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CTENOPHORA Course Name: Zoology B.Sc. 1st Year Paper No. & Title: Z-101B Animal Diversity Topic No. & Title: A-5 Theory Ctenophora - Affinities, Type Study Lecture No. & Title: Lecture - 1 Ctenophora - Affinities, Type Study VIDEO INDEX : (PARTS OF THE VIDEO) 1) Introduction 2) Affinities and systematic position of Ctenophora 3) Examples of Ctenophora types 4) Summary Academic Script Introduction The Ctenophora (Gr., Ktenos-Comb; phores-bearing) are marine, pelagic or free, solitary and biradially symmetrical organisms, with transparent gelatinous bodies. They lack nematocysts, possess ciliary plates in eight rows and have a gelatinous ectomesoderm containing mesenchymal muscle fibers. Their most distinctive feature is the "combs", groups of cilia they use for swimming; they are the largest animals that swim by means of cilia – adults of various species range from a few millimeters to 1.5 m in size. There are peculiae adhesive cells in the tentacles. Skeleton is absent. All the ctenophores have transparent gelatinous bodies and are commonly called as “comb-jellies”. Ctenophores are very common marine animals found in diverse habitat. They are widely distributed being specially abundant in the warmer seas. But some occur in temperate or arctic regions. They are of planktonic habit floating in the surface waters, mostly near shores but few live to depth of even 3000 metres. Ctenophores feed on small marine animals including the eggs and larvae of molluscs, crustaceans and fish. CLASSIFICATION The phylum Ctenophore is divided into two unequal classes: Tentaculata and Nuda. The class Tentaculata includes 4 orders: (i) Cydippida (ii) Lobata (iii) Cestida and (iv) Platyctenea. The class Nuda has only one order: Beroida. AFFINITIES AND SYSTEMATIC POSITION OF CTENOPHORA In the field of taxonomic Zoology, Ctenophora, for a long time, occupies a very important place. The importance is primarily due to its peculiar anatomical organization. Martens (1671) first discovered Ctenophora. Leuckart (1847-1848) included the sponges amongst the ctenophores. But Hatschek (1889) gave a separate status to ctenophores as a group. The ctenophores bear many characters of the Cnidarians, but differ considerably from the other members of the phylum Cnidaria. This group has also some similar features with different animals of diverse phyla, which will be discussed here to judge its systematic position. CNIDARIAN FEATURES - Possession of radial symmetry. - Lack of coelom. - Presence of gelatinous mesoglea. - Presence of ramified coelenterons. - Presence of diffused nerve network. - Presence of statocyst as sense organ. - Absence of organ systems. - Different parts of the body are arranged along and oral-aboral axis. But the Ctenophora differs widely from the cnidarians by the possession of meridional comb-plates, lack of nematocysts and possession of adhesive cells (colloblasts). These have higher and complicated organization of the digestive system, determinate type of development, direct development of muscle cells from the mesenchyme and retention of cilia as locomotor organs in adult. RELATIONSHIP WITH HYDROZOA WITH CTENARIA (ANTHOMEDUSA) SIMILARITIES - Presence of two tentacles, situated at opposite per radii, each is provided with a deep pouch at its base, revealing closely the tentacular sheath of Hormiphora. - Presence of eight radial canals formed by the bifurcation of four inter-radial pouches of the stomach. - The subumbrellar cavity of the ctenaria can be homologized with the stomodaeum of Hormiphora. DISSIMILARITIES - The gullet of Ctenophora is ectodermal in origin. - The tentacles of Ctenaria have no muscular base. - The development of gonad is different and develops from manubrium in Ctenaria. But in Ctenophora gonads develop from meridional canals. - Absence of the characteristic of aboral sense organ in Ctenaria. HYDROCTENA (NARCOMEDUSA). Hydroctena, a trachyline medusa, shows close resemblance with Ctenophora. - Possession of two tentacles with sheath located between the margin and apex of the bell. But the presence of swimming plates in Ctenophora and the presence of velum in Hydroctena remains as important differences to visualize a close relation among them. RELATIONSHIP WITH ACTINOZOA The Ctenophores also possess certain anthozoan features such as: - Ciliated ectoderm of Anthozoa is probably a forerunner of the ciliated band of Ctenophora. - Presence of well developed stomodaeum. - The gut in embryos of both is four lobed, thus, presenting biradial symmetry. - Mesoglea is cellular. - The gonads develop in connection with the endoderm and sexual elements passed out through the mouth. - Both the aboral sense organ and rows of comb-plates of a ctenophore have no parallel parts in an anthozoan. Lasso cells differ structurraly from the nematocysts and tentacles are hollow in Anthozoa, while solid in Ctenophora. RELATIONSHIP WITH SCYPHOZOA The Ctenophores also possess certain Scyphozoa features such as: - The stomodaeum is similar. - Gametes are endodermal in origin. - Coelenteron is four-lobed. But due to the presence of four characteristic oral arms, numerous marginal tentacles and the absence of meridional comb-plates in Scyphozoa, the aforesaid homologies cannot be established. RELATIONSHIP WITH SPONGES Many authors tried to establish the sponges as to be closely related to Ctenophora. Such as: The large central cavity and the osculum of sponges correspond to the coelenteron and mouth of Ctenophora respectively. - Absence of well formed mesoderm in both groups. - Simpler organization in both. But closer examination reveals that these two groups are quite widely apart. The differences are: - Developmentally the osculum of sponges does not correspond to the mouth of Ctenophora. - Presence of inhalant pores and peculiar collar cells in sponges are absent in Ctenophora. - Absence of colloblasts in sponges. - Absence of specialized nervous and sensory structures in sponges. The above differences are sufficient to separate the two groups from each other, though the above relationship is emphasized by Leuckart and some other authors. RELATIONSHIP WITH PLATYHELMINTHES The idea that Ctenophora gave rise to certain bilateria (Polyclad) has been supported by many Zoologists. Platyctenea has been considered to be a connection link between Ctenophora and the Bilateria. Besides, Ctenophora in general shows many structural similarities with the Platyhelminthes and particularly with the Turbellarians. The similarities are: - General ciliation of the body. - The dorsal polar nerve of turbellaria can be compared with the statocyst of Ctenophora. - Origin of the so-called mesoderm is more or less similar. - Primary locomotor organs in the larva (Mullaer’s larva) consist of eight ciliated ridges of ectoderm which can be compared with the Ctenophora meridional comb-plates. - Ctenophora exhibits both radial as well as bilateral symmetry types. The view that the primitive Bilateria have evolved through Platyctenea, has not been accepted. Because a thorough examination of the Platyctenea reveals that it is a Ctenophore which has become extensively modified for sessile habits. It can further there be suggested that Platyctenea is a tissue grade diploblastic animal, whereas Polyclad is an organ grade triploblastic form. Moreover, it is said that amongst the Platyhelminthes, the Acoela is the most primitive group and not Polyclad. The Ctenophora, on the other hand, shows no close similarity with the Acoela. RELATIONSHIP WITH NEMERTINES The larval form of nemertine (Pilidium larva) shows some similarities with Ctenophores. In Pilidium larva locomotion is performed by lobed bands of cilia. The aboral end of the body contains a cup like sense organ. But these similarities are of no use as the anatomical organization in both shows many diversities. Ctenophores lack mesoderm proper. Paired tentacles with basal musculature are present in Ctenophora. The aboral sense organ in both differs widely. EXAMPLES OF CTENOPHORA TYPES: 1) Cestum: Cestum is commonly known as Venus’s girdle. It inhabitats seas specially the Mediterranean. The body is greatly elongated horizontally in the sagittal, and compressed in the lateral plane so that it appears ribbon-like structure. It is usually green, blue or violet coloured. Out of eight comb-plates, four are very small and the other four are continued all along the aboral edge of the body. The base of two principle tentacles are large and are enclosed in sheaths and numerous small tentacles spring from grooves and are continued the whole length of oral edge. The small lateral tentacles line the oral ridge. Mouth lies in the mid-oral edge. The only sense organ, statocyst, is found on the aboral edge. Cestum swims mainly by graceful serpentine movements by the muscular undulations of the body as well as by beating of comb plates. 2) Ctenoplana: Ctenoplana is small marine, solitary planktonic ctenophore reported once of the West Coast of Sumatra and once among the islands to the East of Papua. Its body is nearly circular in outline and flattened dorsoventrally. It measures about 6 mm in diameter. Its dorsal surface is mostly olive green, brown or reddish in color. In the centre of aboral or dorsal surface of its body is a sense organ with a statolith surrounded by a ring of small ciliated tentacles. The mouth lies in the center of ventral or oral surface. Tentacles are two, pinnate and retractile. The organs of locomotion are eight small deeply sunk swimming plates. 3) Coeloplana: Coeloplana is a marine solitary Ctenophore found in the Red Sea and on the Coast of Japan. It resembles in most of the features to Ctenoplana. However, its body is oval and dorso ventrally flattened but elongated in the tentacular plane. It measures about 60 mm in length. Mouth is ventral in position. Tentacles paired and retractile. Statocyst is dorsally placed. Swimming plates are not found. The fertilized eggs are attached to the oral surface of the mother by the sticky secretion and each develop into a typical cydippid larva with comb plates. 4) Beroe: Beroe is commonly known as sea mitres or mitre jelly fish. It is found in great swarms and cosmopolitan in distribution. Its body is thumble-shaped and measures about 10-20 cm in height. It is usually pinkish in color. The rounded aboral end bears the sense organ surrounded by polar fields. The oral end is truncated and it is occupied by a wide mouth. The greater part of the body is occupied by the huge stomodaeum, infundibulum, per radial and infundibular gastrovascular canals, etc. Tentacles are not found in any stage of this life. A small stomach is placed near the sense organ. It is voracious feeder, swallowing crustaceans and sometimes other Ctenophores even larger than itself. 5) Pleurobrachia: Pleurobrachia is a member of the phylum Ctenophora and it is commonly referred to as a sea gooseberry.Pleurobrachia has a pear shaped body about 5-20 mm in diameter, and transparent. The mouth is situated at the center of the oral pole and the opposite or aboral pole is occupied by a complicated and special sense organ. This sense organ is modified statocyst and acts as an organ of equilibrium. From the opposite sides of the broad end hang two long tentacles provided with numerous little tag like process. The surface bears eight equidistant meridional bands or swimming plates or coatae starting from near the aboral pole and extending about two-thirds of the distance towards the oral pole. The body is covered externally by a delicate ectodermal epithelium. The epithelium of stomodaeum is ectodermal and that of infundibulum and its canals are endodermal. Both ectodermal and endodermal epithelia are ciliated. Their bodies are virtually transparent and the many cilia refract the light, producing rainbow-like colors that can give the false appearance of bioluminescence. The branched tentacles can be white, yellow, pink or orange. They have no nematocysts (stinging cells). Instead, the two long extensile branched tentacles are armed with colloblasts. OBJECTIVES To study Ctenophora- Affinity and types study. SYNOPSIS Ctenophora is a phylum of animals that live in marine waters worldwide. Their most distinctive feature is the "combs", groups of cilia they use for swimming, and are the largest animals that swim by means of cilia. Adults of various species range from a few millimeters to 1.5 m in size.Ctenophores are also characterized by the colloblasts found on the surface of their tentacles. These specialized glue cells secrete a sticky substance that helps ctenophores to catch prey. The phylum Ctenophora includes about 150 species which are categorized into two classes- Tentaculata and Nuda. The tentaculata comprises of 4 orders: Cydippida, Lobata, Cestida and Platyctenea. The class Nuda includes a single order: Beroida. The examples of the phylum are: Comb jelly, Beroe, Ctenoplana, Pleurobrachia, etc. FREQUENTLY ASKED QUESTIONS Q1. Both cnidarians and ctenophores have two cell layers, with an acellular, jelly-like layer in between. True or False A1. True Q2. What is an example of Ctenophora? A2. Ctenophores, variously known as Comb jellies, Sea gooseberries. Beroe, etc. Q3. What is Acoela ? A3. Acoela is a taxonomic order of very small invertebrate animals which resemble flatworms. This order is treated either as a group within the flatworms or as one of two classes within the phylum Acoelomorpha, in which case Acoela contains the majority of that phylum's species. QUIZ Q1. An older term that refers to both the Cnidaria and the Ctenophora is ___________, which means hollow gut. Q2. A. Enteron B. Polymorphism C. Gastrovascular D. Coelenterata Of the Eumetazoa listed below, the _______________ are generally considered the most primitive. A. Chordata B. Rotifera C. Ctenophora D. Onychophora Q3. Q4. Q5. Q6. Paragastric cavity of Sponges is lined by A. Choanocytes B. Amoebocytes C. Chromocytes D. Pinacocytes Symmetry of Ctenophores is generally A. Bilateral B. Radial C. Biradial D. Absent Comb plates of Ctenophores possess A. Cilia for locomotion B. Cilia for filter feeding C. Teeth for crushing D. Teeth for filtering Nematocyst is a Group of cell Organ Part of a cell d. Cell ANSWER Q1. An older term that refers to both the Cnidaria and the Ctenophora is ___________, which means hollow gut. A1. Coelenterata Q2. Of the Eumetazoa listed below, the _______________ are generally considered the most primitive. A2. Ctenophora Q3. Paragastric cavity of Sponges is lined by A3. Choanocytes Q4. Symmetry of Ctenophores is generally A4. Radial Q5. Comb plates of Ctenophores possess A5. Cilia for locomotion Q6. Nematocyst is a A6. Cell SUMMARY - Ctenophora commonly known as comb jellies is a phylum of animals that live in marine waters worldwide. Their most distinct feature is the "combs", groups of cilia they use for swimming. - The Ctenophores are characterized by having biradially symmetrical bodies, eight meridional comb-plates on the body,mostly triploblastic, mesenchymal muscles, an aboral sense organ and adhesive cells. - The comb rows of most planktonic ctenophores produce a rainbow effect, which is not caused by bioluminescence but by the scattering of light as the combs move. - The Phylum Ctenophora includes about 150 species which are categorized into two unequal classes- Tentaculata and Nuda. The Tentaculata comprises of 4 orders: Cydippida, Lobata, Cestida and Platyctenea. The class Nuda includes a single order: Beroida. The examples of the phylum are: Pleurobrachia, Bolinopsis, Cestum, Ctenoplana, Beroe, etc. - The phylogenetic status of Ctenophora is controversial. They have possibly diverged from the early trachyline stem from which holds the ancestry of Cnidarians. Tutorials This topic does not contain any tutorials. Assignments This topic does not contain any Assignment. Reference 1). E.L. Jordan and P.S. Verma. (2003). Invertebrate Zoology. 2). Richard C. Brusca, Gary J. Brusca. (2003). Invertebrates Zoology. Links http://en.wikipedia.org/wiki/Ctenophora http://www.ucmp.berkeley.edu/cnidaria/ctenophora http://faculty.washington.edu/cemills/Ctenophores.html http://jellieszone.com/ctenophores.htm GLOSSARY MESOGLEA Mesoglea is the translucent, non-living, jelly-like substance found between the two epithelial cell layers in the bodies of coelenterates. The mesoglea is mostly water. STOMODAEUM The Stomodaeum, is a depression between the brain and the pericardium in an embryo, and is the precursor of the mouth and the anterior lobe of the pituitary gland. POLYCLAD The Polyclad represents a highly diverse clade of free-living marine turbellarian flatworms. They are known from the littoral to the sublittoral zone, and many species are common from coral reefs. Only a few species are found in freshwater habitats. BIRADIAL SYMMETRY Biradial symmetry both radial and bilateral. is one type of body symmetry which is