BERIEEOra OW nm nis theory be and bac ins with th k down. Through tr Elaboration of th that swim with their Ie gave rise to. a" inverted vertebrate ffered from some ma .e nauplius larva of crustaceans and other arthrop ssitional forms, these changes suppe ajor setbacks and we nesses tween chordates and annelids or This argument s' ” ; eis rest from homoplasy rather than hom ology = ID The segmentation and jointed append that are part of an arthropod body are quite unlike the chordate myotomal segmentation e «The main nerve cord of annelids an d arthropods is lid, not hollow as in chordates, embryologically in a fundamentally different way Furthermore, the usual positions for a chordate .s mouth and anus are ventral, whereas an annelid | ‘or an arthropod rolled on its back would turn both mouth and anus uP, pointing skyward {| Inverting an annelid or arthropod to produce a chordate body plan would require migration of the / 7 mouth and anus back ventr ally or formation of new one ventrally beg | © Unfortunately, the embryonic history ¢ chordates also is fundamenta’ ly different in method of coelom formation, derivation of m :, and in basic pattern of early cleav || 2, ¢hordates from Echinoderms | Proposed by Johannes Muller (1860) this theor ised on the comparative studies of larval stages of echinoderms and hemich¢ 4 Torn 1 of hemichordates resembles echinoderm larvae such as Bipinnaria, Auricularia, I sala and Doliolaria, which all possess ciliary I ands and apical tuft of cilia. Johannes Muller, W proposed that echinoderm Jarvae gave rise to chordates by neoteny. Gai y he Neotenous Larva ame sexually mature and uggesting that probably the auricularia larva theory later gave rise to the chordates. evidences, various workers had tried to ¢ or echinoderm larva. It is this similarit chordates. On the basis of Garst di n of | outlining the or echinoderms were the probable chc discovery of fossil echinoderms calle | Evidence : The | mya) further confirms echinoderm showing affinities with both echinoderr in vertebrates the bones are made of hydrat ontological, biochemical ne primitive echinoderm rth an alternative theory affinities, e embryonic alcichordata from Ordovician period (450 asymmetrical animals \cichordates were their skeleton is made of CaCO, whereas is They had a large pharynx with aDIVERSIT ated pharynx for filter fe anal tal ad a po 4 body an oe brian ais during Car ps of ani echinoderm ntable because cemed unace'Pr suggest a phylogenetic aes akin, and peniaradial (F f \¢ followin} hypothesis. ncept) esis (Dipleurula co! ») aricularian Hypoth ; ae embryonic n hypothesis, auricular particular type ©! jarva s (sea cucumbers) sthesis lies on the concept of A perf jovician period Y OF CHORD: js when planktons starfish, sea urchins, such as ; with chordates. They affinity srmed) body symmetry. Gar a dipleurula larva, believed to represent tg f all echinoderm larvae. date characteristics first debuted in this dipleurula larva whi stang proposed that, the chor ymmetry bod adoral band of feeding cilia; and row of cilia, the Near the mouth was an body surface a lor across its la the larva to propel circumoral band, which enable Garstang argued that from such structural ingredients in a, the basic chordate plan emerged the echinoderm la © Inparticular, he stressed on the view that the larval body elongated, becoming increasingly muscular along with the formation of a tail that, together with a notochord, could generate lateral undulations as a means of locomotion * Elongation of the body drew out the circumoral ciliated band and possesses a simple one-way at lary Band — Fig. 7.4 Garstang’s Hypoth« and brought its left and right dorsally, where they met at the midline together with the underlying nerve tract, the antec of the nerve tube. © Garstang; ft ig pointed to the rolling up of the neural ‘ube during vertebrate embryology as an embt Temnant of this phylogenetic event, ; lengthening of the adora the Benning ofan endonice 800b> Grigin of Chordates The larva now lacked the pharyngeal slits to complete its transformation into a full-fledged chordate «Evolutionary changes occurred in the larva with the development of pharyngeal slits to support wy hae suspension feedin : : *Stane Thus the development of pharyngeal slits for suspension feeding and a notochord for muscular locomotion might subsequently have found favor by selection, yielding an echinoderm larva endowed with a full set of chordate characters: notochord, dorsal nerve cord, pharyngeal slits and post anal tal ‘© But froman echinoderm larva, evolution does not look hopefully for the development of a chordate lifestyle ‘© Therefore the changes in the echinoderm larva must have been driven by some immediate | adaptive advantages at the time of their first appearance Probable driving force ew Probably the larva of this ancestral echinoderm spent more and more time in its planktonic stage, fed, and thereby grew in size Pe Larger size is an advantage to escape predation and become established on a substrate once alll metamorphosis begins. the fatal the larva’s body size increased for these or other reasons, this would require compensating changes in two systems, locomotion and feeding vhi 3 y © Thus as an object gets larger, surface and mass increase unevenly relative to each other, Body mass increases to the cube of the linear dimensions, but surface area increases only by the al square of linear dimensions. | © Therefore in a larva that increased in size, surface cilia propelling the larva would not increase enough to cope with the increasing mass. As a result, there would be relatively fewer surface F cilia to move a relatively greater mass. Thus the locomotor surface would fall behind as the larva } got larger, ~ © This situation, favoured the development of ed body, and stiffened bar (notochord). These structures an alternative locomotor system in the form of a segmental swimming musculature, elon placed the faulty ciliary system of locomotion. first supplement and then * Similarly, the mode of feeding had to change, for the same reason, namely, a geometric mismatch te between surface area bearing feeding cilia and body mass requiring nutritional support. ee © The feeding cilia around the mouth sweep suspended particles into the mouth. But as larval size a , body mass outstrips the ability of these surface cilia to meet the nutritional needs. sre, an adoral ciliary band thus probably expanded into an endostyle to improve food transport a * Perforations (slits) in the pharynx would allow one-way flow of a feeding current. Both changes would increase the efficiency of the feeding mechanism. © These feeding structures have been favoured by selective pressures.F CHORDATES pIVERSITY © hic adult, But how tere nto a bent mcnenes xd to transform i ic in pheeal Echinoder™ 1 Cer inoderm had 1 Fans parte evolutionary destin Ces Bt areamyeciere Stra re ndowed with chor” such obstacles iS ech a TET explanation such MT age enbanced was explained by Toma ana the ara tage COPTER ; Bue er ito its benthic «He suggested hat ts 0 oa teeing ies the aaa omer civ Se » adult did not, then time ‘ and the adu modified larva enjoye’ Mere cime spent as an adult = pened at he expense of te sa rva, iteould reproduce, and thus escaPe from | might came sexually mature while still 2 [arvay It es SE te ite ul assis termed paedomorph if yee id to a enti aT Pr of some Larval characteristics in the advil: Ia ig refers to the retention of some larva «The term paedomorphosis refers t pose ea hr weds iste eto ofthe wet OP WT om echinoderm™ ances iuibs Jomorphosis in chordate evolution «The possible significance of pacdomorph cht enjoy the adaptive ped with chordate features mi Oe an arvocess, depart along an independent A paedor hinoderm larva equi ata ter pelagic mobility and, in the proces evolved from echinoderms ancestors via s ted that vertebrates might have J then through urochordate-like, rough hemichordate like, and paedomorphosis. or vor solely eto the blanks: 1. The earliest known chordates were called 2. Auricularia larva are found in — 3. The band of cilia present near the mouth of Dipleurula larva is called 4. The habitat of earliest known chordate was Answers : 1. Ostracoderm, 2. Holothurian, B. Give one word answer. 1. The larvae on which Auricularian hypothesis is based. 2. Who proposed the Dipleurula concept ? 3, The fossil Echinoderm that confirms Echinoderm ancestry of chordates, 4. Retention of some larval feature in adult. Answers : 1. Auricularia larva, 2. W.Garstang, 3. Calcichordata, 4. Paedomorphosis, ‘sentences each. 3. Adoral band, 4. Fresh water I. Paedomorphosis 2. Inverted vertebrate 3. Homoplasy |aoe ____—_pwensrrvorcnonnaes; | 0” se diver, with or without a larval stage wheeas,in J, : \ the development is indirec oe OVER PROTOCHORDATA © Incuchsfdates. forodhordates. the BipAnvanced FEATURE! The advanced features that hel dre the presence of living endoskeleton efficient 1 opment is indirect with a fre s OF VERTEBRATES ip chordates to dominate n, efficient circul: devel over other Phyla (Protochordates) lation, and centralized espiration nervous system. 4. Living endoskeleton column, in which the notochord found ofa vertebrate is the vertebral ts (vertebrae) separated ‘egmented series of st acteristic eplaced by a8 iffer element Teal dises, derived embryonically and evolutionarily from the The defining chi in all chordates has been © by mobile joints (intervertel notochord). otochord into adulthood, . DA few vertebrates have secondarily lost this anatomy retaining the m e Sturgeon and Latimeria. ‘ows in size with rest of the body cendoskeleton that gr “ath of the vertebrate body so that they include such as th All vertebrates possess a living | The living endoskeleton allows an indefinite gro the largest creatures of the world . ii, Efficient respiration ares iv & Allbasal vertebrates breathe through gills. © The int behind the head, as a series of openings from the pharynx to the . «Each gill is supported by a cartilagenous or bony gill arch . The bony fish have three pairs of arches, cartilaginous fish have five to seven pairs, while the «Pec primitive jawless fish have seven . «The vertebrate ancestor no doubt had more arches, as some of their chordate relatives have v. Cent more than 50 pairs of gills. oc «In amphibians and some primitive bony fishes, the larvae bear external gills, branching off from ik the gill arches, These are reduced in adulthood, their function taken over by the gills proper in fishes and by lungs in most amphibians : © Some amphibians retain the external larval gills in adulthood sk gills, the gill arches are formed during fetal development, “7 © While the higher vertebrates lack asis of essential structures such as jaws, the thyroid gland, the larym\ which lay the bi mammals) and the malleus and incus, in mammals. the columella (corresponding to the stapes in ili. Efficient circulation © The circulatory system of chordates is well developed. «The circulatory system is often seen to comprise two separate systems: the cardiovase system, which distributes blood, and the lymphatic system, which circulates lymphind he origin of Chordates st | ‘ 81 The cardiovascular system comprises the blood, heart, and blood vessels The Iympb, Iymph nodes, and lymph vessels Form the lymphatic system, which returns filtered system, which returns filtere blood plasma from the interstitial fluid (between cells) as lymph Blood vascular system is of closed type Blood flows freely into the organs of respiration for exchange of respiratory gasses Single or double form of circulation is exhibited by chordates, nic circulation In single form blood passes through the heart only once before entering the syst as found in most fishes, ¢ In double form of circulation the blood moves through the heart two times before entering the systemic circulation as found in birds and mammals Heart ventral, muscular, contractile and consists of two, three or four chambers, Blood is a fluid consisting of plasma, red blood cells, white blood cells, and platelets that is circulated by the heart through the vertebrate vascular system, carrying oxygen and nutrients to and waste materials away from all body tissues. The red blood corp The efficient blood vascular system forms a suitable medium for different vital activities of the body iv. Presence of paired appendages les contains the respiratory pigment called haemoglobin. ‘© Vertebrates possess two paired appendages * They may be fins or limbs. In fishes fins are present. In tetrapods two pairs of limbs are present * Pectoral and pelvic girdles also developed to support the paired appendages. * The paired appendages help for locomotion in water as well as on land. v. Centralized nervous system The central nervous system of vertebrates is based on a hollow nerve cord running along the stic to vertebrates is the presence of neural crest cells. length of the animal. The unique character These are the progenitors of stem cells, that migrate through the body from the nerve cord during development, to initiate the formation of neural ganglia and structures such as the jaws and skull. No such neural crest cells are found in Protochordates. © The vertebrates are the only chordate group to exhibit cephalisation, the regionalization of brain functions in the head. A slight swelling of the anterior end of the nerve cord is found in the lancelet, though it lacks the eyes and other complex sense organs comparable to those of vertebrates. Other chordates do not show any trends towards cephalisation. vstem branches out from nerve cord to innervate the various systems. © The front end of the nerve tube is expanded by a thickening of the walls and expansion of the central canal of spinal cord into three primary brain vesicles: The prosencephalon (forebrain), * A peripheral nervousDIVERSITY OF CHOR' halon (hindbrain). Vesicles of the forebrain are yg 1 brain) and rhombencephal ae i: mesencephalon (midb " ain) a heres like the cerebral hemispheres in mammals. These are fui) paired, giving rise to hemispheres lik: d i ertebrate groups. differentiated in the various vertebrate grouy shemishrain eco eyes for ound outgrowths from the midbrain, except in Two laterally placed eyes form aro tg th forebrain is well developed and subdivided in most tetrapay this may be a secondary loss. The forebrain is h and some salamanders. while the midbrain dominate in many The resulting anatomy of the central nervous system, with a single hollow nerve cord associat © The resulting anatomy of the central nervou with a series of vesicles (often paired), is unique to vertebrates. PRACTICE QUESTIONS Fill in the blanks: The anterior end of nerve cord becomes specialised to form a Animal that lack true jaws are called 3. The precursor of thyroid gland present in protochordates is called | 4. In the sperm unites with the ovum through vegetal pole. 5. In——— the sperm unites with the ovum through animal pole 6. In— notochord is replaced by vertebral column, 7. Acraniata lack — | 8. In ——__ vertebral column is absent | 9. Gonoducts are absent in but present in — | Answers : |. Brain, 2. Agnatha, 3. Endostyle 4 Protochordates, 5. Euchordates, 6 Euchordates | 7- Cranium, 8. Protochordates, 9, Protochordates and f uchordates, B. Give one word answer. a J; Stff'rod running through the length of the animal 2. Hollow tube of nervous tissue situated above notochord. 3. Prime diagnostic feature of chordate 4 Paired openings leading from pharynx to the exterior @ Space between pharynx and body wall in Protochordates, §- Bony structure that support the paired appendages in vertebrate The chordate group that exhibit cephalisation, hai ba Answers ; | Notochord, 2. § o Te 2. Spinal Cord, 3 Notochord, 4. Gill slits, 5 oe 4 Atrium 5. Cephalisation, 6, Neural ere: ic culation, 9. © SEC f ci sues: oa losed type of circulation, 10, Gnatho |. Fundamental chordate . : 2. Difference between p EUAnetiaeia 7. Double circulation, g characters Totochord, ates and Euchordatan