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The Origin of Species chapter 24 Campbell and Reece Speciation process by which one species splits into 2 or more species  Speciation explains both the diversity of life and the unity of living things.  Speciation : forms bridge between: MICROEVOLUTION MACROEVOLUTION  Evolutionary  Evolutionary change below change above the species level species level  Example:  Examples:  change in allele  origin of new frequencies in group of population over organisms generations  impact of mass extinctions Biological Species Concept  Species: a group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspring members of a species cannot produce viable, fertile offspring with other groups  emphasizes the separateness of species due to reproductive barriers  What holds the gene pool of a species together? Gene Flow: transfer of alleles between populations of same species  exchange of alleles tends to hold populations together genetically  Reproductive Isolation  existence of biological barriers that keep members of 2 populations from interbreeding over long periods of time Reproductive Isolation  hybrids: offspring that result from the mating of individuals from 2 different species or from 2 truebreeding varieties of same species Prezygotic Barriers block fertilization from happening by: 1. impeding members of different species from attempting to mate 2. preventing attempted mating from being completed successfully 3. hindering fertilization if mating was completed successfully  Postzygotic Barriers reproductive barrier that prevents hybrid zygotes produced by 2 different species from developing into viable, fertile adults 1. lethal developmental errors 2. infertility in viable offspring  Types of Prezygotic Reproductive Barriers Habitat Isolation 2 species that occupy different habitats w/in same area may rarely interact example: 1. Types of Prezygotic Reproductive Barriers 2. Temporal Isolation: species breed during different times of day, different seasons, or different years Types of Prezygotic Reproductive Barriers 3. Behavioral Isolation: Courtship rituals used to attract mates are effective barriers Types of Prezygotic Reproductive Barriers 4. Mechanical Isolation:  morphological differences prevent successful completion even if attempted Types of Prezygotic Reproductive Barriers 5. Gametic Isolation:  Sperm of 1 species may not be able to fertilize egg of another: ◦ Reproductive tract hostile to sperm ◦ Sperm does not have enzymes to penetrate zona pellicida of another species Types of Postzygotic Reproductive Barriers Reduced Hybrid Viability: hybrids development or survival is impaired 1. Types of Postzygotic Reproductive Barriers 2. Reduced Hybrid Fertility: hybrids may develop and be healthy but they are not fertile Types of Postzygotic Reproductive Barriers Hybrid Breakdown: Some 1st generation hybrids are fertile but those offspring are feeble or sterile 3. Species There is no single, universally applicable species concept that can define, explain, and identify all species.  There are multiple ways to think about & define species.  Biological Species Limitations unable to use these characterisitics on fossils of extinct species  only applies to organisms that reproduce sexually  only applies where there is no gene flow  Other Definitions of Species These dfns emphasize the unity w/in a species. 1. morphological species concept:  characterizes a species by a structural feature  applies to species that reproduce sexually or asexually  how scientists distinguish most species  disadvantage: subjective  Morphological Species Concept  Problems: ◦ domestic dogs may look very different but are still same species ◦ mouse lemurs look very similar but there are 18 species of them ◦ grey mouse lemur lesser mouse lemur Ecological Species Concept  views species in terms of its niche ◦ the sum of how members of the species interact with the nonliving & living parts of their environment ◦ asexual or sexual species ◦ emphasizes role of disruptive NS as organisms adapt to different environmental conditions Phylogenetic Species Concept defines species as smallest group of individuals that share a common ancestor, forming one branch on the “tree of life”  determining degree of differences is difficult  Species  There are >20 other ways to define species Speciation can take place with or w/out geographic separation Speciation can occur in 2 main ways: 1. Allopatric Speciation 2. Sympatric Speciation  ALLOPATRIC SPECIATION “other country”  Gene flow is interrupted when population is divided into geographically isolated subpopulations  Allopatric Speciation Process Once geographic separation has occurred, the separated gene pools will each have their own mutations  NS & genetic drift may alter allele frequencies in different ways in each subpopulation group  Allopatric Speciation Evidence of Allopatric Speciation There are many studies & examples supporting this type speciation  Indirect support: regions that are isolated or highly subdivided have more species than regions w/out those features  Drosophila Experiment Sympatric Speciation “same country”  occurs in populations in same geographic area  less common than allopatric  occurs if gene flow is reduced by factors like:  ◦ polyploidy ◦ habitat differentiation ◦ sexual selection POLYPLOIDY means extra sets of chromosomes  can occur in animals  gray tree frog (Hyla versicolor)  around Great Lakes  Polyploidy much more common in plants  estimate: 80% of today’s plants species have ancestors that formed by polyploid speciation  2 forms  1. Autopolyploid Individual has >2 chromosome sets all derived from a single species  Plant polyploidy  Plant Polyploidy  tetraploid plant can produce fertile tetraploid offspring by selfpollinating or mating with other tetraploids 2. Allopolyploid 2 different species interbreed making a hybrid  hybrid reproduces asexually  over generations sterile hybrid  fertile polyploid (called an allopolyploid)  Allopolyploids can breed with each other but not with either of their parents so are a new species  rare: 5 new plant species since 1850 documented  Mimulus peregrinus  Allopolyploids include many agricultural crops  Triticum aestivum (bread wheat) has 6 sets chromosomes (2 pair from each of 3 parents), an allohexaploid  1st polyploidy event probably occurred ~8,000 yrs ago as spontaneous hybrid  Top 2 parents bottom: Triticum aestivum Allopolyploids plant geneticists “create” new polyploids making hybrids with desired characteristics  use chemicals that induce meiotic & mitotic errors  Habitat Differentiation  Sympatric speciation can occur when genetic factors enable a subpopulation to exploit a habitat or resource used by the parent population Rhagoletis pomonella North American apple maggot fly  Original habitat was the native hawthorn tree  Habitat Differentiation   As apples mature faster than hawthorn fruit, NS has favored flies with rapid development have an allele that benefits flies that feed off only 1 or the other not both (post-zygotic barrier to reproduction)  The flies feeding on apple trees now show temporal isolation from flies still eating hawthorn fruit (prezygotic restriction to gene flow) Sexual Selection can also drive sympatric speciation:  cichlid fish Pundamilia pundamilia  Cichlid Fish >600 species found in Lake Victoria  originated in past 100 000 yrs  hypothesis : subgroups of original population adapted to different food sources  genetic divergence  female preference for mates may also be a factor: 1 species breeding males have blue back another species has orange back  Breeding Cichlids Colors Sexual Selection Study placed the 2 subspecies in same tank  used monochromatic orange light so both appeared very similar  females bred with either  Conclusion: mate choice by females is based on male coloration so it’s the main reproductive barrier (prezygotic behavior)  Allopatric & Sympatric Speciation are the 2 main modes of speciation Allopatric geographic isolation NS genetic drift sexual selection      Sympatric requires emergence of a reproductive barrier that isolates a subgroup less common polyploidy sexual selection      Hybrid Zones