5/4/05 Biol 473 lecture

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1 5/4/05 Biol 473 lecture animals shown: anomalocaris and hallucigenia 1

2 The Cambrian Explosion MYA THE BIG BANG OF ANIMAL EVOLUTION Cambrian explosion was characterized by the sudden and roughly simultaneous appearance in the fossil record of many diverse animal forms MYA. No other period in the history of life can match this remarkable burst of evolutionary creativity Marks the appearance of abundant life as recorded by an abundance of marine fossil life Marks the appearance of invertebrates with mineralized (calcium carbonate) skeletons [on formerly soft-bodied organisms] The Cambrian Explosion: Before and After What major adaptive features were in place before the Cambrian explosion? 2

3 What major adaptive features were in place before the Cambrian explosion? Before and after the event all life lived in the oceans The major adaptive features that were already present: 1. eukaryotic cell 2. sexual reproduction 3. multi-cellular organisms with soft bodies 3

4 No other period in the history of life can match this remarkable burst of evolutionary creativity BILATERAL ANIMALS WITH LIGHT SENSING ORGANS APPEARED Opabinia (lower right) has five eyestalks! 4

5 hallucigenia and Opabinia Animals appeared in the fossil record with a clearly distinguished front end and back end TRILOBITE The body plans of all major extant animal phyla arose during this time period. Many different lineages acquired complex anatomies and hard parts (the exoskeleton) at the same time 5

6 What factors triggered the Cambrian explosion? The Cambrian record of life is in sharp contrast with that of the preceeding eons The Cambrian appearance of fossils representing diverse phyla has inspired hypotheses about possible genetic or environmental catalysts of early animal evolution: The genetic toolkit hypothesis o The Cambrian explosion was ignited by the evolution of a modern genetic toolkit that was complex enough to facilitate elaborate diversification of body plans The environment/ecological hypothesis o The genetic toolkit was in place well before the Cambrian radiation (that is predated the Paleozoic Era) o The Cambrian explosion was triggered by environmental perturbations and amplified by ecological interactions within reorganized ecosystems 6

7 What is meant by the Genetic Toolkit? Housekeeping genes: genes that encode proteins that function in essential processes in all cells in the body Such as? Roomkeeping genes: Other genes encode proteins that carry out specialized functions in particular cells or issues Such as? The genetic toolkit: genes that govern the construction of the house Or in other words whose protein products determine the overall body plan and the number, identity and pattern of body parts How to identify such genes? What are the Old and new paradigms? 7

8 Old and new paradigms: 1. the geneticist: search for developmental monsters 2. the biochemist: reconstruct in vitro -- obvious problems here WHY? 3. molecular biologist: let some else do the bench work and use genomic sequence to find genes of interest in your organism of interest 8

9 Critical Features of the Toolkit OR What we know so far 1. The toolkit is composed of a small fraction of all genes 2. Most toolkit genes encode transcription factors or components of signaling pathways and act directly or indirectly to control the expression of other genes 3. The spatial expression of toolkit genes is often closely correlated with the region of the animal in which the gene functions 4. Toolkit genes can be classified according to the phenotypes caused by their mutation: o body axis specification o formation and identity of spatial fields o specification of a specific organ (such as the eye) 5. Many toolkit genes are widely conserved among different animal phyla 9

10 Many members of the genetic toolkit are homeobox or zinc finger genes Genome-wide analysis of DNA-binding motifs found in eukaryotic transcription factors 10

11 The homeobox family: a transcription factor family homeodomain is a ~60 amino acid sequence containing many basic residues forms a helix-turn-helix motif that binds specific sequences in DNA the homeodomain is coded for by the homeobox region of the gene Helix-3 contacts the major groove. The specific amino acid sequence of helix-3 determines the DNA binding specificity of the homeodomain protein Protein:DNA complexes: picture and movie gallery 11

12 Homeodomains are highly similar 60 amino acid regions of proteins made by all homeobox gene.. Deviations from the consensus are shown for four fly (top line) and mouse (bottom line) homeodomain proteins. A dash means the sequence matches that of the consensus. All homeobox genes contain the conserved homeobox region. Outside the homeobox, there may be very little sequence conservation between two homeobox genes 12

13 Homeotic genes homeobox originally named for Drosophila homeotic genes: mutations in these genes transform one body part into another genes with a homeobox often are involved as developmental regulators, but possession of a homeodomain does not guarantee a role in development not all mutants are homeotic bithorax mutant phenotype Homeotic genes are part of a hierarchy of regulators that define spatial location in the developing fly embryo If building an embryo, what are the first decisions that need to be made? 13

14 A subset of homeobox genes are called the HOX genes o found in linked clusters o role in specifying location along the AP axis AP/DV genes: homeobox genes, zinc fingers, signal transduction genes Segmentation genes: zinc fingers, homeobox genes HOX gene control the expression of each other and genes necessary for assembling the specialized structures and tissue in each segment 14

15 OK, so what does this tell us about the molecular definition of spatial identity in other animals such as vertebrates? 15

16 16

17 Remarkable observations: 1. Homeobox-containing genes have been found in all metazoan organisms examined as well as in yeast and plants 2. In all metazoans, a subset* of the homeobox genes are organized into gene clusters that are colinear with the Drosophila gene clusters (next figure). Thse clusters are called HOX complexes 3. The relative order of a gene within each vertebrate HOX complex is correlated with its spatial expression along the anteroposterior body axis (next figure) 4. *Orphan homeobox genes are not part of the HOX complex 17

18 In the mouse embryo, four complexes of HOX genes (39 genes in all) occur on four different chromosomes. Not every gene is represented in every complex. The HOX genes are expressed in distinct domains along the AP axis SEE also Watson Box

19 Homeodomain Consensus sequence at top of figure For each box: first line is a fly homeodomain and the second line is a mouse homeodomain. In each box, the homeodomains are more closely related in sequence to each other than they are to other homodomains from the same organism Numbering of mouse genes is different from the preceding figure 19

20 HOX genes are widely conserved among different animal phyla 20

21 The genetic toolkit: genes that govern the construction of the house: or in other words whose protein products determine the overall body plan and the number, identity and pattern of body parts Many toolkit genes are widely conserved among different animal phyla The striking correspondence between the gene clusters in flies and mammals and other animals suggests that they represent the descendants of an ancestral cluster of homeobox genes already present in the common ancestor of insects and vertebrates and other bilateral organisms -- which would have predated the Cambrian explosion 21

22 So what did the genetic toolkit look like for the last common ancestor of all bilaterally symmetric organisms? What other genes are shared by all descendants? Rebuilding Urbilateria: the hypothetical last common ancestor of all bilaterans 22

23 A plausible chronology for animal locomotory abilities, visual function and the occurrence of master control genes. The need for fast locomotion and vision are likely to have originated during the Cambrian explosion, whereas master control genes in general would have been needed much earlier. Bilaterally symmetric animals with priminent eyes appeared suddenly in the fossil record from the Cambrian explosion Eyes were image-forming and vision was binocular 23

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