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Chapter 22 Descent With Modification Classification of Species, a history • Aristotle (384-322B.C.E.) – Species fixed, unchanging, eternal world, absolute – Scala natura • Life forms arranged in linear order of perfection • Carolus Linnaeus (1707 - 1778) – Binomial nomenclature • Homo sapiens Carl von Linné – Nested classification based on similarities • Birds/bats • Systema Natura book Fossils • Remains or traces of organisms from the past • >10,000 years old • Organism dies sediment compression rock strata • Paleontology = study of fossils Fossils buried in strata • Charles Lyell, geologist (1797- 1875) – Uniformitarianism, Earth changes slowly and continually – Change NOT due mainly to catastrophes – Examples of slow processes – Wind, erosion, water flow • Lamarck (1744-1829) – Evolution explains change in fossils over time • Theory of use and disuse • Theory of inheritance of acquired characteristics – Both not correct This is a big tree dwarfed by many years of cutting What size plant will seeds from this Bonsai produce? Charles Darwin (1809-1882) Descent with modification (evolution) Alfred Russel Wallace came to same conclusion just before Darwin published The Origin of Species Voyage of the Beagle Observed plants, animals Darwin’s observations and inferences • 1. Adaptations – Characteristic of organisms enhance survival – Adaptations could lead to new species • Galapagos finches Each on a different island • 2. organisms descend from a common ancestor – Unity of life – Can be represented as a tree – Millions of years Species grouped by evolutionary relatedness! Mechanism of Natural Selection A. Members of a population vary in traits Genetic diversity B. Traits are inherited from parent to offspring C. More offspring produced than environment can support D. Those with beneficial traits survive AND reproduce successfully (survival of the fittest) E. Populations change over time F. Can lead to speciation Note: • Individuals do not evolve, populations do • Only heritable traits are passed on • Environmental factors vary! Evidence for Evolution I. FOSSILS document change in living organisms over time Example Whales have a terrestrial , 4 legged, ancestor Bones suggest a cow- like animal 50 mya II. HOMOLOGOUS STRUCTURES Reflect a common ancestry Shared features 1. Skeletal structures common mammalian ancestor Function/name? Homologous structures (cont.) 2. embryology- shared developmental plan Homologous structures (cont.) 3. Vestigial structures served function in ancestor, function lost, but structure remains Snakes retain vestige of pelvis Ancestor to snakes had legs Homology (cont.) 4. Genetic homology universal genetic code supports evidence for relatedness of species Beta globin amino acid sequence DEVGGEALGRLLVVYPWTQRYFDSFGDLSSASAI Mus musculus DEVGGEALGRLLVVYPWTQRYFDSFGDLSSASAI Rattus norvegicus DEVGGEALGRLLVVYPWTQRFFESFGDLSTADAV Bos taurus • Convergent evolution – Sometimes distantly related species have similar structures – Evolved separately (arose more than once) – Example: mammals marsupial Rodent (placental) III. BIOGEOGRAPHY Geographic distribution of species Affected by continental drift slow movement of continents over time 250 mya Pangaea As continents move, species disperse, environmental changes Example: endemic species found in one location BUT, have relatives nearby Island species colonized by mainland species IV. Direct documentation of evolution Peacock sexual selection http://www.youtube.com/watch?v=T7ftXEQvnyU Direct documentation – Predators influence natural selection – Prey adaptations to avoid predation • Male guppies • Color genes expressed in males only • Predators attracted to color • Sexual selection – Females attracted to colors Observations Pool 1 with few predators: brightly colored males Pool 2 with many predators drab males Add brightly colored males to pool 2 over 15 generations, males more drab Why? Rosemary and Peter Grant, Princeton University • In 1981 an unusually large male finch from Santa Cruz arrived on Daphne Major. • The biologists tagged the bird number 5110, and followed him and his offspring through seven generations total. • In the fourth generation drought killed off all descendants except one male and one female. • These offspring can only breed with each other because they have strange bird song. Finches learn their songs from their father, and the Grants suggest that 5110 sang the songs from his birth home of Santa Cruz then modified his mating song by roughly copying the Daphne Major birds’. • This imperfect copying has, over time acted as a barrier to breeding with other finches . So the immigrant bird’s descendants have bred only with each other for three generations. • New species? The Grants won the Kyoto prize for their life work Theory • Unifies many observations • Supported by all evidence • Predictions stand up Chapter 23 The Evolution of Populations 0rganisms do not evolve • Microevolution – Change in allele frequencies in population over time – Affected by • Natural selection • Genetic drift • Gene flow 23.1 Genetic Variation A. Genetic Variation: Variation within a population Discrete characters one gene Quantitative characters vary along continuum two or more genes • Gene variability in a population average heterozygosity f (Aa) information about genetics of a population • High heterozygosity = high genetic variability • Can range from 0 to 1 Fruit fly has ~ 13, 700 genes On average 1,920 are heterozygous (14%) • Very low heterozygosity in cheetahs indicate severe effects of small population sizes Originated about 4 million years ago and was common in Europe, North America, Asia , Africa. They disappeared about 10,000 years ago from North America when major climatic changes took place. Also disappeared from Europe and most in Asia and Africa vanished. Present populations derived from inbreeding by very few surviving and closely related animals. Every one is nearly identical. B. Genetic Variation: Variation between Populations • Geographic variation, ex. Mus musculus Islands of Madeira – mice brought 15th century several populations have evolved in isolation Portugal Populations descended from single ancestral population, changes in chromosomes (not genes) Geographic cline Graded change along geographical axis Mummichog fish cline in cold adaptive allele Mutation Ultimate source of new alleles Relevant only if in sperm/egg Mutation: Point mutations one base pair change in DNA possible effects: -none – does not change amino acid sequence of protein -harmful – protein function compromised -beneficial - rare Mutations that alter gene number or sequence • Gene duplication – May expand the genome with no negative effects – Mammal ancestor had 1 odor gene duplication • Humans have 1,000 olfactory receptor genes – 60% inactivated in later mutations • Mice have 1,300 Jianzhi Zhang and colleagues (Univ. Michigan) • Paper (handout) • Homework – article analysis Sexual reproduction • Leads to genetic diversity – .crossing over during meiosis – . Random segregation of alleles into gametes – . Independent assortment – (and mutation) Hardy Weinberg Principle • A population that is NOT evolving : (hypothetical population) – No mutations – Random mating – No natural selection – Extremely large population – No gene flow • Change in any one of these change in allele frequencies evolution 23.3 Allele Frequencies (cont.) • Affected by: 1. Natural Selection • Differential success in survival and reproduction • Allele frequencies different in each generation • Adaptive evolution – Some alleles favored over others – These increase in frequency with time Adaptation (at population level) DDT is a pesticide used beginning 1940s banned in 1972 • Example: Drosophila • 1930s wild strain – no DDT resistance gene – zero tolerance to DDT Adaptive evolution Figure: increase in frequency of pesticide resistance in mosquitos after spraying with DDT. A sample of mosquitos was captured at each time indicated and the number that were killed by a standard dose of DDT (4 % DDT for 1 hour) in the laboratory was measured. From Curtis et al. (1978). 2. Genetic Drift • Chance events can cause allele frequencies to vary The founder effect A. A few individuals become isolated from population B. The new population has different allele frequencies than the main population • Founder effect examples: • Few individuals wind up on island during storm • A few individuals are cut off by an earthquake Founder effect Bottleneck effect – Sudden drastic population reduction – Lessens genetic diversity – Example: greater prairie chicken • 1800s – millions in Illinois • 1993 - 50 birds in 2 populations – Low genetic diversity – 50% of eggs hatch Genetic Drift, summary • Significant factor in small populations • Allele frequencies can change wildly with generations • Genetic variation is lost • Harmful alleles may become fixed in population 23.4 Natural Selection is a mechanism that causes adaptive evolution • A blend of chance and sorting as natural selection favors certain alleles. • Relative fitness – Relative contribution individual makes to gene pool of next generation • Note: the organism is subject to natural selection, populations evolve • Directional selection – extreme phenotype most fit – f(alleles) in population shifts in a direction • Example: deer mice – Dark favored, live among dark rocks – Why is this a beneficial trait? Disruptive selection – Individuals of both extremes most fit • Some mice colonized white rocks, some dark • What happened to the intermediate color mice? • Stabilizing selection – Intermediate phenotypes favored Example: birth weight in humans Examples of adaptations • Cuttlefish • Adaptation? Snake adaptation • Sexual selection – Individuals with certain inherited characteristics more likely to obtain mates • Can lead to sexual dimorphism – Differences between males and females • Intrasexual selection – Individuals of same sex compete for mate • Intersexual selection – Mate choice – Showy tail in peacocks advertises fitness to female – Showiness may be risky • Ex. guppies Preservation of genetic variation • Diploidy – Allows recessive alleles to be maintained in population in heterozygotes – Maintains genetic diversity The heterozygote advantage – Sometimes a heterozygote has greater fitness! – Beta globin gene encodes subunit of hemoglobin Genotype Phenotype SS = Ss = ss = Heterozygote advantage • Frequency of s allele highest in areas with malaria Why natural selection cannot fashion perfect organisms 1. Selection can only act on existing variation 2. Evolution builds on what is present in organism already Ex. terrestrial lineage of whales, bats 3. Adapations are compromises Ex. shoulder joint (human) 4. Evolution based on interaction between chance, environment Ex. wind storm kills many insects, including most fit Chapter 24 Origin of Species Speciation • Process by which one species splits into two or more • Unity of life – species evolutionarily related • Microevolution – Change in allele frequencies in populations • Macroevolution – Change over geological time speciation Biological species concept • Species is a group whose members can interbreed (and produce fertile offspring) • Share a gene pool • Reproductive isolation • Pre-zygotic barrier – Sperm from one cannot fertilize egg from another – Why? • Habitat isolation, temporal isolation, behavioral isolation, mechanical isolation Different sperm proteins Mates late winter Mates late summer Misalignment of genital openings • Post-zygotic barrier – Hybrids are infertile – Example: mule May not survive Mechanisms of speciation • Allopatric speciation – A population is divided geographically – Each group continues to evolve, but separately salamanders • Natural selection – Selects for different traits depending on environment • Genetic drift • Speciation occurs - When populations are reintroduced – do not interbreed (reproductive isolation) Sympatric speciation • Gene flow reduced between members of same population/same geographical area • Ex. Polyploidy – Extra set(s) of chromosomes – Not uncommon in plants Sympatric speciation (cont.) • Can be driven by habitat – North American maggot fly lived in hawthorn tree, then, some colonized apple trees. These flies developed faster, now, temporally isolated (mating on apple tree flies is 3 wks earlier) Rhagoletis pomonella. Hybrid zones Both frogs found in the hybrid zone Hybrids have high embryonic mortality and anatomical abnormalities • Hybrid zone outcomes: 1. Reproductive barriers strengthened • Limits hybridization 2. Barriers weakened • Species fuse into single species 3. Stable hybrid zone Genetics of speciation • Identify genes that control traits involved in reproductive isolation • Ex. Japanese snail, Euhadra – Single gene controls shell spiral direction Chapter 25 • History of Life on Earth The Fossil Record • Record of life on earth over time • Incomplete record/limited • Radiometric dating of strata – Decay of radioactive isotopes – Half life = time for 50% of isotope to decay C-14 half life = 5,730 years C-14 C-14 dating not effective > ~75,000 years Measure C-12/C-14 ratio • For older fossils in rocks – Use K-40 (1.3 billion year half life) Key Events in Life’s History Geological eras of life 1. Paleozoic 2. Metazoic 3. Cenozoic The first single celled organisms 3.5 bya Stromatolites Microbes bind thin layers of sediment Large amount of biomass/communities Algal biscuits fossils May have contained cyanobacteria, algae, eukaryotic cells…… Oscillatoria • The first eukaryotes 2.1 bya (older?) – Single celled – Nucleus • Multicellular animals > 535 mya – Soft bodied, small – <1 meter The Cambrian explosion • 535 mya • Cnidaria, Porifera, Mollusca fossils Cambrian explosion lasted ~40 my • Larger animals • Hard shells • Predator/prey evolution 500 mya trilobite The colonization of land by animals • 500 mya • Adaptations to prevent dehydration • Tetrapods ~ 365 mya Mass Extinctions (5) • >50% of animals extinct in “short” period of time 1. Permian extinction – 250 mya – 95% of marine species extinct – Most terrestrial species Cause: Oxygen deficit? – Neil Tyson DeGrasse 2. Cretaceous extinction 65 mya Most marine and terrestrial species including dinosaurs Cause: asteroid hot vapor, debris blocked sunlight? 10 km Consequences of mass extinctions • Once a species is extinct, it does not return! Adaptive radiation • Periods of evolutionary change in which new species adapt to fill different ecological roles • Occurred after each mass extinction • Example: Cretaceous mass extinction 65mya – Mammal radiation Evolution is not goal oriented Chapter 52 Ecology and the Biosphere Ecology • Study of the interactions between organisms (biotic factors) and the environment (abiotic) • Environmentalism – Advocating for the protection of nature • Red kangaroo (Macropus rufus) – Abundant in interior Australia – dry climate – Not found along coast – wet – Hypothesis? Abiotic factor? Biotic? • Moisture, predator, temperature, food, competition, disease Rachel Carson • “ The ‘control of nature’ is a phrase conceived in arrogance, born of the Neanderthal age of biology and philosophy, when is was supposed that nature exists for the convenience of man” • Efforts led to a ban on DDT • Silent Spring (1962) Factors affecting species distribution 1. Dispersal and distribution A. Range expansion Movement of individuals away from area of origin B. Transplants – Intentional or accidental Kudzu, a Japanese invasive vine growing in Georgia, US Brown tree snake (from Australia)has eaten all Guam Flycatcher, Rufus Fantail, Bridled WhiteEye and Micronesian Honeyeater eggs, now eating the Mariana Crow, the Guam Micronesian Kingfisher, the Island Swiftlet and the Mariana Fruit Bat. • 2. Habitat selection and behavior A. Behavior An individual may avoid a certain habitat even though it is suitable Corn borer moth prefers to lay eggs on corn even though larvae can feed on other plants B. Biotic factors that affect species distribution – Parasitism, competition, pollinators, predators, food source… • C. Abiotic factors – Temperature, water, salinity, sunlight, soil – Vary spatially and temporally • Temperature – Cells rupture <OoC – Proteins denature>45oC – Metabolic rate may be affected by T • Water – Dehydration • Salinity – Osmosis – Example: freshwater fish gains water -it is adapted to excrete water. • Sunlight – Photosynthesis • Rocks, soil – pH, minerals, substrate Chapter 54 Community Ecology 1. Competition • Interspecific competition – Individuals of different species compete for a resource – Limits growth and survival • Canadian lynx and fox compete for snowshoe hare Outcomes of competition • Competitive exclusion – Paramecium experiment (G.F. Gause, 1934) P. aurelia + P. caudatum Competition for food source P. caudatum dies Even a slight advantage will lead to disappearance of less fit species P. aurelia P. caudatum • Niche development – Niche = species use of both abiotic and biotic resources (habitat is “address” niche is “profession”) – Example: Anolis lizards in DR Niche (continued) 2 similar species can coexist in same community if niches are different Resource partitioning in Anolis lizards A. distichus prefers sunny twigs A. insolitus prefers shady branches Hypothesis: competition in the past selected for individuals using different sets of resources The fundamental niche of distichus may be sun and shade, but its realized niche is sun – how could this be tested? Anolis lizards, resource partitioning Fundamental and realized niches in barnacle species (Joseph Connell, Scotland, 1961) Balanus cannot survive on high rocks, it dries out during low tide – compare fundamental and realized nice • Character displacement – Observed in sympatric populations (overlapping ranges) – Divergence of body structure and resource use Twig anole Bahamas and Cuba Trunk anole Trunk ground anole 2. Predation • Predator adaptations smell, sight, claws, stingers, teeth, aggressive mimicry • Prey adaptations hiding, herds, alarm calls, cryptic coloration aposematic coloration, Batesian mimicry, Mullerian mimicry Zebra Vervet monkey Aposematic and cryptic coloration Golden poison frog A warning coloration to indicate a toxic defense (alkaloid toxin –batrachotoxin in skin - one frog could kill 20 humans) • Batesian mimicry – A harmless species mimics a harmful one Hereroplanes ornatus (hawk moth larvae) • Mullerian mimicry – Two or more unpalatable species resemble each other – Each species gains additional protection Both untasty • Aggressive mimicry – Predator shares feature with harmless model to attract prey “flower“ lantern tongue “worm” 3. Herbivory • Interaction in which organism eats plant or algae • Adaptations – smell, teeth, digestion • Plant defenses – Chemicals – peppermint, strychnine, nicotine – Spines, thorns 4.Symbiosis • Two that live in direct contact • May be harmful, neutral, or helpful • Parasitism – One organism (parasite) derives nourishment from another (host) – Endoparasite lives within host – Ectoparasite – lives on host plant root • Parasites affect: – – – – Survival Reproduction Population density Behavior • • Lake Superior Island 70,000 ticks per moose hair damage, blood loss increase chance of cold stress, wolf predation • Mutualism – Benefit both species in a relationship. Ex. acacia and ant Central America, ant hollows out large thorns of the acacia plant for nests, feed on secretions from four nectaries at the base of each petiole and on the protein rich tips of the leaves, which together provide an almost complete diet for the ant. The ants protect trees from invertebrate as well as vertebrate herbivores. With movement of the branch, the ants emerge releasing a nasty odor and physically attack the herbivore. • Commensalism – One species benefits, the other is not, but is not harmed Cattle Egret often seen in the company of grazing animals. The grazers stir up insects, which the egret eats. There is no apparent benefit to the cattle. The birds will also follow mowers…… The clownfish slowly accustoms the anemone to the chemical makeup of the fish's skin; this gradual acclimatization prevents the anemone from stinging the clownfish. The fish gets a safe habitate and some food; the anemone gets cleaned clownfish chasing away fish that would harm the anemone, may attract prey. Some scientists do not see any benefit for the anemone and classify this as a commensalism. Each sloth can host 100 moths and 1000 beetles. The entire life cycle of beetle and moth is connected with the sloth, each spending its larval period in sloth dung, and their adult period eating the algae. ? Costa Rica. Tree sloth has algae growing in its fur. Algae help to camouflage the sloth against the lichen-covered tree (note brown fur of baby, not covered with algae). Xenarthra A moth lives only in the sloth's fur and consumes the algae. What is the relationship between the moth and the sloth? Effect of Interaction On: Type of Interaction Species 1 Species 2 (-) (-) Predation Predator ( ) Prey ( ) Parasitism Parasite ( ) Host ( ) Herbivory Herbivore ( ) Plant ( ) Commensalism ( ) ( ) Mutualism ( ) ( ) Competition Species in communities • Species richness = the number of different species in a community • Relative abundance = number of each Community 1: 25A, 25B, 25C, 25D = 4 species of same relative abundance Community 2: 80A, 5B, 5C, 10 D……..? • Trophic structure – Feeding relationships between organisms Food chain Primary producers primary consumers secondary consumers decomposers Food web Linking food chains • Food web links food chains • A species may be in web at different trophic levels Species with a large impact • Dominant species – Most numerous – More biomass In temperate bogs the dominant vegetation is usually species of Sphagnum moss Foundation species primary producer Kelp in kelp forest , CA • Keystone species Beaver are well-known for building dams. These dams create relatively large areas of still water where there once was a small stream. For organisms that live in such still waters the beaver is a wonderful keystone mutualist; for animals that like flowing water it's not such a good deal. • The American Alligator, left, excavates depressions in its habitat that fill with water. During dry times, these gator holes may be the only places with water. Thus, to all the organisms whose survival depends on the water in those holes the alligator is a keystone mutualist. Of course, the gator might eat a few of those things that come to live in its wallow. Community Ecology is useful for understanding pathogen life cycles and controlling disease • Pathogens can alter community interactions • Coral reef communities – White band disease kills corals • Zoonotic disease – Transferred from animals to humans – Direct contact or vector – Parasite life cycle – River blindness and black flies – Avian flu tracking