Traditional Phylogenetic position of flatworms

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Traditional Phylogenetic position of flatworms

Terrestrial triclad Bipalium

PORIFERA

CNIDARIA

CTENOPHORA

PLATYHELMINTHES

P

D

ASCHELMINTHES

NEMERTEA

MOLLUSCA

SIPUNCULA

ANNELIDA

POGONOPHORA

ECHIURA

ONYCOPHORA

ARTHROPODA

CHAETOGNATHA

LOPHOPHORATES

ECHINODERMATA

HEMICHORDATA

CHORDATA, CEPHALOCHORDATA

UROCHORDATES

Pseudoceros crozieri polycladid

PLACOZOA, MONOBLASTOZOA,

RHOMBOZOA

Rupert and Barnes (1994)

The Flatworms – Platyhelminthes- Traditionally with 4 Classes

Trematoda - Parasitic flukes Monogenea - Parasitic flukes (1 gen)

Rugogaster hyrdolagi

Cestoda Taenia pisiformis - Scolex, and anterior suckers of a tapeworm.

Turbellaria

-

A triclad marine flatworm

Phylum Platyhelminthes (Flatworms )

Parasitic or free-living, unsegmented worms

Triploblastic, bilateral symmetrical, spiral cleavage, flattened dorsoventrally

Acoelomates (solid body forms lacking the fluidfilled space, such as coelom, pseudocoel, or hemocoel, between body wall and gut that is seen in more complex metazoans)

Complex but incomplete gut. No gut is some parasitic flatworms (Cestoda)

Cephalization, central nervous system with an anterior cerebral ganglion and sometimes longitudinal nerve cords connected by a ladderlike nervous system

Protonephridia, complex excretory/osmoregulatory structures

Hermaphroditic, complex reproductive structures. Mutually insemination in sexual reproduction . Spermatozoa with two tails

Body plans of triploblastic Metazoans

Acoelomates

Blastocoelomates

Eucoelomates

Class Turbellaria – (Free Living Flatworms)

Bilaterally symmetrical-vary in size from <1 mm to several cm

Cylindrical body in small turbellarians, dorsoventrally flattened in large

Epidermis is ciliated, ciliary locomotion

Secrete solid mucus pellets called rhabdoids

If gut present with one opening (mouth present, no anus), carnivores. Mouth located on ventral surface that opens into the GV cavity.

No circulatory system (diffusion), protonephridia for excretion

Cheliplana sp. (Rhabdocoela)

Connective tissue called parenchyma

Some cephalization

Most likely paraphyletic group

Maiazoon orsaki

Turbellarian vs hydroid epidermis

Rhabdoids (rod-shaped epidermal inclusions) produced by mesenchymal cells

( rhabdites), stored in packets. When released, produce slime (chemical defense)

Gland cell

Gut organization

Digestive system typically a blind sac except where high degree of branching makes return to mouth of wastes difficult

Epithelium consists of gland cells and phagocytic cells

Gut may be simple, unbranched (acoels)- lacks permanent cavity (but with syncitium)

Gut may be highly branched with diverticula – increase surface area for digestion/absorption

Pharyngeal patterns

• Simple pharynx – opening (mouth) directly into digestive syncitium

• Bulbous pharynx – reduction of pharyngeal cavity, separated from parenchyma by muscular septum – can act independently of pressure in body

• Plicate pharynx – muscular pharyngeal tube within a cavity – probably originated as folding of simple

• pharynx

Jawed Proboscis

Cheliplana, Turb.

Dugesia, x-section in the region of the pharynx

Class Turbellaria (Cont’d)

Triclad Dugesia

Most famous example is Dugesia (formerly Planaria ) , a triclad planarian. Most students have worked with them in introductory and invertebrate zoology courses

Oval to elongate in shape

Dorsoventrally flattened, more so with size

Usually quite small (<10 mm) but some very large exceptions ( Rimacephalus arecepta - 60 cm)

Primarily aquatic and most marine, interstitial

• 12 Orders: Acoela, Catenulida, Haplopharyngida,

Lecithoepitheliata, Macrostomida, Nemertodermatida

Polycladida, Proseriata, Tricladida, Rhabdocoela,

Proplicastomata, Proplicastomata. The 2 superorders

Archoophorans and Neoophorans (based on reproductive system) are almost defunct.

Archoophorans- entolecithal (yolk is deposited within the cytoplasm of the ova), development may include planktonic larva, but usually direct

Nucleus

Triclad Bdelloura

Yolk granules

Cell membrane

Class Turbellaria (Cont’d)

Neophorans – ectolecithal (yolk is deposited outside the cytoplasm of the ova), development always direct

Yolk cells

Yolk granules

Most are marine, but some freshwater forms

Yolkless egg

Egg capsule

Larval Forms

4 arms or lobes – Goette’s larva

8 arms or lobes – Mueller’s larva

These eventually resorb arms and develop into small adult worms

Müller's larvae

Müller's larva of

Pseudoceros canadensis

Unidentified Müller's larvae

Archoophoran reproductive plan

Acoels-no separation of gonads by epithelium, ovaries/testes absent

Macrostomida-ovaries/testes present, well developed accessories

Polycladida – multiple branches of ovaries/testes, well developed accessories

Neophorans reproductive plan

Main characteristic – separate vitellarium or yolk producing glands

Brusca and Brusca (2002)

Asexual reproduction;

• Paratomy –formation of 2 or more zooids (buds) along axis of body prior to division

• Architomy – transverse splitting of body into several pieces, each of which regenerates to form a complete individual AFTER fission

Sexual reproduction

Most turbellarians in general are hermaphroditic and have internal fertilization

Dorsoventral muscles may occur and aid in flattening body

Temporary attachment organs allow adhesion to substrate

(duogland organs)

Circular muscles occur below the parenchyma and epithelium - elongation

Longitudinal muscles occur interiorly – contraction in length

Diagonal muscles – twisting

Muscles are usually smooth

Types of Movement

A. Ciliary movement

B. Undulation

C. Muscular creeping

D. Didactically

E. Contraction

G. Peristalsis to mix gut contents

H. Secretion of mucus strings

I. Somersaulting

Dugesia

Detail of the lateral margin of the body of Dugesia showing the adhesive gland on the ventral surface and the differentiation of the cilia between the dorsal and ventral surfaces.

Dorsal

Ventral

Excretion

– Via Protonephridia (major advance over diploblasts) – simple filter cells (flame bulb) with 1 or more internal flagella that draw fluid into cell – wastes empty to the exterior by nephridiopores. Some ammonia excreted but most metabolic waste by diffusion

No protonephridia on Acoela,

Nemertodermatida and some catenulids

Flame cells connected to collecting tubules, protonephridial system in a freshwater turbellarian.

Turbellarian Nervous system

Ladder-like nervous system

Advanced condition

Dugesia

Cephalization of nervous system

Cross linked (segment like)

Anterior end with brain, eyespots and the longitudinal nerve cords

Interstitial turbellarian

Statocyst sensory bristles

Turbellarian nervous systems and organs

Polyclad Planocera

Triclad Crenobia

Inverted pigment cup ocellus (x-sec)

Anterior end of rhabdocoel

Mesostoma (x-sec ) with tactile, chemo- and rheoreceptors

Nervous systems of other Platyhelminthes

Brusca and Brusca (2002)

fluke Probolitrema

Chinese liver fluke

Schistosomiasis

(swimmer’s itch)

Stylochus

Taenia

Tegument organization of flukes and cestodes

Brusca and Brusca (2002)

Class Monogenea (parasitic flukes)

Monogenean flukes do not have a well-developed sucker.

Posterior end with a bulbous structure covered with hooks called an opisthaptor.

Anterior end with buccal sucker (prohaptor)

Most monogeneans are ectoparasites on fish or other aquatic animals, although a few live in the urinary bladders of turtles and frogs.

Their life cycle involves a single host. Eggs hatch into ciliated larvae, which may attach directly to a host or swim freely for a time before attaching.

Hermaphroditic

Adults lack external cilia

Prohaptors and Opisthaptors of Monogeneans

Adhesive organs – adaptations for parasitic life style

Opisthaptor – main organ of attachement

Prohaptor Opisthaptor

Larva of monogeneans

Class Trematoda (digenetic parasitic flukes)

All parasites, most adults parasitize vertebrates, about 9000 species

Body covered with a tegument, a peculiar kind of epidermal arrangement in which the main cell bodies are deep, separated from the cytoplasm that lies next to the exterior by a layer of muscle (but connected to the exterior layer by cellular processes. The tegument lacks cilia in adults.

Exterior layer syncytial (i.e continuous, not broken by cell membranes)

Unlike monogeneans, trematodes have no opisthaptor; with 1 or 2 hookless suckers. Acetabulum. Most are endoparasites.

With a developed alimentary canal (like turbel.), muscular, excretory, and reproductive systems are almost complete.

Complex life cycles, with larval stages parasitizing one or more species that are different from host of adults. Larval stages include miracidium, redia, cercaria, and metacercaria.

Enormous impact on humans (liver flukes, blood flukes causing schistosomiasis).

Human liver fluke

Opisthorchis sinensis

General features of a digenetic fluke

Similar to turbellarian flatworms

• Neophoran

• Diverticulated gut

• Specialized genital organs

Genital pore

Copulating pair of human blood flukeSchistosoma mansoni

Mouth

Oral sucker

Intestine

Ventral Sucker (Acetabulum)

Vitellaria - yolk glands

Testes

Ovary

Miracidium ciliated

Ingested or penetrates snail host

Cercaria

Metacercaria

Redia

Representative Digenean Lifecycles

Sporocyst with redia

Egg - encapsulated, passes in urine, feces

Life cycle of blood fluke

Schistosoma mansoni one of 3 spp, causing schistosomiasis.

Different spp. Affect different organs

(e.g. blood vessels, bladder, liver)

Brusca and

Brusca (2002)

Egg

Cercaria larva

Digenetic Fluke Life stages

Metacercaria

Redia larva

Cercaria larvae

Redia larva

Class Cestoda (tapeworms)

Bodies are long and flat, made up of many segments called proglottids. Each proglottid is a reproductive unit.

Adults lack cilia and their surface is a tegument (as in monogeneans and trematodes), but in cestodes the tegument is covered with tiny projections, microvilli, which increase its surface area and thereby its ability to absorb nutrients from a host.

Digestive tracts are absent completely.

Anterior end with is a specialized segment called a scolex, which is usually covered with hooks or suckers and anchors it to the host.

Proglottids

About 5000 species, all endoparasites.

Require two hosts, with the host of the adult tapeworm a vertebrate.

Intermediate hosts are often invertebrates. Many species inhabit humans.

Proglottids

General features of an intact tape worm

Scolex

Reduced or lost nervous system and gut

Internal parasites

Reproductive machines

Hexacanth larva

Rostellum-on the tip of scolex

Sometimes retractable

Myzorhynchus-Protrucible sucker

3 types of adhesive suckers

Bothria – elongated grooves

Bothridia – 4 leaf-like structures with suckers at anterior ends

Acetabulum – true suckers (4)

Cestode Attachment

Brusca and Brusca (2002)

Phylogeny of Platyhelminthes

Brusca and Brusca

(2002)

2- prohaptor

3-- opisthaptor

4– ventral sucker

5-- tegumental microtriches

6-- scolex

7– loss of digestive tract

8– strobilation of the body

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