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Lecture 3: Origins of Animals
Developmental, molecular and paleontological perspectives
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What is an animal? Multicellular adults, heterotrophic, eukaryotes. Most feed by ingestion of food into a gut Cells lack cell walls, adhere by means of specialised junctions, secrete extra-cellular matrices using collagen. Most animals possess specialised cells for: a) signal transmission - nerves b) contraction for movement - muscles
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Animal Life History Typically a small flagellated sperm fertilises a larger egg to form a diploid zygote. Zygote undergoes cleavage. Formation of a blastula Blastula undergoes gastrulation during which embryonic tissue layers form Many animals have distinct larval stage
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Figure 32.1 Early embryonic development (Layer 1)
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Figure 32.1 Early embryonic development (Layer 2)
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Figure 32.1 Early embryonic development (Layer 3)
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Figure 32.2 A choanoflagellate colony
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Figure 32.3 One hypothesis for the origin of animals from a flagellated protist
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Figure 32.4 A traditional view of animal diversity based on body-plan grades
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First major split Parazoans (no true tissues) versus
eumetazoa (all other animals)
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Second major split Radial versus bilateral symmetry
Cnidaria (jellyfish and anemones) and Ctenophora (comb jellies) versus rest of animals Bilateral symmetry associated with cephalisation (development of a head) Some bilateral animals have acquired radial symmetry (e.g., some echinoderms)
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Figure Body symmetry
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Third split: body cavities
Acoelomate (no cavity) Pseudocoelomate (partially lined with mesoderm) Coelomate (cavity completely lined with mesoderm)
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Fourth split: Protostomes vs. Deuterostomes
Different type of cleavage Different origin of coelom Different fate of blastophore (becomes either mouth or anus)
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Figure 32.4 A traditional view of animal diversity based on body-plan grades
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Figure 32.8 Animal phylogeny based on sequencing of SSU-rRNA
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Lophophorate animals Bryozoans, brachiopods, and phoronids
Mouth surrounded by hollow tentacles (lophophore)
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Figure 32.9 A trochophore larva
Larva found in molluscs and annelid worms
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Arthropods and nematodes peridically shed their cuticle.
Figure Ecdysis Arthropods and nematodes peridically shed their cuticle.
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Figure 32.12 Comparing the molecular based and grade-based trees of animal phylogeny
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Why are animals so successful and so complex?
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What genes can tell us Sequence comparison tells us about evolutionary relationships Can also tell us about homology and development Question: what does genomics tell us about animal evolution?
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HOX genes Regulate development of segmentation in animals
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Gene and genome duplications
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The Cambrian Explosion Myth or reality?
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Cambrian Explosion Nearly all major animal phyla appear in Cambrian rocks (545 to 525 million years ago) Many weird and wonderful creatures appear suddenly in the fossil record Question: Is the fossil record giving us a faithful picture of what happened.
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First appearance of animals in the fossil record
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Nemiana > Sea anemonea? Algal? < Spriggina Annelid Worm?
Arthropod? Cyclomedusa > Benthic Polyp 1-5 mm in diameter
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< Dickinsonia Annelid Worm? Cnidarian? Charnia > Sea pen- up to 1m in length
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Burgess Shale and the Cambrian Explosion
British Colombia, Canada Discovered in 1909 by Charles Walcott Deep water deposit Around 515 million years (10 million years after Cambrian explosion)
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Stephen J. Gould “Wonderful Life”
Major body plans laid down very quickly Much more diversity in Cambrian than we see today Survival of a subset of phyla, due to luck more than anything else
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Did the Cambrian explosion really happen?
"Cambrian explosion" model Molecular data Today Molluscs Annelids Arthropods Echinoderms Jawless fish Gnathostomata Molluscs Annelids Arthropods Echinoderms Jawless fish Gnathostomata 500 Myr 1000 Myr 1500 Myr
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Fossil record by itself
A B C 10 Millions of years 20 at face value, group is about 10 Myr old
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Fossils + molecular data
B C A B C 1 1 % 2 2 3 3 4 4 calibration molecular divergence 1% per 10 million years
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A B C Extrapolation 10 20 Millions of years If rate of molecular evolution has been constant, then group is about 40 Myr old 30 40
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“Phylogenetic Fuse” Molecular dating suggests origins prior to diversification. Keep in mind serious concerns about molecular dating. Cambrian phylogenetic fuse Cooper & Fortey 1998 TREE
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Summary Major divisions of animals defined by fundamental body plans
Molecular data challenges some traditional views about relationships Much debate about when animals first evolved
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