le C] ee lass Thaleacea has been divided ber bs a Order—1. Dotiolida Characters : (i) Body ba Gi) Fight muscle bs Gi) Larva with a tail Example—Doliolum. \ds, each band forms complete ring, yomida Forms a well organized colony Order—2. Py Sharacters : (i) (ii) No larval stage found. Example—Pyrosoma. ‘Order—3. Sal opme, ‘ent Characters : (i) Rot ansparent body. (i) Muscle bands incomplete (iii) No larval stage Example—Salpa C Retrog: O Introduct In the ontogeny of species, met Basen ee er passonl pear in the larva, gradually retrograde and disappea yeh a life history refers to retrogressive metamorph' zs been clearly established by the life history of an Ascidia indicates sequential progressive changes. y in the life history of Ascidian, character r in the adult josis and O Definition : Retrogressive metamorphosis is a developmental process i hich a highly developed tadpole larva retrogrades its characters nerated sessile adult 4g metamorphoses into a degi evelopment of Tadpole I O Life history and De varva: small hermaphrodite, eges and ion takes place outside Fig. horionic cells marine, ater and fertili esof { surrounded by Adults are sessile. 1.1: Extemal featur berms are discharged in wa « body. Ovum is yolk free and test cells. Cleavage is holoblastic and equal (embryogenesis) @ fish like creature by invagination. After two gastrulation is which is known as is produced s of developm «idian tadpoleBook of Zoology Modern Approach TO A TEM! Ie Ascidian Tadpol ren 4 trunk, slender taper 2nd-1-6 © Characters of (@) Uniformly small, vol tail, trunk containing the rudimen 1 tail, ost tissu future tadpole. ig covered by thin test er nnterior end a ro present at Yn which there is a iid) Taree adhesive papillae are A an nicely nsory vesicle is prest ' Mo areca ‘ {iv) In the trunk there is m with a pigmented cup, fo otolith and on the dorsal wall tere is associated with 2 F formed (©) The ovellus consists O ory retinal cells oll about a dozon of pigmented sensory Fe! Tail F Tail Consists ofa sing) epidermal sheath supporting a cutiggs fin and contains the following struety i) A hollow neural tube extend slong the dorsal side from the poste, wall of the sensory vesicle to the fip the tail (vii) A notochord extend: the centre of the tail from beneath posterior wall of the vesicle through tip consisting of 40-42 vacuolat le column. in a sin (viii) On either side of the notochen a band of striated muscle tissue, of 18 cells, derived band consistir rtilization and Development of an Asc mesodem, are found. These are acti continuously as a single contractile unit. Other cells of the tissue migrate ventrally to m the pericardium, heart and mesenchyme. Pe (xT dimentary canal is distinguishable into pharynx, oesoph: nd intestine not yet functional, because the mouth is covered by tunic and the larva is a non feed: organism. (x) The atrial cavity and a sin atrial pore are developed. O Tactic movement of the Lar For several hours the todpole larva becomes ly active swimmer and at the fi stage it becomes positively phototropic and negatively » urface. Within a day or two depending on the the bottom eotropic and so proceeds to the Si condition tropism is re they sw @ suitable surface to which it attaches by and tum to any dark place and finds the adhesive papillae, looses the tail, develops a | large pharynx and continues post larval growth) O Post Lary 1 Development : The post larval organs according to the ir degree of vy sen classified by Bicsard Cloney (1978 f vulnerabiity has been classified 83) as (4) Transitional tarval organ, (b) Pp; organ Tospective Juvenile organ, (c) Larval Juve (@) ad) Q) @) (4) (5) (6) a) Re metanClassification of Chordata (a) Transitional Larval Organ (T:L.O.) : TI z - ofc ‘gan ( ) : These organs are (2) Caudal musculature, j (3) Visceral ganglion and nerve. i (4) Nerve cord. lula (5) Papillac, (6) Outer cuticular layer of tunic. 1 by] (7) Sensory vesicle and sensory organ, vole pean uo organs are required for larval settlement and Subsequently lost during “ular )_ Prospective Juvenile Organ (P.0.) + ‘These organs ate ture, (1) Atrial siphon and atrium. ding (2) Branchial basket, oesophagus, stomach and intestine, erior (3) Pyloric gland. P of (4) Heart. (3) Neurohypophysis or cerebral ganglion and neural gl ugh Remark : Organs are fully differenciated, 1 the functional histogenesis. the (©) Larval Juvenile Organ (Ll.0.) : cells (1) Epidermis. (2) Tunic and mesenchyme derivatives including blood. hong Remark : This group includes orga each larval phases of development. id. become functional following several days of These organs are — ins and tissues that function through larval and post from Q Significance of Ascidian Tadpole in Habitat Selection : As described above, the tadpole in its second phase of activity becomes positively geotropic and negatively phototropic. This is guided by ocellus and otolith and ultimately attaches to a hard surface by adhesive papillae. So selection of clean hard surface at the bottom of the sea is of great importance for further metamorphosis. On this pretext two views are eding. important for consideration. Berrill's View (1955) : He has emphasised the very great importance of habitat selection in the life of Ascidians and consequently the high selective value of the possession of this Particular type of non feeding larva. Cloney's View (1978) : Presumebly its short life in the chordate stage is sufficient to ensure distribution and the simple nervous system serves to find a place in which to live. stine, > first e sea jm to. Importance of Habitat Selection (Good body’: experiment — 1961) : es by In an experiment to show the importance of habitat selection he took two population A owth 0d B in the month of June and maintained these two population at a depth of four feet. He again took two population D and F in the month of August and maintained them at a fepth of seven feet. ; ; ; ed bY Explanation : The number of animals that could survive was much less in the population and F after a period of about a month and a half, ‘The explanations for these less survivals vyenilé| tre (O If the larvae settle at any substratum other than clean, hard and rocky substratum ‘ey fail to develop further.+ Gi) Sufficient algal growth on the sub: On Wit Population ‘of Nereida, Serpulids and other col Ac COUn, | for their reduction in number. 2nd-1-8 ‘Modem Approach To A Text Book of Zoology stratum prevents adhernce and competiti Jonies of Ascidians on the same substratum, O09 1000" 5006 500 | = (By) E 100" J 100 Ww Days ba a JUNE AUGUST Good body's experiment Q Dual Organization in Tadpole Larva : The ascidian egg developsasadual system. A tadpole on one sideandas sessile adult on the other An ascidian tadpole consists of two sets of tissues which are distinct from one another. (1) Larval tissue : Fully matured and differentiated cells irrespective of number and size, (2) Residual tissue : Which will form the adult, are present also but far from being histologically matured. Q Retrogressive Metamorphosis : It is encountered in two forms namely — (1) Loss of Organs : Organs which are lost during metamorphosis are — () Notochord. i) The dorsal tubular nerve cord dwindles into a ganglion and neural gland. (iii) Tail with fin. (4) Ocellus with otolith. Development of Organs : (1) Alimentary canal is well developed, pharynx with endostyle and major divisions of the alimentary canal but not yet functional as the mouth is covered with test. (2) Stigmata greatly increased, atrium well developed with single atrial opening. (3) Reproductive organs developed. (4) Heart and pericardial cavity well formed. Remark : Organs which will be present in the adult now take up the position through a rotation of 90° from the point of their attachment. Mouth opening is also shifted. QO The events of metamorphosis : The events are summerized by Richard Cloney (1983), (1) Secretion of adhesive substance by the papillae. (2) Evertion and the reaction of the adhesive papillae, (3) Gradual contraction of the tail, (4) Loss of the larval cuticular layer of the tunic,Classification of Chordata (5) Rotation of organs through 90°, (6) Elaboration of branchial basket. (7) Perforations of siphons, (8) Retraction of sensory vesicle, (9) Release of organ rudiments from an atreste 2nd-1-9 d_ state of development. Ort funnel Ganglia Endostyle Stigmata, Notoghord Fig. 1.3: Retrogressive metamorphosis of an Ascidian tadpole ),(©) & (©) — Entire extemal features of larva, (a/1), (b/1) & (c/1) — Their sagittal veiws. O Causes for the Retrogression : Various opinions have They are— (1) Gravel's view (1935, '44) : Metamorphosis is conditioned by two factors as— (1) Ageing of the larva, (2) The swimming activity. He finally said that the larva produces a substance which greatly accclarates metamorphosis and the effect is directly proportional to the concontration of the substance secreted. (2) Glesen and Anslow (1949) : According to them concentration of calcium ions (Ca**) released within the tissue are responsible for retrogression. (3) Berrill's view (1955, '75) : It was formerly thought that exhaustion of notochord tnd tail is due to phagocytic activity of the cell. But Berrill said that there is no phogocytosis and shrinkage of notochord and tail. As a whole notochord and muscles play an important tole. They secrete some active substance which forces the progressive shrinkage of the tail, Utilization and exhaustion of the nerves of the tail is due to ageing factors. As there is no ‘cretory organ or in the larva toxic metabolites including CO, are deposited in the tail region, ‘2d their concentration held the shrinkage of the tissue. Conclusion : It is evident from the above discussion that chordate features of Ascidia Y appear during larval stage. Had there been no larval stage showing chordate features determination of systematic position of the adult Ascidians would have been very difficult, Notonly that how a dual organization can be shown by a larva has been clearly examplitied Tetogressive metamorphos! in tadpole. ‘= been put forward to explain the factors underlying this metamorphosis