Phylum Platyhelminthes
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Phylum Platyhelminthes (flatworms). A phylogeny of Animalia. Platyhelminthes. Bilateria. Phylum Platyhelminthes. The Flatworms Part 1: Free-living Flatworms. Emerging Patterns in Evolution. Bilateral symmetry Dorsal & ventral, anterior & posterior

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A phylogeny of

Animalia

Platyhelminthes

Bilateria


Phylum platyhelminthes

Phylum Platyhelminthes

The Flatworms

Part 1: Free-living Flatworms


Emerging patterns in evolution
Emerging Patterns in Evolution

  • Bilateral symmetry

    • Dorsal & ventral, anterior & posterior

    • Associated with locomotion on a solid surface

  • Evolution of a head end: cephalization

    • Possess a head (leading end with sense organs)

  • Internal fertilization


Bauplan
Bauplan

  • Dorsoventrally flattened

    • Lack circulatory system for transport‑ must rely on diffusion for gas exchange

  • Triploblastic

    • Embryo has 3 cell layers

      • Ectoderm (cnidarians)

      • Endoderm (cnidarians)

      • AND NOW, introducing THEMESODERM!

  • Acoelomate - Hey, where’s my cavity?


Phylum Platyhelminthes – tissue organization

  • Triploblastic = 3 tissue layers

    • Ectoderm (outer)Endoderm (inner)Mesoderm (between)


Muscles and movement
Muscles and Movement

Hydrostatic Skeleton


Bauplan1
Bauplan

  • Nervous system

    • Anterior brain

      • Paired ganglia and nerves

    • Pairedventral nerve cords

      • Longitudinal

      • Lateral

  • Incomplete gut

    • No anus

    • Food exits via mouth


Phylum Platyhelminthes - digestive system

  • Gastrovascular cavity

  • Protrusible pharynx in some taxa

  • Digestion is both intracellular and extracellular


Bauplan2
Bauplan

  • Excretory system & osmoregulation

    • Diffusion across body wall

    • Protonephridia

      • Specialized excretory organs

      • Probably mainly for water balance - produces urine- a filtrate of extracellular fluid

      • May also help osmoregulate


Phylum platyhelminthes excretory system
Phylum Platyhelminthes- excretory system

  • “Protonephridial” system: blind‑ending tubes with flame cells & fenestrae at ends, connected with excretory pores

  • Probably mainly for water balance produces urine- a filtrate ofextracellular fluid

  • Flatworms have extracellular fluid compartmentcells protected from external environment

    • Cnidaria lack this- cells are in contact with water (in freshwater cnidaria, each cell has prominent contractile vacuoles)


Protonephridial osmoregulatory/excretory system

“flame cell”

fenestrae

duct


Bauplan3
Bauplan

  • Lack cuticle or other rigid skeletal support

  • Lack fluid-filled body cavity

  • Spermatozoa possess a pair of flagella that arise from the paired centrioles

  • Epidermal cells are typically multiciliated.

  • Neoblasts---stem cells that give rise to the other cell types in the body.



Life history
Life History

  • Most are simultaneous hermaphrodites

    • Function as male and female simultaneously

      • Transfer sperm and receive sperm simultaneously

      • Monoecious

    • Most are not self-fertile

      • (A few exceptions)

    • Most are oviparous (= produce shelled eggs that are released and hatch outside the body).

  • Most can also reproduce asexually by budding or transverse fission


Asexual reproduction by transverse fission budding
Asexual reproduction by transverse fission (budding)

  • Interesting in 2 ways:

    • Another flatworm class (cestoda) is segmented ‑ they grow by serial reproduction of body parts

    • If you cut a turbellarian in half, it can regenerate

  • Regeneration involves undifferentiated cells in mesoderm called neoblasts


Class Cestoda(tapeworms) all parasitic

Class Monogenea(monogenetic trematodes, gill flukes) all parasitic

Class Trematoda(digenetic trematodes, flukes) all parasitic

Class Turbellaria(turbellarians)mostly free-living

Platyhelminthes- 4 classes traditional


Platyhelminthes
Platyhelminthes

  • Inability of flatworms to synthesize fatty acids and sterols may explain their need for establishing symbiotic relationships

  • The parasitic classes of flatworms are linked by a synapomorphy- the tegument(see next slide)

    • Clade Neodermata


Turbellaria

“Neodermata” The parasitic flatworms)

(Tegument)


Classification
Classification

  • Phylum Platyhelminthes

    • Classes

      • Acoelomorpha:

        • Acoels (free-living) + Neodermata (parasites)

      • Turbellaria: free-living flatworms

      • Trematoda: flukes

        • All are parasites

      • Cestoda: tapeworms

        • All are parasites

      • Monogenea? (Maybe belong with cestoda)

        • Ectoparasites of fish



Acoels are not platyhelminthes
Acoels are not Platyhelminthes??? form of the gastrovascular cavity

  • rRNA analysis (see cladograms) shows that Acoela is basal to other bilateral clades

  • Digestive syncytium instead of a gut

  • Netlike nervous system, lack brain

  • Kinked cilia

  • Acoel eggs cleave only once and the two resulting cells immediately generate many small cells.

  • Link to more info on Acoela: http://devbio.umesci.maine.edu/styler/globalworming/


18S rDNA-based maximum-likelihood tree of 61 metazoan species

From:Ruiz-Trillo et al. (1999) Acoel flatworms: earliest extant bilaterian metazoans, not members of Platyhelminthes. Science 283:1919-1923

Platyhelminthes


Class turbellaria
Class Turbellaria species

  • Order Acoela

    • Small flatworms with no permanent gut cavity.

    • Free-living, marine and brackish water.


Rhabodocoel turbellarians
Rhabodocoel turbellarians species

  • Diverse and important predators on meiofauna in freshwater ecosystems

  • Recent research on predation on freshwater bivalves


Mesostoma – species a viviparous rhabdocoel – note the embryos within the uterus


C. Barnhart species

Mesostoma – close-up showing the babies

Mom’s oral sucker

Babies

Mom’s eyes


Macrostomum – species predator on juvenile bivalves


Eggs of species Macrostomum


Bipalium

A land planarian common in greenhouses world-wide – causing problems in Britain


Bipalium species feeding on an earthworm


Class turbellaria1
Class Turbellaria species

  • Order Tricladida

    • Gut with three branches

    • Free living

      • Freshwater

        • Planarians such as Dugesia

      • A few inhabit terrestrial (moist) habitats

    • Neodermata

      • Exclusively parasitic


Class turbellaria2
Class Turbellaria species

  • Marine species (beautiful colors)

    • Significant members of coral reef ecosystems

      • Some are major predators of colonial ascidians (sea squirts)

      • Others are pests of commercial clams and oysters

    • Some live symbiotically with/on a variety of reef invertebrates


Bdelloura species - a marine triclad that is commensal with horseshoe crabs

(most turbellarians are not parasites)


Class turbellaria3
Class Turbellaria species

  • Locomotion

    • Usually by cilia on a layer of mucus

      • Dugesia can move at about 1.5 mm/sec

    • Peristaltic waves can achieve higher velocities

      • Marine flatworms can use this to swim gracefully

      • Also use cilia same way as protists do



Class turbellaria4
Class Turbellaria species

  • Locomotion

    • Terrestrial planarians

      • Glide smoothly on the substrate by the action of powerful, closely spaced cilia in a special medial ventral strip (creeping sole), on a thin coat of mucus secreted on the substrate by glands opening into the creeping sole

      • Planarians that migrate on plants or objects above the ground sometimes lower themselves to the ground on a string of mucus.


Class turbellaria5
Class Turbellaria species

  • Body construction

    • Lab notes

      • Live Dugesia

      • Slides of whole planaria and cross sections


Class turbellaria6
Class Turbellaria species

  • Respiration

    • Diffusion

      • Must be less than 0.5 -1 mm thick for diffusion to be effective. Flatworms are flat by necessity


Class turbellaria7
Class Turbellaria species

  • Excretion

    • NH4 excretion mostly by diffusion through epidermis

    • Protonephridia

      • Mostly osmoregulatory but may also help with excretion

      • Pair of longitudinal canals

      • Open to outside through two dorsal pores

      • Tributaries to excretory canals highly branched, ramify throughout the body;


Class turbellaria8
Class Turbellaria species

  • Protonephridia

    • One end of the tubule opens through a small pore to the exterior.  The other end of the tube ends blindly within the body in a spherical structure containing long cilia - these are called flame cells

    • Excess water (and possibly wastes) enters the flame cell system and is propelled through the tubules toward the outside by the beating of the cilia (the "flame").



Class turbellaria9
Class Turbellaria species

  • Protonephridia

    • Branches terminate in blind flame cells.

      • Flame cells have slits that penetrate the cell

      • Slits are crossed by filaments or a membrane that reduce the effective pore size

      • Slits act as an ultrafilter to keep back large proteins

      • Filtrate of mesenchymal intercellular fluid enters tubule.

        • Inorganic and organic materials actively reabsorbed in tubule; remainder is excreted


Class turbellaria10
Class Turbellaria species

  • Feeding Ecology

    • Most turbellarians are carnivorous predators or scavengers.

      • Carnivores feed on organisms that they can fit into their mouths, such as protozoans, copepods, small worms, and minute mollusks.

      • Some species use mucus that may have poisonous or narcotic chemicals to slow or entangle prey.

      • Some have specific diets and feed on sponges, ectoprocts, barnacles, and tunicates.


Class turbellaria11
Class Turbellaria species

  • Feeding Ecology

    • Several species have commensal relationships with various invertebrates and few actually border on being parasitic because they graze on their live hosts.

    • Land planarians devour earthworms, slugs, insect larvae, and are cannibalistic. Prey are located by chemoreceptors located in a single ciliated pit under the head or in a ciliated ventral groove.


Class turbellaria12
Class Turbellaria species

  • Feeding Ecology

    • Land planarians

      • Struggling prey are held to the substrate and entangled in slimy secretions from the planarian.

      • A few species have symbiotic algae that supply the worm with carbohydrates and fats and the worm supplies the algae with nitrogen waste products and a home.


Class turbellaria13
Class Turbellaria species

  • Feeding and Digestion

    • The pharynx is protruded from the mouth and into the prey.

      • The pharynx and gut cells produce digestive enzymes that breakdown food extracellularly.

      • Nutritive cells in the gastrodermis then phagotize partially digested material that is distributed throughout the body.

      • Because these worm lack a circulatory system, larger species have extensive anastomosing guts to aid in distribution.

      • Since these worms have incomplete guts, all waste must pass back out of the mouth.


Class turbellaria14
Class Turbellaria species

  • Nutrition

    • Planaria store food in digestive epithelium and can survive many weeks shrinking slowly in size without feeding.

    • They are capable of utilizing their own tissues such as reproductive tissue for food when reserves are exhausted.

    • Lab animals often tend to shrink in size when not fed properly

      • Liver or egg yolk

      • Dugesia feeds various invertebrates, including mosquito larvae


Class turbellaria15
Class Turbellaria species

  • Digestive system

    • Ventral mouth

    • Muscular pharynx

    • A blind intestine (details depend on order)

      • Tricladida — 10-20 mm long

        • Tripartite gut; one main anterior branch and two main posterior branches with numerous blind pockets off all three. Dugesia

      • Polycladida — up to 5 cm long

        • Gut has numerous branches ramifying throughout the body.



Class turbellaria16
Class Turbellaria species

  • Nervous system and sensory organs

    • Dorsal, anterior eyes

    • Ciliated pits behind auricles on head are probably chemosensors

    • Dorsal, bilobed brain underlays eyes

    • Primitively 3-4 pairs of longitudinal nerve cords

      • Reduced to two longitudinal ventral nerve cords

      • Run down length of body with numerous cross connections and branches in most groups.


BRAIN species

Auricle

Cerebral ganglion

Paired nerve cords


Retinular cells species

Light sensitive region

Pigment cups


Class turbellaria17
Class Turbellaria species

  • Reproduction

    • Hermaphroditic

      • Worms are male and female at same time

      • Most do not self fertilize

    • Fertilization is internal

      • Stab penis through body wall in marine flatworms

        • Penis Fencing

      • See film notes


nervous system species

male

female

gut


Class turbellaria18
Class Turbellaria species

  • Development

    • Turbellarians have either direct development or produce a pelagic larva.

    • Polyclads often produce a pelagic Muller's larva that settles to the bottom and goes through metamorphosis in a few days.

      • This larva has eight ventrally directed ciliated lobes, which it uses to swim.


Metamorphosis of Muller’s larva into a free-living Turbellarian (above)

A living larvae (right)


Class turbellaria19
Class Turbellaria Turbellarian (above)

  • Asexual reproduction

    • Architomy

      • Type of fission in which the worm divides into two fragments without prior differentiation of new parts.

      • Transverse cleavage just posterior to the pharynx divides the worm into an anterior, nearly normal, worm with head, mouth, pharynx and most of the gut, and an incomplete, headless posterior mass of tissues which must replace its missing parts.

      • Following division, the anterior end behaves normally but the posterior end remains immobile until regeneration is complete and the missing parts replaced.


Class turbellaria20
Class Turbellaria Turbellarian (above)

  • Additional sketches from R. Fox.


The dorsal epidermis contains numerous Turbellarian (above)secretory vesicles and rod-shaped membrane enclosed secretions, the rhabdites (rhabd = rod). Rhabdites are synthesized by epidermal gland cells submerged below the basal lamina into the parenchyma. When expelled at the surface, rhabdites absorb water and expand to become sticky mucus which may help trap small invertebrate prey.


Ecology and evolution
Ecology and Evolution Turbellarian (above)

  • Many flatworms are brightly colored

    • Warning coloration (aposomatic coloration)

    • These flatworms are believed to be toxic or distasteful

  • Other species are mimics of toxic nudibranchs (sea slugs)


Ecology and evolution1
Ecology and Evolution Turbellarian (above)

  • Evolution of mimicry

    • Imagine that the nudibrach (a type of shell-less mollusc) Phyllidia is the 'model’

      • Has extremely toxic and distasteful secretions which deter fish from eating it

      • Fish learn to avoid the Phyllidia shape and color and so Phyllidia are somewhat protected from predation

      • Any animal that looks like Phyllidia will gain some protection from predation by 'tricking' predatory fish into thinking they are a Phyllidia .


Ecology and evolution2
Ecology and Evolution Turbellarian (above)

  • Evolution of mimicry

    • Imagine that the nudibrach Phyllidia is the 'model’

      • If a flatworm species has a vaguely similar shape and color to Phyllidia then those individuals that look most like a Phyllidia are most likely to escape fish attack

      • Gradually the surviving flatworms of each generation will become more and more like Phyllidia simply because only those with the genes to look like Phyllidia will survive.


Phyllidia Turbellarian (above)- a sea slug (Phylum Mollusca)



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