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3/22/2016 LINES OF DESCENT FROM COMMON ANCESTRY EK 1.B.1: Organisms share many conserved core processes and features that evolved and are widely distributed among organisms today. EK 1.B.2: Phylogenetic trees and cladograms are graphical representations (models of evolutionary history that can be tested Illustrative examples are titled in blue Overview: Investigating the Tree of Life • Phylogeny is the evolutionary history of a species or group of related species • The discipline of systematics classifies organisms and determines their evolutionary relationships – Systematists • Use fossil, molecular, and genetic data to infer evolutionary relationships • Reconstruct phylogeny (evolutionary history) of a group of organisms – This history is illustrated in phylogenetic trees. – Phylogenetic trees identify likely relationships among species. Classification • A hierachical classification will group species into broader taxonomic categories. • Species that appear to be closely related are grouped into the same genus. – For example, the leopard, Panthera pardus, belongs to a genus that includes the African lion (Panthera leo) and the tiger (Panthera tigris). – Biology’s taxonomic scheme formalizes our tendency to group related objects. Binomial Nomenclature • Under the binomial system, each species is assigned a two-part Latinized name, a binomial. – The first part, the genus, is the closest group to which a species belongs. – The second part, refers to one species within each genus. – The first letter of the genus is capitalized and both names are italicized – For example, Linnaeus assigned to humans the scientific name Homo sapiens. Taxonomy • The identification & naming of species – Genera are grouped into progressively broader categories: family, order, class, phylum, kingdom & domain. • Each taxonomic level is more comprehensive than the previous. – As an example, all species of cats are mammals, but not all mammals are cats. • The named taxonomic unit at any level is called a taxon. – Example: Pinus is a taxon at the genus level, the generic name for various species of pine trees. – Mammalia, a taxon at the class level, includes all the many orders of mammals. 1 3/22/2016 Phylogenetic trees • Phylogenetic trees reflect the hierarchical classification of taxonomic groups nested within more inclusive groups. Building Trees • Analyzing the taxonomic distribution of homologies enables us to identify the sequence in which derived characters evolved during vertebrate phylogeny. • Two approaches to Phylogenetic Systematics – Phenetics • Determines taxonomic affinities on measurable similarities and differences. – Cladistics • classifies organisms according to the order in time that taxonomic groups diverge. Phylogeny • A phylogeny is determined by a variety of evidence including fossils, molecular data, anatomy, and other features. • Most systematists use cladistic analysis, developed by a German entomologist Willi Hennig to analyze the data. • A phylogenetic diagram or cladogram is constructed from a series of dichotomies. Cladistics • Primary criterion is common ancestry • Each branch or clade can be nested within larger clades • A clade consists of an ancestral species and all its descendants • Groups that do not fit this definition are unacceptable in cladistics (although further investigation may show their usefulness ) • Primary criterion is common ancestory • Clades-ancestral species with all of its descendents • Monophyletic, paraphyletic, and polyphyletic groupings Some Rules • Determining which similarities between species are relevant to grouping the species in a clade is a challenge. • It is especially important to distinguish similarities that are based on shared ancestry or homology from those that are based on convergent evolution or analogy. • In convergent evolution, natural selection shapes analogous structures based on ecological roles. • As a general rule, the more homologous parts that two species share, the more closely related they are. – Adaptation can obscure homology and convergence can create misleading analogies. • Also, the more complex two structures are, the less likely that they evolved independently. – For example, the skulls of a human and chimpanzee are composed not of a single bone, but a fusion of multiple bones that match almost perfectly. 2 3/22/2016 Figuring Stuff Out • Systematists must sort through homologous features or characters to separate shared derived characters from shared primitive characters. • A shared derived character is unique to a particular clade. Ex. Feathers • A shared primitive (ancestral) character is found not only in the clade being analyzed, but older clades too. Ex. Teeth • All species exhibit a mix of both types of characters and they provide information about evolutionary relationships. Homology and Analogy • Homology – likeness attributed to shared ancestry • Analogy – similarities due to convergent evolution, which is the acquisition of similar characteristics in species due to sharing similar ecological roles, NOT a common ancestor! Traits Used In Classification • Morphological Characteristics – Visible, measurable traits that distinguish groups of organisms from one another – Reflect genetic differences – Are preserved in the fossil record • Using Molecular Methods • The application of molecular methods and data for comparing species and tracing phylogenies has accelerated revision of taxonomic trees. – If homology reflects common ancestry, then comparing genes and proteins among organisms should provide insights into their evolutionary relationships – The more recently two species have branched from a common ancestor, the more similar their DNA and amino acid sequences should be • These data for many species are available via the internet Using DNA • Molecular systematics makes it possible to assess phylogenetic relationships that cannot be measured by comparative anatomy and other non-molecular methods. – This includes groups that are too closely related to have accumulated much morphological divergence. – At the other extreme, some groups (e.g., fungi, animals, and plants) have diverged so much that little morphological homology remains. • Most molecular systematics is based on a comparison of nucleotide sequences in DNA, or RNA. – Each nucleotide position along a stretch of DNA represents an inherited character as one of the four DNA bases: A (adenine), G (guanine), C (cytosine), and T (thymine). – Systematists may compare hundreds or thousands of adjacent nucleotide positions and among several DNA regions to assess the relationship between two species. – This DNA sequence analysis provides a quantitative tool for constructing cladograms with branch points defined by mutations in DNA sequence. 3 3/22/2016 Molecular Clocks • The timing of evolutionary events has rested primarily on the fossil record. • Recently, molecular clocks have been applied to place the origin of taxonomic groups in time. – Molecular clocks are based on the observation that some regions of genomes evolve at constant rates. – For these regions, the number of nucleotide and amino acid substitutions between two lineages is proportional to the time that has elapsed since they branched. What does the Tree of Life look like??? • No one REALLY knows…some scientists even predict a “ring of life” • One thing we can be sure of…. …there are a LOT more singe-celled organisms than multicellular. Relatedness of all Eukaryotes • • • • Cytoskeleton Membrane-bound organelles Linear chromosomes Endomembrane system How accurate are those molecular clocks? • Proportional differences in DNA sequences can be applied to access the relative chronology of branching in phylogeny, but adjustments for absolute time must be viewed with some caution. – No genes mark time with a precise tick-tock accuracy in the rate of base changes. – Genes that make good molecular clocks have fairly smooth average rates of change. – Over time there may be chance deviations above and below the average rate. • The molecular clock approach assumes that much of the change in DNA sequences is due to genetic drift (chance events) and is selectively neutral. Relatedness of Domains • DNA & RNA are carriers of genetic information through transcription, translation and replication. • Major features of the genetic code are shared by all modern living systems. • Metabolic pathways are conserved across all currently recognized domains. Cladograms & Speciation • Phylogenetic trees and cladograms illustrate speciation that has occurred, in that relatedness of any two groups on the tree is shown by how recently two groups had a common ancestor. 4 3/22/2016 Phylogenetics & Relatedness • Morphological similarities • DNA/protein sequence similarities Phylogenetics are dynamic • Use Biological data • Mathematical & computer models • Current & emerging knowledge 5