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This document provides a historical overview of comparative vertebrate anatomy (CVA), beginning with early Greek physicians' observations and studies, continuing up to modern developments and evolving concepts of biological systems.

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History of CVA Aristotle (384-322 BC) The Beginning Father of Biology 500 BC ○ A Greek philosopher and polymath, Greek Physicians and Philosophers Be...

History of CVA Aristotle (384-322 BC) The Beginning Father of Biology 500 BC ○ A Greek philosopher and polymath, Greek Physicians and Philosophers Begin Aristotle made significant Systematic Observation through Animal contributions to many fields, Dissection including biology. He laid the ○ Greek physicians and natural foundations for comparative philosophers such as Alcmaeon, anatomy with his detailed studies of Croton, began cutting open animals animals. for naturalistic research, marking the Herophilus (c. 335-280 BC) and beginning of systematic observation Erasistratus (c. 304-250 BC) (500 BC) ○ Hellenistic physicians who were ○ Among earliest Greek physicians among the first to conduct detailed and natural philosophers, anatomical human dissections. Their works observations of animals were contributed greatly to early anatomy, scattered. (from 500 BC onwards) though most of their writings have ○ Xenophanes of Colophon described been lost. the fossilized remains of small Galen (c. AD 129-c. 216) marine animals and plants. ○ A prominent Greek physician, 400 BC Galen’s work in medicine and Early Comparative Anatomy and Animal anatomy dominated the medical field Dissection by Alcmaeon, Calcidius and for many centuries. He dissected Democritus animals to understand the human ○ Calcidius (fl.c. AD 400), Alcmaeon of body, as human dissection was not Croton (fl.c. 500 BC) was the first to common during his time. cut open the animal body for Physiologus (c. AD 140) purposes of naturalistic research ○ Although not a person, ○ Democritus of Abdera (c. 460-400 “Physiologus” is a Christian text that BC), may have directly compared originated in Alexandria, Egypt. It fish to other (land) animals, reporting compiles descriptions of animals and enigmatically that “common fish became widely influential in have no backbones (vertebrae)” Christian Europe 360 BC Nemesius of Emesa (c. AD 400) Plato’s Timaeus Explores Animal Anatomy, ○ A Christian bishop whose work “On Paving the Way for Aristotle’s Studies the Nature of Man” synthesized ○ Plato discusses the anatomical Greek philosophical and medical structure of various animals in his traditions particularly those of work Timaeus, laying the conceptual Aristotle and Galen, with Christian groundwork for later studies by thought. Aristotle. Constantine the African (d. 1098) 384 - 322 BC ○ A Muslim physician from Tunisia Aristotle’s Foundational Research in who converted to Christianity and Comparative Anatomy - Establishes became a monk in Italy. He Systematic Study of Animal Anatomy to translated many Arabic medical texts Enhance Understanding of Human Anatomy into Latin, helping to revive ○ Aristotle conducts research in Greco-Arabic knowledge in Europe. comparative anatomy, dissecting Averroes (Ibn Rushd) (1126-1198) and vivisecting a wide variety of ○ A Muslim philosopher from Spain, animals. History of Animals (HA), Averroes wrote extensive Parts of Animals (PA) AND commentaries on Aristotle’s works, Generation of Animals (GA) influencing both Islamic and establish a systematic approach to Christian scholars. the study of animal anatomy, Michael Scot (c.1175-c. 1235) (classifies and highlights the ○ A Scottish scholar who translated importance of studying animal Aristotle’s biological works from anatomy to understand human Arabic into Latin, contributing to the anatomy better) revival of Aristotelian science in ○ Herophilus and Erasistratus at the Europe Museum of Alexandria, make Albertus Magnus (c. 1193-1280) significant advancements in ○ A Dominican friar and scholar, understanding the nervous system Albertus Magnus wrote and perform systematic human comprehensive works on natural dissections and vivisections science, including biology. He used The Middle Ages Michael Scot’s translations and Ulissi Aldrovandi added his own observations. ○ Monstrorum historia - a comparative Gersonides (Levi ben Gershom) teratology (1288-1344) 1628 ○ A Jewish philosopher from William Harvey Provence, Gersonides wrote a “A new theory that the animal heart pumps super-commentary on Averroes’ blood which circulates through the arteries works, further advancing the study of back to the heart through the veins” Aristotle’s biology. Harvey’s Motion of the Heart and Blood Mondino de Luzzi (c. 1275-1326) (Exercitatio Anatomica de Motu Cordis et ○ An Italian physician who authored Sanguinis in Animalibus) was published “Anatomia Mundini,” the oldest 1662 - 1664 known manual on human dissection. Rene Descartes His work marked a significant step Descartes’ Treatise on Man, conception of forward in the study of human animals as mechanical beings was anatomy in medieval Europe. published in 1664 Renaissance 1660 The term “Renaissance” is French for Robert Boyle “rebirth.” It earned this name because it (Develops Various Techniques to Preserve marked the start of a new era in art, reviving Anatomical Specimens) classical styles from Ancient Greek and ○ Esp. in glass containers with alcohol Roman times while incorporating modern (tissues, vessels, and organs) techniques. 1661 - Biologists’ Invention of The Renaissance was also a time of Microscope (including Antoni significant discoveries and innovation van Leeuwenhoek used this Dramatical Cultural Events Changes in Europe to publish actively and Fall of Constantinople which led Byzantine discover the anatomical scholars to Western Europe, bringing structures within organisms) ancient Greek knowledge and texts Evolutionary Ideas and CVA The Gutenberg Bible revolutionized 18th Century printing, enabling mass production and 1719: Peter the Great established one of dissemination of texts, including anatomical the first government-sponsored large public diagrams museums 1450s to 1476 1735: Systema Naturae (pamphlet) Theodoros Gaza of Thessalonica 1749: Comte de Buffon critique Linnaeus in A scholar who had immigrated from Histoire Naturelle Constantinople ○ Claims that Linnaeus was being too Translated the anatomical works of Aristotle abstract with his groupings being from Greek into Latin, making it more mere conventions accessible and publishing its printed edition 1758: Systema Naturae becomes a two in Venice in 1476 volume tome by its tenth edition 1543 1707 - 1778 - Carolus Linnaeus made an Andreas Vesalus advancement made an advancement in giving De Humani Corporis (Fabrica) was comparative anatomy a better system in naming published with detailed and anatomical and describing life forms illustrations Everything belongs to either the plant or ○ Fabrica pointed out the comparison animal kingdom of human anatomy from Galen’s text Kingdoms were divided into orders, then reports on monkeys “by presenting genera-Consisting of different species anatomical differences between at a Argued that plants or animals should be high degree of precision and setting given twofold names the stage for further detailed ○ One Latin word for its specie and monographs” genus Pierre Belon 1763-1793: John Hunter, a surgeon, ○ L’histoire naturelle de poissons created a collection of specimens that marins - a monograph on fish sought to lay out the anatomy and Conrad Gesner physiology of animals ○ Historiae animalium libri IV - a 1793: the Paris Museum of Natural History massive four - volume - is founded encyclopaedia on animals 1731-1802: Erasmus Darwin was inspired Thomas Moufet by both de Buffon and Linnaeus. He ○ Insectorum sive minimorum developed a more involved theory on how animalium theatrum - a monograph one species can evolve into another on insect 19th Century 1802: Suggestion of Evolution ○ Disagreed on whether he adequately 1809: Jean Baptiste Lamarck published considered developmental Zoological Philosophy processes (Cosans, 2009) ○ Argued complex species evolved ○ Debated with Thomas Huxley about from simpler ones the similarities between the human 1818: Etienne Geoffroy Saint-Hilaire brain and the ape brain published Philosophical Anatomy Huxley ○ Different animals share a common ○ Structural comparisons proved that anatomical plan the human brain and the ape brain 1830: clash between Cuvier and Geoffroy were similar 1830s: Richard Owen developed the Owen empirical approach to philosophical ○ Different in relation to their size anatomy 1890s ○ Promoted the term homology for Sigmund Freud anatomical analogies Geoffroy had Darwinian Idea found in different species ○ Higher animals have parts 18th-19th Century reminiscent of earlier stages of their 1724-1832: Johann Goethe developed an evolutionary history approach that built on the analysis of ○ Inspired comparative anatomy and Immanuel Kent and explored the possibility was therefore applied to human of evolution psychology Advanced a dynamic typology focused on Beyond just being a unified logical being, how the order of an organism emerges from the human mind made decisions from an dynamic movement of its developing parts interplay that expresses impulsive needs, 769-1832: George Cuvier offered an and a more social Ego, and a Super Ego exhaustive exploration of animal that seeks to follow evolutionarily recent morphology and won the reputation as one human society of the greatest comparative anatomists Started his scientific career in Germany of ○ Animal body can be seen as having Ernst Wilhelm von Brucke two types of function: ○ Darwin’s evolutionary theory was ‘Animal’ function: executed beginning to encourage traditional by neuromuscular system biological research programmes to and lets organisms move include the study of the around the world phylogenetic history of animal ‘Vital’ functions: executed by species in the onto\genetic internal organs and development of the individual maintains animal’s basic life organisms composed thereof 1844: The Theory of Evolution gains 1879 popularity through the publication of Robert ○ Freud immersed himself completely Chambers’ Vestiges of the Natural History in evolutionary neurology of Creation ○ Published a paper on the spinal 1858: Henry Gray published The Anatomy ganglia and cord of a species of Textbook which will be used in schools and lampreys hos[itals Century and a half after Darwin’s Origin 1859: Charles Darwin’s Origin of Species Intellectual movements shifted the focus of convinced many scientists that evolution comparative anatomy and its view of occurs by a process called natural selection evolution 1793-1874: Robert Grant takes up evolution 1st half of the 20th century 20th Century ○ Ideal morphologists Genealogical ○ Included Adolf Naef Arrangement of groups within each class Sought to advance an To appear more natural account of anatomy in the Amount of differences may differ due to the tradition of ideal morphology different degrees of modification the species of thinkers (Goethe and have undergone Owen) ○ Expressed by forms ranked under Gave a conceptually rigorous different genera, families, sections or understanding of the order evolution of animal and plant Theory of Evolution by Charles Darwin forms based on what can be Richard Owen directly observed in ○ Agreed with Darwin that evolution anatomical patterns as they occurred are seen today ○ Evolution is an undisputed fact ○ The mechanisms in Darwin’s Origin Thomas Kaufman and Rudolf were inadequate Raft published a monograph Embryology and on the study of embryos, Identification of more genes, and evolution morphological patterns were (Kaufman and Raft, 1983) also key to understanding ○ 1987 anatomical diversity (Rieppel Leo Buss - The Evolution of et. al., 2013) Individuality 1950 Questions about how Willi Hennig developmental (zoologist) processes evolved ○ Worked on (Buss, 1987) mosquitoes ○ 1990s and other Advances in techniques for invertebrates studying molecular ○ Foundations developmental genetics of a Theory of increased the interest in the Phylogenetic study of the interplay Systematics between developmental - Revolutionize comparative anatomy by biology and evolution putting it on a more rigorous foundation ○ 2000 grounded in the epistemology of logical Society for Integrative and empiricism (Rieppel, 2007) - Cladistics Comparative Biology formed Early 20th century a Division of Evolutionary ○ Tension among biologists who Developmental Biology focused on embryology (Amudson, 2005) Those that compared animals as field naturalists Subdisciplines of CVA Those that studied genetics In Animal Biology (Zoology), the in the laboratory sub-disciplines were categorized as pure ○ 1930s and 1940s zoology (deals with theoretical and Some biologists synthesized conceptual knowledge about the discipline), a perspective that combined which includes organismal zoology and population genetics and systematic zoology and applied zoology, evolution and displaced which deals with the application of concepts embryology (Smocovitis, from pure/basic zoology. 1996) Hailed as having a developed a complete vision of biology during the Centennial Celebration of Darwin’s Origin in 1959 1980s ○ Re-emphasis of the importance of incorporating developmental processes in an account of life ○ 1982 Webster and Goodwin - published sweeping critique Organismal Zoology - Vertebrate Zoology of the synthesis perspective Comparative Vertebrate study of structure, function and Comparative anatomy still Anatomy range of variation in structure needed to be open to and function among vertebrates. developmental concepts of pre-Darwinian rational (a) Morphology study of the form or shape of the vertebrate body. morphology such as those considered by Geoffroy, (b) Gross Anatomy study of internal structures of Goethe, and Owen, as well vertebrate body (dissection) as to the concepts of the Physiology Study of bodily Process early 20th century embryologist Hans Driesch Embryology Study of the development of (Webster and Goodwin, animals from fertilized egg to 1982) birth ○ 1983 Ecology Study of interrelationships of 1. Dorsal - back or upperpart (human vertebrates to their physical and anatomy: posterior) biological environments. 2. Ventral - underpart (human anatomy: anterior) Zoogeography study of distribution of animals in different continents of the world 3. Lateral - sides (left and right) based on constructed geologic 4. Anterior/Cephalic/Cranial - head end realms. (human anatomy: superior) Wallace’s Zoogeographic 5. Posterior/Caudal - tail end (human Realms: anatomy: inferior) 1. Oriental 2. Palearctic 6. Median - middle 3. Nearctic 4. Neotropical ❖ Central-Peripheral - part of a system 5. Ethiopian nearest the middle of the body or contrarily 6. Australian closest to the surface Vertebrate Paleontology Study of fossil vertebrates ❖ Proximal-Distal - part nearest the main mass or axis of the body or contrarily part Ontogeny Branch of biology that deals with farther from the main mass or axis. {NOTE: development of an organism, anatomical/behavioral feature The term is used with segmented structure from early to mature stage. such as limbs; in the case of 3 or more segments, the middle segment will be distal Phylogeny Evolutionary History of a Taxon relative to the segment before it, but Systematic Zoology - Vertebrate Systematics & Taxonomy proximal relative to the segment next to it. - Focus on taxonomic diversity and phylogenetic ❖ Superficial-Deep - commonly used in relationships of animals (vertebrates) myology (muscles); superficial means “near Ichthyology Study of Fishes the surface”, deep means “far from the surface” Herpetology Study of Amphibians and ❖ Extrinsic-Intrinsic - used in describing the Reptiles position of a set muscles (musculature); ie. Ornithology/Aviology Study of Birds extrinsic muscle of the eye lies external to the eyeball, while the intrinsic muscles lie Mammalogy (Theriology) Study of mammals internal to it. (a) Primatology Study of monkeys, apes, man ❖ Superior-Inferior - above and below BASIC MOVEMENT (b) Physical Study of evolutionary origin of Anthropology human species and racial Flexion vs Extension differences Describe movements that affect the angle between two parts of the body Applied Vertebrate Zoology ○ Flexion - describes a bending Economic Zoology Application of zoology movement that decreases the angle knowledge for the benefit of between a segment and its proximal mankind segment Animal Biotechnology Study of biological techniques in ○ Extension - opposite of flexion, doing animal genetic describing a straightening movement manipulation and biochemical that increases the angle between research for economic, medical body parts and legal purposes. a. Animal cell culture genetic engineering b. Cloning Immunobiology Veterinary Medicine (Zootherapeutics) Animal Husbandry Science of breeding and caring for animals Aquaculture study of culturing fishes and other aquatic animal resources of economic importance. Body Regions and Anatomical Positions Anatomical Planes: 1. Frontal Plane 2. Sagittal Plane 3. Transverse Plane Anatomical Positions: Abduction vs Adduction when depressing a car pedal or Refer to motions that move a structure away standing on the tiptoes can be from or towards the center of the body described as plantar flexion. ○ Abduction refers to a motion that pulls a structure or part away from the midline of the body. ○ Adduction - refers to a motion that pulls a structure or part toward the midline of the body, or towards the midline of a limb. b. Pronation vs Supination Refer to rotation of the forearm so that in the standard anatomical position of the palm or sole is facing anteriorly (supination) or posteriorly Elevation vs Depression (pronation) The terms elevation and depression refer to i. Pronation - at the forearm is a movement above and below the horizontal rotational movement where the hand ○ Elevation - refers to movement in a and upper arm are turned inwards superior (upward) direction ii. Supination - of the forearm occurs ○ Depression - refers to movement in when the forearm or palm are an inferior (downward) direction, the rotated outwards. Supination of the opposite of elevations. foot refers to turning of the sole of the foot inwards. c. Inversion vs Eversion Refer to movements that tilt the sole Location Specific Movements of the foot away from (eversion) or a. Flexion and Extension of the Foot towards (inversion) the midline of the i. Dorsiflexion - occurs where the body. toes are brought closer to the shin. ○ Eversion - the movement of This decreases the angle between the sole of the foot away the dorsum of the foot and the leg. from the median plane. For example, when walking on the ○ Inversion - the movement of heels the ankle is described as the sole towards the median being in dorsiflexion. plane. (e.g., inversion ii. Plantar Flexion - the movement describes the motion when which decreases the angle between an ankle is twisted) the sole of the foot and the back of the leg. For example, the movement upon which an organism was built Evolutionary Morphology Past evolutionary changes is inscribed in animal structure ○ The external environment must serve certainly brings to bear evolutionary pressures on its survival - anatomical features convey adaptive benefits Examples: 1. Tiktaalik - “first tetrapods 2. Archaeopteryx - “dinosaur bird” Functional Role - action or property Biological Role - how the part is used Comparative Vertebrate Anatomy (Basic Concepts) Comparative Morphology Vertebrates - Significance - Structure Functional Morphology ○ Relates a structure to its function Homocercal (homo - same; kerkos - tail) Heterocercal (homo - different; kerkos - tail) Historical Predecessors - Morphology Thomas Huxley Preadaptation ○ Monographs on Comparative The structure of behavior possesses the Anatomy necessary form and function before the ○ Darwin’s bulldog biological role arises that it eventually ○ Proposed connections between the serves. development of organisms and their ○ Traits that have enabled a evolutionary histories phenotype to meet a new ○ Vertebrate Embryology environmental challenge before it ○ Homology materializes Eg. lungs were present Georges Cuvier in fishes long before they function on ○ Founder of Comparative Anatomy land (Functional Morphology) Historical Predecessors - Evolution ○ Claimed that ancient animal Jean Baptiste de Lamarck mummies were structurally identical ○ Philosophie Zoologique “Theory of to modern species Evolutionary Descent” ○ “Parts were adapted to perform ○ Theory of Use and Disuse specific functions.” ○ Theory of Acquired Characteristics Etienne Geoffroy Saint-Hilaire ○ Theory of Perfection ○ Doctrine of the Unity of the Organic “A progressive change in Plan of the Animal Kingdom species along an ascending ○ Philosophical Anatomy scale from the lowest on one ○ Different animals share a common end to the most complex and anatomical plan and made the perfect on the other hand.” identification of the manner a given Charles Darwin part had been altered in evolution ○ The Origin of Species Richard Owen ○ Mechanisms for evolutionary change ○ Made use of homology High reproductive potential ○ Species were immutable Competition ○ Correspondence between parts Survival of the fittest (homologies) could not be left ○ Mechanism evolutionary change without explanation Carolus Linnaeus ○ Archetypes ○ Devised a system for naming plants Kind of bio blueprint - a and animals supposed underlying plan ○ “Species were unchangeable, created originally as we find them today.” ○ Perfected adaptations of animals Evolutionary Change Evolutionary Convergence Diversity of Species Two unrelated organism which occupy the Alfred Russel Wallace same environment whether concurrently or ○ Explained evolutionary change by million years apart natural selection Acquire similar morphological features as ○ “Why do some die and some live?” adaptation to the environment ○ “Modification of form is admitted to Produces look alike features that are not the be a matter of time” result of inheritance from a common ○ “On the whole, the best fit lived from ancestor the effects of disease - most healthy Symmetry escaped.” Describes the way in which an animal’s body meets the surrounding environment Segmentation/Metamerism A body or structure built of repeating or duplicated sections-segment (metamere) Morphology Concepts Similarities In different organisms, corresponding parts may be considered similar to each other by three criteria - ancestry, function and Cephalization appearance (Kardong, 2018) Tendency for the anterior end of the body to Pierre Belon become distinctly separated and French physician and naturalist of the 16th differentiated from the rest of the body as century; compared human and bird head ○ Concept: architectural plan of the Phylogeny creator Evolutionary history of a taxon What about cladistics? Method of hypothesizing relationships among organisms Serial Homology Applied to metameric structures in a single organism; represents a pattern of segmental equivalence (segments of the craniate head) Boyden Not all look alike are homogenous; same organ in different animals under every variety of form and function Monophyletic Group/Clad: includes common ancestor Metamorphosis and all descendants Transition: birth (or hatching) to adult stage Passes through morphologically distinct stages (change in form) Paraphyletic Group: Includes a common ancestor but one or more descendants is omitted (eg. birds are excluded from reptiles) Paedomorphosis Truncated development; descendants mimics ancestral juvenile; decreasing rate; postpones the start of development Polyphyletic Group: Does not share a common ancestry Peramorphosis (eg. Uniting birds and mammals based on Extended development; increasing rate; endothermy) starts development early (larva undergoes morphological changes) Heterochrony 1. Paedomorphosis - “juvenile features” a. Progenesis - acceleration of sexual maturation while body development slows down or remains unchanged b. Neoteny - slowing down of somatic Ontogeny development while reproductive Developmental history of organism development proceeds at the usual Embryogenesis, post-embryonic stages, pace aging and death c. Post Displacement - delay in the Von Baer’s Law onset of somatic development, Organisms which exhibit a similar leading to retention of juvenile architectural pattern of both structure and characteristics development 2. Peramorphosis - “hyper-adult” a. Hypermorphosis - extension of development before the typical stopping point b. Acceleration - faster development of certain features —> more pronounced traits at adulthood c. Predisplacement - early onset of development —> more developed features Phylogeny Blastopore “mouth” Blastopore “anus” Monophyletic GROUP/CLAD: includes a common Spiral cleavage Radial cleavage ancestor and all descendants schizocoelom entercoelom Ectodermal skeleton Mesodermal skeleton Mollusks, annelids, Echinoderms, arthropods hemichordates, chordates Chordate Characteristics Paraphyletic GROUP: recent common ancestors, but not all descendants are included The Big 5! Notochord Polyphyletic Pharyngeal slits GROUP: recent common ancestors but Endostyle include distant relative Dorsal hollow nerve cord Postnatal tail Taxonomic Relationship of Protochordates and Vertebrates Cladistic Versus Traditional View of Reptilian Phylogeny Chordates (+) notochord in the embryonic stage Craniates - chordates with neurocranium Vertebrates - chordates with vertebrae The Craniate Body Plesiomorphic (primitive) Develops similar patterns of embryonic development Phylum Chordata Regions: head, trunk and post anal tail Living Chordates Main Support Cephalochordates Urochordates Notochord Vertebrae Vertebrates First to appear Provide more Protostomes Deuterostomes in craniate rigid support embryos Components: “First mouth” “Second mouth” Becomes centrum, neural incorporated in arch, spinal the floor of the cord, basal skull processes, Exemption: hemal arch hagfish (Myxine (sharks and glutinosa) fishes) Significance: serves as a common ancestry between craniates and protochordates Pharynx Vital part of the craniate embryo Produces the gills of fishes Lungs of tetrapods Skeleton and musculature of the jaws Endocrine glands that regulate the metabolism of cell Give rise to middle ear cavity in tetrapods Contributes to immune system during the fetal life Pharyngeal Pouches Arises as diverticula of the endoderm of the foregut; persist as gill slits in fishes and amphibians; close permanently in other vertebrates Pharyngeal Arches Serves as support in the gill slits of fishes Dorsal Hollow CNS Brain and spinal cord ○ Cranial and spinal nerves connect the CNS ○ PNS nerves - ganglia and plexuses Craniate vs Vertebrate Craniate Vertebrate Cranium Vertebral column Three part brain Two part brain Single semicircular Two semicircular canal canals Unicellular sense Multicellular organs Lateral line system Neural crest and its multicellular neuromast derivation Has a number of tissue cartilage specialization

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