CHAPTER 22 Carl Linnaeus (1707-1778) – founder of taxonomy (scientific name) grouped similar species into same genus Georges Cuvier (1769-1832) – catastrophism – different species in layered rock due to catastrophic events like floods James Hutton (1726-1797) – gradualism – profound change is a cumulative product of slow but continuous process; ex. Rivers making canyons Charles Lyell (1797-1875) – uniformitarianism – geological process have not changed throughout Earth’s history Jean Baptiste Lamarck (17441829)- thought acquired characteristics can be passes on to offspring Worked on the HMS Beagle in 1830’s Observed and collected thousands of different species Galapagos Islands (west of S. America) most interesting “The Galápagos tortoise (or Galápagos giant tortoise), is the largest living tortoise, endemic to nine islands of the Galápagos archipelago. Adults of large subspecies can weigh over 300 kilograms (660lb) and measure 1.2 meters (4 ft) long. Although the maximum life expectancy of a wild tortoise is unknown, the average life expectancy is estimated to be 150-200 years.” Source: en.wikipedia.org/wiki/Gal%C3%A1pagos_tortoise Darwin read Lyell’s Principles of Geology and felt age of earth was much older than previously thought 1844 Darwin wrote essay on the origin of species 1858 – Alfred Wallace sends manuscript to Darwin about Natural Selection Lyell presented Wallace’s paper as well as Darwin’s 1844 essay to scientists 1859 The Origin of Species published by Darwin Descent with modification Natural selection (the mechanism) Individuals vary within a population Traits inherited All species are capable of overproduction Many offspring do not survive Individuals with inherited traits that give them a better chance of surviving and reproducing tend to leave more offspring than those without those traits Unequal ability of individuals to survive will lead to favorable traits in populations over generations. A population evolves, not an individual! Acquired characteristics may be adaptable but are not inherited!** The environment does not create a best fit characteristic, but selects for it! Goldendoodle and a liger AP:April. 29, 2005 ST. THOMAS, Barbados - It's male. But what is it? A zonkey? A deebra? That's the debate in Barbados since a zebra gave birth to a foal sired by a donkey. Biogeography – geographical distribution of species Ex. Islands with similar species to mainland Fossil record – transitional forms Comparative Anatomy – homologous structures among different organisms Vestigial organs –marginal, if any importance, remnants of structures that once served a function Whale pelvis and leg bones and human appendix Comparative Embryology – most vertebrates share common early development (gill slits) Molecular Biology – similar overall DNA, similar proteins (ex. Cytochrome c) Chapter 23 Genetic variation that makes evolution possible Mutations Change in DNA sequences Average 1 in 100,000 genes per generation Sexual reproduction Crossing over Independent assortment Fertilization Population – localized group of individuals belonging to the same species Species – organisms that can interbreed and produce fertile offspring Gene pool – total genes in a population If all members of a population are homozygous for the same allele, that allele is fixed. Gene frequency - two or more alleles for a gene, each having a relative frequency (proportion) in the gene pool Example: A = pink a = white 1000 plants = 200 white + 800 pink 800 pink = 340(AA) + 460(Aa) Find A’s frequency From AA: 340 x 2 = 680 From Aa: 460 x 1 = 460 680 + 460 = 1140 1140/2000 = .57 = .6 The 2000 is total number of alleles for 1000 plants Find a’s frequency From aa: 200 x 2 = 400 From Aa: 460 x 1 = 460 400 + 460 = 860 860/2000 = .43 = .4 Find A’s frequency 1- .4 = .6 Frequencies of alleles and genotypes in a population’s gene pool remain constant over the generations unless acted upon by agents other than random sexual recombination Hardy-Weinberg tells us what to expect if a population is NOT evolving!! Random Gamete production Probability of picking 2 A from example .6 x .6 = .36 Probability of picking 2 a from example .4 x .4 = .16 Probability of picking Aa from example (aA or Aa) (.4 x .6) + (.6 x .4 ) = .48 The sexual process of meiosis and random fertilization maintain the same allele and genotype frequencies over generations. In the example: p = .6 = A q = .4 = a p+q=1 p = 1 - q and q = 1 - p p2 + 2pq + q2 = (AA)+ (2Aa) + (aa) = 1 1 1. Very large population size 2. No gene flow (genes entering or leaving a population) 3. No net mutations 4. Random mating 5. No natural selection These mean NO EVOLUTION! Microevolution – generational change in a population’s frequencies of alleles or genotypes 1. Genetic Drift – rapid changes in a gene pool of a small, isolated population due to chance Flip coin 10 times: may get 7 heads and 3 tails Flip coin 1000 times: unlikely to get 700 head and 300 tails a. Bottleneck Effect – genetic makeup of a small surviving population is unlikely to be representative of original population Northern elephant seals nearly extinct due to hunting in late 1890’s which caused little genetic variation at 24 different loci b.The Founder Effect – occurs when a few individuals colonize a new habitat; the smaller the sample size, the less the genetic makeup of the colonists will represent the gene pool of the large population they left 2. Gene Flow – a population may gain or lose alleles through genetic exchange due to immigration and/or emigration Example: wind blowing pollen 3. Mutations- change in an organism’s DNA For any one gene locus, mutation does not have much effect on population unless mutation is a benefit and allows for more offspring Example: bacteria resistant to antibiotics Mutations – can only be passed on to offspring when they occur in cells that produce gametes On rare occasions, a mutant allele may be beneficial Ex. Houseflies resistant to DDT A single bacterium can make a billion cells in 10 hours so mutations can change whole populations quickly. 4. Nonrandom mating – individuals choosing mates Inbreeding – mating between closely related partners (extreme example is self-fertilization) Increases homozygous offspring, decreases heterozygous offspring Assortive mating – individuals select partners like themselves 5. Natural Selection – populations consist of varied individuals and some of these variants leave more offspring then others White flowers easily seen by herbivorous insects so more pink survive to make more pink plants Of all the agents of microevolution that change gene pools, only natural selection is likely to adapt a population to its environment. Neutral Variation – variation that appears to have no selective advantage Example: fingerprints Natural selection is the mechanism for adaptive evolution. Darwinian fitness – the relative contribution that an individual makes to the gene pool of the next generation Relative fitness – the contribution of a genotype to the next generation compared to other alternative genotypes for same locus The highest relative fitness a genotype can have is 1 If plants with white flowers average 80% as many offspring as pink flowered plants, their relative fitness is 0.8 Stabilizing selection– acts against both of the extreme phenotypes and favors intermediates Examples: human birth weight in 3-4Kg range horseshoe crab Directional selection– shifts towards one extreme phenotype (often during periods of environmental change) Examples: bacteria resistant to antibiotics light vs. dark moths in England Diversifying (disruptive) selection – when environmental conditions favors both extremes of a phenotypic range Examples: finch population with 2 bill sizes butterfly with 2 distinct morphs Sexual dimorphism – distinction between the secondary characteristics of male and females Sexual selection – selection process leading to sexual dimorphism Recessive alleles persist in environment due to heterozygotes The rarer a recessive allele, the greater degree of protection a hybrid offers (especially if recessive allele is harmful) Heterozygote advantage – when a heterozygote has greater survivorship and reproductive success than any homozygote Example: those who are carriers for sickle cell anemia are resistant to malaria Inbreeding can cause excessive homozygous conditions Frequency-dependent selection – the reproductive success of any one morph declines if that phenotypic form becomes too common Female African swallowtail butterflies mimic several noxious species This would be less effective if only one species was imitated. Right and left mouthed fish (cichlids) have different shaped mouths for approaching prey and eating scales Chapter 24 Macroevolution – the origin of new taxonomic groups Speciation – the origin of new species How does one species split into two????? Species - population(s) whose members interbreed in nature and produce fertile offspring The biological species concept is based on interfertility rather than physical similarity Barriers that prevent different species from interbreeding: Reproductive isolation – barriers that prevent two species from producing viable, fertile offspring 1. Prezygotic – impede mating between species by hindering the fertilization of ova 2. Postzygotic – impede mating between species by preventing the zygote from developing into a viable, fertile adult a. b. Habitat Isolation – living in different habitats within same area • Example: snakes in water vs. land Behavioral Isolation – special signals to attract mates (probably most important barrier) • Example: fireflies using different blinking signals c. Temporal Isolation – breeding during different seasons or years • d. Example: skunks mating in summer vs. late winter Mechanical Isolation – cannot mate due to anatomical differences • Example: flowers with different structures for different pollinators Gametic Isolation – gametes unable to fuse together to make zygote e. • Example: sperm not surviving vaginal environment a. Reduced Hybrid Viability – zygotes/embryos aborted (miscarriage) • b. • Example: frogs (Ranus) Reduced Hybrid Fertility – offspring end up being mostly sterile Example: horses mating with donkeys to make sterile mules Hybrid Breakdown – offspring are fertile, but next generation is sterile c. • Example: cotton Extinct organisms Asexual organisms Too rigid: dogs and coyotes Gene flow through subspecies Morphological – physical features Recognition – mating adaptations Cohesion –phenotype (genes and adaptations) Ecological – live and do (niches) Evolutionary – sequence of ancestral and descendant populations 1. 2. Allopatric – a geographic barrier isolates populations blocking gene flow Sympatric – intrinsic factors alter gene flow (like nonrandom mating) Geographical barriers – mountains forming, canyons forming, climate changing land Example: pupfishes (Cyprinodon) in springs of Death Valley CA and OR (drying caused separated “pools” in which speciation occurred) Peripheral isolate already different from original population (ex. phenotypic extremes) Genetic drift at work because smaller population size Different natural selection in new environment Mate choice in two species of Lake Victoria cichlids: females chose mates that have same color as themselves. Under monochromatic light, females chose both colors equally because they look the same. (Nonrandom mating causes sympatric speciation) Adaptive divergence – when 2 populations adapt to different environments, they accumulate differences in their gene pools Reproductive barriers may evolve coincidentally causing the populations to differentiate into 2 species Two populations get back together and interbreed = no new species Two populations get back together and do not interbreed = 2 new species Hybrid Zone = where 2 populations get back together and interbreed to make hybrids only around the region where they overlap Alleles specific to yellow-bellied toads decrease from 100% in areas where only they are found, to 50% in hybrid area, to almost 0% in fire-bellied area. Speciation can occur rapidly or slowly and can result from changes in few or many genes Punctuated equilibrium – describes periods of apparent stasis “punctuated” by sudden change CHAPTER 25 Life originated between 3.5 and 4 billion years ago Stromatolites – fossilized mats that contain banded domes of sedimentary rock (3.5 bya) and contain prokaryotes Oldest rocks are 3.9 billion (Greenland) To produce simple cells via chemical/physical processes and natural selection: 1. Abiotic synthesis of small organic molecules (ex. amino acids) 2. Joining of smaller molecules into macromolecules (ex. proteins) 3. Packaging macromolecules into protobionts where internal and external environments are different 4. Origin of self-replicating molecules that lead to inheritance A.I. Oparin and J.B.S. Haldane (1920’s) – early atmosphere of earth favored chemical reactions that could produce organic compounds Low oxygen = a reducing (electron adding) atmosphere Energy from lightning and higher UV radiation needed to make bonds Stanley Miller and Harold Urey (1953) – made amino acids from H20, CH4, NH3, H2 and electricity Now we wonder where this occurred – atmosphere or deep sea vents? Proteinoids – proteins formed from abiotic means (no enzymes) Need a substratum like hot sand, clay, or rock Vaporization would concentrate monomers on substratum Metals in substratum act as catalysts to bind monomers together Aggregates of abiotically produced molecules Not capable of precise reproduction, but maintain different internal conditions than external environment Liposomes – found to form spontaneously and are made of lipids Can have membrane potential and undergo osmotic pressure When a RNA strand is added to a solution of RNA nucleotides, small sequences can be made using strand as template and base pairing Thomas Cech (1980s) found ribozyme (enzyme that is not a protein) which catalyzes RNA synthesis RNA can make a variety of shapes due to different sequences This could lead to natural selection of certain shapes (sequences) Weak binding of amino acids to strand of RNA allows protein to be made (this happens today in rRNA/protein interactions) Packaging of RNA genes and their products within a membrane a great milestone! Laboratory experiments prove that life could have evolved in the “primordial soup”, but cannot prove that it did. First bacteria able to survive extreme heat so life could have evolved near deep sea vents and volcanoes Extraterrestrial source? Line between protobionts and live cells blurry Radiometric dating – absolute dating Half-life – the amount of time that it takes for 50% of the original sample to decay Carbon-14 has a half life of 5,730 years so its used for younger fossils Uranium-238 has a half-life is 4.5 billion years so its used for older fossils Not temperature sensitive Earth is approximately 4,600,000,000 years old Precambrian – (4.6 bya to 542 mya) Only bacteria for a billion years Towards end of era there were some eukaryotes which included algae and softbodied invertebrates (some multicellular) Gradual increase of oxygen caused by photosynthetic bacteria (2.7 – 2.2 bya) Oxygen revolution followed with great increase in O2. Why? Maybe chloroplasts??? Larger prokaryotes engulfed smaller prokaryotes (ancestor of mitochondria and chloroplasts) for origin of eukaryotes Evidence for endosymbiosis Inner membranes of both organelles have ETC like prokaryotes Both organelles replicate like binary fission, have ribosomes, and circular DNA like prokaryotes Many genes move to nucleus (transposons) Eukaryote genome “chimera” like – mixture of prokaryotic genes and cell parts Paleozoic – (542 – 245 mya) Cambrian explosion of animals Mostly marine life Colonization of land by plants and later animals First amphibians, reptiles and insects Shallow seas Extinction of marine life at end Pangaea formed at end Mesozoic – (245 – 65 mya) Flowering plants appear Pangaea breaks up Small mammals appear Extinction of dinosaurs as well as many other organisms (65mya) Cenozoic – (65 mya – present) Major radiation of mammals Humans appear 500,000 years ago Long periods of slow change punctuated by briefer intervals when turnover of species was extensive Mass extinctions Explosive adaptive radiation Survivors became adapted to vacant niches left by extinctions Examples: dozen or more in fossil record Two most studied Permian Extinctions (end of Paleozoic) Cretaceous Extinctions (end of Mesozoic) Claimed 90% of marine life Occurred in less than 5 million years Possible causes: Pangaea forming, Siberian volcanoes caused global warming, reduced temperature differences causing low O2 in oceans Claimed half of marine species and most dinosaurs Possible causes: Continental drift (volcanoes etc.) Asteroid hitting earth on the Yucatan coast of Mexico Chicxulub crater approximately 180 km in diameter Evidence supporting Chicxulub: Thin layer of iridium in rock layers (from ET debris) Dust cloud blocks sun and makes acid rain Extinction rates in N. America more severe and occurred faster Extinction rates not uniform across the globe Adaptive Radiation 4.6 bya – Origin of earth 4 bya – first prokaryotes Oxygen increases due to photosynthesis by cyanobacteria (2.7 billion) 2.1 bya - first eukaryotes 1.2 bya – first multicellular organisms Snowball earth – possible severe ice age that ended ~570 mya which allowed explosion of life 570 mya – oldest animal fossils 500 mya – plants and symbiotic fungi colonize land 66 mya – dinosaurs extinct 5 mya – apelike humans 500,000 years ago – first “humans” Animals more like fungi than plants Most of life on earth has been aquatic Evolution is NOT goal oriented! Evolution is like tinkering—it is a process in which new forms arise by the slight modification of existing forms Fig. 25-25 Recent (11,500 ya) Equus Pleistocene (1.8 mya) Hippidion and other genera Nannippus Pliohippus Pliocene (5.3 mya) Hipparion Neohipparion Sinohippus Megahippus Callippus Archaeohippus Miocene (23 mya) Merychippus Hypohippus Anchitherium Parahippus Miohippus Oligocene (33.9 mya) Mesohippus Paleotherium Epihippus Propalaeotherium Eocene (55.8 mya) Pachynolophus Orohippus Key Hyracotherium Grazers Browsers CHAPTER 26 Phylogeny – evolutionary history of a species or related species Incomplete record Minerals replace organic material Hard parts leave fossils Some tissues preserved Molds made Relative dating (older fossils in bottom layers of rock) Domain Kingdom Phylum Class Order Family Genus Species Scientific Name = Genus species Homo sapiens Any level is a taxon (pl. taxa) Example: phyla and order etc. Monophyletic – when a single common ancestor gave rise to all species within that taxon (ideal) Polyphyletic – members of a taxa are derived from 2 or more common ancestors Paraphyletic – when a taxon excludes species that share a common ancestor Homology – shared likeness due to common ancestry Analogy – shared likeness due to convergent evolution Convergent evolution – species from different evolutionary branches may come to resemble each other due to similar ecological roles and natural selection The ocotillo of southwestern N. America (left) looks like Alluaudia of Madagascar (right). Species diverge only when changes occur in nucleotide sequences Species that are phylogenetically closely related have more similar nucleotide sequences The number of differences in nucleotide bases between homologous sequences is a measure of evolutionary distance Clocks calibrated by graphing differences in sequences against known events in fossil record Assumes constant mutation rates Natural selection would alter mutation rates •Most widespread strain of HIV •Estimated to jump to humans in 1930’s •Based on DNA sequences from 1980 – 1990’s Use PCR DNA may be contaminated with bacterial DNA or other DNA Even with DNA cloned, cannot make dinosaurs until we understand the developmental steps involved Phenetics – based on measurable similarities and makes no phylogenetic assumptions Cladistics – classifies according to the order in time that branches arose along a dichotomous tree Clade – an evolutionary branch Outgroup – a species that is relatively closely related to the group of species being studied, but is clearly not as closely related as any study group members are to each other Synapomorphies – shared derived characteristics Characteristics that are homologous and evolved in an ancestor that is common to all species on one branch of a fork, but not common to other branch Parsimony – find the simplest explanation Cladistics accepts only monophyletic taxa Example: birds are more closely related to crocodiles than snakes and lizard are to crocodiles (birds and crocodiles have synapomorphies not present in snakes and lizards) Class Aves and Class Reptilia wrong cladistically because birds should be in same group as crocodiles On other hand both mammals and birds have 4chambered hearts and yet birds are more closely related to reptiles (not mammals) Four chambered heart evolved more than once