Processes of Evolution Chapter 12 12.1 Impacts/Issues Rise of the Super Rats When humans tried to eradicate rats with warfarin, natural selection favored individuals with a mutation for warfarin resistance Video: Rise of the super rats 12.2 Making Waves in the Gene Pool Individuals in a population share the same traits (phenotype) because they share the same genes (genotype) Gene pool • All of the genes in a population Alleles and Traits Alleles of the same genes are the main source of variation in a population • Traits with two distinct forms are dimorphic • Traits with several distinct forms are polymorphic • Traits with continuous variation may have interactions of several genes or be influence by environment Mutation is the source of new alleles Sources of Variation in Traits Phenotypic Variation in Humans Mutation Revisited Mutations are the original source of alleles, but many are lethal or neutral Lethal mutation • Mutation that drastically alters phenotype; usually causes death Neutral mutation • A mutation that has no effect on survival or reproduction Allele Frequencies Microevolution (change in allele frequencies) is always occurring in natural populations Microevolution • Small-scale change in allele frequencies of a population or species Allele frequency • Abundance of a particular allele among members of a population Genetic Equilibrium Genetic equilibrium • Theoretical state in which a population is not evolving (allele frequencies do not change) Only occurs if five conditions are met: • Mutations never occur, population is infinitely large, population is isolated from gene flow, mating is random, all individuals survive and reproduce equally Processes of Microevolution Genetic equilibrium does not occur in nature because processes that drive microevolution are always in play • • • • Mutation Natural selection Genetic drift Gene flow Animation: Adaptation to what? Animation: How to find out if a population is evolving 12.3 Natural Selection Revisited Natural selection occurs in different patterns depending on species and selection pressures • Directional selection • Stabilizing selection • Disruptive selection Directional Selection Directional selection • Mode of natural selection in which phenotypes at one end of a range of variation are favored • Allele frequencies shift in a consistent direction in response to selection pressure Examples: peppered moths, rock pocket mice, antibiotic-resistant bacteria Directional Selection Directional Selection in Peppered Moths Predation pressure favors moths that are best camouflaged when the environment changes Fig. 12-4a, p. 219 Fig. 12-4b, p. 219 Fig. 12-4c, p. 219 Fig. 12-4d, p. 219 Directional Selection in Rock Pocket Mice Mice with coat colors that do not match their surroundings are more easily seen by predators Stabilizing Selection Stabilizing selection • Mode of natural selection in which intermediate phenotypes are favored and extreme forms are eliminated Example: sociable weavers Stabilizing Selection Stabilizing Selection in Sociable Weavers Body weight in sociable weavers is a trade off between starvation and predation Disruptive Selection Disruptive selection • Mode of natural selection that favors extreme phenotypes in a range of variation • Intermediate forms are selected against Example: African seedcrackers Disruptive Selection Disruptive Selection in African Seedcrackers African seedcrackers tend to have either a large bill or a small one – but no sizes between 12.4 Factors That Affect Variation in Traits Interactions within species, and among genes and the environment, also influence natural selection • • • • Sexual selection Balanced polymorphism Genetic drift Gene flow Sexual Selection Sexual selection leads to forms of traits that enhance reproductive success Sexual dimorphism is one outcome Sexual selection • Some individuals of a population outreproduce others because they are better at securing mates Examples of Sexual Selection Balanced Polymorphism In balanced polymorphism, nonidentical alleles for a trait are maintained in a population Balanced polymorphism • Maintenance of two or more alleles for a trait in some populations • Occurs when environmental conditions favor heterozygotes over homozygotes Balanced Polymorphism and Sickle Cell Trait Genetic Drift Genetic drift can lead to the loss of genetic diversity (fixation) Genetic drift • Random change in allele frequencies in a population over time, due to chance alone Fixed • Refers to an allele for which all members of a population are homozygous Genetic Drift in Flour Beetles Genetic Drift and Bottlenecks Genetic drift is pronounced in small or inbreeding populations, such as those that occur after an evolutionary bottleneck Bottleneck • Reduction in population size so severe that it reduces genetic diversity Inbreeding • Nonrandom mating among close relatives Genetic Drift and the Founder Effect A bottleneck can lead to the founder effect Founder effect • Change in allele frequencies that occurs after a small number of individuals establish a population The Founder Effect in Old Order Amish Populations of Old Order Amish in Pennsylvania are moderately inbred – 1 in 200 is homozygous for the recessive allele that causes Ellis vanCreveld syndrome Gene Flow Gene flow counters the effects of mutation, natural selection, and genetic drift in a population Gene flow • The physical movement of alleles into and out of a population, as by individuals that immigrate or emigrate 12.5 Speciation Individuals of sexually reproducing species can interbreed successfully under natural conditions, produce fertile offspring, and are reproductively isolated from other species Speciation • Process by which new species arise from existing species Four Butterflies, Two Species Reproductive Isolation Reproductive isolation typically occurs after gene flow stops Divergences then lead to speciation Reproductive isolation • Absence of gene flow between populations Different species form and . . . Prezygotic mechanisms Individuals reproduce at different times (temporal isolation). Physical incompatibilities prevent individuals from interbreeding (mechanical isolation). Reproductive Isolating Mechanisms Individuals live in different places so they never meet up (ecological isolation). Individuals ignore or do not get the required cues for sex (behavioral isolation). Mating occurs and . . . No fertilization occurs (gamete incompatibility). Zygotes form and . . . Postzygotic mechanisms Hybrid embryos die early, or new individuals die before they can reproduce (hybrid inviability). Hybrid individuals or their offspring do not make functional gametes (hybrid sterility). Interbreeding is successful Fig. 12-15, p. 227 Mechanical Isolation Animation: Albatross courtship Allopatric Speciation In allopatric speciation, a geographic barrier interrupts gene flow between populations Genetic divergences then give rise to new species Allopatric speciation • Speciation pattern in which a physical barrier that separates members of a population ends gene flow between them Allopatric Speciation in Snapping Shrimp Sympatric Speciation With sympatric speciation, populations in physical contact speciate Polyploid species of many plants (and a few animals) originated by chromosome doublings and hybridizations Sympatric speciation • Pattern in which speciation occurs in the absence of a physical barrier Sympatric Speciation in Wheat 2 Triticum monococcum (einkorn) 14AA 1 Unknown spontaneous species of chromosome T. turgidum (emmer) doubling Triticum × 14BB 14AB 28AABB 3 × T. tauschii (goatgrass) 14DD T. aestivum (common bread wheat) 42AABBDD Fig. 12-19, p. 229 Animation: Sympatric speciation in wheat Sympatric Speciation in Cichlids Different Speciation Models Animation: Models of speciation Animation: Temporal isolation among cicadas Stasis With stasis, a lineage changes very little over evolutionary time Stasis • Macroevolutionary pattern in which a lineage persists with little or no change over evolutionary time Stasis: The Coelacanth Coevolution Coevolution occurs when two species act as agents of selection upon one another Coevolution • The joint evolution of two closely interacting species; each species is a selective agent that shifts the range of variation in the other • Examples: predator and prey, host and parasite, pollinator and flower Coevolved Species Madagascar orchid and its pollinator Extinction Permanent loss of a species is extinction Extinct • A species that has been permanently lost Mass extinction • Simultaneous extinction of many lineages Adaptive Radiation An adaptive radiation is a rapid diversification into new species that occupy novel niches Adaptive radiation • A burst of genetic divergences from a lineage gives rise to many new species Key innovation • An evolutionary adaptation that gives its bearer the opportunity to exploit a particular environment more efficiently or in a new way Adaptive Radiation of Hawaiian Honeycreepers Fig. 12-23 (right), p. 232 Fig. 12-23a, p. 232 Fig. 12-23b, p. 232 Fig. 12-23c, p. 232 Fig. 12-23d, p. 232 Fig. 12-23e, p. 232 Fig. 12-23f, p. 232 Fig. 12-23g, p. 232 Fig. 12-23h, p. 232 Fig. 12-23i, p. 232 Fig. 12-23j, p. 232 Fig. 12-23k, p. 232 Evolutionary Theory Many biologists disagree about how macroevolution occurs Dramatic jumps in morphology may result from mutations in homeotic or other regulatory genes Macroevolution may be an accumulation of many microevolutionary events, or an entirely different process Animation: Allopatric speciation on an archipelago 12.7 Organizing Information About Species Taxonomy • Science of naming and classifying species • In traditional taxonomy, species are organized into a series of ranks (taxa) based on their traits • Such systems do not necessarily reflect evolutionary relationships Taxon (taxa) • A grouping of organisms Linnaean Classification System Ranking Versus Grouping Cladistics is a set of methods that allow us to reconstruct evolutionary history (phylogeny) Phylogeny • Evolutionary history of a species or group of species Cladistics • Method of determining evolutionary relationships by grouping species into clades Cladistics Cladistics groups species into clades on the basis of shared characters Character • Quantifiable, heritable characteristic or trait Clade • A group of species that share a set of characters Cladistic Analysis The result of a cladistic analysis is an evolutionary tree diagram in which a line represents a lineage Evolutionary tree • Type of diagram that summarizes evolutionary relationships among a group of species Cladograms In evolutionary trees called cladograms, a line (lineage) can branch into two sister groups at a node, which represents a shared ancestor Cladogram • Evolutionary tree diagram that shows a network of evolutionary relationships among clades Sister groups • Two lineages that emerge from a node on a cladogram Clades Every branch of a cladogram ends in a clade Ideally, each clade is a monophyletic group Monophyletic group • An ancestor and all of its descendants hagfishes animals with a skull lampreys cartilaginous fishes ray-finned fishes lobe-finned fishes lungfishes amphibians amniotes (reptiles, birds, and mammals) animals with a backbone and a skull animals with a swim bladder or lungs, a backbone, and a skull animals with four limbs,* a swim bladder or lungs, a backbone, and a skull animals with four membranes around their eggs, four limbs,* a swim bladder or lungs, a backbone, and a skull *Snakes are included in these clades because their ancestors had four legs. Fig. 12-25b, p. 234 Classification Systems Evolutionary trees are revised as new information is gathered Two different ways to organize life’s diversity • Six-kingdom classification system • Three-domain classification system Two Major Classification Systems Animation: Current evolutionary tree