reality Populations in nature never meet the conditions of Hardy–Weinberg equilibrium— All biological populations evolve. The model is useful for predicting approximate genotype frequencies of a population. Specific patterns of deviation from Hardy– Weinberg equilibrium help identify mechanisms of microevolutionary change. Microevolution • • • • • • • How does it work? Microevolution is driven by five processes. Small sample size. Non random mating. Mutation. Gene flow. Natural selection. Short time scale events (generation-togeneration) that change the genotypes and phenotypes of populations Allelic frequency will remain constant generation to generation unless disturbed by mutation, migration, nonrandom mating, natural selection, or genetic drift. These are sources of microevolutionary change. Sources of genetic variability 1. Mutations • Changes in: – Nucleotide base pairs – Arrangement of genes on a chromosome – Chromosome structure • Only mutations in gametes are inherited • Most mutations are silent: – Only a small % of the DNA is expressed – Mutations that are expressed are usually harmful – Mutations do NOT cause evolution, but natural selection needs the variations which mutations create Sources of genetic variability 2. Gene Flow Genetic exchange due to the migration of fertile individuals or gametes between populations. Also called migration — is any movement of individuals, and/or the genetic material they carry, from one population to another. Gene flow includes lots of different kinds of events, such as pollen being blown to a new destination or people moving to new cities or countries. If gene versions are carried to a population where those gene versions previously did not exist, gene flow can be a very important source of genetic variation. Sources of genetic variability 3. Sexual Reproduction NON-RANDOM MATING • Mating in which a given member of a population is not equally likely to mate with any other given member. • Sexual selection is a form of nonrandom mating. This is mating based on phenotype, based on the choices made by the female of the species. • The male will increase the proportion of his alleles in the next generation. The most common cases of nonrandom mating involve inbreeding – mating between individuals of similar genotypes, either by choice or due to environmental factors such as location. Sexual reproduction con’t • Inbreeding does not change genetic variation, but increases the allele frequency of homozygous genotypes. • Inbreeding depression is seen in some cases, where inbred individuals have lower fitness than non-inbred individuals. • – fitness – relative ability of a genotype to contribute to future generations. – fertility declines and high juvenile mortality associated with “unmasking” harmful recessive alleles can reduce fitness for inbred individuals. – hybrid vigor leads to higher relative fitness for hybrids by increasing heterozygous allele frequency. Self-fertilization is the most extreme case of inbreeding. Sexual Reproduction con’t • Assortive mating – a type of nonrandom mating where mates are (sexually) selected based on phenotypes – really is an aspect of natural selection. – positive assortive mating – selection for the same phenotype; works like inbreeding for the genes governing that phenotype, and for loci closely linked to those genes. – negative assortive mating – selection for the opposite phenotype. • less common than positive assortive mating. • leads to a decrease in homozygous genotypes for the genes governing the selected phenotype, and for loci closely linked to those genes. Population-environment interactions 1. Small sample size: Genetic drift Consequences of small population size: genetic drift – – • Genetic drift tends to decrease genetic variation within a population. Consider taking a small sample of individuals from a larger population. • Genetic drift tends to increase genetic variation between If only two individuals were picked populations. they almost certainly won’t reflect the allele frequency in the larger NOTE: Genetic drift is a major factor population (in many cases, they in evolution, especially when can’t even possibly do so). populations are split, but does The same holds true for 3, 4, or 5 NOT involve natural selection. individuals. As the selected sample gets larger it becomes more likely that the sample reflects the allele frequency in the larger population. Genetic Drift con’t • Two reasons of genetic drift – Bottleneck effect – Founder’s effect • A small sample of alleles is likely to yield a gene pool that is different from the distribution found in the larger population. Genetic Drift con’t • Bottleneck effect – The change in allele frequency in a population due to change following a sharp reduction in the population size. • Founder effect – When a small portion of a population migrates/separated to another area, starting a new population. Genetic Drift con’t If there is a drastic change (disease, migration or natural catastrophe) in the survival of a large population, there is little change in the frequency of that population's alleles. In a smaller population, drastic change can result in loss of an entire allele. Bottleneck effect Bottleneck effect Cheetahs survived a drastic bottleneck after the last ice age, 10,000 years ago. Bottleneck effect • Consequences: low genetic diversity and depressed reproduction. • Examples; – Golden hamster; most captive hamsters descended from one wild litter found in Syria in the 1930’s. – European Bison descended from 12 animals. – Video of founder and bottle neck effect. Founder effect • When small groups of individuals in secluded areas (eg islands) found larger populations. • In humans this often leads to genetic diseases due to inbreeding. • Example Cohn’s syndrome a fatal genetic disease common in the Amish community in the USA. • The Amish are a reclusive religious select descended from a few 100 migrants from Europe in the 18C. • Such diseases are common in historically isolated communities. • Another example are Darwin’s Finches. Here the result over a longer time period was new species. Population-environment interactions 2. Natural Selection • Traits of those who are more successful in reproducing (survival of fittest) will become more widespread in a population, the alleles that bring about these traits will increase in frequency from one generation to the next. • Natural selection is environment oriented and thus selected adaptive traits change according to changes in the environment. • Natural selection can only ‘work’ if there is preexisting variation within the population. Artificial selection • Artificial selection: Humans choose traits they prefer in animals or plants and breed to achieve those “desirable” individuals. Natural Selection con’t • Natural selection is the only one of the five agents of microevolution that consistently works to adapt organisms to their environment. – Genetic drift is random. – Mutation has a negative effect, or no effect. – Gene flow doesn’t necessarily bring in genes that are better suited to the environment. – Non random mating doesn’t have anything to do with matching individuals to environment. Three modes of Natural Selection • When natural selection operates on characters that are polygenic and continuously variable, it can proceed in one of three ways. • stabilizing selection • disruptive selection • directional selection Stabilizing Selection • Intermediate forms are favored over extreme forms. Example: Human baby birth weight. • Infant deaths are higher at the extremes of birth weight, children most likely to survive have an average birth weight. Example: Directional Selection • When natural selection moves a character towards one of its extremes. • Example Video - Mutation behind the colour change Disruptive Selection • When natural selection moves a character to both its extremes. • Occurs much less frequently in nature. • Examples in nature are Darwin’s finches and the Peppered moth Microevolution • The following video give examples of the five processes that contribute to microevolution. 1. Small sample size. 2. Non random mating. 3. Mutation. 4. Gene flow. 5. Natural selection. • Video of microevolution. Microevolution in humans • Wisdom teeth: When we had to • Lactose tolerance: The ability to chew roots, nuts and eat raw meat digest lactose in our mothers milk we wore out our molars so we had was lost when we were weaned. an extra set –Wisdom teeth that Cattle, goats and sheep were erupted after puberty. Now our domesticated 10,000 years ago food is easier to chew and our providing milk. 8,000 years ago a jaws and mouths are smaller with mutation occurred in Europe that no room for the extra teeth. allowed people to digest lactose as adults. This gave them a • Now 35% of people are born with nutritional advantage. This no wisdom teeth. mutation spread as far a India. Today 95% of people of north European descent have this mutation. Similar mutations also arose in Arabia and sub-Sharan Africa Microevolution in humans • The human genome is still • Extra Artery in the arm: The changing as we adapting to our median artery forms in the post hunter-gatherer foetus and supplies blood the environment: Over 40,000 years fore arm and hand. It vanishes approximately 1800 genes have once the radial and ulnal become more common, many arteries form. But now 1 in 3 are involved in fighting diseases people keep this artery for life. such as malaria and We do not know yet if this tuberculosis. When we adopted gives us any advantage. our agrarian life style 10,000 years ago the population • The links below are to popular increased as did the incidence articles in the press on of disease. This increased the microevolution. selection pressure for increased disease resistance. https://nypost.com/2020/10/09/more-humans-born-with-an-extra-artery-as-part-of-microevolution/ https://www.latimes.com/archives/la-xpm-2009-feb-08-sci-evolution8-story.html https://www.mentalfloss.com/article/30795/5-signs-humans-are-still-evolving How Evolution Works? MACROEVOLUTION MACROEVOLUTION • Macroevolution is essentially the formation of new species (speciation) and accompanying events. What are species? • relatively easy to define for sexual organisms, hard for asexual organisms • biological species concept (for sexual organisms) – one or more populations whose members are: • capable of interbreeding • able to produce fertile offspring • reproductively isolated from other such groups What is species con’t • Asexual species – definition based on biochemical differences (think DNA sequence) and morphological differences; no solid rules. • Also includes use of “race,” “subspecies,” and “strain” designations. • In asexual species, microevolution over time directly leads to macroevolution (speciation). Macroevolution– causes of speciation There are two basic mechanisms via which speciation can occur: • Allopatric speciation (geographical isolation) • Sympatric speciation (reproductive isolation) Polyploidy (extra sets of chromosomes) is a major factor in sympatric speciation in plants Polyploidy (extra sets of chromosomes) is a major factor in sympatric speciation in plants. There are different mechanisms for producing polyploids. 1. Autopolypoidy – multiple sets from one parent species. The resulting tetraploid plant can produce fertile offspring. These are reproductively isolated from the 2n plants of the original population. This is because triploid (3n) offspring from a 4n x 2n cross have reduced fertility. Examples Naturally occurring polyploidy has occurs in Soybean and cabbage. This is called the triploid block. So in one generation we have reproductive isolation. Bananas are a famous example of a sterile triploid crop. Fruit are produced without fertilisation and are seedless. Polyploidy is a major factor in sympatric speciation in plants – cont. 2. hybridization + allopolyploidy – closely related species produce a hybrid that must double its chromosome number to reproduce successfully; a new, viable hybrid species is thus formed. 1. Species A undergoes a doubling in the chromosome number during gamete formation. 2. Gametes from both species then hybridize during reproduction. 3. The resulting hybrid has the full complement of chromosomes from the one parent (Species A) but not the other. 4. In the next generation the hybrid gamete is not reduced. And the plant can not self pollinate. 5. On hybridization with a gamete from species B a fertile hybrid id produced. This form of polyploidy greatly increases genetic diversity in the new hybrid. It is a major force in plant diversification eg wheat. 1 4 2 3 5 Example of Allopatric speciation Video of Darwin’s Finches Food choice in finches Macroevolution con’t Sympatric Speciation • The basis of macroevolution in sexual species is microevolution coupled with reproductive isolation (ways of preventing gene flow between species) ---Reproductive isolation can occur in a variety of ways • Reproductive isolating mechanisms can be classified as either prezygotic or postzygotic. • Prezygotic barriers – prevent fertilization (zygote formation) between gametes from two species. • Postzygotic barriers – reproductive isolation after fertilization has occurred. Pre-zygotic barriers – prevent fertilization Habitat isolation (or ecological isolation) – isolation by differences in habitat occupied at the time of mating. Temporal isolation – isolation by differences in timing of mating; Ex: different mating season in some skunks, diffent flowering time in some plants, different mating dates in some frogs Behavioral isolation – differences in behavior that cause reproductive isolation Ex: mating calls, courtship patterns, and other mating rituals. Ex: Tigers vs Lions Pre-zygotic barriers – prevent fertilization con’t • Mechanical isolation – differences in physical structure make mating impossible. Ex: two species of dragonfly flower differ in flower color and shape which attract different pollinator animals (hummingbird vs bumbble bee) so they can not cross pollinate • Gametic isolation – mating occurs, but the sperm and egg can not fuse; Ex: two species of sea urchins Post-zygotic isolation : after fertilization has occurred Hybrid inviability: The most common type of postzygotic barrier • Zygote formed from the mating of two species does not develop normally, the embryo is aborted. Hybrid sterility – a zygote of a hybrid proceeds through normal development, but is reproductively sterile. • mostly due to problems in meiosis Hybrid breakdown – a zygote of a hybrid proceeds through normal development, but F2 generation is sterile Ex: some hybrids of cotton, rice and sunflower species The slow process of evolutionary change • • • • • • Darwin realised two things: • producing of different breeds of farm animals, dogs, and pigeons etc. was an accelerated version of natural selection. • for natural selection to result in speciation in nature requires many thousands of years. In this he was influenced by the work of the geologists Hutton and Lyell (see first lecture). Fossil evidence from the geological record supported this. This is the result of the slow speed of micro and macro evolution. Ultimately, over 1000’s years, small differences between populations (microevolution) diverge and are amplified when migration between them is reduced or stopped by allopatric or sympatric isolation. This allows macroevolution to occur. This is where geologic time scales become important in explaining evolutionary change. Darwin realized these changes required long periods of time. The geological timescale is a 'calendar' of events in the Earth's history. It shows major geological and climactic events, and how these events affected the emergence and disappearance of species over time. These timelines are products of several scientific fields such as paleontology, biogeography, plate tectonics geology, and biology). •The study of glaciers left over from the ice ages has provided an important line of evidence for continental drift. • Glacial sediments from South America, Africa, India, Madagascar, Arabia, India, Antarctica and Australia showed evidence of having once been joined together, suggesting the existence of the supercontinent Gondwana. Continental Drift English naturalist Charles Darwin (1809–1882) Almost 30 years of detailed observations and evidence collected by Darwin led to the publication of the book ‘ The Origin of Species’. This is the base used to explain the mechanisms in nature which leads to EVOLUTION. The theory of evolution by natural selection, first formulated in Darwin's book "On the Origin of Species" in 1859, is the process by which organisms change over time as a result of changes in heritable physical or behavioral traits. Changes that allow an allow an organism to better adapt to its environment will help it will help it survive and have more offspring. The Origin of Species EVOLUTION accounts for both the unity and diversity of life. In many cases, features shared by two species are due to their descent from a common ancestor. Differences are end product of natural selection modifying the ancestral origin in different environments. It is this combination of unity and diversity in life that we will consider in future lectures. Evidences of Unity & Diversity of Life Evidences come from; Fossil record Homologous structures Vestigial structures Biochemical evidence Embryological development Genetics Fossil Records What does the Fossil Record tell us about organisms? Looks (size, shape, etc.). Where or how they lived. What other organisms they lived with. What time period they lived in (based on location in rock layers). What order living things came in (based on location in rock layers). Transitional forms. Organisms that were intermediate (between) two other major organisms. Example: Evolution of Horse Homologous Structures Homologous structures: Homologous structures develop from the same tissues, but have different forms with different functions. Same origin -- different form/function The similarity is due to having derived from the same common ancestor. Example: Bone structure of arms and legs in all vertebrates Vestigial Structures Todays animals may have structures that serve little or no function remnants of structures that were functional in ancestral species evidence of change over time EXAMPLES: some snakes & whales show remains of the pelvis & leg bones of walking ancestors EXAMPLES: Wings that do not fly EXAMPLES: In Human: Wisdom teeth, tailbone (coccyx), muscles around the ear, appendix, Vestigial Structures Human hand con’t During embryo development we have extra lizard-like muscles in our hands. They disappear before birth. They are a relic from when reptiles transitioned to mammals 250 million years ago. Source; https://www.bbc.com/news/health49876827 Biochemical Evidence • • • • DNA is present as the heritable genetic material in all living forms. All proteins in all living organisms contain the combinations of same 20 amino acids. Cytochrome C (respiration) protein structure. Hemoglobin (gas exchange) protein structure. Evolutionary relationships among species are documented in their DNA & proteins. Closely related species have sequences that are more similar than distantly related species. Embryologic development • Similar embryological development in closely related species • all vertebrate embryos have a gill pouch at one stage of development • fish, frog, snake, birds, human, etc. They all have gills because our common ancestor lived in the ocean. What else tells us we originated in the ocean? We all have a salt water space suit. EVOLUTION EVENTUALLY GENERATED DIVERSE CLASS OF LIFE NEXT LECTURE Evolution is based on survival of the fittest. Why does one species not dominate all others? What evolutionary mechanism stops one species from dominating all others? How is evolutionary balance maintained? It could all depend on a children’s game. How a children's game explains evolutionary balance This game played between two players originated in China over 2000 years ago and has evolved in to the game “Rock, paper, scissors” now widely played. Hand signals represent rock, paper or scissors. Rock breaks scissors: Rock wins Scissors cut paper: Scissors wins Paper wraps rock: Paper wins In this game, no single strategy can dominate as the opponent can also change their strategy to counter it. In game theory this means we have an equilibrium or balance. This came to interest biologists who were interested in explaining evolutionary balance. Rock, Paper, Scissors If evolution favours survival of the fittest. Why is there no permanent winner? • Populations of species continually change. • This is because species interact with each other through competition for resources, or predation of one by another. • Within species, competition is usually for access to mates. • This process results in evolutionary balance between and within species. Competition between bacteria • Eschericia coli bacteria found in the digestive tracts of animals are an example. • Strain C produces the antibiotic Colchicine to which it is also resistant. Both production and resistance have an energetic cost so its growth is slow. • The susceptible strain, Strain S, grows quickly but is killed by strain C • STRAIN C WINS! But no. A resistant strain, Strain R, develops. It is not killed by Strain C and only has the energetic cost of resistance so grows more quickly than strain C and out competes it and dominates. • But, then Strain S which grows more quickly than strain R makes comeback as there is less Colchicine in the environment and starts to take over by out competing strain R. • ROCK, PAPER, SCISSORS. https://www.quantamagazine.org/biodive rsity-may-thrive-through-games-of-rockpaper-scissors-20200305/ Within species competition for mates • The male side blotch lizard found in California plays the same game but this time the prize is access to females. • There are three “strains” of this small lizard. • Orange throated: Aggressive males keep large numbers of females and attack and drive off male rivals. • Blue throated: Solitary males are monogamous. Males warn each other of approaching rivals and group together to defend their females. • Yellow throated. Males look like females and enter rival territory as orange throated males can not watch all their females • https://aeon.co/videos/how-multicoloured-sideblotched-lizards-put-game-theory-into-evolutionaryaction Implications of this evolutionary balance • When two species compete one will drive the other to extinction. • Introduce a third species and you start to see a game of rock, paper, scissors. • The more species the more complex the game. Computer simulations of bacteria have used over 70 species. • One species cannot be good at everything. So there is always competition. • This competition also changes local environments. This dynamic situation maintains and increases biodiversity. • When an external force or barrier is introduced (e.g. pollution, deforestation) that reduces competition between some species, one species may be driven to extinction. • This may produce a cascade effect with some species now able to drive others to extinction. • To prevent extinction we may have to protect whole ecosystems. How might this game effect us? • In 1900 there were 1.6 billion people on earth. • Now there are 7.8 billion. • Advances in medicine, public health (water and sewage treatment, vaccination) and agriculture protect us from diseases and famine that so often devastated populations. • By this we reduced the restraint on population growth. • Our rock broke natures scissors. • Will nature change its strategy? • Are virus’s like COVID-19, the paper that beats our rock? https://slideplayer.com/slide/5281291/ VIRUSES: Virus (Latin for poison) NOT AN ORGANISM NO CELL STRUCTURE NOT ALIVE OUTSIDE HOST CAN NOT REPLICATE IN THE ABSENCE OF A CELL HAVE GENETIC MATERIAL THUS IN GENERAL THEY ARE NEITHER LIVING NOR NON-LIVING----SO WHAT ARE THEY? They are infectious particles obligate intracellular parasites Can be crystallized Size range of 20-250 nm (1 nanometer (nm) = 0.000000001 meter) Their genetic material codes for from 3 genes to as many as 2000 Virus Structure Nucleic acid enclosed in a capsid (protein coat) and, sometimes, a lipid –rich envelope (derived from host cell membrane). Genetic material can be DNA or RNA, linear or circular, single- or double-stranded. VIRAL CLASSIFICATION 1. Nature of the nucleic acid: RNA or DNA 2. Symmetry of the capsid 3. Presence or absence of an envelope 4. Dimensions of the virion 5. Type of host organism 6. Replication type RdRp = RNA dependent RNA polymerase VIRUS GENOME Their genomes may consist of double-stranded DNA, single-stranded DNA, double-stranded RNA, or single-stranded RNA, depending on the specific type of virus. A virus is called a DNA virus or an RNA virus, according to the kind of nucleic acid that makes up its genome. Replication in RNA viruses ONLY RNA viruses carry the RNA dependent RNA polymerase Virus classification based on how mRNA is produced during virus replication The Baltimore system: This is the most commonly used classification system. Devised by Nobel Laureate Peter Baltimore VIRUS COAT: The protein coat that encloses the viral genome is called a capsid. Depending on the type of virus, the capsid may be rod-shaped (more precisely, helical), polyhedral, or more complex in shape. Capsids are built from a large number of protein subunits called capsomeres, but the number of different kinds of proteins is usually small. Some viruses carry a few viral enzyme molecules within their capsids. Classification by morphology • Naked – ecosahedral & helical • Enveloped ecosahedral and helical • Complex –many proteins with mixed ecosahedral and helical structures VIRAL ENVELOPE: Some viruses have accessory structures that help them infect their hosts. many viruses found in animals, have viral envelopes. These envelopes are derived from membrane of the host cell, but in addition to host cell phospholipids and proteins, they also contain proteins and glycoproteins of viral origin. Some envelopes drive from nuclear cell membrane like in herpesvirus (dsDNA, found latent in cell nucleus like prohage, stress activates it and cause blisters after viral production). VIRUS-HOST RELATION: Viruses are obligate intracellular parasites; that is, they can reproduce only within a host cell. An isolated virus is unable to reproduce. Viruses lack the enzymes for metabolism and have no ribosomes or other equipment for making their own proteins. Each type of virus can infect only a limited range of host cells, called its host range. This host specificity depends on the evolution of recognition systems by the virus. Viruses identify their host cells by a "lock-and-key" fit between proteins on the outside of the virus and specific receptor molecules on the surface of the host cell. Some viruses have host ranges as broad as several species or as small as single species. Viruses of eukaryotes are usually tissue specific. Viruses of prokaryotes are referred to as bacteriophages (phages). Viral replication shown variation with respect to host type and viral genome type. VIRAL REPLICATION STEPS 1) ENTRY TYPES: naked nucleic acid fusion of envelop with host membrane phagocytosis of the capsid 2) DISASSEMBLY release of capsid release of envelop and capsid. 3) SYNTHESIS RNA genome –viral RNA dependent RNA polymerase RNA genome---viral reverse transcriptase DNA genome----Host DNA polymerase 4) REASSEMBLY in nucleus in cytoplasm 5) EXIT lysis of host cell membrane Exocytosis 1) ENTRY 1) ENTRY HOST CELL MEMBRANE HOST CYTOPLASM 2) DISASSEMBLY 3) SYNTHESIS 4) REASSEMBLY 3) SYNTHESIS HOST NUCLEUS 5) EXIT HOST CELL MEMBRANE Bacteriophage Reproductive Cycles Recognition of viral coat/envelope by host cell membrane/cell wall receptor proteins Entery of whole virus or just virus genome into host cell Removal of protein coat and envelope Copy of viral genome and synthesis of viral proteins Assembly of viral particals Removal of viruses from the host cell ADSORPTION OF BACTERIA VIRUS (Bacteriophage) T4 bacteriophage attacking E. Coli Reproduction of Bacteriophages Lytic Cycle – culminates in the death of the host cell similar to general reproductive cycle of viruses phages that reproduce only by the lytic cycle are called virulent phages Lysogenic Cycle – does not destroy the host cell viral DNA (prophage) is incorporated into host cell’s chromosome & replicated when the bacterium reproduces occasionally a prophage will exit the host chromosome and initiate the lytic cycle to produce new viruses phages that reproduce using both types of cycles are called temperate phages Video – lytic and lysogenic cycles •WHAT DOES LYSOGENY PROVIDE The prophage represses phage replication making the host immune to further bacteriophage attack as the lytic cycle is blocked. Viral genome multiplies with the host genome. Prophage genes may code for proteins which provide new traits to the host (ex: the botulism toxin (botulinum) is the gene product of prophage in Clostridium botulinum bacteria) (SEE NEXT FIGURE). Virus may gain new trait from bacteria when prophage exits the host genome when it starts the lytic cycle.(SEE NEXT FIG). Prophage promoter may activate certain genes of the host (ex: activation of proto-oncogenes), leading to cancer. This occurs only in animal cells. Viral entry in to a host cell ADSORPTION OF ANIMAL VIRUS Video examples Penetration uncoating sythesis assembly release ANIMAL VIRAL REPRODUCTIVE CYCLE AFTER ADSORPTION 23 Some viruses are released by lysing host cells (non-enveloped or naked) Some viruses do not kill host cells, but bud out of membrane Budding viruses contain host covering (enveloped) Video Influenza virus ORIGIN OF VIRUSES Viruses infecting cells from the three domains of life, Archaea, Bacteria and Eukarya, :, suggesting that viruses originated very early in the evolution of life. The origin and evolution of viruses is mostly unknown. Viruses have never been detected as fossil particles, probably because they are too small and too fragile to succumb to fossilization processes. Even in fossilized biological materials such as plant leaves or insects in amber, preserved nucleic acid sequences of viruses have never been detected. Hypothesis for their origin are generally speculation. The origin and evolution of viruses. There is little scientific evidence to support any unifying theory. The main hypothesis is the theory of ‘‘cell origin’’, (escape hypothesis) which assumes that viruses reflect their origin from cell DNA and/or RNA, which acquired the ability to auto-replicate, create extracellular virions, exist and function independently (example of origin from plasmid and or transposons). VIRUSES & CANCER Viruses may cause cancer Often undetected Most particles of viruses do not induce cancer Cancer might develop long after viral infection Cancers do not seem contagious like viruses Proto-Oncogene Cancer causing alteration to cellular proto-oncogenes Proto-Oncogene activated by various agents ○ Mutagenic chemicals ○ High energy radiation ○ Viruses Oncoviruses – viruses that can cause cancer by activating proto-oncogenes during lysogenic state . VIRUSES & CANCER Viruses may cause cancer Often undetected Most particles of viruses do not induce cancer Cancer might develop long after viral infection Cancers do not seem contagious like viruses Proto-Oncogene Cancer causing alteration to cellular proto-oncogenes Proto-Oncogene activated by various agents ○ Mutagenic chemicals ○ High energy radiation ○ Viruses Oncoviruses – viruses that can cause cancer by activating proto-oncogenes during lysogenic state . VIRUSES & CANCER Human papillomavirus (HPV) is a group of viruses that are extremely common worldwide. They are transmitted during sex. 70% of all cervical cancer is caused by HPV. Vaccines that protect against HPV 16 and 18 are recommended by WHO. HOW TO COMBAT VIRUSES ? •Immune respose via B- lymphocyte and T- lymphocyte cells •Vaccines are harmless variants or derivatives of pathogens that stimulate the immune system to mount defenses against the actual pathogen. The antibiotics that help us recover from bacterial infections are powerless against viruses. Dead vaccine: viruses are treated with physical (heat, UV or Xray radiation) or chemical agents (hydrolytic enzymes, formaldehyde, lipid solvent) to damage the genome. Capsid stays intact and trigger immune response Live vaccines: continuous cultivation of virus for long times may spontaneously create mutant with slow replication rate. •Drugs: most antiviral drugs interfere with viral nucleic acid synthesis DNA or RNA polymerases reverse transcriptases Emerging Viruses Appear suddenly (ex) HIV, Ebola virus, West Nile virus, coranovirus (cause of SARS, severe acute respiratory syndrome) Processes that contribute to the sudden emergence of viral diseases: Mutation of existing viruses (ex: flu epidemics). Spread of existing viruses from one host species to another. Spread of a viral disease from a small, isolated population to large populations causing epidemics. PRIONS Stanley Prusiner (1942) • Discovered Prions • 1997 Nobel prize for Physiology or Medicine. • Awarded for his discovery of prions “a new biological principle of infection. What are prions? Prions are normal constituents of cells. The prion protein PrPc can under go a conformational change to PrP-sc (PrPscrapie). This is named after the degenerative disease found in sheep. Prion diseases usually transmitted via contact with nerve tissues (brain or spinal cord) Examples of diseases causes by PrPSc Scrapie in sheep ○ Exact mechanism of transmission among sheep is unknown Bovine spongiform encephalitis in cows (BSE) ○ Possibly transmitted to cows from feed that was fortified with bone marrow from sheep Diseases in Humans ○ Kuru in old New Guinea tribes Cannibalistic rituals—contact with brain matter ○ Creutzfeldt-Jakob disease (CJD) Known cases include - Contamination during corneal transplants (CJD) - Contamination from eating beef from cows with BSE (vCJD) Video Prion diseases How did BSE jump from cattle to humans After butchering the carcase remaining meat, connective tissue, cartilage and nerve tissue was mechanically removed from bones. This resulting “meat” was used in low cost meat products such as burgers and sausages. By 2001 in the UK, the use of bones was banned in mechanically recovered meat as was its use in the human food chain. Postulated Mechanism Host cells have a neuron glycoprotein PrPC . This is nearly identical to the prion protein. The incoming prion protein PrPsc can modify PrPC converting it to PrPsc. This gives abnormal folding patterns This abnormal protein loses function and protease resistant. Neurological symptoms begin. PrPc PrPSc PrPC PrPSc Mad Cow Disease PRION formation PrPSC Structure High in beta-sheet content (>40%) Partial resistance to proteinase K digestion Can form aggregated fibrous or amyloid structure Involves refolding two helices into beta-sheets KINGDOM EUBACTERIA Classifying Prokaryotes • All prokaryotes were once in the Kingdom Monera. • Recently biologists divided them into two different Kingdoms; the Eubacteria and the Archeabacteria. • Prokaryote comes from the Greek "before" and "nut" or "kernel“. • Prokaryotes appeared over a billion years before the Eukaryotes. Prokaryotes lack of true nuclei lack the extensive compartmentalization by internal membranes small circular DNA extrachromosomal DNA= plasmids Contain cell wall Plasmids • Most consisting of only a few genes. • In most environments, prokaryotes can survive without their plasmids because all essential functions are programmed by the chromosome. • Endow the cell with genes for resistance to antibiotics, for the metabolism of unusual nutrients not present in the normal environment. • Plasmids replicate independently of the main chromosome. • Can be readily transferred between partners when prokaryotes conjugate Types of Bacterial Plasmids Based on their function, there are five main classes: 1. Fertility-(F)plasmids: Conjugation. 2. Resistance-(R) plasmids: Containing antibiotic or drug resistant gene(s). 3. Col-plasmids: Contain genes that code for colicines proteins that can kill other bacteria. 4. Degradative plasmids: Enable digestion of unusual substances, e.g., toluene or salicylic acid. 5. Virulence plasmids: Turn the bacterium into a pathogen. REPLICATION by BINARY FISSION Origin of replication Actin fibres move the two origins of replication to opposite ends of the cell. While the chromosome is replicating the cell elongates. Bacterial growth • A single prokaryotic cell in a favorable environment will give rise by repeated divisions to a colony of offspring via binary fission. Lag phase: adaptation of bacteria to medium, start of synthesis to prepare to division Exponential phase: Fastest growth by continous divisions Stationary phase: Amount of survivors and amount of death are equal. Accumulation of waste. Death phase: Amount of dead cells exceeds the survivor. BACTERIAL GROWTH cont • GROWTH as applied to prokaryotes refers more to the multiplication of cells and an increase in population size than to the enlargement of individual cells. The conditions for optimal growth--temperature, pH, salt concentrations, nutrient sources, and so on--vary according to species. • In an environment with unlimited resources, the growth of prokaryotes is effectively exponential: One cell divides to form 2, which divide again to produce a total of 4 cells, then 8, 16, and so on, the number of cells in a colony doubling with each generation. Many prokaryotes have generation times in the range of 1-3 hours. Some species can double every 20 minutes (ex: E. coli) in an optimal environment. Potentially from 1 cell to over 1 million cells in 20 generations. • Most prokaryotes have diameters in the range of 1-5 µm, compared to 10-100 µm for the majority of eukaryotic cells. GENERAL BACTERIAL MORPHOLOGY Different arrangements of coccus and bacillus type of bacteria in colony CAPSULE & SLIME LAYER All bacteria secrete some sort of glycocalyx , an outer viscous covering of fibers extending from the bacterium. The glycocalyx is usually a viscous polysaccharide and/or polypeptide If it appears as an extensive, tightly bound accumulation of gelatinous material adhering to the cell wall, it is called a capsule If the glycocalyx appears unorganized and more loosely attached, it is referred to as a slime layer The functions of glycocalyx: Antiphagocytic Aid Attachment of bacteria to different surfaces Protect against drying when exposed to dry conditions Trap nutrients Help colonization with each other • Outside cell wall • Made of filaments of flagellin protein • Attached to a protein hook • Anchored to the wall and membrane by the basal body Flagella Rotate flagella to run or tumble Move toward or away from stimuli (taxis) Figure 4.8 Flagella • Outside cell wall • Made of filaments of flagellin protein • Attached to a protein hook • Anchored to the wall and membrane by the basal body Rotate flagella to run or tumble Move toward or away from stimuli (taxis) The motor is drive by a proton gradient making it a nanotech hydrogen engine. Flagella Arrangement Figure 4.7 & Pili FLAGELLA MOTIONS Figure 4.9 Enviromental conditions determine the direction of the movement • In a relatively uniform environment, prokaryotes may wander randomly • In a heterogeneous environment, prokaryotes are capable of movement toward or away from a stimulus= Chemotaxis A positive chemotaxis= moving toward food or oxygen or light A negative chemotaxis= moving away from some toxic substance Pili & Fimbriae • Thin proteins extended from plasma Membrane _ pilin protein • Short protein appendages – smaller than flagella The key difference between pili and fimbriae is that pili are found in gram-negative bacteria, whereas fimbriae are found in both Gram-negative and gram-positive bacteria. • Adhere bacteria to surfaces - Attachment to surface and each other (mostly fimbriae) • F-pilus (Sex pilus); used in conjugation – Exchange of genetic information • Flotation; increase buoyancy – swipping env. around CELL WALL • Maintains the shape of the cell. • Provides physical protection, prevents the cell from bursting in a hypotonic environment. Like other walled cells, however, prokaryotes plasmolyze (shrink away from their wall) and may die in a hypertonic medium, • Contain a unique material called peptidoglycan, CELL WALL- Peptidoglycan • Polymer of disaccharide N-acetylglucosamine (NAG) & N-acetylmuramic acid (NAM) • Linked by polypeptides Figure 4.13a Peptidoglycan Β 1-4 Glycosidic linkage Figure 4.13b, c Gram positive cell wall Gram negative cell wall GRAM STAINING • One of the most valuable tools for identifying specific bacteria is the Gram stain, which can be used to separate many species into two groups based on differences in peptidoglycan content of the cell walls. • Gram-positive bacteria have simpler walls, with a relatively large amount of peptidoglycan. • Gram-negative bacteria have less peptidoglycan and are structurally more complex. An outer membrane on the gram-negative cell wall contains lipopolysaccharides, carbohydrates bonded to lipids. Gr (-) bacteria are mostly pathogenic because of the lipopolysaccharide layer. VIDEO OF GRAM STAIN The gram stain indicates a high peptidoglycan content of the cell wall. As indicated by the blue stain of the positive cells. Gram-Positive cell walls • Teichoic acids: – Lipoteichoic acid links to plasma membrane – Wall teichoic acid links to peptidoglycan • Provide antigenic variation to avoid host defences. TEICHOIC ACID Gram positive bacteria only Glycerol, Phosphates, & Ribitol containing types Attachment surface for viruses Gram-Negative Outer Membrane • Lipopolysaccharides, lipoproteins, phospholipids. toxic, helps protect the pathogens against the defenses of their hosts. more resistant to antibiotics because the outer membrane impedes entry of the drugs. O -polysaccharide antigenic site (recognised by host immune system). Lipid A is an endotoxin. • Forms the periplasm between the outer membrane and the plasma membrane. • Protection from phagocytes, antibiotics. • Porins (proteins) form channels through membrane. Gram-positive cell walls Gram-negative cell walls • Thick peptidoglycan • Thin peptidoglycan • Teichoic acids • No teichoic acids • Outer membrane • (lipopolysaccharide) Damage to Cell Walls • Lysozyme (enzyme) digests disaccharide in peptidoglycan. • Penicillin (antibiotic) inhibits peptide bridges in peptidoglycan. Protoplast is a wall-less cell of plant, bacteria, fungi. BACTERIAL SPORES • The ability of some prokaryotes to withstand harsh conditions is impressive. Some bacteria form resistant cells called endospores • Boiling water is not hot enough to kill most endospores in a relatively short length of time. • To sterilize media, glassware, and utensils in the laboratory, microbiologists use an appliance called an autoclave, a pressure cooker that kills even endospores by heating to a temperature of 120°C. • In less hostile environments, endospores may remain dormant for centuries or more. If placed in a hospitable environment, they will hydrate and revive to the vegetative (colony-producing) state. For example, in 2000, researchers revived a bacterial spore that had apparently been encased for 250 million years in a salt formation within caverns in New Mexico. Bacterial ‘striptease’ evades antibiotics • Antibiotics attack the bacterial cell wall. • This weakens it resulting in the enclosed cell bursting. • Scientists have observed some bacteria, found in urinary tract infections common in old people, releasing themselves from their cell wall when they come into contact with antibiotics. • This is very risky as they may burst if in a low osmotic potential environment. • These ‘naked’ bacteria may be attacked by the immune system but some may survive to regrow their cell wall and reinfect once the antibiotic has gone. • In the video you can see the moment bacteria strip off their cell wall as they lose their clear (in this case rod-like) structure and become bigger and more flimsy. Video: Bacterial striptease Source: https://www.bbc.com/news/health-49826085 The developmental cycle of the Endospore. https://www.youtube.com/watch?v=NAcowliknPs Bacterial endospores. Phase microscopy of sporulating bacteria demonstrates the refractility of endospores, as well as characteristic spore shapes and locations within the mother cell. Figure. An anthrax endospore. This prokaryote is Bacillus anthracis , the notorious bacterium that produces the deadly disease called anthrax in cattle, sheep, and humans. 4 Kg of powdered toxin is enough to kill whole human population. Bacterial Toxins Endotoxin (Ex: Salmonella) components of Gr (-) bacterial cell wall effect the host when bacteria die and components are disintegrated cause muscle pain and fever Exotoxin (Ex. Cholera, botulism) it is secreted out by bacteria cause disease even in the absence of bacteria inactivated upon sterilization very small amounts can be a potent toxin Definition of autotrophs and heterotrophs • Autotrophs: Organisms that can produce their own food, using materials from inorganic sources. The word “autotroph” comes from the root words “auto” for “self” and “troph” for “food.” An autotroph is an organism that feeds itself, without the assistance of any other organisms. • Heterotrophs: Organisms that cannot manufacture their own food by carbon fixation and therefore derives their intake of nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are secondary and tertiary consumers. Prokaryotes can be grouped into four categories according to how they obtain energy and carbon Nutrition refers here to how an organism obtains two resources from the environment: energy and a carbon source to build the organic molecules of cells. •Photoautotrophs are photosynthetic organisms =light energy to drive the synthesis of organic compounds from carbon dioxide. (ex: cyanobacteria) •Chemoautotrophs need only CO2 as a carbon source, but instead of using light for energy, these prokaryotes obtain energy by oxidizing inorganic substances (hydrogen sulfide (H2S), ammonia (NH3), ferrous ions (FeS)), mode of nutrition is unique to certain prokaryotes. •Photoheterotrophs can use light to generate ATP but must obtain their carbon in organic form. •Chemoheterotrophs must consume organic molecules for both energy and carbon. This nutritional mode is found widely among prokaryotes, protists, fungi, animals. Mode of Nutrition Energy Source Carbon Source Source Types of Organisms Autotroph Photo-autotroph Light CO2 Photosynthetic organisms, including cyanobacteria; plants; certain protists (algae) Oxidizing Inorganic chemicals CO2 H2S, FeS, NH3 Certain prokaryotes (for example, Sulfolobus ) Photoheterotroph Light Certain prokaryotes Chemoheterotroph Organic compounds Organic compounds Chemoautotroph Heterotroph Organic compounds Saprobes (dead organism eaters), parasites (live within or on live host), biodegradants(petroleu m and some plastics), nitrogen fixers Bacterial Symbiosis Symbiosis means living together, thus require a symbiont and a host. Mutualistic; both side equally benefit. (Ex: nitrogen fixing bacteria) Commensalistic; one symbiont benefits and the other neither benefit nor damaged. (Ex: bacteria living in intestine, mouth or genitals) Parasitic; one symbiont benefits extensively while the other may even get harmed. (Ex: disease causing bacteria = pathogens) mutualistic commensalistic Clostridium difficile obtained from a human gut. Source: Wiggs, 2007. Parasitic Gene Transfer in Bacteria • transformation – alteration of a bacterial cell’s genotype by the uptake of naked, foreign DNA from the surrounding environment • transduction – virus carry bacterial genes from one host cell to another as a result of aberrations in the viral reproductive cycle • conjugation – direct transfer of genetic material from one bacterial cell to another TRANSFORMATION Heat shock treatment Conjugation ability to conjugate (form sex pili & donate DNA) is controlled by the presence of an F plasmid that contains a special piece of DNA called an F factor DNA donors (F+ cells) contain the F plasmid; DNA recipients (F− cells) lack the F plasmid only the F plasmid DNA is transferred the F+ cell can convert the F− cell to an F+ cell Video – Bacterial Conjugation • chromosomal genes can also be transferred when the F factor is integrated into the bacterium’s chromosome – cells with F factor built into the chromosome are called Hfr cells Video - Bacterial Conjugation Video – General Transduction Video – Specialized Transduction