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Evolution's Witness
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Table of Contents

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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