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PLATYHELMINTHES

Introduction
- group of some 34,000 species
- more than 80% of all describes platyhelminth species are parastites
- includes one class of mostly free-livig individuals (the tubellarians)
and three classes of exclusively parasitic individuals (the
monogeneans, trematodes, and cestodes)
- oldest known flatworm fossil is approximately 40 million years old;
was found about 10 years ago, preserved in a fossilized tree resin
(amber)

General Characteristics
- most conspicuous feature is that they are flat
- no uniquely defining characteristics (synapomorphies)
- all are acoelomate and bilaterally symmetrical Eyes
- development is protostome-like; cleavage is spiral and at least in - one or more pairs
some species, determinate - anterior
- mouth forms before the anus and forms from the blastopore
- have conspicuous anterior brain which is connected to at least one Statocyst
pair of longitudinal nerve cords - only about 10% of the species have this
- only a single pair of nerve cord is present (most advanced species); - provides feedback about body orientation
these cords are always located ventrally
- they have no anus Body
- food enters and unmetabolized wastes leave the digestive system - outer surface is ciliated; often more so on the ventral surface than on
through a single opening the dorsal surface
- no specialized circulatory system (although a few possess - most species move at least partly by secreting mucus from the ventral
hemoglobin) surface and beating the ventral cilia withing this viscous mucus
- no specialized respiratory system
- gas exchange is accomplished by simple diffusion across the body Rhabdites
surface - aggregation of small and cylindrical on the body surface
- metabolic wastes probably move out of flatworms mostly by diffusing - related to rhabdoids
across the general body surface - function is uncertain; they release a thick mucus that coats the
animal's body, possibly in response to attempted predation or to
Parenchyma tissue desiccation
- a loose collection of cells where their mesodermal layer embryo
develops Locomotion
-involves subtle waves of muscular contraction along the animal's
Protonephridia ventral surface
- a series of specialized organs that consist of a group of cilia projecting - pedal waves are unidirectional, moving from the anterior of the worm
into a fine-meshed cup posteriorly
- creates negative pressure, drawing fluid through the mesh cup and - the musculature of the body wall includes fibers running longitudinally,
into the protonephridial tubule circumferentially, dorsoventrally, and diagonally; this movement is
called looping
FORMS OF PROTONEPHRIDIA - for effective looping locomotion, the flatworm must be able to adhere
locally to the substrate to prevent sliding backward while the pulling
and pushing forces are being generated
Flame cell
- the beating cilia in the cup, which is likened to the flickering of a flame
Solenocytes
- single flagellum found within the cup

REPRODUCTION
- majority of the flatworms are simultaneous hermaphrodites
- sperm exchange and egg fertilization can occur when any individual
encounters another individual of the same species
- they cannot fertilize themselves

Clades
Neodermata (new skin)

CLASS TURBELLARIA
- only about 15% of all flatworm species

General Characteristics
- typically less than 1cm long
- prone to dehydration
- most are aquatic: most are marine; some are found in freshwater; a
few are terrestrial, although these are restricted to very humid areas
- most aquatic are benthc; they live in or on the ocean, lake, pond, or
river bottom
- most 4,500 are free-living; about 150 are commensal or parasitic with
other invertebrates

Nervous system
- consists of a coelenterate-style, diffuse nerve net in the most primitive
turbellarian species
- has a cerebral ganglion-a primitive but distinct brain
- has one to three (rarely four) pairs of longitudinal nerve cords
Paired secretory cells
- flatworms typically posses a large number of paired secretory cells
(duo glands)
- located on the ventral surface and opening to the exterior
- one cell produces a viscous glue while the other cell presumably
secretes a chemical that breaks this attachment to the substrate

Digestive System
- most are active carnivores; some ingest detritus and algae; a number
harbor algal symbionts
- fairly simple
- differences in the structure of the mouth parts and the gut, along with
differences in reproductive morphology, are key elements in dividing
the class into its 12 constituent orders
- bear a simple mouth opening on the ventral surface
- no well-formed gut cavity
- may be straight, three-branched, or multi-branched
- in more advanced species, mouth is often borne at the end of a
protrusible pharynx
Regeneration
- in other species, mouth possesses a separate proboscis that spears
- many possess remarkable regenerative powers that go far beyond the
food items and then transfers them to an adjacent mouth opening
ability to repair wounds
- digestion of food is initially extracellular through secretion of enzymes
- regeneration is accomplished through the activities of neoblasts
- particles are later phagocytosed, and digestion is completed
- many reproduce routinely by asexual fission, an ability closely linked
Intracellularly
to their great regenerative capabilities

Neoblasts
- undifferentiated cells
- unique to turbellarians
- has remarkable versatile developmental plasticity

Reproductive System
- both male and female reproductive organs are found within a single
individual
- male system is particularly complex
- when triclad flatworms mate, each individual typically inserts its penis
into the female opening of the other member of the pair, so sperm
transfer is reciprocal
- copulation may occur by hypodermic impregnation, in which the
stylets of the penis pierce the body of the partner
- the egg of each animal is released after fertilization and generally
develop directly into miniature flatworms within a protective capsule
- there is no free-living larval stage in the life cycle
- developing embryo gives rise to a short-lived microscopic CLASS CESTODA
free-swimming larval stage; most commonly a Muller’s larva
Defining Characteristics
- small anterior hooked attachment organ (scolex)
- division of body into segments (proglottids) arising from anterior end,
behind the scolex
- absence (loss) of digestive tract
General Characteristics Egg development
- total length may exceed 10-12m - after an oncosphere hatches, it lyses (dissolves) its way through the
- most are members of the subclass Eucestoda and commonly known intestinal wall, taking up residence as an encysted form in the coelomic
as tapeworms space or in specific organs and tissues of the host
- they are endoparasitic; they primarily parasitize vertebrates, - in genus Taenia, oncosphere typically produces a single resting
inhabiting regions of the host digestive tract individual called a cysticercus or bladder worm
- an estimated 135 million people worldwide have tapeworm infections - in some species, the resting stage divides asexually many times
within the intermediate host to produce a large, sometimes deadly,
Body hydatid cyst
- a nonciliated tegument covers the cestodes - asexual replication is common among taeniids (family Taeniidae) but
- the tegument contains numerous nuclei, but these are not separated is rarely encountered among other cestode species
by cell membranes; that is, the tegument is syncytial - further development is arrested until the intermediate host is eaten by
- outer surface is outfolded into numerous cytoplasmic projections a different host (w/c could be the final host or another intermediate
- it has no mouth or digestive tract of its own at any point in its life cycle host)
- the complete sequence from egg to adult is achieved only if the
Scolex fertilized cestode eggs reach the appropriate intermediate hosts, which
- an anterior end in turn must be ingested by the appropriate final or definitive host
- studded with hooks and/or suckers that are used to maintain position - fishes, cows, pigs, dogs, and sometimes birds may serve as
within the host’s gut intermediate host
- most species have this but relatively few lack this, placing them within - humans often serve as acceptable final and intermediate hosts
a second subclass called the Cestodaria - most invertebrate intermediate hosts are arthropods

Neck
- a region that lies just posterior to the scolex
- the essence of tapeworm existence
- a seemingly endless series of sections called proglottids bud from the
neck of most cestodes at a rate of several per day

Proglottid
- rarely more than 3-5 mm long
- involved primarily with the process of sexual reproduction
- each proglottid contains both male and female reproductive systems
(hermaphrodite)
- each proglottid contains perhaps 50,000 eggs
- in some species, the posteriormost proglottids break off periodically
- in other species, mature proglottids burst open, releasing the fertilized
eggs into the host’s gut CLASS MONOGENEA

Reproduction Defining Characteristics


- eggs are generally fertilized by sperm from a neighboring cestode, but - posterior attachment organ (haptor [=opishaptor]) including sucker
they can be fertilized by sperm from the same individual--or in fact, and complex attachment hooks and sclerites
from the same proglottid - larva (oncomiracidium) bearing 3 bands of cilia and usually 1 or 2
- estimated 2000 to 4000 proglottids are produced per individual in pairs of eyes
many specis, and an incredible number of fertilized eggs (tens of
thousands, or even hundreds of thousands) are produced per day per General Characteristics
cestode - usually parasitic on the skin or gills of fishes; ectoparasites
- eggs leave the host’s body along with the feces (ecto = G: outside)
- generally, eggs cannot take up residence in the definitive (final) host - attaches primarily by means of suckers, hooks, and complex sclerites
immediately; they must first enter an intermediate host, or in some located at its posterior end
species, a series of intermediate hosts - most species show a very high level of host specificity; typically
occupy highly specific sites within a host
Ex: one species live only at the base of a fish’s gill filament while
members of another species are found only near the tips of the same
filaments
- only about 8,000 species have been described so far
- about 25,000 species are though to exist, which would make this the
most speciose group in the Platyhelminthes

Haptor
- highly specialized posterior attachment organ (hapto = G: fastened)
(= opisthapto; opistho = G: behind, at the near)

Prohaptor
- anterior adhesive organ consisting of suckers and adhesive glands
that aids in attachment

Life cycle
- there are no intermediate hosts, so the life cycle generally involves
the following stages:

Taxonomic Status
Oncosphere - has been uncertain for some time
- a larva that hatches out when a fertlized eggs is ingested by the - long thought to be trematodes (flukes) since they closely resemble as
appropriate intermediate host adults
- has muscles, flame cells, and most significantly, 3 pairs of hooks with - some argued that monogeneans are more closely related to the
which it attached to the wall of the host’s digestive tract cestodes because of the similarity between the cestode oncosphere
- oncus = a barb or hook and the monogeneans oncomiracidium
- others say that monogeneans are sufficiently dissmilar from either
cestodes or trematodes to merit categorization as a separate class
Life cycle
CLASS TREMATODA
- each fertilized egg generally gives rise to a single, free-living, ciliated
miracidium larva
General Characteristics
- all are parasitic
Miracidium
- most reach adulthood only as parasites in or on vertebrate host
- gutless
- flukes are placed into one major group with over 6,000 species, the
- almost always eaten by a mollusc, most commonly a snail
digenetic trematodes (also known as digeneans)
(intermediate host)
- one much smaller group with fewer than 100 species, the
- must generally enter a particular species of molluscan host to develop
aspidogastreans (also called aspidobothreans)
further
- a single miracidium generally gives rise to, on average, tens of
The successful parasite must
thousands of cercariae
1. Reproduce within the definitive host;
- host penetration is accomplished by secretions from several glands
2. Get fertilized eggs or embryos out of the host;
located anteriorly in the miracidium
3. Contact and recognize a new, appropriate host;
- miracidium then dedifferentiates into a mother sporocyst stage
4. Obtain entrance into the host;
5. Locate the appropriate environment within the host;
6. Maintain position within the host;
Mother sporocyst
7. Withstand what is often a rather anaerobic (oxygen-poor)
- most of the miracidial structures (including the external cilia) are lost,
environment;
except for the protonephridia
8. Avoid digestion or attack by the host’s immune system, and
- lives in the molluscan blood circulatory system (the hemocoel)
9. Avoid killing the host, at least until reproduction has been achieved
- no mouth is present at this stage of development
- grows within the host by taking up dissolved nutrients from the
Body
surrounding body fluids
- outer body layer of adult trematodes, like that of the cestodes and
- each sporocyst are numerous balls of cells
monogeneans, is an unciliated syncytial tegument
- each of these germ balls develops either into another sporocyst (the
- trematode body closely resembles that of a turbellarian
daughter of sporocysts) or into another larval stage, the redia
- has a mouth opening and a blind-ended digestive tract that is bilobed
- the rediae or daughter sporocyst migrate to their host’s digestive
(with a few exceptions)
gland or gonad
- body is never segmented
- ingests the host’s tissues and blood through its mouth
Rediae
- active feeders
Schistosomiasis
- possessing a mouth and a functional, blind-ended gut
- deadly disease prominent in many regions of the world
- some rediae larvae serve as a soldier caste instead of replicating
- results from an infection by trematodes known as “blood flukes”
themselves, attacking and killing the larvae of other trematode species
- more than 200 million persons in 77 countries are estimated to suffer
withing the same host, and even consuming rediae from other clones of
from it, making it the second most prevalent disease in the world (next
the same species
to malaria, which is caused by a protozoan)

THE DIGENEANS

di=two
gena=birth
- exclusively endoparasitic
- has a mouth and a blind-ended gut, usually two-branched, and often
has both an anterior and a ventral sucker for maintaining position within
the host
- most are hermaphroditic; although some gonochoristic--have sepatae
anatomically distinct sexes
- digenetic flukes always require at least one intermediate host before
reaching the final host
- host location among digenetic trematodes is generally an active
process, mediated by highly specialized, free-living larval stages
- typically show little specificity for the second intermediate host, which
includes both vertebrate and invertebrate species
Ex. One species in the genus Echinostoma can use either certain
freshwater snails or the larval stage of certain amphibians as second
intermediate hosts
- yet the miracidia of that species can successfully infect only snaols of
one species or of several closely related species
Daughter sporocyst
- do not feed
- although they probably absorb soluble nutrients from their
surrounding

Cercariae
- a germ ball; another anatomical distinct larval stage that are within
most rediae larvae or daughter sporocyst
- leave the “mother” by means of a birth canal if escaping from redia or
by lysing through the body wall if escaping from sporocyst
- free-living
- noncliated but they can swim actively by means of a muscular tail
- usually possess at least one sucker anteriorly; many species also
have a ventral sucker
- transform into adults once they encounter the next host, attach it, and
enzymatically penetrate the host tissues
- in other cases, cercaria encysts on submerged vegetation likely to be
eaten by the definitive host
- penetration by the cercaria larva into the next host is accompanied by
detachment of the cercarial tail, either before or after the cercaria
enters the host’s body
- once in the second intermediate host, the cercaria transforms, in
many species, into an encysted waiting stage, the metacercaria

Metacercariae
- most of the specifically larval organs degenerate
- may survive for many months in the intermediate host while remaining
fully infective
- once eaten by the definitive host, the juvenile trematode migrates
through the digestive tract to take up residence in the proper,
species-specific location, where adulthood is reached
Summary of Digenean Life Cycles
1. Typically includes five disctine and unique developmental stages
(miracidium, sporocyst, redia, cercaria, and metacercaria) and typically
two or more intermediate hosts
2. The first intermediate host almost always must be a mollusc: of the
approximately 10,000 known trematode species, all but two require a
gastropod or bivalved mollusc as the first intermediate
3. Species specificity fo the first intermediate is extremely high
4. Specificity for subsequent intermediate hosts and for the definitive
host is far less
5. The digenean life cycle includes asexual replication of
developmental stages, something not found in any turbellarian species,
and that asexual replication occurs entirely within the molluscan
intermediate host
6. Finally, most species can reach adulthood only in a vertebrate host,
usually after a period of encystment in the environment or in a previous
host.

THE ASPIDOGASTREANS = (Aspidobothreans)

Defining Characteristic
- large ventral sucker divided by septa, generally forming a row of
suckers

General Characteristics
- a mere 80 species
- bear similarities to monogeneans and digeneans, but they fit neatly
into neither group
- most species have a simple life cycle involving a single host (like
monogeneans)
- the host is always a mollusc (freshwater mussels or gastropods) but
otherwise, they generally show very little host specificity
- never have the posterior attachment organ (haptor)
- the entire ventral surface of the body is modified to form a powerful
attachment sucker
- some species require an intermediate host to complete the life cycle;
these species reach adulthood in fished or turtles
- never exhibit asexual replication within the intermediate molluscan
host

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