PHYLUM ACANTHOCEPHALA
Beware: scanned document.
-
Historical account
-
A. Acanthocephala not noticed until beginning of the 18th century,
not distinguished until Koelreuther (1771) proposes the name Acanthocephala
.
-
B. Muller (1776) unaware,of Koelreuther's work calls them Echinorhynchus.
-
C. Rudolphi (1809) formally gives them the name Acanthocephala.
-
D. Many species were described in the 19th century and all were placed
under one generic name- Echinorhynchus.
-
E. Position of Acanthocephala was very uncertain; placed among flatworms,
roundworms, chaetognatha, and finally lumped into the Aschelminthes.
-
F. Hyman removes Acanthocephala from the Aschelminthes making them
into a separate phylum.
-
Definition of the phylum- Endoparasitic, pseudocoelomate, and wormlike
Bilateria, lacking a digestive tract, with the anterior end formed into
an invaginated proboscis armed with hooks.
-
External features:
-
Body is elongated, becoming tapered at both ends, with size ranging from
1.5mm-.5m in length.
-
Body wall is covered with a thin epicuticle of muccopolysacchorides and
a noncelbular cuticle (1 micron or less).
-
Body is divided into two major regions- the presoma and the trunk.
-
Presoma is composed of the proboscis- a retractile organ armed with hooks
and spines, and the neck- an unarmed region posterior to the proboscis.
-
Trunk is the major bulk of the animal, surface may be smooth, wrinkled,
or irregularly ringed; some genera arm the trunk with spines.
-
Proboscis serves as an attachment organ within the hosts intestine or for
locomotion.
-
Contains alternating rows of hooks and spines in a definite pattern.
-
Hooks are larger structures with roots sunken deep in proboscis walls;
spines are smaller structures lacking roots.
-
Proboscis is invaginable and can be withdrawn into a muscular sac - the
proboscis receptacle.
-
Proboscis musculature-
-
retractors: attach to base of proboscis sheath and run posteriorly dividing
into ventral and dorsal retractors; function to withdraw proboscis into
receptacle.
-
protractor: originate in a circle at the base of neck and attach to posterior
end of sheath; function to extend proboscis out of receptacle.
-
Sense Organs are generally deficient and are of a tactile nature.
-
Proboscis contains a terminal, small pit with a nerve ending.
-
Male bursa and penis have genital ganglion which terminate into bulbous
or spherical sense organs.
-
Usually lack coloration but may appear red, orange, yellow or brown due
to absorption of food.
-
IV. Internal Features: cuticle, epidermis, dermis
-
A. Body wall made up of epicuticle, and muscle layer.
-
B. Syncytial epidermis is composed of three layers.
-
1. Thin outer layer of parallel radial fibers and pores extending into
the middle layer.
-
2. Middle layer of randomly arranged fibers.
-
3. -Very thick inner layer of radial fibers and a series of channels known
as lacunae;s without definite walls but having
-
C. Lacunae- fluid filled channe3 a definite pattern; thought to be
a food distributing system. Fluid moves only with body movements and system
does not connect onto any body structures. circular
-
D. Musculature is a syncytial fibrous network of thin outer muscles
and inner longitudinal muscles.
-
E. Pseudocoel is found in trunk region and slightly into presoma.
-
F. Inner layer epidermis extending into the trunk pseudocoel forms
a pair of projections called leignisci that act as rese~iriors for the
fluid of the lacunar system.
-
G. Nervous system-
-
1. "Brain" is a cerebral ganglionic mass located along the ventral border
of the proboscis sheath.
-
2. System has two main lateral nerve bundles wrapped in a muscular sheath
called a retinacula.
-
3. Male has two genital ganglia connected by a nerve ring.
-
H. Digestive system- mouth, anus, and digestive tube are completely
lacking, nutrients taken in via pores and diffusion.
-
I. Circulatory system is absent.
-
J. Excretory system consists of a mass of flame bulb protonephridia
which empty into a common sac (bladder). The bladder leads to the sperm
duct in the male and the terminal portion of the uterus in the female.
-
K. Reproductive system is unique in Acanthocephala because the organs
are suspended within a ligament sac.
-
1. Ligament sacs are hollow tubes of connective tissue w/wo muscle fibers
that run the length of the body.
-
a. Females have two sacs, a dorsal and a ventral.
-
b. Males have one sac, a dorsal enclosing the testes.
-
2. All are dioecious with females generally longer that males.
-
3. Male system:
-
a. Two testes, two sperm ducts that unite in the genital sheath to form
the vas deferens which ends in a muscular cup, the bursa.
-
b. Penis projects into the bursa which is eversible to grasp female.
-
c. Cement glands are present and-drain into the sperm duct.
-
4. Female system:
-
a. Fragmented ovaries are unattached in liagament sac forming free floating
ovarian balls.
-
b. Ligament sac ruptures when ova are ripe sending them into the pseudocoel.
-
c. Uterine bell uses peristaltic contractions to collect developing eggs
and pass them to the uterus.
-
d. Vagina is also present.
-
V. Reproduction:
-
A. Sexual:
-
1. Internal fertilization; male bursa is wrapped around posterior of the
females, cirrus is introduced into the gonopore, sperm migrate up the vagina,
uterus, uterine tube. Male cement glands secrete a cement plug sealing
up the gonopore preventing sperm escape.
-
2. Eggs develop in the pseudocoel until a larval stage provided with rostellum
and hooks is reached; hard shell forms.
-
3. Uterine bell contractions pass larvae to exterior.
-
4. Expelled eggs are highly resisitant and remain viable for months.
-
VI. Embryology:
-
A. Elliptical egg enclosed in four membranes.
-
B. Spiral determinate cleavage.
-
C. Four blastomeres are displaced due to elliptical shape.
-
D. Cleavage is not synchronous, stereoblastula is formed at 34 cell
stage, syncytium forms.
-
E. Gastrulation by in wandering of nucleii, not definite.
-
F. Acanthor larvae forms, development stops until ingested by proper
invertebrate host.
-
VII. Life cycle:
-
A. Vertebrate host- eggs containing partially developed acanthors are passed
out in feces and are ingested by intermediate host.
-
B. Intermediate host- usually an arthropod
-
1. Acanthor ruptures from egg, penetrates gut wall and lodges in the hemocoel.
-
2. Acanthor rounds up develops proboscis and loses hooks, rostellum larvae
now called acanthella.
-
3. Once all the features of the adult are developed the animal is termed
a cystacanth; gonads are nonfunctional.
-
C. Second vertebrate ingests the intermediate host, cystacanth develops
to sexual maturity in the intestine.
-
VIII. Ecology and Physiology:
-
A. Primarily anaerobic though they can survive in the presence of 02.
-
B. Eggs can withstand temp. of -10 to 45 degrees C. and are viable up to
three years.
-
C. Number of worms per host is large: 1000 in a duck intestine.
-
D. Reproductive capacity is high, up to 10,000,000 eggs per female.
-
E. Very injurious to host as proboscis hooks cause damage to intestinal
tissues.
-
F. Glycogen stored in body wall and lacunar fluid,fatty substances deposited
in epidermis and female reproductive system (ovaries).
-
IX. Classification:
-
1. Order Archiacanthocephala- intestinal parasites of terrestial hosts
primarily birds and mammals. Invertebrate hosts are roaches and grubs.
Proboscial spines concentrically arranged, possess protonephridia
-
2. Order Palaeacanthocephala- parasites of fish, aquatic birds and mammals.
Invertebrate host are crustaceans. Proboscis spines in alternating radial
rows, no protonephridia.
-
3. Order Eoacanthocephala- parasites of fishes and reptiles. Invertebrate
host are crustaceans. Proboscis hooks are radially arranged, no protonephridia,
cement glands in male are syncytial.
-
X. Phylogeny:
-
A. Acanthocephala closegy associated to Platyhelminthes:
-
1. Armed proboscis found in Cestodes.
-
2. Body wall musculature is distinctly flatworm-like.
-
3. Presence of flamebulb protonephridia.
-
4. Embryology is strongly Cestode- like.
-
B. Acanthocephala closely associated to Aschelminthes.
-
1. Body structure, presoma + trunk, seen in many groups.
-
2. Superficial segmentation found in several Aschelminthe groups rotifers,priapulids.
-
3. Pseudocoel is present although formation is not typical.
-
4. Flamebulb protonephridia.
References
Barnes, R. 1980. Invertebrate Zoolpgy. Philadelphia:Saunders Company.
pp. 309-311.
Cheng, T. 1973· General Parasitology. New York: Academic Press.
PP· 54~5-563-
Cleave, H. 1952. Speciation and Formation of Genera in Acanthocephala.
Systematic Zoology. Vol. 1, No. 2.
Hyman, L. 1951. The Invertebrates: Acanthocephala, Aschelminthes, and
Entoprocta. Vol. 3. New York: McGraw Hill. pp.459-531.