Phylum Platyhelminthes

The Flatworms Part 1 Free-living Flatworms

Emerging Patterns in Evolution

Evolution of a head end: cephalization

Mesoderm

Kidney
Internal fertilization

Bauplan

Dorsoventrally flattened
Triploblastic
Embryo

has 3 cell layers

Ectoderm (cnidarians) Endoderm (cnidarians) AND NOW, introducing MESODERM!

Acoelomate - Hey, wheres my cavity?

endoderm

Bauplan

Bilaterally symmetrical
Nervous system
Anterior

brain Paired ventral nerve cords

Longitudinal

Incomplete gut
No

anus Food exits via mouth

Bauplan

Excretory system & osmoregulation

Diffusion

across body wall Protonephridia

Specialized excretory organs May also help osmoregulate

Life History

Most are simultaneous hermaphrodites

Function

as male and female at the same

time Transfer sperm and receive sperm at same time Most are not self-fertile

(A few exceptions)

Classification

Phylum Platyhelminthes
Classes

Turbellaria: free-living flatworms

Trematoda: flukes All are parasites

Cestoda: tapeworms All are parasites Monogenea? (Maybe belong with cestoda) Ectoparasites of fish

Class Turbellaria

Free living
Freshwater

Planarians such as Dugesia

few inhabit terrestrial (moist) habitats

Class Turbellaria
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 on a variety of reef invertebrates

Class Turbellaria

Locomotion
Usually

by cilia

Dugesia can move at about 1.5 mm/sec

Peristaltic

waves can achieve higher

velocities

Marine flatworms can use this to swim gracefully

Class Turbellaria

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 Turbellaria

Body construction
See

lab notes

Live Dugesia Slides of whole planaria and cross sections

Respiration
Diffusion

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

Class Turbellaria

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 Turbellaria
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").

Excretory pores

Class Turbellaria

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 Turbellaria

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 Turbellaria

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 Turbellaria

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 Turbellaria

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 Turbellaria

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 Turbellaria

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.

Polycladida digestive system

Class Turbellaria

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
Auricle

Cerebral ganglion

Paired nerve cords

(connectives)

Retinular cells Light sensitive region

Pigment cups

Class Turbellaria

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 See film notes

nervous system
male female

gut

Class Turbellaria

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 Mullers larva into a free-living Turbellarian (above)

A living larvae (right)

Class Turbellaria

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 Turbellaria

Additional sketches from R. Fox.

The dorsal epidermis contains numerous 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

Many flatworms are brightly coloured

Warning coloration (aposomatic coloration) These flatworms are believed to be toxic or distasteful

Other

species are mimics of toxic nudibranchs (sea slugs)

Ecology and Evolution

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 colour 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 Evolution

Evolution of mimicry
Imagine

that the nudibrach Phyllidia is the

'model
If a flatworm species has a vaguely similar shape and colour 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 - a sea slug

(Phylum Mollusca)

Explain how this scenario fulfills the 3 conditions necessary for evolution to occur.