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Bellringer: Grade FRQ’s Grade the writing examples (Scale 0-3 pts) Discuss with shoulder partner Now grade with a rubric Discuss with shoulder partnerThis should give you ideas on how to writefor AP Biology TestsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Chapter 23 NotesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 23-13Original populationOriginalEvolvedpopulation population(a) Directional selectionPhenotypes (fur color)(b) Disruptive selectionCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings(c) Stabilizingselection

The Key Role of Natural Selection in AdaptiveEvolution Natural selection increases the frequencies ofalleles that enhance survival and reproduction Adaptive evolution occurs as the matchbetween an organism and its environmentincreasesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 23-14(a) Color-changing ability in cuttlefishMovable bones(b) Movable jawbones insnakesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Because the environment can change,adaptive evolution is a continuous process Genetic drift and gene flow do not consistentlylead to adaptive evolution as they can increaseor decrease the match between an organismand its environmentCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Sexual Selection Sexual selection is natural selection formating success It can result in sexual dimorphism, markeddifferences between the sexes in secondarysexual characteristicsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 23-15Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Intrasexual selection is competition amongindividuals of one sex (often males) for matesof the opposite sex Intersexual selection, often called matechoice, occurs when individuals of one sex(usually females) are choosy in selecting theirmates Male showiness due to mate choice canincrease a male’s chances of attracting afemale, while decreasing his chances ofsurvivalCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

How do female preferences evolve? The good genes hypothesis suggests that if atrait is related to male health, both the maletrait and female preference for that trait shouldbe selected forCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 23-16EXPERIMENTFemale graytree frogSC male graytree frogLC male graytree frogSC sperm Eggs LC spermOffspring of Offspring ofSC fatherLC fatherFitness of these half-sibling offspring comparedRESULTSFitness Measure19951996Larval growthNSDLC betterLarval survivalLC betterNSDTime to metamorphosisLC better(shorter)LC better(shorter)NSD no significant difference; LC better offspring of LC malessuperior to offspring of SC males.Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Preservation of Genetic Variation Various mechanisms help to preserve geneticvariation in a populationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Diploidy Diploidy maintains genetic variation in the formof hidden recessive allelesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Balancing Selection Balancing selection occurs when naturalselection maintains stable frequencies of two ormore phenotypic forms in a populationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Heterozygote Advantage Heterozygote advantage occurs whenheterozygotes have a higher fitness than doboth homozygotes Natural selection will tend to maintain two ormore alleles at that locus The sickle-cell allele causes mutations inhemoglobin but also confers malaria resistanceCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 23-17Frequencies of thesickle-cell allele0–2.5%Distribution ofmalaria caused byPlasmodium falciparum(a parasitic unicellular eukaryote)Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin % 12.5%

Frequency-Dependent Selection In frequency-dependent selection, thefitness of a phenotype declines if it becomestoo common in the population Selection can favor whichever phenotype isless common in a populationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 981 ’82 ’83 ’84 ’85 ’86 ’87 ’88 ’89 ’90Sample yearCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Neutral Variation Neutral variation is genetic variation thatappears to confer no selective advantage ordisadvantage For example,– Variation in noncoding regions of DNA– Variation in proteins that have little effect onprotein function or reproductive fitnessCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Why Natural Selection Cannot Fashion PerfectOrganisms1. Selection can act only on existing variations2. Evolution is limited by historical constraints3. Adaptations are often compromises4. Chance, natural selection, and theenvironment interactCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

You should now be able to:1. Explain why the majority of point mutationsare harmless2. Explain how sexual recombination generatesgenetic variability3. Define the terms population, species, genepool, relative fitness, and neutral variation4. List the five conditions of Hardy-WeinbergequilibriumCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

5. Apply the Hardy-Weinberg equation to apopulation genetics problem6. Explain why natural selection is the onlymechanism that consistently producesadaptive change7. Explain the role of population size in geneticdriftCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

8. Distinguish among the following sets of terms:directional, disruptive, and stabilizingselection; intrasexual and intersexualselection9. List four reasons why natural selection cannotproduce perfect organismsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Chapter 24The Origin of SpeciesPowerPoint Lecture Presentations forBiologyEighth EditionNeil Campbell and Jane ReeceLectures by Chris Romero, updated by Erin Barley with contributions from Joan SharpCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

What you need to know The difference between microevolution andmacroevolution. The biological concept of species. Prezygotic and postzygotic barriers that maintainreproductive isolation in natural populations. How allopatric and sympatric speciation are similarand different. How an autopolyploid or an allopolypoloidchromsomal change can lead to sympatric speciation. How punctuated equilibrium and gradualism describe2 different tempos of speciation.Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Overview: That “Mystery of Mysteries” In the Galápagos Islands Darwin discoveredplants and animals found nowhere else onEarthVideo: Galápagos TortoiseCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Speciation, the origin of new species, is at thefocal point of evolutionary theory Evolutionary theory must explain how newspecies originate and how populations evolve Microevolution consists of adaptations thatevolve within a population, confined to onegene pool Macroevolution refers to evolutionary changeabove the species levelAnimation: MacroevolutionCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Concept 24.1: The biological species conceptemphasizes reproductive isolation Species is a Latin word meaning “kind” or“appearance” Biologists compare morphology, physiology,biochemistry, and DNA sequences whengrouping organismsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Biological Species Concept biological species concept - states that aspecies is a group of populations whosemembers have the potential to interbreed innature and produce viable, fertile offspring;they do not breed successfully with otherpopulations Gene flow between populations holds thephenotype of a population togetherCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-2(a) Similarity between different species(b) Diversity within a species

Reproductive Isolation Reproductive isolation is the existence ofbiological factors (barriers) that impede twospecies from producing viable, fertile offspring Hybrids are the offspring of crosses betweendifferent species Reproductive isolation can be classified bywhether factors act before or after fertilizationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-4Prezygotic barriersHabitat IsolationTemporal ic barriersBehavioral IsolationMechanical IsolationGametic IsolationMatingattempt(c)(d)(e)(f)Reduced Hybrid ViabilityReduced Hybrid FertilityHybrid )(i)(j)(b)(k)(l)

Fig. 24-4aPrezygotic barriersHabitat IsolationTemporal empt(c)(d)(b)Mechanical IsolationBehavioral Isolation(e)(f)

Fig. 24-4iPrezygotic barriersGametic IsolationPostzygotic barriersReduced Hybrid Viability Reduced Hybrid FertilityHybrid )(i)(j)(k)(l)

Fig. 24-4bPrezygotic barriersHabitat IsolationIndividualsofdifferentspeciesTemporal IsolationBehavioral IsolationMechanical IsolationMatingattempt

Fig. 24-4jPostzygotic barriersPrezygotic barriersGametic IsolationReduced Hybrid ViabilityFertilizationReduced Hybrid FertilityHybrid BreakdownViable,fertileoffspring

Prezygotic barriers block fertilization fromoccurring by:– Impeding different species from attempting tomate– Preventing the successful completion ofmating– Hindering fertilization if mating is successful– Include: Habitat isolation, behavioralisolation, temporal isolation, mechanicalisolation, gametic isolationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Habitat isolation: Two species encounter eachother rarely, or not at all, because they occupydifferent habitats, even though not isolated byphysical barriersCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Temporal isolation: Species that breed atdifferent times of the day, different seasons,or different years cannot mix their gametesEastern spotted skunk(Spilogale putorius)Western spotted skunk(Spilogale gracilis)Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Behavioral isolation:Courtship rituals and otherbehaviors unique to aspecies are effectivebarriersCourtship ritual of bluefooted boobiesVideo: Albatross Courtship RitualVideo: Giraffe Courtship RitualVideo: Blue-footed Boobies Courtship RitualCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Mechanical isolation:Morphologicaldifferences can preventsuccessful matingBradybaena with shellsspiraling in oppositedirectionsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Gametic isolation:Sperm of one speciesmay not be able tofertilize eggs of anotherspeciesSea urchinsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Postzygotic barriers prevent the hybridzygote from developing into a viable, fertileadult:– Reduced hybrid viability– Reduced hybrid fertility– Hybrid breakdownCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Reduced hybridviability: Genes of thedifferent parent speciesmay interact and impairthe hybrid’sdevelopmentEnsatina hybridCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Reduced hybrid fertility: Even if hybrids arevigorous, they may be sterile DonkeyHorseCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin CummingsMule (sterile hybrid)

Hybrid breakdown:Some first-generationhybrids are fertile, butwhen they mate withanother species or witheither parent species,offspring of the nextgeneration are feebleHybrid cultivated rice plants withor sterilestunted offspring (center)Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Limitations of the Biological Species Concept The biological species concept cannot beapplied to fossils or asexual organisms(including all prokaryotes)Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Other Definitions of Species Other species concepts emphasize the unitywithin a species rather than the separatenessof different species The morphological species concept definesa species by structural features– It applies to sexual and asexual species butrelies on subjective criteriaCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The ecological species concept views aspecies in terms of its ecological niche– It applies to sexual and asexual species andemphasizes the role of disruptive selection The phylogenetic species concept: defines aspecies as the smallest group of individuals ona phylogenetic tree– It applies to sexual and asexual species, but itcan be difficult to determine the degree ofdifference required for separate speciesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Concept 24.2: Speciation can take place with orwithout geographic separation Speciation can occur in two ways:– Allopatric speciation– Sympatric speciationCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-5(a) Allopatric speciation(b) Sympatric speciation

Allopatric (“Other Country”) Speciation In allopatric speciation, gene flow isinterrupted or reduced when a population isdivided into geographically isolatedsubpopulationsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Process of Allopatric Speciation The definition of barrier depends on the abilityof a population to disperse Separate populations may evolveindependently through mutation, naturalselection, and genetic driftCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-6A. harrisiA. leucurus

Evidence of Allopatric Speciation Regions with many geographic barrierstypically have more species than do regionswith fewer barriersCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-7Mantellinae(Madagascar only):100 speciesRhacophorinae(India/SoutheastAsia): 310 speciesOther Indian/Southeast Asianfrogs100608012402003Millions of years ago (mya)132IndiaMadagascar88 mya65 mya56 mya

Reproductive isolation between populationsgenerally increases as the distance betweenthem increasesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Degree of reproductive isolationFig. 24-82.01.51.00.50050200250100150Geographic distance (km)300

Barriers to reproduction are intrinsic;separation itself is not a biological barrierCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Sympatric (“Same Country”) Speciation In sympatric speciation, speciation takesplace in geographically overlapping populationsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Polyploidy Polyploidy is the presence of extra sets ofchromosomes due to accidents during celldivision An autopolyploid is an individual with morethan two chromosome sets, derived from onespeciesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-10-12n 64n 12Failure of celldivision afterchromosomeduplication givesrise to tetraploidtissue.

Fig. 24-10-22n 64n 12Failure of celldivision afterchromosomeduplication givesrise to tetraploidtissue.2nGametesproducedare diploid.

Fig. 24-10-32n 64n 12Failure of celldivision afterchromosomeduplication givesrise to tetraploidtissue.2nGametesproducedare diploid.4nOffspring withtetraploidkaryotypes maybe viable andfertile.

An allopolyploid is a species with multiplesets of chromosomes derived from differentspeciesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-11-1Species B2n 4Unreducedgametewith 4chromosomesMeioticerrorSpecies A2n 6Normalgameten 3

Fig. 24-11-2Species B2n 4Unreducedgametewith 4chromosomesMeioticerrorSpecies A2n 6Normalgameten 3Hybridwith 7chromosomes

Fig. 24-11-3Species B2n 4Unreducedgametewith 4chromosomesMeioticerrorSpecies A2n 6Normalgameten 3Hybridwith 7chromosomesUnreducedgametewith 7chromosomesNormalgameten 3

Fig. 24-11-4Species B2n 4Unreducedgametewith 4chromosomesMeioticerrorSpecies A2n 6Normalgameten 3Hybridwith 7chromosomesUnreducedgametewith 7chromosomesNormalgameten 3Viable fertilehybrid(allopolyploid)2n 10

Polyploidy is much more common in plantsthan in animals Many important crops (oats, cotton, potatoes,tobacco, and wheat) are polyploidsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Habitat Differentiation Sympatric speciation can also result from theappearance of new ecological niches For example, the North American maggot flycan live on native hawthorn trees as well asmore recently introduced apple treesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Sexual Selection Sexual selection can drive sympatric speciation Sexual selection for mates of different colorshas likely contributed to the speciation in cichlidfish in Lake VictoriaCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-12EXPERIMENTNormal lightP.pundamiliaP. nyerereiMonochromaticorange light

Allopatric and Sympatric Speciation: A Review In allopatric speciation, geographic isolationrestricts gene flow between populations Reproductive isolation may then arise bynatural selection, genetic drift, or sexualselection in the isolated populations Even if contact is restored betweenpopulations, interbreeding is preventedCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

In sympatric speciation, a reproductive barrierisolates a subset of a population withoutgeographic separation from the parent species Sympatric speciation can result frompolyploidy, natural selection, or sexualselectionCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Concept 24.3: Hybrid zones provide opportunitiesto study factors that cause reproductive isolation A hybrid zone is a region in which members ofdifferent species mate and produce hybridsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Patterns Within Hybrid Zones A hybrid zone can occur in a single band whereadjacent species meet Hybrids often have reduced fitness comparedwith parent species The distribution of hybrid zones can be morecomplex if parent species are found in multiplehabitats within the same regionCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-13EUROPEFire-belliedtoad rangeHybrid zone0.99Allele frequency (log scale)Yellow-bellied toad,Bombina variegataYellow-belliedtoad rangeFire-bellied toad,Bombina bombina0.90.50.10.014020301001020Distance from hybrid zone center (km)

Hybrid Zones over Time When closely related species meet in a hybridzone, there are three possible outcomes:– Strengthening of reproductive barriers– Weakening of reproductive barriers– Continued formation of hybrid individualsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Reinforcement: Strengthening ReproductiveBarriers The reinforcement of barriers occurs whenhybrids are less fit than the parent species Over time, the rate of hybridization decreases Where reinforcement occurs, reproductivebarriers should be stronger for sympatric thanallopatric speciesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-15Sympatric malepied flycatcher28Allopatric malepied flycatcherPied flycatchers24Number of femalesCollared flycatchers20161284(none)0Females mating OwnOtherwith males from: species speciesSympatric malesOwnOtherspecies speciesAllopatric males

Fusion: Weakening Reproductive Barriers If hybrids are as fit as parents, there can besubstantial gene flow between species If gene flow is great enough, the parent speciescan fuse into a single speciesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-16Pundamilia nyerereiPundamilia pundamiliaPundamilia “turbid water,”hybrid offspring from a locationwith turbid water

Concept 24.4: Speciation can occur rapidly or slowlyand can result from changes in few or many genes Many questions remain concerning how long ittakes for new species to form, or how manygenes need to differ between speciesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Time Course of Speciation Broad patterns in speciation can be studiedusing the fossil record, morphological data, ormolecular dataCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Patterns in the Fossil Record The fossil record includes examples of speciesthat appear suddenly, persist essentiallyunchanged for some time, and then apparentlydisappear Niles Eldredge and Stephen Jay Gould coinedthe term punctuated equilibrium to describeperiods of apparent stasis punctuated bysudden change The punctuated equilibrium model contrastswith a model of gradual change in a species’existenceCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-17(a) Punctuated patternTime(b) Gradual pattern

Speciation Rates The punctuated pattern in the fossil record andevidence from lab studies suggests thatspeciation can be rapid The interval between speciation events canrange from 4,000 years (some cichlids) to40,000,000 years (some beetles), with anaverage of 6,500,000 yearsCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-18(a) The wild sunflower Helianthus anomalusH. anomalusChromosome 1Experimental hybridH. anomalusChromosome 2Experimental hybridH. anomalusChromosome 3Experimental hybridKeyRegion diagnostic forparent species H. petiolarisRegion diagnostic forparent species H. annuusRegion lacking information on parental origin(b) The genetic composition of three chromosomes in H.anomalus and in experimental hybrids

Fig. 24-18a(a) The wild sunflower Helianthus anomalus

Fig. 24-18bH. anomalusChromosome 1Experimental hybridH. anomalusChromosome 2Experimental hybridH. anomalusChromosome 3Experimental hybridKeyRegion diagnostic forparent species H. petiolarisRegion diagnostic forparent species H. annuusRegion lacking information on parental origin(b) The genetic composition of three chromosomes in H.anomalus and in experimental hybrids

Studying the Genetics of Speciation The explosion of genomics is enablingresearchers to identify specific genes involvedin some cases of speciation Depending on the species in question,speciation might require the change of only asingle allele or many allelesCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-19

Fig. 24-20(a) Typical Mimulus lewisii(b) M. lewisii with an M. cardinalis flower-color allele(c) Typical Mimulus cardinalis(d) M. cardinalis with an M. lewisii flower-color allele

You should now be able to:1. Define and discuss the limitations of the fourspecies concepts2. Describe and provide examples of prezygoticand postzygotic reproductive barriers3. Distinguish between and provide examples ofallopatric and sympatric speciation4. Explain how polyploidy can causereproductive isolation5. Define the term hybrid zone and describethree outcomes for hybrid zones over timeCopyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 24-4a Habitat Isolation Temporal Isolation Prezygotic barriers Behavioral Isolation Mating attempt Mechanical Isolation (a) (c) (e) (f) (b) (d) Individuals

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