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Fossil Record patterns
-anagenesis
-adaptive radiation
-convergent evolution
-Dollo’s Law (of irreversibility)
Fossil Record rates
-phyletic gradualism
-punctuated equilibrium
Anagenesis
the gradual evolution of a single species within a single lineage over time, where the population changes so much that it becomes a new species while the ancestral form is replaced or considered extinct
Adaptive radiation
an evolutionary process in which organisms diversify rapidly from a single ancestral species into a multitude of new forms, particularly when an environmental change opens up new resources, niches, or ecological opportunities.
Dollo’s Law
once lost, a trait is not regained
Macroevolution
-evolution above the species level
-observed in the natural world and is recorded in the fossil record
Different macroevolutionary patterns
-convergent evolution
-adaptive radiations
-phenotypic plasticity
-morphological stasis
-Dollo’s Law (of irreversibility)
Phenotypic plasticity
a single genotype that may produce different phenotypes in different environments
Why are transitional forms so informative
-very rare
-these fossils have characteristics that are intermediate between 2 distinct taxonomic groups (fish and tetrapods)
Phyletic gradualism
-gradual change over a long period of time
-contains most transitional fossils (tiktaalik, dinos)
“saltation”
Punctuated equilibrium
-short periods of rapid change and macroevolutionary events during which new taxa arise (speciation)
-while morphology isnt changing, DNA still accumulates mutations over time
EX: stickleback fishes
“radualism”
What happens to stickleback fish when they change to salt to freshwater?
Freshwater: they lose their pelvic spines (increases fitness bc dragonfly larvae cant grab them)
Saltwater: gain pelvic spines (increases fitness bc they cant be swallowed by predator fish)
Neutrally evolving; molecular clocks
Neutrally evolving DNA sequences evolve in a clocklike manner
DNA substitutions is proportional to time
What is evolutionary developmental biology?
a field of biology that compares how different animals and plants grow from a single cell into an adult, helping scientists understand how new body shapes and traits evolve
What are typical model organisms used in Evo-Devo?
fruit flies
chicks
What is the genetic toolkit?
-genes in the genome that are used to form and pattern the body
-occurs typically during embryonic life
-deeply evolutionary conserved
-transcription and signaling factors
What are cis and trans acting components of the toolkit?
transcription factors (Hox) that regulate the expression of other genes
Signaling factors (sonic hedgehog) that mediate short and long-range interactions between cells
How does CRE variation in different organisms correlate to changes in gene expression and morphology?
alters gene expression in a modular, tissue-specific manner without disrupting the core pleiotropic functions of developmental proteins
What is a gene developmental gene network?
a complex biological blueprint made of interacting DNA segments, molecular regulators, and transcription factors that control how an organism develops from a single cell into a structured body plan
What is the main objective of evo-devo?
-to study the mechanisms that produce evolutionary changes in body plans
Fossil Record Pattern 1
Homology
correspondence or similarity of structures and function in different taxa due to shared common ancestory
Fossil Record Pattern 2
Homoplasy due to similar ecological/environmental pressures
ex: vetebrate and octopus eye, hyenas and dogs
Fossil Record Pattern 3
Dollo’s Law supported
-Due to natural selection and developmental constraints associated with a developmental toolkit expression
Dollo’s Law violated: common lizards and reproductive mode
Fossil Record Pattern 4
Anagenesis- “against growth”
-change in the trait (x-axis) as time (y-axis) proceeds from past towards the present
-there is no speciation event (no bifurcation
Fossil Record Pattern 5
Adaptive radiation
“bushy” and have polytomies
Rates of fossil record
Gradual vs punctuated
Von Baer’s developmental hourglass phylotypic stage conserved shows:
species specific changes occur later in development
The 3 Evo Devo principles
-deep homology of conserved toolkit loci (Hox)
-Pleiotropy (Pitx1 sticklbacks)
-Tinkering via modular CREs
What does Evo Devo investigate and compare?
Embryonic development in different organisms to better understand how these processes evolved and shaped the diversity of body plans
Evo Devo provides:
-a mechanistic understanding for the origin of traits (wings, lungs, feathers)
-insights into repeated patterns of evolution observed in different times and different lineages
-a mechanistic understanding of how traits are modified through time consistent with gradual vs punctuated models
Early ideas on embryonic development
Preformation-embryos are “preformed” and basically unroll and grow
-EPIGENESIS-embryo starts “from scratch” every generation (CORRECT HYPOTHESIS)
Who was the first to examine and describe embryos?
Aristotle
proposed epigenesis
Who was the first to describe homology in embryos?
Rathke
-studies how fish, salamanders, frogs, birds, mammals developed
-noticed striking similarities in the development of the different vetebrates
Darwin embraced:
That there is strong evidence of common ancestry and descent with modification in embryos
Model organism systems:
-must be able to get embryos
-rear in the lab
-genome resources
Camera eyes, independently evolved in cephalopds and vertebrates are an excellent example of:
Convergent evolution
The vertebrate camera eye, occurs _____ in phylogeny
Early, ~518 MYA
In all metazoans, there is a small and deeply conserved set of genomic loci that:
-act during embryonic life
-pattern the body plan
-can be “tinkered” with to produce diversity of body plans
They are referred to as “toolkit” genes because they “build body plans”
Von Baer’s Hourglass
add pic
Evo-Devo principle 1
Deep Homology
Experimental of drosophilia eyeless or the mouse orthology pax6 shows the function of eyeless/pax6 is conserved in developmental of analogous compound and single lens eyes
Evo Devo principle 2
Pleiotropy
Mouse pax6 is responsible for eye, brain, and pancreas patterning and development in embryonic life. pleiotropy is defined as a single genomic locus that is responsible for different traits
Evo devo 3
Modular CREs
Different CRE modules give specific and separate instructions to Pax6
They are bound by transcription factor proteins to control gene transcription/expression
Where, when, how quantity instructions for transcription and protein
Pax6 is a:
toolkit locus that acts in embryonic life to pattern the eyes of metazoans
Distalles (D11)
all species share deep homology of D11
Pleiotropy
genetic phenomenon where a single gene influences multiple, distinct, and seemingly unrelated phenotypic traits
Evo-devo helps us understand:
-difference between structural homology and deep homolgy
-Repeated evolution (convergent evolution) : pleiotropy, development bais/constraint
-rapid vs gradual evolution
-evolution of novel traits
Difference between structural homolgy and deep homolgy
-structures in adults of very distantly related species have NO structural homology
-all distantly related species share deep homology for the toolkit locus or loci (D11, Hox, pax6)
A mouse that doesnt have pax6 doesnt have eyes, but fruit fly protein can be used to give the mouse eyes. What evo-devo priniciple is this an example of?
DEEP HOMOLOGY
A genetic network that includes the Distalless (Dll) gene triggers outgrowths during
the development of starfish and insects. There is NO structural homology of the different outgrowths that serve as appendages. What evo-devo principle is this?
Deep homology
All species share deep homology of D11
Pleiotropy also _______ evolution
Constraints/limits
-range of variation of a single genomic locus is limited because it is responsible for many traits
Pitx1
Pleiotropic master regulator
Where is pitx1 found?
stickleback fishes
-it is expressed in several regions of the stickleback embryo, corresponding to different adult structures (pelvis, eye, jaw, pupitary)
What evo devo principle is pitx1?
Pleiotropy
In freshwater stickles, the CRE module that gives specific instructions to Pitx1 is _____
DISABLED
Pitx1 isnt expressed in these embryonic cells and adults dont have a pelvic spine
Can you lose the Pitx1 genomic locus?
NO because it is pleiotropic
it is lethal to the embryo
Von Baer’s developmental hourglass “variable” region is an example of what principle?
3
Von Baer’s “conserved” region is an example of what principles?
1, 2, 3
Deep homology and pleiotropy are extremely evolutionarily
conserved in metazoans. How did phenotypic diversity occur? How
do species-specific characteristics occur?
-modular CREs “permit” sequence variation that only affects one specific set of instructions but doesnt influence other instructions
-the flexibility of genetic variation WITHIN a CRE module explains diversity withing a species and between different species
Bmp4 is a pleiotropic toolkit locus that:
patterns different bone and cartilage structures
Heavy deep beaks: have high bmp4 expression in beak embryonic cells
Small, pointy beaks: almost undetectable bmp4 expression in beak embryonic cells
The CRE module of bmp4 that give instructions for HOW MUCH bmp4 expression in embryonic beak cells will show sequence variation in different finch species
Tbx4 and Tbx5 establish :
fore vs hindlimb
Shh (sonic hedgehog) establishes :
zone of polarizing activity
Fgfs establish:
apical ectoderm ridge
The timing and spatial pattern of gene activity differs in different species. Why?
Caused by species-specific differences in CRE modules
There are a few RARE examples of mutation in protein-coding DNA sequences of a toolkit genomic locus that cause phenotypic diversity
UBX is the gene
The sequences have changed leading to different embryonic patterning of the body plan
Example of a novel trait
Butterfly spots
How do evo devo principles explain evolution of novel traits?
-modular CREs can place a pleioptropic master regulator in a new spatial and/or temporal context in the embryo without interfering with the ancestral spatial and temporal contexts
-once a pleitropic master regulator toolkit locus in a cell, it will activate the GRN, composed of thousands of genomic loci\
-in closely related taxa that have acquired a new CRE module, further diversity can occur by sequence diversity in a given CRE
A fossil lineage shows a gradual change in body size through time, but the
lineage never splits into two branches. Which macroevolutionary pattern does
this best represent?
Anagenesis
A single ancestral lineage rapidly gives rise to many descendant lineages
occupying different environments. Which pattern is most consistent with the
diagram?
Adaptive radiation
Two distantly related lineages independently evolve similar saber-toothed
morphologies. What concept best explains this pattern?
Convergent evolution
Two structures look similar because their ancestors inherited the same
underlying feature from a common ancestor. Which term applies?
Homology
A lineage loses a complex anatomical trait. Based on Dollo’s Law, which
prediction would be most appropriate?
Regaining the same complex trait will NOT occur
A fossil sequence contains numerous transitional forms showing relatively
continuous morphological change through time. Which model does this
support?
Phyletic gradualism
A fossil lineage shows long periods with little morphological change followed by
relatively rapid change associated with speciation. Which model best fits?
Punctuated equilibrium
In a punctuated-equilibrium diagram, most morphological change is
concentrated around branching/speciation events. What does this pattern
indicate?
Substantial phenotypic change can occur during speciation events
A lineage appears to have a gap in the fossil record between two forms.
According to the presentation, what is an important interpretation of such a gap
under punctuated equilibrium?
The gap can mean rapid evolutionary change with speciation.
Researchers compare neutrally evolving DNA sequences between two lineages.
The number of substitutions is used to estimate how long ago they diverged.
Which concept is being applied?
Molecular clock
Embryos of different species show the greatest conservation during an
intermediate developmental stage, while species-specific differences become
more pronounced later. Which concept matches this pattern?
Developmental Hourglass
A single gene influences eye, brain, and pancreas development. A mutation
affecting that gene therefore produces effects in several traits. What term
describes this?
Pleiotropy
A mutation changes gene expression in the pelvic region of a fish while leaving
expression of the same gene in other tissues relatively unaffected. What feature
of gene regulation makes this possible?
Modular CREs
Freshwater sticklebacks have reduced pelvic structures because a
pelvic-specific regulatory element is disabled, while other Pitx1 functions
remain. Why is this mechanism useful evolutionarily?
It changes one developmental context without disrupting other functions
Embryonic beak cells in finches with deep, heavy beaks show higher Bmp4
expression than those with small, pointy beaks. What does this demonstrate?
Differences in gene expression for development
Fruit fly Pax6 can rescue eye development in a mouse lacking functional Pax6.
What concept is most directly illustrated?
Deep homology
A conserved developmental toolkit gene is brought into a new tissue or
developmental context through a new regulatory element, contributing to a
novel trait. What process is this?
Cooption
The python limb-bud diagram shows Fgf2 and msx expression in mesenchyme
but no AER, and Dlx is not detected in the limb bud. What does this evidence
most directly illustrate?
Protein-coding genes never change during evolution
Hox genes encode:
transcription factors