Principles of Parasitology

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Notes prepared by:
J. R. Barta, PhD
Dept. of Pathobiology, OVC
University of Guelph
Guelph, Ontario, Canada
jbarta@uoguelph.ca
519-824-4120 ext. 54017

All contents Department of Pathobiology, 2011

Coourse Ovverview and

a Resou
urces
Introducttion to the Principless of Parasittology Nottes
The Notes: Theese notes aree designed to
t supplemennt the inform
mation provided in lectuures and
laboratory sessio
ons. A few of the notes will contaain informatiion coveringg all aspects about a
parasite includin
ng the clinicaal problem associated
a
w infectionn and treatm
with
ment; other nootes will
contain more lim
mited inform
mation.
These no
otes should not
n be regarrded as conttaining inforrmation that is all-inclussive and
encyyclopedic in scope. Hoowever, the notes, in adddition to material
m
coveered in lectuures and
laboratory sessio
ons, provide all the requuired informaation for thee purpose of examinationns in the
courrse. In study
ying for thosse exams, sttudents shouuld be guidedd by the goaals and objecctives of
the course
c
as id
dentified in the
t Course Outline andd by the empphasis on thhe various topics as
given in lecturess and labs.
The notees will be ideentified withh respect to the
t parasitess and host grroups under study as
folloows:

Arrthropods

Caarnivores

Protists

Ruminants

Trrematodes

Horses

wine
Sw

Cestoddes

N
Nematodes

Birds

Humans

A NOTE
N
ABOU
UT LABOR
RATORY SA
AFETY
Safeety in the laaboratory is a priority at
a all times. In order too ensure
safeety of all paarticipants inn laboratory activities, you
y must folllow the
safeety proceduures/guidelinees providedd by the instructor, teaching
assistant or technicians at all times. In the paraasitology labboratory,
studeents will be expected to safely handlle unfixed feeces or other material thaat may be pootentially
infecctious to hum
mans or anim
mals. You must
m wear suuitable laborratory outerw
wear (lab coaat or lab
coveeralls) at all times. Dispoosable glovees and other safety-relateed items thatt will be proovided to
you are to be used when inddicated. Failuure to observve safety reqquirements may
m necessittate your
remooval from the
t
laboratoory. For obbvious persoonal safety and gastroonomic reaasons, a
laboratory in which
w
you will
w handle unfixed fecees, animal bodies and//or body paarts, and
poten
ntial pathog
gens is no pllace for open
n food or drrink! You WILL
W
be askeed to put aw
way food
and or drink in the lab and refusal to follow
fo
this reeasonable ru
ule will resu
ult in your exxpulsion
from
m the laborattory session.
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Principless of Parasitoloogy

LEARNING RESOURCES:
Course Coordinator/Instructor J. R. Barta - No formal office hours are posted but I am
readily available for out-of-class assistance. For specific questions, email is convenient
(jbarta@uoguelph.ca) and is checked regularly. Otherwise, drop by 4826 Pathobiology (PAHL)
or call (519-824-4120 ext. 54017) to arrange an appointment. Do leave a message with a return
telephone number (and best time to reach you) if you are unable to contact me directly and I will
return the call as soon as possible.
Course Notes These are largely a complete set of parasitology notes for the course; there is no
prescribed textbook for the course. Additional material, particularly regarding appropriate
treatments for parasitic infections (this information changes regularly and therefore the most
current information may be available only after the notes have been prepared), may be given
solely in lectures or laboratories. A glossary of parasitological and other terms will be made
available through the courses website.
Review Images and Lectures A compilation of images that are used in the notes and lectures
as well as some additional images have been digitised and are available through the course site.
All PowerPoint lectures will be made available as an additional resource associated with each
taxonomic grouping within the course website.
Glass Slides - A set of glass slides of parasite forms and lesions used in the laboratories will be
available from Nathalie Lemieux, Department of Pathobiology, Room 1814/1813.

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Principles of Parasitology

Introduction to Parasites and Parasitology

Introduction
This lecture is intended to highlight the major groups of parasitic organisms as well as the
strategies used by all parasites to survive in the host and techniques used by parasites to move
between hosts.

Relationships between Host and Parasite

Symbiosis Levels of interdependency between species
Increasing
potential
for harm
to host

Phoresis (traveling together no physiological dependence)

Mutualism (benefit to both organisms)
Commensalism ecto and endo (host not harmed other species gains)
Parasitism (parasite lives at the expense of its host)

Life Cycle Survival Strategies

Types of Infections
Cyclo-developmental - a single parasite develops from a single infective organism by
maturation, e.g. ascarid nematodes, many cestodes
Cyclo-propagative - a number, generally a large number, of progeny are produced from a
single infective organism, e.g. malarial parasites, trematodes
Duration of Infections
Acute or self-limiting - these infections are similar in many ways to bacterial or viral
infections. Infections are acquired and are rapidly resolved in most cases, generally with a
species or strain-specific immunity developed that protects against subsequent infections
with the same parasite. Examples include coccidia and cryptosporidia.
Prolonged or chronic these infections are more typical of parasitic infections. Many
helminths and some tissue cyst-forming protists are known to cause prolonged infections
(i.e. tens of years or 'for life'). Encysted or arrested stages of a number of parasitic protists
can remain in hosts for many years. The ability to form a long-lasting resting phase is
especially common in intermediate hosts that will be eaten by the final or definitive host.
Fecundity
Most parasites are prodigious egg or cyst producers as an adaptation to the relatively low
chance of survival for their progeny. As an example, a single ascarid worm may produce
20,000 or more eggs per day and may live for many years.
Over-dispersion in the Host Population
A distribution of parasites amongst hosts is said to be overdispersed, or aggregated, if
parasites are found to co-occur in particular hosts more often than if the parasites were
distributed at random amongst all hosts. Macroparasites (large parasites such as worms as
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Principles of Parasitology

opposed to microparasites such as protists) are typically overdispersed in their host

populations, so that the majority of hosts harbour few or no parasites while a few hosts
harbour large parasite burdens. This distribution serves to limit negative effects of carrying
parasites to a small number of hosts. The health of the population (although not the
heavily infected individuals) is generally not affected by the infections being carried.
Modifications to Host Behaviour to Promote Transmission
A number of parasites will modify the behaviour of their hosts to enhance their survival
and/or chance of transmission to the next host. This is particularly evident in intermediate
hosts (both invertebrate and vertebrate) that are eaten by a definitive host of the parasite.
Some examples of this include coccidia (some Sarcocystis spp. that reduce the speed and
efficiency with which their rodent intermediate hosts move away from the centre of an
open area and hide along walls or in crevices) and acanthocephalan worms (intermediate
stages of a one species changes the colour of its Daphnia sp. intermediate host and
reverses its negative phototaxis -- these changes greatly enhance the chances that the
intermediate host will be ingested by the definitive host, a duck.

Major Groups of Parasites

Starting from smallest to largest, the major groups of parasites that are frequently encountered
are grouped into 5 major groups. These are:
Protists (=Protozoa)
- amoebae, microsporidia, myxozoa, flagellates, apicomplexan protists
Trematodes
- monogenetic and digenetic flukes
Cestodes
- tapeworms of various forms
Nematodes
- roundworms of various types
Arthropods
- ticks and mites, insects, crustaceans, pentastomes (tongueworms)

Over the next few pages, these major groups will be introduced. The purpose of this section is
NOT that you will learn these groups in any detail at this time. Instead, this information is
provided now so that you have an overview to whet your appetite for these interesting
creatures before more detailed notes.

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INTRODUCTION TO THE PARASITOLOGY NOTES

Most of the notes describing parasites will be provided in the following format:
A.

Taxonomic Classification

Phylum
Class
Order
Super Family
Family
Genus species (common name)
The taxonomic grouping as identified above will be given wholly or in part for each parasite.
You are expected to know only the genus name of the parasite and, if given, also the common
name. For some parasites, the species name is important to know and these will be used in
lectures and labs. In these cases, you will be responsible for the species names as well.
B.

Host Spectrum

A parasite may be capable of developing completely in only one type of host, e.g., dogs, and is
considered highly host-specific. Other parasites may develop in any one of a number of
species of hosts e.g., cattle, sheep, horses, and thus have a broad host spectrum.
A parasite may have a single host (monoxenous) or require more than one host for complete
development (heteroxenous). These hosts are identified as a definitive (final) host or an
intermediate host. A parasite may enter or be on other hosts in which there is no development
and such hosts are either a paratenic or mechanical host.
Definitive or Final Host (DH) - the host(s) in which sexual reproduction occurs
Intermediate Host (IH) - the host(s) in which the immature stages or asexual replication
occurs
Paratenic Host (PH) - a host in which the parasite does not develop but which can serve in the
dissemination or transport of the parasite. Such hosts are usually part of the food chain of other
PHs, IHs or DHs
Some parasites use arthropodan hosts as Vectors. A Vector is an invertebrate such as a flea,
non-biting fly, mosquito, etc, that transmits (vectors) a parasite from one host to the next. There
are two types of vectors that are differentiated based on their relationship with the parasite.
Biological Vector these are invertebrates that form an essential role in the life cycle of the
parasite and in which the parasite grows or develops by invasion of the vectors body. The
parasite is biologically dependent on the invertebrate vector for nourishment and survival.
Widely known examples include mosquitoes transmitting malaria or tsetse flies transmitting
African sleeping sickness (trypanosomiasis).
Mechanical Vector - a host, usually an arthropod, that transmits, usually externally on its
mouth parts, the parasite from one host to another and in which there is no growth or
development.
C.

Site of Infection

In the Definitive Host - the organ where the parasite undergoes sexual reproduction or where
the sexually mature parasite is found. That site is often referred to as the predilection site.
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The parasite may be found in other organs of the DH as it migrates to attain the predilection
site.
Endoparasite - A parasite in internal organs of the host including within the alimentary tract.
Ectoparasite - A parasite in or on the skin of the host
D.

Morphology

For multicellular parasites, gross features will be stressed e.g. size, shape, color, other unique
morphological characteristics e.g., holdfast organs. Wherever possible, emphasis will be on
features that can be seen by eye. In many instances, identification of a parasite is possible only
on microscopic examination and this will be required for protistan (unicellular) parasites.
The microscopic characteristics of parasitic stages found in body fluids, e.g., microfilariae in
blood, or in excretory products (feces, urine), e.g., eggs, larvae, trophozoites, are important for
identification of parasites.
E.

Life Cycle

The life cycle of a parasite refers to the development of the parasite through its various forms
or stages. This process can be simple or complex and the following points should be noted:
1. Host(s) involved and the stage(s) found in them
2. Stage(s) in the external environment and the development there
3. Infective stage of the parasite
4. Source of the infection or the place or material where the DH contacts the infective stage
of the parasite and becomes infected
5. Route(s) of entry into, and the migratory route(s), in the hosts
6. Factors influencing transmission of the parasite from host to host
Prepatent period (PPP), i.e., the developmental time in the host, from invasion to when
parasite forms e.g., eggs, cysts, larvae, can be found in body fluids or excretions
The patent period is the duration of time that eggs or larvae are detected in body fluids or
excretions

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Principles of Parasitology

F.

Pathogenesis

G.

Clinical Signs

H.

Lesions

The pathologic mechanisms, lesions and clinical signs associated with disease may be a result
of both the parasite and the host reaction to it.
1. The parasite - its physical presence, activity or production of toxic products may be
injurious to the host. The extent of the injury can be influenced by size and numbers of the
parasites, whether the parasites invade, develop and migrate in cells, tissues, organs, ducts,
vessels or other body channels, and how the parasite feeds or derives its nutrition
2. The host response - this will depend on whether the host had previous exposure to the
parasite, type of host response e.g., inflammatory, immune, the host's compensatory
physiological mechanisms, nutritional status and age of the host
3. A parasite may be non-pathogenic but may transmit an agent which incites disease
I.

Diagnosis

J.

Treatment

Therapies for parasitic infections are often being updated. As a result, current therapies for
parasites will be discussed in lectures and laboratory sessions in most cases.
K.

Control

L.

Public and Human Health Significance

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Protistss (= Protoozoa)
A.

Taxxonomic Cla
lassification

Kingdom
m Protozoa
Phyllum Sarcomaastigophora
C
Class
Zoomastigophora (flagellates)
C
Class
Rhizopoda (amoebaae)
Phyllum Apicom
mplexa
C
Class
Sporozo
oasida (cocccidia, malariaa)
C
Class
Piroplaasma (Babesiia, etc.)
Phyllum Microsp
pora
C
Class
Microsp
poridea (Nossema, Glugeea)
Phyllum Ciliopho
ora
C
Class
Ciliateaa (ciliates)
B.

Hoost Spectrum
m

Any anim
mal.
Some meembers, partiicularly cocccidia of the genus
g
Eimerria, are highlly host speciific.
Others, such as cocciidia of the geenus Cryptosporidium, are
a more inddiscriminate in choice.
C.

Sitte of Infectio
on

Most sitees, e.g. intesttine, tissues such as musscle, brain annd liver, andd blood.
Some aree intracellulaar while otheers live free within
w
the inntestine or blloodstream.
D.

Moorphology

Unicellullar
Some havve flagella or
o cilia, used for movemeent
Some havve sexual rep
production, others have no sexual sttages
E.

Liffe Cycle

Some havve a cyst staage (infectivee stage) withh the cyst waall renderingg them protecction againstt the
external environment
e
t.
Simple or
o complex liife cycle thatt involves a variety of sttages and host animals.
Some inffect hosts dirrectly, otherss rely on a vector
v
(most often an arthhropod) for transmission
t
n.

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Protistss (= Protoozoa)
PHYLUM
M OR
SUBPHYL
LUM

ASS
CLA

O
ORDER

Sarcomastigophora

Zoom
mastigophora

Diiplomonadida

SUBORDER
R

FAMILY
Y

Trrichomonadida
Kiinetoplastida

Apicomplexaa

opoda
Rhizo

Reetortamonadida
Am
moebida

Sporo
ozoasida

Euucoccidiorida

Eimeriorina

Eimeriidae

Cryptosporridiidae
Sarcocystiddae
matinae
Toxoplasm

Lankesterellidae
Haemosporina

Piropllasma

Pirroplasmorida

Microspora

osporidea
Micro

Miicrosporida

Ciliophora

Ciliateea

Trrichostomatida

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GEN
NUS
Giardiia
Hexam
mita
Trichoomonas
Histom
monas
Trypannosoma
Leishm
mania
Chilom
mastix
Entam
moeba
Iodam
moeba
Naegleeria
Eimeriia
Isospoora
Caryosspora
Tyzzerria
Cryptoosporidium
Sarcoccystis
Frenkeelia
Toxoplasma
Neospora
Besnoiitia
Hamm
mondia
Cystoiisospora
Lankessterella
Atoxopplasma
Plasmodium
Haemooproteus
Leucoccytozoon
Hepatoocystis
Babesiia
Theileria
Cytauxxzoon
Nosem
ma
Glugeaa
Balanttidium

P
Principles
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P

Illustrrations of Vaarious Protiistan Parasittes

Gam
monts (male, low
wer; female, uppper) of a
Haem
moproteus sp., a malaria-like blood parasitee of
birdss.

A cyst of Balantidium
B
co
oli (a ciliate) inn the feces of
swine. This is a low levell zoonotic threat.

Braddyzoites (merozzoites) of Toxop

oplasma gondiii freed
from
m a cyst in the brain
b
of a mousse. This organism is
the causative agent of toxoplasmoosis.

Cryptosporridium parvum
m, a cosmopolittan protistan
parasite off mammals, inccluding humanss.

Oocyysts of bovine coccidia,

c
Eimeeria species.
The monkkey face of a trophozoite
t
of a Giardia sp.

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Treematodaa
A. Taxxonomic Cla
lassification
Phylum Platyhelmint
P
thes (flatworrms) Class Trematoda
T
(fflukes)
Order Monogenea
M
Order Aspidogastre
A
a
Order Digenea
D
A
Alaria
spp.
C
Cryptocotyle
sp.
D
Dicrocoelium
m sp.
F
Fasciola
hepa
atica
F
Fascioloides
magna
H
Heterophyes
sp.
N
Nanophyetus
salmincola
O
Opisthorchis
sp.
P
Paragonimus
s sp.
P
Paramphistom
mum spp.
P
Platynosomum
m sp.
S
Schistosoma
spp.
B.

Hoost Spectrum
m

Definitivve Host (DH

H)
Monogenneans and Aspidogastreaans - fish, am
mphibians, reeptiles; chieffly ectoparassites, e.g. gillls of
fishes
Digeneanns mammaals, includingg humans
Intermed
diate Host(ss) (IH)
Monogenneans and Aspidogastreaans - none
Digeneanns - one or more
m
dependiing on the sppecies of fluuke; a snail iss always the first IH andd this
is usuallyy an aquatic or semi-aquuatic snail; eaach species of
o fluke requuires a speciffic species of
o
snail for developmen
nt; snails are fussy aboutt their habitaat and are lim
mited geograpphically - heence
a also limitted geographhically; som
me flukes requuire more thhan
trematodde infections in animals are
one IH annd the one or
o more IH foorm part of the
t food chain to the DH
H

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C.

Site of Infection

Digenea
The migration of the fluke may involve several organ systems.
Adult flukes are found in many organ systems with each species of fluke having a specific site
for maturation
D.

Morphology

Digenea
Adult - unsegmented, spiny cuticle in some, two holdfast organs or suckers, one anterior and
one ventral near the middle of the body or at the posterior end. Internal anatomy: mouth,
anterior sucker, esophagus, two blind ceca; single ovary, coiled uterus, vitellaria (yolk glands),
paired testes, genital pore which is the convergence of the male and female ducts
Immatures - miracidium, sporocyst, redia, cercaria, metacercaria in that order and found either
free or in the IH
Egg - oval, operculate, shell medium thick and smooth on the outside, yellow to golden brown
in many species, embryo varies from a segmenting embryo to a fully developed miracidium; in
most cases the contents usually completely fill the shell
E.

Life Cycle

Digenea
1. Indirect life cycle
2. Eggs in feces, generally hatching only when in water
3. Egg with a miracidium swallowed by a snail or miracidium hatches from egg and penetrates
the snail
4. Asexual generations in the snail (sporocyst, redia).
5. Cercariae emerge from the snail and encyst as metacercaria on vegetation or after
penetrating another IH.
6. Excystment of ingested metacercariae occurs in the intestine of the DH and the juvenile
fluke migrates to the anatomic site where maturation occurs

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Illustraations of Vaarious Tremaatode Parasiites

Lungg flukes (Paraggonimus kellicootti) in a dog luung.

Egg of the canine fluke, Alaria

A
americaanum the
larval form
m of this fluke is
i potentially zoonotic and is
found in am
mphibians in so
outhern Ontariio.

Paraagonimus kelliccotti freed from

m the lung.

Adults of Alaria
A
america
anum attached to
t the small
intestine off a fox. Only th
he lighter colouured posterior
region is visible.

o lung fluke (P
Paragonimus kellicotti).
k
Egg of

Adult Clonnorchis (Opisth

horchis) sinenssis, the
Chinese livver fluke of humans.
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Fasccioloides magnaa (liver flukes)) encysted in a liver.

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C
Cestoda
A. Taxxonomic Cla
lassification
Phylum Platyhelmint
P
thes (flatworrms)
C
Class
Cestodaa (tapeworm
ms)
Anopllocephala sppp.
Diphyyllobothrium
m sp.
Dipyliidium caninuum
Echinococcus sppp.
Hymeenolepis spp.
Mesoccestoides sppp.
Monieezia spp.
Spirom
metra spp.
Taenia spp.
Thysa
anosoma sp.
B. Hoost Spectrum
m
Tapeworrms with few
w exceptions have both definitive
d
andd intermediaate hosts
Definitivve Host (DH
H) - most aduult tapeworm
ms are highlyy host specifi
fic and each species has its
i
own DH
Intermed
diate Host (IH)
(
- for moost tapeworm
ms there is only
o
one IH. The IH form
ms part of thhe
foodd chain of thee DH or can be accidentally ingestedd. Hymenolep
epis nana, a tapeworm
t
off
man and rodentss, is unique and
a does nott require an IH.
I
C. Sitte of Infectio
on
Adult - principally
p
th
he small inteestine, but allso bile and pancreatic
p
duucts and largge intestine
Immaturres - abdominal cavity, liver,
l
musclee, but also crranial cavityy and eye, wiith each speccies
of taapeworm hav
ving a preferrred site
D. Moorphology
Adult - some
s
a few mm
m long, buut generally in
i 10s - 100ss cm and divvided into a scolex,
s
neckk and
strobbila
-the scolex is thee anterior poortion of the tapeworm, is
i a minute holdfast
h
orgaan and contaiins
four suckers and
d sometimes an anterior projection,
p
thhe rostellum
m, which mayy be armed i.e.
i
withh hooks, and may be retraactable into the body of the scolex. Scolices of some tapewoorms
may have longitu
udinal groovves (bothria))
- thee neck is a sm
mall narrow germinal areea adjoining the scolex and
a from whhich the strobbila
is buud off
- thee strobila is a chain of segments. Eacch segment matures
m
as itt moves proggressively
furthher away from the scolexx; anterior seegments are immature; middle
m
segm
ments have
increeasingly welll-developedd male and feemale reprodductive organns and becom
me
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reproductively active or mature; terminal segments are egg laden or gravid and with
degenerating reproductive elements - there is no mouth or alimentary system, nutrients are
absorbed directly through the integument
- each segment has one or two sets of reproductive organs (single ovary, coiled uterus,
vitellaria, yolk glands, testes, one or two genital pores which are the convergence of the
male and female ducts)
Egg - in most tapeworms of domestic animals there is no true shell. Where there is a true shell,
it is operculate. The embryo is fully developed, infective when shed and contains six hooks.
It is referred to as a hexacanth (six hooks) larva, embryo or onchosphere. It is surrounded
by embryonic membranes called the embryophore, which is usually spherical but can be
variously shaped
Immatures - a hexacanth (six hooks) embryo is the first stage larva. In the IH the first stage
larva matures to the second stage larva of which there are several types. In some
tapeworms, there is a third stage larva
- The type of second stage larva formed is associated with the type of IH used in the life
cycle.
1. IH is a mammal
Tapeworm genus - Taenia
A. Cysticercus - a white fluid-filled bladder up to the size of a pea and containing an
invaginated scolex. The cyst is surrounded by a connective tissue capsule formed by the
host.
B. Strobilocercus - a partially developed cysticercus with the scolex evaginated and a small, 1
cm, strobila connected to a fluid-filled bladder
C. Coenurus - a fluid-filled cyst, up to 5 cm, with many invaginated scolices and surrounded
by a connective tissue capsule formed by the host. Each scolex has the potential of forming
a mature tapeworm
Tapeworm genus - Echinococcus
A. Hydatid Cyst - a fluid-filled cyst, usually 5-10 cm, but can be up to 50 cm in diameter and
in which there is endogenous budding of multiple brood capsules, each containing many
scolices. Brood capsules may rupture and release the scolices to form many small round
bodies called hydatid sand in the hydatid fluid. The hydatid membrane is surrounded by an
inflammatory connective tissue capsule. Exogenous budding is rare; when it occurs it is
never infiltrative and is confined by the connective tissue capsule.
B. Alveolar Hydatid - similar to a hydatid cyst, but not confined by a capsule and with
exogenous budding which infiltrates surrounding tissue. The hydatid is a diffuse growth
composed of many compartments each filled with a gelatinous matrix and many brood
capsules. As the hydatid grows like a malignant neoplasm, the center dies and is filled with
necrotic tissue.
2. IH a terrestrial arthropod
Tapeworm genera Dipylidium, Moniezia, Anoplocephala
Cysticercoid - a small, 1 mm, white, solid cyst containing a scolex and with a tail-like
appendage
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3. IH a crustacean
Tapeworm genus - Diphyllobothrium
Procercoid, Plerocercoid - solid, ribbon-like. In tapeworms with crustaceans as the first IH, the
hexacanth larva is ciliated and referred to as a coracidium.
E. Life Cycle
1. Usually indirect life cycle
2. Gravid segments or free eggs are passed in the feces. Segments may be motile
3. Segments dehydrate and rupture to liberate the eggs
4. In most tapeworms, the eggs are ingested by the IH; in some, a ciliated hexacanth larva
hatches from the egg and swims to penetrate the IH
5. The larva in the IH migrates to a specific site and matures to the second stage. This is the
infective stage for most tapeworms of domestic animals and man. Where there is a
requirement for a third stage larva this occurs in the second IH.
6. The DH ingests that part of the IH containing the infective stage and the juvenile tapeworm
matures and attaches to the intestine wall.

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16

Principles of Parasitology

Illustrations of Various Cestode Parasites

Egg of a taeniid tapeworm in the feces of a dog. The

paired hooks of the hexacanth embryo are visible
within the egg.
Adult tapeworms strobilas (chain of segments) of
Taenia taeniaeformis in the small intestine of a cat.

A coenurus of Taenia multiceps. This sac-like

structure contains many developing scoleces
(progenitors of the long strobilas that will be found in
the vertebrate gut).

The scolex (holdfast) of a Taenia species showing the

prominent hooks at the anterior end (the rostellum)
and the four suckers around the base of the rostellum.

A complete strobila of an Echinococcus species from

scolex on the right side to gravid (egg-bearing)
segment on the left. These small tapeworms of dogs
shed eggs that can infect humans and cause hydatid
disease (hydatidosis).
Cysticercus (a larval cestode) of Taenia pisiformis
found within the liver of a rabbit.

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17

Principles of Parasitology

Neematoda
A.

Taxxonomic Cla
lassification

Phylum Nematoda
N
(rroundwormss)
O
Order
Rhabdiitida (rhabdiitids)
O
Order
Strongy
ylida (stronggylids)
O
Order
Oxyuriida (oxyurids, pinwormss)
O
Order
Ascarid
dida (ascaridds)
O
Order
Spiruriida (spiruridss, filarids)
O
Order
Enoplid
da (trichuridds, Dioctophyyme)
B.

Hoost Spectrum
m

Most nem
matodes have a strong hoost preferencce; some aree highly hostt specific
Some nem
matodes req
quire intermeediate hosts
C.

Sitte of Infectio
on

This variies with the species

s
of neematode. Most
M of the sppecies are fouund in one or
o more sectiions
of thhe gastrointestinal tract
D.

Moorphology (F
Figure)

Adult
- <1 mm
m to many cm
c long - thee largest is about
a
1 m, cyylindrical in cross sectioon, unsegmennted,
sexees separate an
nd each withh alimentary, excretory and
a nervous systems
- in crooss section, the
t nematodde appears ass an outer tubbe separatedd from inner tubes by a
secoondary body cavity or pseudocoelom
m. The outer tube consistts of an inelaastic outer laayer,
the cuticle,
c
and internally
i
a hypodermis
h
and longituddinal musclees. Nerve truunks and
excrretory ducts are
a found in the outer tubbe. The innner tube conssists of the allimentary annd
reprooductive sysstems
- the allimentary traact has an annterior oral opening,
o
whiich leads intoo a mouth orr buccal capssule,
esopphagus, and intestine.
i
Inn the male, thhis tract endss in a terminnal cloaca annd in the fem
male a
term
minal or subteerminal anuss
- the reeproductive system in feemales consists of two tuubular structuures, each with
w an ovaryy
leadiing to a uteru
us which joiin at a comm
mon vagina and
a vulva; inn males, there is a single tube
withh a testis lead
ding to a vas deferens annd ejaculatorry duct in a cloaca
c
in which, in somee
malees, there are one or two cuticular
c
exttrusible orgaans, spicules,, used for dillating the vuulva
durinng copulatio
on
- the chharacteristics of the cuticcle, the oral opening, buuccal capsulee and reproduuctive elemeents
are useful
u
for ideentification of
o the nemattode. In thiss course, the emphasis will
w be placedd on
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Principles
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gross features, those that can be seen by eye, e.g., size and colour. These features and
location of the worm in the host can lead to a fairly accurate identification. Occasionally, it
will be necessary to examine some microscopic features.
Egg
-

spherical to oval; the shell may have up to three layers; the stages of the embryo that are
commonly seen in fresh feces are the one or two cell stage, the morula consisting of 16-64
cells, the vermiform embryo or partially developed larva, and the fully developed larva. The
characteristics of the egg as seen in fresh feces or urine, its size, shape, color, form of the
shell and development of the embryo, are important in diagnosis
Larva
- for some nematodes, a larva is found in the feces, blood or skin and its characteristics are
important for diagnosis
E.
1.

2.
3.

4.

5.

6.

7.

Life Cycle
The stages include an egg, five larval stages (L1, L2, L3, L4, L5) with each stage separated by
a molt, and the L5 maturing without a further molt to an adult. The L3 is usually the
infective stage
Some nematodes can complete the entire life cycle within a host, but generally there is a
parasitic phase and a phase outside of the host
Eggs are usually liberated in the feces of the host, but eggs may be seen also in the urine. In
some nematodes, larvae are found in the feces and may be seen also in the blood or skin of
the host
The life cycle may be direct or indirect. Most nematodes of domesticated animals use a
direct life cycle. In a direct life cycle, only one host is required for development of the
nematode. In an indirect life cycle, a definitive or final host (DH) and an intermediate host
(IH) are required
In some nematodes with a direct cycle, the egg hatches in the external environment to give
an L1. The L1 and subsequently the L2 and L3 are free-living stages. The L1 and L2 are
found in feces or soil and are microbivorous i.e., they feed on microbes and microbial
products. The L3 is the infective stage, is non-feeding and will move away from the fecal
mass when it is wet or when it disintegrates. When the leafy parts of the vegetation are wet
with dew or rain, L3s can migrate upwards in moist films of water. When the vegetation is
dry, the larvae are found in the moist environment of the mat of the vegetation
In some nematodes with a direct cycle, the egg does not hatch in the external environment
but the embryo develops within the egg to the infective stage. The infective form is the egg
containing a larva and this is generally the L3
Development of the free-living stages is controlled by temperature and moisture.
Temperatures of 18-26C and high humidity are favourable for development. Many eggs
and larvae will not survive freezing nor temperatures above 40C or arid conditions.
However, the egg and the L3 are the stages that are more resistant to environmental
extremes. The shell of the egg and an extra cuticle on the L3 allows for protection. The
extra cuticle on the L3 is the uncast cuticle of the L2 - the molt from the L2 to the L3 is an
incomplete one.

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Principles of Parasitology

8.

In an indirect life cycle, the egg or L1 in the feces is ingested by an IH in which development
proceeds to the infective stage. The L1 in the blood or skin of the DH is ingested by a biting
arthropod. The length of time for development of the parasite in the arthropod is controlled
by temperature
9. There are several ways by which an animal can be infected with nematodes. The most
common route of infection is by ingestion of the infective form or oral transmission. In
nematodes with a direct cycle, the host consumes the infective form with feed or water. If
the infective form is the infective egg, the egg hatches in the gastrointestinal tract of the host
to liberate the L3. Other routes of infection are per cutaneous transmission when infective
larvae penetrate the skin, transplacental transmission when larvae cross the placenta to infect
the fetus, and transmammary transmission when larvae are ingested by neonates with milk
as they suckle the dam. In nematodes with an indirect life cycle, the definitive host is
infected after consuming the infective form with the IH which is in a predator-prey
relationship with the DH, by scavenging on infected carcasses or by accidental ingestion of
the IH. Biting arthropods that are IH inject or deposit the L3 in/on the skin of the DH
10. An animal can be infected also when it ingests a paratenic host containing the infective
form. The paratenic host is infected after consuming the infective form either as a freeliving stage or as an encysted form in an IH
11. In the DH, development proceeds from the L3 to the adult. The early parasitic stages may
migrate and develop in several organs before attaining the site where they mature to adults.
Some common migratory routes are:
Tracheal migration - Infective larvae penetrate the wall of the gastrointestinal tract and proceed
to the liver by way of the portal vein, then through the caudal vena cava to the right side of
the heart, pulmonary arteries and the lung. Larvae attain the lungs also following skin
penetration and the venous system. Larvae in the capillaries in the lung break out through
the alveoli, migrate up the bronchial tree and the trachea to the mouth, are swallowed and
mature in the gastrointestinal tract
Somatic migration - Larvae arrive in the lung as in tracheal migration, but remain in the blood
stream, bypassing the alveoli to attain the left side of the heart. Larvae in the arterial blood
are sprayed to all parts of the body and lodge in many tissues, primarily skeletal muscle,
where they do not develop but may remain alive for long periods
Mucosal migration - Infective larvae enter the mucosal pits in the stomach or abomasum or
spaces between villi and within crypts in the intestine or may penetrate the mucosa. Larvae
develop for a period in those sites before returning to the lumen to mature to adults
12. Prepatent Period (PPP) - this is the length of time from infection of an animal (the DH) with
the infective stage, through the development of the parasite to the adult and to that point
where eggs or larvae produced by the adults are found in the feces (or larvae in blood) of the
animal. Development of a nematode may be suspended or stopped at some point in the life
cycle and the developmental time will be prolonged. Usually this stoppage occurs early in
the parasitic phase and larvae are referred to as hypobiotic, arrested, dormant or inhibited.
Stimuli which induce hypobiosis or cause resumption of development have not been clearly
defined. In some nematodes, climatic factors influence infective larvae in the external
environment to become hypobiotic when they enter the host. Hormonal and other stimuli
from the host may influence the resumption of larval development
PATH*3040

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Principles of Parasitology

Generalized nematode anatomy

Buccal Cavity
Oesophagus
Nerve Ring
Excretory Pore
Intestine
Tubular Oviduct
Oesophagus

Tubular Testes
Uterus
Pseudocoelum
Body Wall

Bursa
Spicule
Anus
Cloaca

Dorsal Nerve
Cuticle
Hypodermis
Intestine

Ovary
Muscle Cells
Pseudocoelum

Excretory Canal

Uterus with Eggs

PATH*3040

Ventral Nerve

21

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Principles of Parasitology

Illustraations of Vaarious Nemaatode Parasiites

A larrge mass of asccarid nematodees, Toxocara caanis,

remooved at necropssy from the sm
mall intestine off a
pupppy.
Egg of an ascarid
a
nemato
ode showing thhe lumpy
protein coaat that protects the developingg larva.

Adullt whipworms, Trichuris vulppis - any guess where

the common name came from??

Typical egggs of Trichuriss vulpis (an enooplid

nematode the whipworm
m of dogs) shoowing the
typical bipolar plugs.

Hypertrophhied right ventricle of a caninne heart

showing a large number of
o adult heartw
worms,
Dirofilariaa immitis.
PATH*30440

The business
b
end of a strongylid nematode,
n
the
buccal cavity of thee canine hookw
worm, Ancylosttoma
caninnum, showing its
i prominent teeth.
t
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Artthropodaa
A.

Taxxonomic Cla
lassification

Phylum Arthropoda
A
(arthropods)
(
)
Class Inssecta (insectss)
Class Araachnida (tick
ks, mites, scorpions, spidders)
Class Cruustacea (cop
pepods, crayffish, crabs)
Class Penntastomida (tongueworm
(
ms)
B.

Hoost Spectrum
m

Any anim
mal
Some meembers are highly
h
host sppecific
Others inndiscriminan
nt in choice
C.

Sitte of Infectio
on

Most are parasites off the skin eithher on or in;; some are innternal
e.g., botss, warbles, to
onguewormss
D.

Moorphology

Jointed appendagesa
paired, bilatterally symm
metrical
Chitinous exoskeleto
on - rigid porrtions unitedd by flexible portions
Body seggmented - a number
n
of seegments; som
me may havve legs
- segmenntation has diisappeared in some mem
mbers and soome stages inn the life cyccle have no llegs
Sexes sepparate
E.

Liffe Cycle

An egg a series off stages or innstars each separated by a molt ann adult.

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Exxamples of Arthropodan
A
n Parasites

Cheyletiella sp.) found on a cat.

c
Mitees: Fur mites (C

Lice: Hem
matopinus suis removed
r
from a pig.

g
to the haiir of its host.
Lice a niit (louse egg) glued

Fliess :A brachyceraan fly a deer fly Chrysopss

speciies.
Ticks: Derrmacentor and
dersoni (male, l.;
l female, r).

Flies: Gasterophilus inteestinalis larvae from a horse.

PATH*30440

Mitees: The sarcopttic mange mite, Sarcoptes scaabiei.

24

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