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Evolution's Witness
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Table of Contents Table of Contents Foreword: Russell Fernald PhD Acknowledgments Introduction Prologue Molecular genesis Hadean 4600-3750 million years ago Chapters 1. The age of first cellular life Archean 3750-2500 million years ago Prokaryotes Early cells--protobionts Cellular Evolution First life First witness Phylum Cyanobacteria The road to cellular success Retinal Photolyases and cryptochromes Sunlight and blue light Beginning to organize Genetic machinery--the toolkit Prokaryotic gifts Further organization 2. The age of complex cellular life Proterozoic Early life (2500-543 million years ago) Inception of Eukaryotes Nucleated Kleptomaniacs Euglena gracilis Erythropsidium Bridging the Gap to Metozoa Pre-Metazoa Volvox carteri 3. Eukaryotes organize and metozoans arise Neoproterozoic Cryogenian--850-650 million years ago Ediacaran--650-543 million years ago 1000-543 million years ago Multicellular animals Trichoplax adhaerens Corals Diploastrea heliopora Goniopora species Cubozoan Jellyfish Tripedalia cystophora Ciliary and Rhabdomeric Photoreceptor Cells Sensory input The eye and the brain Which came first? The making of an eye Photoreceptors The Crystalline lens Extraocular Muscles and other structures(Adnexa) Metazoans and their eyes 4. Early animals prepare the ground Ediacaran Period 650-543 million years ago A major genetic step Urbilaterians Another major genetic step Wormy beginnings Genetic development of eyes Platyhelminthes (polyclad flatworms) Pseudoceros dimidiatus Annelids Platynereis dumerilii Sabellids Megalomma sp How Photopigment Creates a Signal Uribilaterians Brittle stars Ophiocoma wendtii Mollusks take the stage Kimberella quadrata Acanthochiton communis Chitons Tonicella lineata Small Shelly Fauna 5. Vision's big bang blazes the trail Early Paleozoic Era Cambrian Explosion 543-490 million years ago The Burgess Shale Invertebrates in ascendancy What is Adenosine Triphosphate (ATP)? The first eye Trilobites Olenellus Phacops rana milleri Phacops rana crassituberculata Trilobite eyes The f-Number Erbenocile erbeni More invertebrates and their eyes appear Opabinia Anomalocaris Waptia Adoraia Alta Ecolsozoa and Lophotrachozoa Tardigrades Echiniscus testudo Onychophora--The Velvet Worm Epiperipatus biolleyi Onychophoran Eye 6. The age of Arthropods: A major phylum begins Early Phanerozoic Paleozoic Era Cambrian Period 543-490 million years ago Compound eye development Horseshoe Crab Limulus polyphemus Apposition Eyes Mantis shrimp-stomatopod Odontodactylus Scayallurus Evolution of the Apposition Eye to Superposition Eye Refracting Superposition Compound Eyes Long-bodied Decapod Crustaceans Spot prawn Pandalus platyceros Reflecting Superposition Eyes Tasmanian rock lobster Astacopsis gouldi Mysid shrimp Dioptromysis paucispinosa The Crabs Hermit Crab Strigopagurus strigimanus Parabolic Superposition Eye 7. Vertebrates gain a foothold Cambrian 543-490 million years ago Phylum Chordata Hagfish Epatratus stoutii First true vertebrates Conodonts Building an eye Southern Lamprey Geotria australis Short-headed lamprey Mordaciidae Tapetum Why Color vision? 8. Shelly fauna rule the sea Paleozoic Era Ordovician 490-445million years ago Mollusa Bivalvia Scallops Aequipecten irradians Giant Clam Tridacna gigas Cephalopoda Nautilus Gastropoda Conchs Strombus galeatus 9. The piscine eye develops Silurian 445-415 million years ago Vertebrate expansion Placoderms Dunkleosteus Elasmobranchs - Taeniura lymma Scalloped hammerhead shark Sphyrna lewini 10.The piscine eye matures Early Devonian Period 415-362 million years ago Bony fish appear Anatomy of Piscine eye The outer coats of the eye Sclera Cornea Spiny Puffer Diodon holocanthus The Flyingfish Hirundichthys albimaculatus The Lens Crystalline lens Koran angelfish Pomacanthus semicirculatus Accommodation Matthiessen's Ratio Cuatro Ojos Anableps anableps Oil droplets Bluefin Tuna Thunnus thynnus Extraocular muscles Northern Stargazer Astroscopus guttatus Silverstripe Mudskipper Periophthalmus argentilineatus Striped Marlin & White Marlin Tetrapturus albidus & audax Retina Photoreceptor design Double Cones Twin Cones Retinal vascularization Retinomotor Pigment movement Argenta Neurology and optics The African Butterfly Fish Pantodon buchhlozi Sanddab Citharychthys sp Habitat expansion Barreleye Macropinna microstoma Javelin Spookfish Bathylychnops exilis Stylephorus chordatus Malacosteus niger Sandlance Limnichthyes fasciatus Piscine camouflage Bullethead parrotfish Chlorurus sordidus 11. Insects arise to fly Paleozoic Era Early Devonian 415-398 million years ago Arthropods come ashore The Superposition Eye Mayflies Baetidae sp. The apposition eye Dragonflies The Green Darner Anax junius The Blue Dasher Pachydiplax longipennis Flame Skimmer Libellula saturata 12. Stealth, Speed and Predation Paleozoic Era Devonian 415-362 million years ago Arachnida Wolf spider Lycosid sp. Jumping spiders Portia fimbriata Net-casting spider Deinopis subrufa 13. The age of Tetrapods and Terrestrials Late Devonian Period 385-362 million years ago, Vertebrate Animalia comes ashoreEL. Vertebrate quest for land Early tetrapodian eyes Lungfish Neoceratodus forsteri Tapetum The challenge of a terrestrial environment Frogs Phyllomedusa bicolor Eyelids and the lacrimal system Extraocular muscles Tear Glands Cornea and lens Retina and vision Neurologic changes North American Wood Frog Rana sylvatica Consolidation of the assumption of land 14.Terrestrial life flourishes Carboniferous 362-299 million years ago Permian 299-251 million years ago Vertebrates Synapsids and their eyes Diapsids and their eyes Invertebrates Giant Ceiba Beetle Euchroma gigantean Lantern Click Beetle Pyrophorus phosphorescens Tiger Beetle larvae Tetracha carolina Whirligig Beetle Gyrinidae Permian extinction 15. Reptiles push the ocular envelope The age of reptiles Mesozoic Era Triassic 251-208 million years ago Synapsids Sauropsids Anapsids Diapsids Adnexa of the reptilian eye Eyelids Nictitans Lubricating glands Extraocular muscles Reptilian eye Cornea Uvea Crystalline lens Parson's Chameleon Calumma parsonii Iris Tokay Gecko Gekko gekko Retina Basilisk Basiliscus galeritus Lepidosaurs Tuatara Sphenodon punctatus Archosaurs 16. March of the Archosaurs Mesozoic Era Triassic 251-208 million years ago Jurassic 208-145 million years ago Archosaurs The Order Testudines (turtles, terrapins, and tortoises) Turtle Eye Leatherback Turtle Dermochelys coriacea Red-eared Slider Trachemys scripta elegans Crocodilians American Alligator Alligator mississippiensis Archosaurian sisters Ichthyosaurs Ophthalmosaurus 17. Dinosaurs and their companions Mesozoic Era Triassic 251-208 million years ago Jurassic 208-145 million years ago Pterosaurs Dinosaurs Sauropods Theropods 18. Cephalopods change direction Mesozoic Era Jurassic 208-145million years ago Coleoids Cephalopod lens Extraocular muscles Coleoid Expansion Pacific Giant Octopus Octopus dofleini Rhabdomeric arrangement Humboldt Squid Dosidicus gigas Histioteuthidae sp Histioteuthis heteropsis Giant Cuttlefish Sepia apama Pacific Bobtail Squid Euprymna scolopes 19. Snakes arise from the ground Cretaceous (145-65million years ago) Fossorial lizards Recreating an eye Rowley's Palm viper Bothriechis rowleyi Banded sea krait Laticauda colubrine 20. The Age of Birds - The eye is taken to great heights Mesozoic Era Cretaceous (145-65 million years ago) Tertiary Period (65-2million years ago) Birds arise Globe morphology Globose Globe Flattened Globe Australian Darter Anhinga melanogaster Tubular Eye Western Screech Owl Megascops kennicotti Orbital size and contents Visual fields American Woodcock Scolopax minor Papuan Frogmouth Podargus papuensis Eyelids Nictitans How the nictitans works Peregrine falcon Falco peregrinus Cornea Iris and pupil Accommodation Uvea: Choroid, ciliary body, iris Retina Visual Processing Oil Droplets Macular design Area Centralis Single fovea Infula Flamingo Phoenicopterus ruber Convexiclivate fovea-Single deep fovea Bifovate retina White-fronted Bee Eater Merops bullockoides African Fish Eagle Haliaeetus vocifer Green and Rufous Kingfisher Chloroceryle inda Pecten Bush Stone Curlew Burhinus grallarius Pileated Woodpecker Dryicioys pileatus Neurologic evolution Barn Owl Tyto alba 21. Pollinators Co-Evolve Mesozoic and Cenozoic Eras Jurassic 208-145 million years ago Cretaceous 145-64 million years ago Tertiary 65-2 million years ago The Earth in bloom A bounty for insects Caddisfly Trichoptera Lepidoptera Tobacco hornworm moth Nicotiana sp Hummingbird hawkmoth Mcroglossum stellatarum The Union Jack Delias myisis Afocal Apposition Eye Ommatidial adaptations Social Hymenoptera Wasp, Bees, Ants, and Sawflies Fairy wasp Gonatocerus ashmeadi Carpenter Bee Xylocopa micans Apposition Eye Resolution An Evolutionary Twist Megalopota genalis True Flies and a new eye Diptera Stalk eyed fly Achias longividens Neural superposition eye Brachyceran flies Robber flies Diogmites sp Trioria interrupta Male Scale insect oidae sp. in amber Puto albicans 22. Mammalia diversifies Mesozoic and Cenozoic Eras Mid to late Jurassic 161 to 145 million years ago) Cretaceous 145-65 million years ago Early to mid Tertiary Period 65-56 million years ago) Mammals thrive Monotremes Platypus Ornithorhynchus anatinus The descent of color vision Echidna Tachyglossus aculcatus Zaglossus bruijni Marsupials Fat-tailed Dunnart Sminthopsis crassicaudata North American Opossum Didelphis virginiana Brush-tailed Possum Trichosurus vulpecula 23. The Age of Mammals Mesozoic and Cenozoic Era Late Cretaceous 100-65 million years ago Tertiary 65-1.8 million years ago Quaternary1.8 million years ago to present Mammals extend their dominance Placental Mammals Eastern tube-nosed fruit bat Nyctimene robinsoni Primate tuning of color vision? Black and White ruffed lemur Varecia variegata variegata Tarsier Tarsier spectrum 24. Planktonic soup evolves Cenozoic Era Tertiary 65 to1.8 million years ago Quaternary 1.8 million years ago to present Tiny aquatic Arthropods Testing the limits of eye size Copepods Copilia quatrata Pontella scutifer Transformation of larval eyes Vent shrimp Rimicaris exoculata 25. Mammals return to the sea Tertiary 65-1.8 million years ago Quaternary 1.8 million years ago to present Aquatic Mammals Bottlenose Dolphin Tursiops truncatus Harbor seal Phoca vitulina 26. The visual witness and a conscious brain Cenozoic Era Quaternary period 1.8 million years ago-present The human eye The direction of the visual witness Appendices A The human eye-a camera style eye B Extraocular muscles Medial rectus Superior oblique Retractor bulbi Trochlea C Retinal Vascularization Evolution of retinal vascularization Mammalian forms of retinal vascularization D Evolution of the cornea and ocular coats Safeguarding precious contents E Accommodation F Crystalline lens G Photoreceptor cells H Neurologic evolution in birds References by Chapter Glossary Index

About the Author

Ivan R. Schwab M.D. is currently a professor at the University of California, Davis where he has worked as an Ophthalmologist for over twenty years, and was on the faculty at West Virginia University for seven years before coming to UCD. His strong interest in biology and natural history has led him to investigate a diverse range of topics including ocular stem cells, bioengineered tissues for the eye and comparative optics and physiology. He has published extensively in these fields, with three previous books to his credit, and he was the winner of the 2006 IgNobel for Ornithology. He has combined those interests with one in evolution to produce this text on the evolution of the eye.

Reviews

This book shows what can be achieved by combining insights from sensory physiology with anatomy, phylogeny and the fossil record across the widest range of organisms to document the evolution of a biological system. It demonstrates how the study of living forms can successfully be used to interpret fossil ones, and vice versa. The book's magnificent sweep is all-encompassing, and remarkably up-to-date for such a cross-disciplinary work. It's rare for one person to have such a broad background, but Schwab has achieved this, such that the work provides an example for future studies of the kind. -Prof. Jennifer A. Clack, ScD, FRS Professor and Curator of Vertebrate Palaeontology, University Museum of Zoology An astounding erudite and exciting visual description of eye evolution, something every inquisitive biologist, veterinarian, neuroscientist, or ophthalmologist should have in their library. This book contains 400 illustrations that define and refine the text providing a unique look at the how the eye was achieved. It is truly one of a kind. Bruce E. Spivey, M.D., M.S., M.Ed. President, International Council of Ophthalmology Schwab's lavishly illustrated book documents the amazing proliferation of eyes across the animal kingdom, in all their variations and all their splendour, and it explains the pathways by which these eyes have evolved. Aimed at the non-specialist but intelligent reader, the book begins with the early evolution of life on earth, and sets the scene for the advent of eyes that took place some 500-600 million years ago (mya). By combining fossil evidence with information from extant "primitive" organisms, Schwab explains current ideas about the simple animals that were present during the Ediacaran period (around 600 mya), about the light-detection mechanisms and the genetic machinery that they possessed, and about the split of these primitive animals into two major divisions - comprising on the one hand most invertebrates and on the other hand our own line that led to vertebrates and mammals. (cont'd below) High on the agenda is the invention, around the time of the Cambrian explosion in body forms (more than 500 mya), and the subsequent re-invention, of the two radically different forms of eye: the camera-style eye (as we have) with a single lens, and the compound eye, with multiple repeated units each having its own lens or mirror. Using beautiful images, Schwab charts the myriad variations on these two themes that have been employed by countless species, extinct and extant, over the course of 500 million years of global experimentation, refinement, and rejection. The result is a stunning book that will serve both to introduce non-specialists to the concepts of evolution and eye evolution and also as a reference work for experts. Trevor D Lamb Professor of Neuroscience John Curtin School of Medical Research The Australian National University Canberra, Australia The evolution of the eye was unquestionably one of the most important innovations in the history of life. Ivan Schwab has synthesized a huge array of disparate information to provide us with an indispensable guide through the complexities of visual systems throughout the animal kingdom. Richard Fortey FRS FRSL "Which leads to a just published book -- a most remarkable book -- that needs to be in the library of every school and college, and maybe in every household ...There aren't many books like this one, transformative books that provide a wondrous experience -- especially for young people -- just by turning the pages of mesmerizing illustrations of the evolution of the eyes that are looking at the book... A deep majestic event: human eyes looking at a book explaining the evolution of human eyes. (The only event that I know of with more majesty is the human brain contemplating its own evolution.)" -- Huffington Post "Evolution's Witness: How the Eyes Evolved is an outstanding book. It can be highly recommended. Every neuroophthalmologist should strongly consider adding this book to their library. It would also be valuable for ophthalmologists, neurologists, and neurosurgeons interested in the field of evolution." -- Walter M. Jay, MD, Neuro-ophthalmology "An all-inclusive glossary and chapter-specific references accommodate diverse readers of this captivating text. Appendices detailing evolution of ocular muscles, retinal blood supply, cornea, lens, accommodation, and photoreceptors prove in valuable for specialists and generalists alike. Evolution's Witness, in whole or part, is an essential mainstay for multiple disciplines, irreplaceable for vision scientists and eye care professionals." -- Jeff C. Rabin, The Journal of the American Academy of Optometry "Evolution's Witness can serve a variety of uses. For the scholar of vision, it is a comprehensive reference but also a source of inspiration and ideas for new ways to study myriad aspects of vision. For the instructor, it would function well as a textbook in a seminar, since it is laid out in a didactic and accessible way, with clear and engaging writing. With its sumptuous illustration, it would make a great coffee-table book for anyone with an interest in biology. The book is also an implicit riposte to the fallacy that eyes are too complex to have evolved by natural selection, so it will also be useful for those engaged in countering creationist agendas. The book has helpful appendices on detailed anatomical matters that are discussed at many points in the book." -- Daniel Graham, Perception "This comprehensive and beautifully illustrated text should sit on the shelves of all students, graduates and academics with an interest in evolution, adaptation, neural plasticity and natural history. For those more intimately working in the field of visual neuroscience (including optometrists and ophthalmologists), it provides a unique and holistic perspective that provides the context in which we are all working. The book will also emphasize how inadequate the human visual system is in many ways and how much we can learn from nature. Therefore, I highly recommend you to witness the 'inimitable contrivances' of the eye in evolution and appreciate the exquisite complexity of how different organisms form an optical image of the world and convert this to a neural image that is recognizable by the central nervous system." -- Clinical and Experimental Optometry

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