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What are the 2 historical points where significant increases in the toolkit gene content evolved?
regulatory changes over gene invention and conserved toolkits
Homologs
genes/proteins/structures that share a common evolutionary origin
Orthologs
genes in different species that evolved from 1 ancestral gene via speciation event
Paralogs
genes within the same genome that arose via gene duplication event
Pseudogenization
1 gene copy accumulates loss of function mutation and becomes a non-functional pseudogene
Subfunctionalization
original gene had many functions/domains following duplication, each copy accumulates mutations disabling a subset of original functions
Neofunctionalization
1 copy retains ancestral function under constraint, redundant copy accumulates mutations, allowing it to acquire a completely novel function/expression
The evolutions of hox genes by the number
Cnidaria posessed 3 primitive hox genes → Bilateria 7/8 → protostomes ~10 → deuterostomes 14 → vertebrates up to 39
7 major ancestral Hox genes: 5 anterior, 1 central, 1 posterior
Living fossils
Extant organism that closely resembles ancient fossilized ancestors → allows inferences on genetic and physiological traits on common ancestors, morphological stasis, and deciphering the genetic toolkit
Evolution of Hox clusters
Invertebrate chordates (1 hox cluster) → 1R duplication split into 2 clusters, 2R duplication split into 4 clusters (a,b,c,d) and present in jawed vertebrates
Why are the outer genes the oldest in Hox clusters?
The 3’/5’ ends represent the ancestral pair whereas the center genes were added via duplication
What is the Parahox cluster?
sister gene to hox cluster: derived from endoderm
Colinearity: expressed anterior (Gsx), central (pdx), posterior (Cdx)
Clade
organism group including 1 common ancestor and all descendants
Basal
describes lineage that branches off earlier (more ancestor-like)
Outgroup
taxon closely related to, but outside of, studied organismal group
Tetraploidization events
Events where organism’s genome is doubled, resulting in 4n chromosomes
How is BLAST used as a tool to find orthologs and paralogs in animal genomes?
Aligns each search against target sequences and generates similarity/E (signifiance) scores to prove homology and not just similarity
Subfunctionalization of Gsb vs. Gsb-n
Gsb: retained for epidermal segment polarity and ectoderm cuticle patterning
Gsb-n: neural specialization within developing CNS
Who has the most toolkit genes of Cnidarians, Fruit flies, and humans?
Humans (39)