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cladogenesis vs anagenesis
- cladogenesis: branching or multiplication of lineages (each of which evolves by anagenesis)
- anagenesis: evolution within species along its own path
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species definition
groups of actually or potentially interbreeding populations which are reproductively isolatedd from other such groups - several biological differences between teh populations greatly reduce gene exchange between them
this biological species concept takes in account teh individudal and geographic variation and is good for sexually reproducing and extant organisms (not asexual, not fossils)
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sibling species
reproductively isolated populations that are difficult or impossible to distinguish by morphological features, but are different in ecology, behavior, chromosomes, or other such characters
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hybrid zones
genetically distinct populations meet and interbreed to a limited extent, but in which there exist partial barriers to gene exchange (hybridizing entities may be calledd semi-species)
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phylogenetic species concept
teh smallest monophyletic group of common ancestry
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geneological species concept
species are exclusive groups of organisms, where all the members of the group are more closely related to one another than to any organism outside teh group
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isolating barriers
barriers to gene flow - isolating mechanisms - biological differences
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speciation
the origin of two species from a common ancestral species - consists of the evolution of biological barriers to gene flow
how can two different populations form without intermediates?
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prezygotic vs postmating vs. postzygotic barriers
- prezygotic - prevent or reduce the likelihood of transfer of gametes to members of other species
- postmating - features that prevent successful formation of hybrid zygotes even if mating takes place
- postzygotic - reduced survival or reproductive rates of hybrid zygotes
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premating barriers
- ecological isolation - seasonal isolation (breed at different times), isolatedd by habitat (wont even meet),
- behavioral isolation - sexual isolation or ethological isolation
specific mate recognition system: the signals and responses between potential mates - one sex (usually the females) will not respond to inappropriate signals
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postmating barriers/prezygotic
- mechanical - copulatory isolation: genitalia may differ between related species
- conspecific sperm precedence - if the female mates with both a heterospecific male and a conspecific - the conspecific males sperms will be successful in fertilizing her eggs
gametic isolation - games of different species fail to unite
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postzygotic barreirs
- hybrid inviability
- hybrid sterility
- multiple isolating barriers
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specific mate recognition system
signals/responses between potential mates - one sex will not respond to inappropriate signals
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hybrid inviabilty
a postzygotic barriers - structural differences between teh chromosomes that cause segregation of some anueploid gametes during meiosis or by differences between teh genes of teh two parents
Haldanes rule
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haldanes rule
- about hyrbid sterility
- sterility usually affects teh heterogametic sex (teh one with the two different sex chromosomes)
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genetic bases for barriers to gene flow (3)
- single genes
- epistatic interactions of genes
- chromosome differences - alterations of chromosome structure and differences in the number of chromosome sets --> can cause aneuploidy! (or differences between the genes of the parent populations)
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which is more strong, premating or postzygotic isolation in a population/species that has recently diverged
premating!
prezygotic isolation = stronger among sympatric than among allopatric pairs of taxa
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which gender does postzygotic isolation evolve more rapidly?
in males! hybrid sterility/inviability is almost always seen in males only (female sterility seen when taxa are much older)
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coalescent theory
loss of all teh ancestral lineages of DNA sequence variants except one - coalescence to a common ancestral gene copy occurs in each species
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hybridization
occurs when offspring are produced by interbreeding between genetically distinct populations - may be the source of new adaptations or even new species
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primary hybrid zones vs. secondary hybrid zones
- primary: originate in situ as natural selection alters allele frequencies in a series of more/less continuously distributed populations (may correspond to a sharp change in an environmental factor)
- secondary: formed when two formerly allopatric populations that have become genetically differentiated expand so that they meet and interbreed
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tension zones
hyrbid zones when populations meet at secondary hybrid zones and have low intrinsic fitness (b/c of heterozygote disadvantage/breakdown of coadapted gene complexes)
hybrids have low fitness irrespective of variation in environmental conditions
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4 fates of hybrid zones
- may persist indefinitely
- may favor alleles that enhance prezygotic isolation - may result in full reproductive isolation - extinction of hybrids
- alleles that improve fitness of hybrids may increase in frequency - postzygotic barreirs may break down - merging of species
- some hybrids may become reproductively isolated from teh parent forms and become a third species
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3 modes of speciation
- allopatric speciation - evolution of reproductive barriers in populations that are prevented by a geographic barrier from exchanging genes (gene flow = 0)
- parapatric speciation - spatially distinct populations between which there is some gene flow, diverge and become reproductively isolated
- sympatric speciation - evolution of reproductive barreirs within a single, initially randomly mating population (gene flow is maximum = 0.5)
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vicariance vs. peripatric speciation
- two forms of allopatric?
- vicariance - divergence of two large populations
- peripatric speciation - divergence of a small population from a widely distributed ancestral form
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allopatric speciation
- populations that are geographically separated by a physical barrier (reduces gene flow)
- argued that it is the prevalent mode of speciation - at least in animals
if the barrier was removed or new species disperse over the barrier --> may become sympatric but without exchanging genes - they may form a hyrbidzone if incomplete reproductive isolation has occurred
the more geographically distant the populations, the more genetically different they were and the less likely they were to mate
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vicariant allopatric speciation
as more allelic differences accumulate - later mutations have more opportunities to cause incompatibility and lower the fitness of hybrids
thought to be caused by natural selection, which causes the evolution of genetic differences that create prezygotic and/or postygotic incompatibility
- mutation/selection proceed independently - speciation is a side effect
- when they come back into contact the hybrids are sterile so natural selection favors prezygotic isolation
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ecological speciation
evolution of barriers to gene flow caused by divergent ecologically based selection
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peripatric speciation
- founder effect speciation - restricted distributions, isolated, peripheral to the parent species, highly divergent - genetic change can be rapid when only a few individuals move!
- snowballing of genetic change -- reproductive isolation
- foundation for the idea of punctuated equilibrium
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parapatric speciation
- speciation in neighboring populations
- higher rate of gene flow - force of selection is stronger to create genetic differences for reproductive isolation
- occurs when gene flow is much weaker than divergent selection for different gene combos
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sympatric speciation
- controversial - can it really happen?
- biological barrier to gene exchange arose within an initially randomly mating population wihtout any segregation of the species (speciation despite initially high gene flow) - how are the intermediates reduced then?
- diversifying selection
- divergent adaptation to two distinct resources (ecological speciation - or temporal speciation)
- 2 models of mating - assortative (individuals prefer mates that match own phenotype) and trait preference (genes control female preference and male trait)
- recombination = huge obstacle!
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polyploidy
- organism with more than two complements of chromosomes - isolated by postzygotic barriers
- instantaneous speciation by single genetic event
- may be important in plant evolution
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autopolyploid vs. allopolyploid
- auto - formed by union of unreduced gametes from genetically and chromosomally compatible individuals that belong to the same species
- allo - derivative of a diploid hybrid between two species (or two ancestors that form hybrids that are sterile b/c of genetic/chromosomal incompatibility)
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recombinational speciation
- hybrid speciation
- hybridization can give rise to distinct species with same ploidy as their parents
- generates diverse gene combos on which selection can act
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rate of speciation meanings
- transition time (time for speciation) - time for reproductive isolation to evolve once process has started
- biological speciation interval - average time between origin of a new species and when that species speciates again
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what are rates of speciation affected by?
generation time, envirionmental factors, mating systems
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specific coevolution
two species evolve in response to each other
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diffuse/Guild coevolution
when several species are involved - effects are not independent
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escape-and-radiate coevolution
species evolves a defense against enemies and is thereby enabled to proliferate into a diverse clade
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coevolution of enemies and victims
- could continue indefinitely - unending escalation of evolutionary arms race
- could result in stabel equilibrium
- cause continual cycles (or irregular fluctations) in the genetic composition of both species
- may leadd to extinction of one/both species
- gene for gene interactions
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gene for gene interactions
- model of enemy-victim coevolution
- example: host as R that confers resistance to the parasite, but in the parasite v confers infectivity (ability to infect/grow in host with R allele) - can become frequency dependent
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virulent parasites
those that reduce the survival/reproduction of their hosts
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evolution of virulence - 3 theories
- greater virulence is expected to evolve in parasite species in which multiple infection is frequent
- if hosts rapidly become immune to the parasite slection favors rapid reproduction to outrun the host's immune system
- if horizontally transmitted - their fitness does not depend on reproduction of host - more virulent in horizontal transmitted parasites
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mutualisms
- interactions between species that benefit individuals of both species
- symbiotic - individuals are intimately associated for much of their lifetimes
- reciprocal exploitation
- always a chance of cheaters - selection favors protective mechanisms to punish cheaters
- important basis for adaptation and complexity - provides one or both partners with new capabilities
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evolution of competitive interactions
- competition is one of hte major causes of adaptive radiation - divergence in resource use (competition for resoruce within species)
- density-dependent diversifying slection - two species
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character displacement
divergence in response to competition between species - process of divergence that is due to competition
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ecological release
- a species/population exhibits greater variation in resource use and phenotypic characters if it occurs alone that if it coexits with competing species
- alleviation of competition may enhance rates of speciation/diversification
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apparent competition
- if a predators presence increases with the abundance of one prey, another species of prey may suffer heavier predation and decline
- one or both prey may be selected to diverge from the other in characteristics of susceptibility to the predator (can also converge if it lowers risk of predation)
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Batesian mimicry
a mimic resemples a dangerous species (the model)
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mullerian mimicry
two or more dangerous species are co-mimics (or co-models) that jointly reinforce aversion learning by predators
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comparative genomics
- comparative study of whole genomes
- to reveal major life history events
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c-value paradox
finding htat organismal complexity and total genome size dod not display a tight correlation
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codon bias
- a way in which protein-coding genes evolve
- measure of the departure from equality of the frequency of synonymous codons for a given amino acid in proteins
- codon usage is uneven
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gene dispensability
the measure of how dispensable certain genes are (can be deleted from genome or not)
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translational robustness
ability of an amino acid sequence to maintain its proper protein folding in the face of ongoing mutations
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origin of new genes methos
lateral gene transfer, from noncoding regions, exon shuffling, gene chimerism, motif multiplication/exon loss, gene duplication
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lateral gene transfer
horizontal gene transfer between lineages - can arise in the origin of new genes
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exon shuffling
- produces truly new genes -
- domain accretion - new genes are produced by the addition of domains to teh beginnings/ends of ancestral genes
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chimeric gene
one that consists of pieces derived from two or more different ancestral genes (by exon shuffling, or by retrotransposition - can result in pseudogenes)
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multigene families
larger groups of genes, related to one anotehr by clear ancestry/descent, have diverse functions that have a common theme
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gene duplication
new genes arise as copies of pre-existing genes
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paralogous vs orthologous genes
- paralogous - members of multigene families, originated from a common ancestral gene by gene duplication
- orthologous - found in different species have diverged from a common ancestral gene by phylogenetic splitting at the organismal level
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copy number variants
emerging as an important source of variation in the human genome
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possible fates of duplicate genes
- may diverge in sequence and function
- one copy may remain functional while the other becomes nonfunctional pseudogene (or deleted)
- undergo gene conversion
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gene conversion
when sequence information from locus is transferred unidirectionally to other membres of hte gene family so that all acquire essentially the same sequence - causes concerted evolution of the gene family - causes the production of the same gene product from multiple loci
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neofunctionalization
one of the gene duplicates retains original function and the other acquires a new function because of fixation of new mutations
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subfunctionalization
each gene duplicate becomes specialized for a subset of functions originally performed by teh ancestral single-copy gene
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evolutionary developmental biology (evodevo)
seeks to understand teh mechanisms by which development has evolved in terms of developmental processes and evolutionary processes
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Hox genes
- regulate teh patterning of specific body structures- homeobox genes (with homeobox- in the gene), homeodomian - in the protein)
- morphological differences is caused by changes in expression of genes that the Hox genes regulate
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homologous features
- thsoe that are inherited, with more/less modification froma common acnestor in which teh feature first evolved (synapomorphies)
- similarity in position, structural features, intermediate forms,
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serially homologous
share same developmental genetic machinery in their ontogeny but are not historically homologous within a species
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biological homology
a features may be homologous among species at oen level of organization but not at another level
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regulatory modularity
- with different enhancers on genes
- enables evolutionary changes in development of specific tissues/bodyd structures
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exaptations
novel uses of pre-existing morphological traits
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recruitmenta nd co-option
refers to teh evolution of novel functions for preexisting genes and developmental pathways
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heterochrony
evolutionary changes in the timing of development
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allometry
differential growth rates of different parts of the body
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developmental constraints on evolution
- phsyical constraints (properties of biological materials do not permit them to evolve)
- selective (functional) constraints - some features may not appear b/c they are always disadvantageous
- genetic contraints - varying degrees of tolerance for variation
- developmental constraints - bias on the production of various phenotypes caused by the structure, character, composition, or dynamics of the developmental system
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developmental/genetic constraints can explain some evolutionary patterns
- absence of features in certain lineages
- directional trends
- parallel evolution of traits in independent lineages
- reduction of morphological variation of traits (canalization - most highly advantageous phenotype is mroe reliably produced)
- morphological stasis over long periods
- similarities in embryological stages among higher taxa
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explanations of stasis in punctuated equilibrium
- internal genetic/developmental contraints
- stabilizing selection for a constant optimum phenotype (more common explanation)
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habitat tracking
teh shifting of the geographic distributions of species in concert with teh distribution of their typical habitat
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punctuated equilibrium vs. saltation (by jumps) evolutionary theory
- punctuated equilibrium - passed through intermediates but it was so fast taht the fossil record makes it appear like a discontinuous change
- saltation - intermediates never existed - mutatnt individuals differ drastically from their parents
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evolvability
idea that the production of genetic and phenotypic variation may be structured in a way that makes adaptive evolution more likely
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trend vs. progress
- trend: persistent directional change in teh average values of a feature over a course of time
- progress: implies betterment
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passive trend
- lineages in teh clade evolve in both directions, but if there is a boundary on one side, teh variation can only expand in the other direction - mean increases but some lineages remain near the ancestral value
- the boundaries may be due to functional/developmental genetic constraints
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driven (active) trend
changes within lineages in one direction are more likely that changes in the other direction
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trends in history of life?
- efficiency/adaptedness
- complexity
- diversity/disparity
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disparity
increases in phenotypic variation among species
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theistic evolution
GOd established natural laws (like natural selection) and then let the universe run on its own without further supernatural intervention
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special creation
each species was created independently by GOd essentially in its present form
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scientific creationism
attacks on, and scientific disputes of evolution
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intelligent design
- do no invoke special creation
- argue that many biological phenomen are too complicated to have arisen by natural processes - can only be explained by an intelligent designer (ID is a scientific not religious concept)
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hypothesis vs fact vs theory
- hypothesis: proposition
- fact: hypothesis that has become so supported by evidence
- theory: big idea that encompasses other ideas/weaves them into coherent fabric
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evidence for evolution
- fossils
- phylogenetic and comparative studies
- genes and genomes (molecular clocks)
- biogeography
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social darwinism
evolutionary science that was misappropriated to jsutify racism and domination - genetic inferiority for superior people
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coevolution
the joint evolution of ecologically interacting species, each of which evolves in response to selection imposed by the other(s)
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virulence and infectivity
- virulence - effect on host
- infectivity - transmission
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domain accretion
- formation of truly new genes through exon shuffling
- new genes produced by addition of domains to the beginnings/ends of ancestral genes
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chimeric genes
consists of pieces derived from two or more different ancestral genes - by exon shuffling or by retrotransposition
results in processed pseudogenes sometimes
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motif multiplication
can give rise to new genes ith new functions - multiplication of specific motifs
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orthologous vs paralogous genes
- orthologous: found in different species ahve diverged from a comon ancestral gene by phylogenetic splitting at the organismal level
- paralogous: like members of multigene families - have a common ancestral gene by gene duplication
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developmental pathways/circuits
the genes that regulate morphogenesis function in heirarchies or netowrks
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regulation of gene expression is achieved by...
- enhancers for each gene (bind to one or more transcription factors)
- genes have different enhancers - regulatory modularity
- changes in enhancers = phenotypic adaptations
- or in teh transcription factor itself
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modularity
- the degree to which teh development of different body structures is independent
- individual structures/units seen as modules
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polyphenism
non-genetically based polymorphism in which environmental differences cause development to produce a small set of (usually two) alternative states
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living fossils
have changed so littler since millions of years ago
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phylogenetic conservatism
- how there are living fossils - they have changed so little since millions of eyars ago
- hypotheses for it include stabilizing selection or internal contraints
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niche conservatism
- logn dependence of related species on the same resources/environmental conditions
- caused because maybe another species will prevent a species from shifting/expanding or because stabilizing selection or ancestral character states because a majority of the population occupies the ancestral environment, not the new one
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the irreducible complexity argument
- the argument against evolution that a complex feature cannot function effectively except with coordination of all componenets - so intermediates couldn't have worked
- and the entire complexity couldn't have arisen by a single mustational step
- BUT intermeditaes are function (like the eye)
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facilitated variation
- teh core processes of protein activity and cell/organ development, with the establishment of regulatory interactions among ancient genes causes variation to arise that facilitates for evolution
- facilitates the evolution of complex characters
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