A Quick Review from Earlier in the Course. to the Extent That We Saw Development It Was As a Constraint
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A quick review from earlier in the course. To the extent that we saw development it was as a constraint - The number of cervical vertebrae in mammals - Which digits are lost in lineages that evolve fewer digits Almost all mammals have 7 cervical vertebrae, it is not for a functional reason (swans have 22-25, ducks 16) -Hox genes are involved in axial patterning & regulating cell proliferation -Human children with embryonic cancers have a 125x risk of cervical ribs -Human children born with cervical ribs have a 120x risk of early childhood cancer -Taxa with more cervical vertebrae have lower metabolic and cancer rates (birds, reptiles, amphibs, manatees, sloth) -Pleiotropy of Hox genes prevents the evolution of novel # of cervical vertebrae in taxa with cancer risk Galis, F. 1999. Why do almost all mammals have seven cervical vertebrae? Developmental constraints, Hox genes, and cancer. J. Exp. Zool. / Mol. Dev. Evol. 285: 19-26. http://wwwbio.leidenuniv.nl/~galis/ Patterns of digit loss in tetrapods correlate 3 4 2 1 5 inversely with order of digit development 3 4 2 1 5 lost last lost first (perrisodactyls, horses in particular) Patterns of digit loss in tetrapods correlate 3 4 2 1 5 inversely with order of digit development Skink limbs 3 4 2 1 5 Chick limbs lost last lost first (skinks and chickens too) Patterns of digit loss in tetrapods correlate 3 4 2 1 5 inversely with order of digit development 1 2 3 4 5 1 2 3 4 5 lost last lost first (caudata) Exception that proves the rule A quick review from earlier in the course. To the extent that we saw development it was as a constraint - The number of cervical vertebrae in mammals - Which digits are lost in lineages that evolve fewer digits But the study of the connection between development and evolution goes far deeper than this. Phenotype: what an organism looks or acts like. Phenotypic evolution is usually what we care about the most. Genotype: the DNA of an organism - The phenotypes of organisms are determined (in part) by their genotypes. - Heredity is of genotypes not phenotypes Variation GENOTYPIC Heredity Selection PHENOTYPIC We must understand both genotypes and phenotypes; and their relationship. The process of development connects genotypes and phenotypes. Phenotype Development Genotype The relationship between genotype and phenotype is sometimes called the genotype:phenotype map. Studying this has been a long-standing part of biology in general; evolutionary biology as well. We will look at 3 approaches: 1. Comparative embryology - Haeckel, 1800s 2. Allometric studies - Gould & D'Arcy Thompson, 1900s 3. Molecular genetics - 1990s + The process of development connects genotypes and phenotypes. Phenotype Development Genotype The relationship between genotype and phenotype is sometimes called the genotype:phenotype map. Studying this has been a long-standing part of biology in general; evolutionary biology as well. We will look at 3 approaches: 1. Comparative embryology - Haeckel, 1800s 2. Allometric studies - Gould & D'Arcy Thompson, 1900s 3. Molecular genetics - 1990s + First approach - Haeckel, 1800s The first evolutionary approach to development was by Ernst Haeckel in the 1800s. His idea was that ontogeny recapitulates phylogeny; the developing organism develops through its previous evolutionary stages until it reaches its final form. By this model, evolution works by adding more developmental steps onto the end of the ancestral developmental program. Haeckel saw evidence for this in the similarity of developing mammal embryos to fish and amphibians. Next slide - Haeckel's famous drawing First approach - Haeckel, 1800s First approach - Haeckel, 1800s The first evolutionary approach to development was by Ernst Haeckel in the 1800s. His idea was that ontogeny recapitulates phylogeny; the developing organism develops through its previous evolutionary stages until it reaches its final form. By this model, evolution works by adding more developmental steps onto the end of the ancestral developmental program. Problems with Haeckel's model: - Adult features of ancestors not really seen (as predicted by Haeckel) - Some species skip developmental steps (e.g., direct developing sea urchins) - Development of traits doesn't proceed at same relative rates in all organisms (heterochrony) - Some species appear to cut development short instead of adding terminal steps (paedomorphic) These weaknesses were known fairly quickly; but better tools (math, DNA) to explain development had to wait until 20th century. The process of development connects genotypes and phenotypes. Phenotype Development Genotype The relationship between genotype and phenotype is sometimes called the genotype:phenotype map. Studying this has been a long-standing part of biology in general; evolutionary biology as well. We will look at 3 approaches: 1. Comparative embryology - Haeckel, 1800s 2. Allometric studies - Gould & D'Arcy Thompson, 1900s 3. Molecular genetics - 1990s + Second approach - allometry, 1900s As organisms grow (develop) different parts of them grow at different rates - this leads to different shapes. Maybe small changes in the relative growth rates (due to genotypic changes) can cause big differences in the final form (phenotype)... We can plot the sizes of height traits over time (age) and the resulting curve is termed an ontogenetic trajectory. length Second approach - allometry, 1900s We can plot the sizes of traits over time (age) and the resulting curve is termed an ontogenetic trajectory. Different developmental rates or durations would result in different final shapes head head body body Second approach - allometry, 1900s Hypermorphosis 4 types of Acceleration developmental change: - Hypermorphosis - Progenesis Log(head) - Acceleration - Neoteny 2 phenotypic results: Progenesis - Peramorphosis - Paedomorphosis Neoteny Log(body) Second approach - allometry, 1900s Many salamanders have gills when juveniles and then as they mature they lose the gills. Some species of salamanders have gills as adults - neoteny causing paedomorphosis juvenile Ambystoma mabeei adult Ambystoma mabeei adult Necturus spp. Second approach - allometry, 1900s Irish elk antlers are HUGE But they are proportional to the size of the body along a height:antler allometric curve - hypermorphosis creating peramorphosis Antler size Shoulder Height Second approach - allometry, 1900s Second approach - allometry, 1900s A more sophisticated prediction of relative growth rates can be done by using a grid and altering growth in X and Y directions in each grid. (D'Arcy Thompson) This approach can help us understand the mola mola, one of the weirdest fish in the world Second approach - allometry, 1900s A more sophisticated prediction of relative growth rates can be done by using a grid and altering growth in X and Y directions in each grid. (D'Arcy Thompson) Pufferfish and the mola mola are in the same order, fairly closely related. Similarities in the skeleton are apparent. Second approach - allometry, 1900s A more sophisticated prediction of relative growth rates can be done by using a grid and altering growth in X and Y directions in each grid. (D'Arcy Thompson) Pufferfish and the mola mola are in the same order, fairly closely related. Developmental mutations that change rates of growth in different regions can "transform" one into the other. Second approach - allometry, 1900s If we reverse previous head:body ratio; head starts relatively large and grows slower than body - neoteny would therefore lead to relative larger head for the body Second approach - allometry, 1900s If we reverse previous head:body ratio; head starts relatively large and grows slower than body - neoteny would therefore lead to relative larger head for the body Selection for increased intelligence (via larger relative brain size in adults) could act via neoteny leading to paedomorphosis - the adult looking like the younger version of ancestral species. Do human heads look like juvenile chimpanzees/human ancestors? Thompson's grid? Second approach - allometry, 1900s Second approach - allometry, 1900s Human faces may have arisen via neoteny from ancestral primate Cause or correlation? Other correlations? The process of development connects genotypes and phenotypes. Phenotype Development Genotype The relationship between genotype and phenotype is sometimes called the genotype:phenotype map. Studying this has been a long-standing part of biology in general; evolutionary biology as well. We will look at 3 approaches: 1. Comparative embryology - Haeckel, 1800s 2. Allometric studies - Gould & D'Arcy Thompson, 1900s 3. Molecular genetics - 1990s + Third approach - Molecular, 1990s Molecular developmental evolution Using modern molecular tools to look directly at genotypic differences and their effect on phenotype - specifically how when and where genes are expressed is altered. Regulatory sequences control gene expression during development. Mutations in these sequences can change the function of a protein. biochemical function vs evolutionary function It is now thought that the evolution of regulation may be a more potent force for evolution than evolution of protein biochemical function. What is a gene anyway? Third approach - Molecular, 1990s Transcription factors (trans elements) are proteins that bind to binding sites in the DNA (cis elements), turning the gene on or off. The sequence of each binding site is critical to the binding of the trans elements. Evolution of these binding sites can lead to changes in gene expression during development. The trans