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Ancylostoma caninum

Ancylostoma caninum is a species of nematode which

principally infects the small intestine of dogs.[1][2][3] The
result of A. caninum infection ranges from asymptomatic
cases to death of the dog; better nourishment, increasing age, prior A. caninum exposure or vaccination are all
linked to improved survival.[2][4][5][6] Other hosts include
carnivores such as wolves, foxes and cats with a small
number of cases having been reported in humans.[1][2]
Warm and moist conditions are important to allow survival of A. caninum during the free-living stages of
its life cycle and for this reason it is largely restricted
to temperate, tropical and sub-tropical regions.[3][7] In
parts of the world where these climatic requirements
are met such as Sri Lanka, southeast Asia and Malaysia
A. caninum egg.
A. caninum is the main cause of hookworm disease in

is a unique feature of Strongylida members, thus making it a useful means for identifying members of this
suborder; it is also used to distinguish members within the
suborder due to dierences in bursa appearance between
species.[2] The vulva of A. caninum females is located at
the boundary of the second and nal thirds of the body.[1]
The teeth of A. caninum are found in the buccal capsule
and divided into three sets.[1][2] Two ventral sets form a
lower-jaw equivalent, while a further set projects from
the dorsal side and loosely equates to an upper-jaw.[2]
Each ventral set has three points with those furthest to
the sides being the largest.[1][8] While the ventral sets are
prominent, the dorsal set is hidden deeper in the buccal
A. caninum bends its head end upward (dorsally) which
has in the past been noted as a potential source confusion
when determining how the hookworm is oriented.[2] If it
has recently ingested blood A. caninum is red in colour,
if not it appears grey.[1] A. caninum has an alimentary
canal made up of an esophagus, intestine and rectum
the esophagus is highly muscular reecting its role in
pulling intestinal mucosa into the body when it feeds.[2][3]
Esophageal and anal rings of A. caninum are the source of
nerve bres that extend throughout the body to innervate
sensory organs including amphids and phasmids.[1][3]

Mouth and teeth of A. caninum.

A. caninum females are typically 1416 millimetres

(0.550.63 in) long and 0.5 mm (0.02 in) wide, while
the males are smaller at 1012 mm (0.390.47 in) in
length and 0.36 mm (0.01 in) in width.[2][3] On males
a copulatory bursa exists which, during copulation, attaches the female via ~0.9 mm long spine-like spicules positioned on three muscular rays.[1][2][3] As with other ne- Eggs are laid by the females typically when at the 8-cell
matodes, the sperm lack agella.[1] The copulatory bursa stage.[3] Eggs are 38-43 M in width with thin walls.[3][5]



Freezing, heating above 37 C (99 F), drying or exposing A. caninum to sunlight all give reduced survival of
the free-living stage with rates of infection rising with
temperature provided 37 C is not exceeded.[1][2] A. caninum is therefore largely restricted to warm, moist climates though infections are seen in the USA and southern
Canada where the temperature is sub-optimal.[2] Specic
niches are also able to satisfy the environmental requirements of A. caninum despite not necessarily being in the
tropics, such as mines.[2]


Life Cycle
Transmission via the Environment

3.2 Direct transmission

It is also possible for direct transmission between hosts.
Larvae having accessed through the skin may avoid exit
via the lungs and remain in circulation for transport
around the body.[3] At the uterine artery of a pregnant female the larvae are able to cross the placenta to cause prenatal infection of foetuses.[3] Larvae of an infected foetus
will move to the liver until birth at which point migration
continues with movement to the intestine via the circulation and lungs as previously described.[3] Alternatively
A. caninum larvae evading exit from the circulation at the
lungs may instead be carried to the mammary glands and
transmitted from the mother in her colostrum or milk to
her pups; infection then proceeds in an identical manner
to infection by ingestion from the environment.[3][5] Infected bitches have been found to only rarely give prenatal transmission to pups while the likelihood of nursing
of pups causing transmission (via the lactational route) is
much higher.[12]

4 Pathogenesis

Eggs are excreted from host in the feces and typically

hatch within a day on moist, warm soil giving larvae with 4.1
a non-living cuticle layer.[1][2][5] By four or ve days the
larvae have moulted twice and are now able to infect a
host.[2] Migration occurs from the faeces into the surrounding soil.[2] Two routes of infection from the environment exist. The rst route involves penetration of skin
at hair follicles or sweat glands, especially between the
footpads where contact with soil is frequent and the skin
is thinner than otherwise.[2] Secretion of a protease by
A. caninum is thought to aid this process.[2] The larvae
then migrate through the dermis of the skin, enter the
circulatory system and are carried to the lungs.[2] A. caninum larvae exit the blood at the lungs, move from the
alveoli up through the trachea and are swallowed to end
up in the intestine.[2]
The second and more common route to the small intestine is by direct ingestion of A. caninum by the host, but
the subsequent process is identical in either case.[2][3] It
is during this third stage of the larva that male or female
reproductive organs become established.[3] Larvae of this
stage have been shown to secrete a molecule (Ac-asp-2)
related to venom allergens in response to host-specic signals; this is thought to have a possible role in helping with
the infection process.[9] A third and nal moulting occurs to give the mature form of A. caninum which then
feeds on mucosa and blood of the small intestine wall.[2]
The trigger of feeding is understood to be a receptormediated response, however the detail of this process has
yet to be established.[10] Sexual reproduction also occurs
in the intestine to give a further round of eggs to complete
the cycle.[2] Females are thought to produce a pheromone
which attracts males and are able to produce approximately 10,000 eggs per day.[1][11]

Damage during migration to intestine

Pair of A. caninum hookworms.

Ancylostomum caninum larvae cause damage to the host

at the point of entry through the skin leaving a wound vulnerable to secondary infections.[2] As the larvae migrate
through the skin an inammatory response, dermatitis,
is often stimulated which can be exacerbated in hosts
which give hypersensitive responses.[2][5] Further damage is caused when the larvae leave the circulation and
enter the lung with the amount of damage dependent on
the extent of the infection; pneumonia and coughing are
common consequences.[2]

4.2 Damage once in intestine

Once in the gut A. caninum attaches to and ingests the
mucosal lining along with some consumption of blood;

up to 0.1mL in 24hrs.[2][5] In a 24hr period A. caninum typically feeds from six sites.[2] This damage to
the mucosa compromises the bodys defences and can
result in secondary infections by microbes.[7] A group
of anticoagulant proteins called AcAPs (A. caninum anticoagulant proteins) which inhibit a range of blood
coagulation factors such as Xa are utilised by A. caninum
to help in the feeding process by preventing clotting and
increasing blood loss.[13][14] These AcAPs are among the
most powerful natural anticoagulants that exist and are a
key reason for anemia being caused and blood being observed in the faeces of infected hosts.[5][14] Blood losses
peak just prior to egg production by the females because
this is when their requirements for food are greatest; the
amount that they are eating is also peaking and so maximal damage to the intestine is being caused.[3]


Analysis of faeces is the denitive method by which a

suspected A. caninum infection is conrmed.[2] The faeces are sampled and the characteristic ovular, thin-shelled
eggs of A. caninum looked for.[5] Absence of eggs in faeces does not rule out infection; a signicant delay of at
least 5 weeks exists between initial infection and excretion of eggs in the faeces (larvae must fully mature and
reproduce before eggs can be laid).[3][5] In fact, pups frequently die before passing of eggs in the faeces begins.[5]
Using the number of eggs in stool samples as an indicator of the extent of infestation requires care to be taken
because females have been shown to produce fewer eggs
each when the overall number of worms increases.[15]
Signs and symptoms expected to be observed together
with A. caninum eggs in the faeces are lethargy, weight
loss, weakness, roughness of the hair coat and pale
mucous membranes indicative of anemia.[2][5] Well-fed,
older dogs with smaller infestations may present few
or even none of these symptoms.[2][5] Diarrhoea is rare
but stools are typically black due to the blood-derived
haemoglobin present in them.[2]

Dog kennel: keeping kennels clean reduces risk of A. caninum


health complications to the pups.[5] When infection of

a pregnant bitch is known or suspected fenbendazole or
ivermectin can be administered to the bitch to help avoid
transmission to the pups.[5]
Canines have been seen to develop signicant resistance
to A. caninum naturally with age; this protection develops faster in bitches than in dogs and fully mature bitches
show substantially greater resistance than fully mature
dogs.[4] Specically the age-related resistance means A.
caninum takes longer to reach sexual maturity in older
animals and fewer larvae fully develop.[11]

7 Vaccination

The disease resulting from such A. caninum infection

is referred to by the general term hookworm disease
Numerous vaccines have been developed with varying
or the more specic ancylostomiasis and ancylostomosis
success against A. caninum. Use of an enzyme important
diagnoses which recognise the genus of the causative
in the worms feeding process is popular with one examnematode.[2]
ple being AcCP2, a protease, which when used to vaccinate dogs gives a strong antibody response, lowering of
numbers of eggs found in stools and a decrease in intestinal worm size.[16] These eects are attributed to reduced
6 Prevention and Control
AcCP2 activity upon antibody binding.[16] A similar apA clean environment minimises the risk of A. caninum proach has been taken using another A. caninum digestive
it failed to give statistically signifinfection; this can include regularly washed concrete or enzyme, AcGST1, but
gravel in kennels instead of soil.
Bitches are typically
checked prior to using them for breeding purposes for An alternative approach has been to disrupt the migranematodes such as A. caninum and birth and suckling tory ability of A. caninum, this was done so successfully
can be restricted to sanitised areas to lower the risk of using the AcASP1 protein of A. caninum which gives


increases in antibody levels of all subclasses and a reduced worm burden.[6] Other studies using the same vaccine have shown statistically signicant 79% reductions
in worm burden resulting from this approach.[18]
Animals with prior exposure to A. caninum show enhanced resistance but careful removal of all worms
from the previous infection results in loss of this
improvement.[11] Studies in mice show resistance due to
past exposure can protect against otherwise lethal worm
doses and that this is a general form of resistance - defence is oered against subsequent infections via either
mouth or skin.[19]


Drugs used in treatment of A. caninum infections of dogs include: dichlorvos, fenbendazole,

ubendazole, mebendazole, nitroscanate, piperazine,
pyrantel, milbemycin, moxidectin, diethylcarbamazine,
oxibendazole, and ivermectin.[5]

In Humans

In inappropriate hosts such as humans A. caninum is able

to enter the skin but cannot proceed into the circulation
and on to the intestine; instead the disease dermal larva
migrans results, caused by movement of the nematode
within the skin and which can persist for several months
without intervention.[1]


11 References
[1] Saeed, Sophia (2003). "Ancylostoma caninum". Animal
Diversity Web. Retrieved March 20, 2013.
[2] Marquardt W; Demaree; Grieve (2000). Parasitology and
Vector Bology (2nd ed.). Harcourt Academic. pp. 370
376. ISBN 0124732755.
[3] Olsen W (1986). Animal Parasites: their life cycles
and ecology (3rd ed.). Dover. pp. 399416. ISBN
[4] Miller (1965). Inuence of Age and Sex on Susceptibility of Dogs to Primary Infection with Ancylostoma caninum". The Journal of Parasitology 51 (5): 701704.
doi:10.2307/3276142. PMID 5857264.
[5] Peregrine, Andrew (March 2012). Hookworms in Small
Animals. The Merck Veterinary Manual. Retrieved
March 20, 2013.
[6] Ghosh K, Hotez, P (January 1999).
AntibodyDependent Reductions in Mouse Hookworm Burden
after Vaccination with Ancylostoma caninum Secreted
Protein 1. J Infect Dis. 180 (5): 16741681.
[7] Cheng T (1986). Parasitology and Vector Bology (2nd
ed.). Academic Press. pp. 93, 508. ISBN 0121707555.
[8] Ruppert E (1994). Parasitology and Vector Bology (6th
ed.). Saunders College Pub. p. 293. ISBN 0030266688.
[9] Hawdon J, Narasimhan S, Hotez P (April 1999).
"Ancylostoma secreted protein 2: cloning and characterization of a second member of a family of nematode
secreted proteins from Ancylostoma caninum". Molecular and Biochemical Parasitology 99 (2): 149165.
doi:10.1016/S0166-6851(99)00011-0. PMID 7872431.

While access to the intestine is not possible via this route,

it can occur via ingestion; in a report of 93 enteritis cases [10] Hawdon J, Schad G (February 1990). Serum-Stimulated
Feeding In vitro by Third-Stage Infective Larvae of the
in northern Queensland, Australia which were possibly
Canine Hookworm Ancylostoma caninum". The Journal
caused by A. canium infection, all those interviewed deof Parasitology 76 (3): 394398. doi:10.2307/3282673.
scribed behaviour consistent with A. caninum exposure
PMID 2112598.
and a colonoscopy of one patient gave positive identication of an adult A. caninum worm.[20][21] Since then work [11] Herrick C (September 1928). A Quantitative Study of
Infections with Ancylostoma caninum in Dogs. Am. J.
has shown A. caninum can easily go unnoticed or fail to
Epidemiol. 8 (2): 125157.
be preserved in specimens making the true incidence of
infection in humans likely to be higher than is ocially [12] Burke T, Roberson E (February 1985). Prenatal and lacrecorded.[22]
tational transmission of Toxocara canis and Ancylostoma
caninum: Experimental infection of the bitch before pregnancy. International Journal for Parasitology 15 (1): 71
75. doi:10.1016/0020-7519(85)90104-3.


Economic Burden

[13] Stanssens P, Bergum P, Gansemans Y, et al. (March

1996). Anticoagulant repertoire of the hookworm Ancylostoma caninum". PNAS 19 (5): 21492154. PMC

The animals aected by A. caninum infection are not

used for food or labour purposes thus the economic burden from animal illness is low.[1] Work showing that hu- [14] Cappello M, Vlasuk G, Bergum P, et al. (June
man A. caninum infections are likely underestimated and
1995). "Ancylostoma caninum anticoagulant peptide: a
misdiagnosed indicates the economic impact of A. canhookworm-derived inhibitor of human coagulation facinum through human work days lost may be underestitor Xa. PNAS 92 (13): 61526156. doi:10.1016/0140mated and signicant.[22]
6736(90)91186-E. PMID 7597095.

[15] Sarles M (November 1929). The Eect of Age and Size

of Infestation on the Egg Production of the Dog Hookworm Ancylostoma caninum". The American journal of
hygiene 10 (3): 658666.
[16] Loukas A, Bethony JM, Williamson AL, et al. (May
2004). Vaccination of dogs with a recombinant cysteine protease from the intestine of canine hookworms diminishes the fecundity and growth of worms. J. Infect.
Dis. 189 (10): 195261. doi:10.1086/386346. PMID
[17] Zhan B, Liu S, Perally S, et al. (October 2005).
Biochemical characterization and vaccine potential of
a heme-binding glutathione transferase from the adult
hookworm Ancylostoma caninum". Infect. Immun. 73
(10): 690311. doi:10.1128/IAI.73.10.6903-6911.2005.
PMC 1230892. PMID 16177370.
[18] Ghosh K, Hawdon J, Hotez, P (May 1996). Vaccination
with Alum-Precipitated Recombinant AncylostomaSecreted Protein 1 Protects Mice against Challenge
Infections with Infective Hookworm (Ancylostoma
caninum) Larvae. J Infect Dis. 174 (6): 13801383.
[19] Kerr K (June 1936). Studies on Acquired Immunity to
the Dog Hookworm Ancylostoma caninum". Am. J. Epidemiol. 24 (2): 381406.
[20] Prociv P (June 1990). Human eosinophilic enteritis
caused by dog hookworm Ancylostoma caninum". The
Lancet 335 (8701): 12991302. doi:10.1016/01406736(90)91186-E. PMID 7597095.
[21] Croese J, Loukas A, Opdebeeck J, et al. (January 1994).
Occult enteric infection by Ancylostoma caninum: a previously unrecognized zoonosis. Gastroenterology 106
(1): 312. PMID 8276205.
[22] Walker N, Croese J, Clouston A, et al. (March 1995).
Eosinophilic enteritis in northeastern Australia. Pathology, association with Ancylostoma caninum, and implications. The American Journal of Surgical Pathology
19 (3): 328337. doi:10.1016/S0166-6851(99)00011-0.
PMID 7872431.


External links

Cartoon illustration of the A. caninum life cycle in

dogs, cats and humans.




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