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''Ancylostoma ceylanicum'' is a
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has c ...
roundworm The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...
belonging to the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''
Ancylostoma ''Ancylostoma'' is a genus of nematodes that includes some species of hookworms. Species include: : ''Ancylostoma braziliense'', commonly infects cats, popularly known in Brazil as ''bicho-geográfico'' : ''Ancylostoma caninum'', commonly infec ...
''. It is a
hookworm Hookworms are intestinal, blood-feeding, parasitic roundworms that cause types of infection known as helminthiases. Hookworm infection is found in many parts of the world, and is common in areas with poor access to adequate water, sanitation, an ...
both of humans and of other mammals such as
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
s,
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s, and
golden hamster The golden hamster or Syrian hamster (''Mesocricetus auratus'') is a rodent belonging to the hamster subfamily, Cricetinae. Their natural geographical range is in an arid region of northern Syria and southern Turkey. Their numbers have been d ...
s. It is the only
zoonotic A zoonosis (; plural zoonoses) or zoonotic disease is an infectious disease of humans caused by a pathogen (an infectious agent, such as a bacterium, virus, parasite or prion) that has jumped from a non-human (usually a vertebrate) to a human. ...
hookworm species that is able to produce
symptom Signs and symptoms are the observed or detectable signs, and experienced symptoms of an illness, injury, or condition. A sign for example may be a higher or lower temperature than normal, raised or lowered blood pressure or an abnormality showin ...
atic infections in humans, with the majority of cases being in
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
.


Discovery and history

Since the description of ''Ancylostoma ceylanicum'' by
Arthur Looss Arthur Looss (16 March 1861 – 4 May 1923) was a German zoologist and parasitologist. Looss was born in 1861 in Chemnitz, and was educated both there and in Łódź, Poland. Thereafter, he studied at the University of Leipzig, where he received ...
in 1911, and '' A. braziliense'' by Gomes de Faria in 1910, the two species were considered
synonymous A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
because of their apparent similarities in almost all respects. In 1913, comparison of specimens from human,
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
,
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
and
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'' native to Africa and India. It has a muscular, broad-chested body; short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphi ...
infections in India led to the conclusion that they were definitely identical. In 1915 Gomes de Faria described the
anatomical Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
structures and concluded that the two were distinct species. Until 1921 ''A. ceylanicum'' was accepted as a valid species. However, in 1922 Gordon made an exhaustive comparison from specimens collected in
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
,
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
and India, and he failed to identify any distinction. Other
parasitologist Parasitology is the study of parasites, their hosts, and the relationship between them. As a biological discipline, the scope of parasitology is not determined by the organism or environment in question but by their way of life. This means it f ...
s also became convinced that the two names were synonymous. In 1951 Biocca made an elaborate study of different hookworms in the collection of
London School of Hygiene and Tropical Medicine The London School of Hygiene and Tropical Medicine (LSHTM) is a public research university in Bloomsbury, central London, and a member institution of the University of London that specialises in public health and tropical medicine. The inst ...
, the
Liverpool School of Tropical Medicine The Liverpool School of Tropical Medicine (LSTM) is a higher education institution with degree awarding powers and registered charity located in Liverpool, United Kingdom. Established in 1898, it was the first institution in the world dedicated ...
and personal collections. He finally identified the defining characters between the two for classifying them as distinct species, which eventually gained general acceptance.


Structure

The adult hookworms are white and about 6–10 mm long. They are generally stouter than ''A. braziliense''. The anterior end is bent dorsally, which gives the body a characteristic "hooked" or J-shaped appearance, hence the common name hookworm. Females have a tapered narrow posterior end, while males have a feathery posterior end owing to their copulatory
bursa ( grc-gre, Προῦσα, Proûsa, Latin: Prusa, ota, بورسه, Arabic:بورصة) is a city in northwestern Turkey and the administrative center of Bursa Province. The fourth-most populous city in Turkey and second-most populous in the ...
. Hookworm species are not easily distinguished from the gross appearance. The key diagnostic feature is the appearance of their mouths. Unlike other hookworms, ''A. ceylanicum'' has a mouth with cutting plate with a sharp dorsal end that looks like a tooth and a less distinct sharp ventral end.


Life cycle

The Infective
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e quickly undergo
moulting In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
to shed their sheath either upon
ingestion Ingestion is the consumption of a substance by an organism. In animals, it normally is accomplished by taking in a substance through the mouth into the gastrointestinal tract, such as through eating or drinking. In single-celled organisms ingest ...
by the
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman * Michel Host ...
or upon burrowing into the host's skin. If ingested, they pass through the stomach into the
intestine The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans ...
and attach themselves to the
mucosa A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It is ...
. If they have burrowed through the skin, they invade the subcutaneous blood vessels, are carried to the
lung The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of t ...
s, and then move to the intestine via
trachea The trachea, also known as the windpipe, is a Cartilage, cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends ...
,
oesophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
and
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
. In either case, the larvae develop into the final 3rd stage in the intestinal wall. The 4th stage larvae appear in 47 hours after oral infection. The 5th stage immature worms appear 6 days after infection. They reach
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...
on and after 2 weeks after infection. Unlike the hookworms ''
Necator americanus ''Necator americanus'' is a species of hookworm (a type of helminth) commonly known as the New World hookworm. Like other hookworms, it is a member of the phylum Nematoda. It is an obligatory parasitic nematode that lives in the small intestine ...
'' and ''
Ancylostoma duodenale ''Ancylostoma'' is a genus of nematodes that includes some species of hookworms. Species include: : ''Ancylostoma braziliense'', commonly infects cats, popularly known in Brazil as ''bicho-geográfico'' : ''Ancylostoma caninum'', commonly infec ...
'', ''A. ceylanicum'' can competently infect and thrive within not only human hosts but other mammalian hosts as well (such as dogs, cats, or golden hamsters). Because it may primarily infect non-human mammals, and infect humans only opportunistically when its mammalian hosts are in close contact with people, ''A. ceylanicum'' is termed a zoonotic hookworm. This same trait makes ''A. ceylanicum'' a uniquely useful hookworm, since it can be studied in the laboratory while infecting golden hamsters, unlike most or all strains of '' N. americanus'' or '' A. duodenale'', and thus can also be used as a test organism for possible drugs or vaccines aimed at preventing human infections.


Pathogenicity

''Ancylostoma ceylanicum'' attaches itself to
capillary A capillary is a small blood vessel from 5 to 10 micrometres (μm) in diameter. Capillaries are composed of only the tunica intima, consisting of a thin wall of simple squamous endothelial cells. They are the smallest blood vessels in the body: ...
beds in the
small intestine The small intestine or small bowel is an organ in the gastrointestinal tract where most of the absorption of nutrients from food takes place. It lies between the stomach and large intestine, and receives bile and pancreatic juice through the p ...
of a host where it feeds on blood and causes
anaemia Anemia or anaemia (British English) is a blood disorder in which the blood has a reduced ability to carry oxygen due to a lower than normal number of red blood cells, or a reduction in the amount of hemoglobin. When anemia comes on slowly, th ...
. In hamsters anaemia is most severe between the 13th and 60th days of infection, and is accompanied by significant loss in body weight. Experimental infection of hamsters shows increased
antibodies An antibody (Ab), also known as an immunoglobulin (Ig), is a large, Y-shaped protein used by the immune system to identify and neutralize foreign objects such as pathogenic bacteria and viruses. The antibody recognizes a unique molecule of the ...
, peripheral cellular immune suppression, which is characterized by a reduction in the total
white blood cell White blood cells, also called leukocytes or leucocytes, are the cell (biology), cells of the immune system that are involved in protecting the body against both infectious disease and foreign invaders. All white blood cells are produced and de ...
count,
neutropenia Neutropenia is an abnormally low concentration of neutrophils (a type of white blood cell) in the blood. Neutrophils make up the majority of circulating white blood cells and serve as the primary defense against infections by destroying bacteria ...
and
lymphopenia Lymphocytopenia is the condition of having an abnormally low level of lymphocytes in the blood. Lymphocytes are a white blood cell with important functions in the immune system. It is also called lymphopenia. The opposite is lymphocytosis, which re ...
. The most serious effects are manifested in children and women of childbearing age displaying chronic intestinal
blood loss Bleeding, hemorrhage, haemorrhage or blood loss, is blood escaping from the circulatory system from damaged blood vessels. Bleeding can occur internally, or externally either through a natural opening such as the mouth, nose, ear, urethra, vag ...
which may result in
iron deficiency Iron deficiency, or sideropenia, is the state in which a body lacks enough iron to supply its needs. Iron is present in all cells in the human body and has several vital functions, such as carrying oxygen to the tissues from the lungs as a key ...
, anaemia and
hypoalbuminemia Hypoalbuminemia (or hypoalbuminaemia) is a medical sign in which the level of albumin in the blood is low. This can be due to decreased production in the liver, increased loss in the gastrointestinal tract or kidneys, increased use in the body, or ...
. The long-term effects include impaired physical, intellectual and cognitive development of children, increased mortality in pregnant women and their infants and reduced physical capacity.


Epidemiology

''Ancylostoma ceylanicum'' infection is found in
Cambodia Cambodia (; also Kampuchea ; km, កម្ពុជា, UNGEGN: ), officially the Kingdom of Cambodia, is a country located in the southern portion of the Indochinese Peninsula in Southeast Asia, spanning an area of , bordered by Thailand t ...
,
Malaysia Malaysia ( ; ) is a country in Southeast Asia. The federation, federal constitutional monarchy consists of States and federal territories of Malaysia, thirteen states and three federal territories, separated by the South China Sea into two r ...
,
The Solomon Islands Solomon Islands is an island country consisting of six major islands and over 900 smaller islands in Oceania, to the east of Papua New Guinea and north-west of Vanuatu. It has a land area of , and a population of approx. 700,000. Its capita ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
,
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
, India,
Sri Lanka Sri Lanka (, ; si, ශ්‍රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...
, South Africa,
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
,
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
,
Fiji Islands Fiji ( , ,; fj, Viti, ; Fiji Hindi: फ़िजी, ''Fijī''), officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists ...
and
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia, at the junction of the East and South China Seas in the northwestern Pacific Ocean, with the People's Republic of China (PRC) to the northwest, Japan to the nort ...
.


Diagnosis

Infection is detectable from a
stool sample A stool test is a medical diagnostic technique that involves the collection and analysis of fecal matter. Microbial analysis (culturing), microscopy and chemical tests are among the tests performed on stool samples. Collection Stool samples shoul ...
of the host. Eggs can be microscopically analysed. However, there is no clear-cut distinction between different hookworms due to their physical similarity, and species are often mistaken for one another. The Kato-Katz method and molecular techniques such as
polymerase chain reaction The polymerase chain reaction (PCR) is a method widely used to rapidly make millions to billions of copies (complete or partial) of a specific DNA sample, allowing scientists to take a very small sample of DNA and amplify it (or a part of it) t ...
(PCR) can be used to distinguish hookworm species, but are not always effective at doing so. Sequencing of
ribosomal RNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal ...
gene is often reliable but time consuming. A PCR coupled with high resolution melting-curve (HRM) was found to be highly sensitive in discriminating ''A. ceylanicum'' from other hookworms.


Treatment

Ivermectin Ivermectin (, '' EYE-vər-MEK-tin'') is an antiparasitic drug. After its discovery in 1975, its first uses were in veterinary medicine to prevent and treat heartworm and acariasis. Approved for human use in 1987, today it is used to treat inf ...
is highly effective even at the low dose of 100 μg per kg, and
pyrantel Pyrantel is a medication used to treat a number of parasitic worm infections. This includes ascariasis, hookworm infections, enterobiasis (pinworm infection), trichostrongyliasis, and trichinellosis. It is taken by mouth. Side effects include n ...
is also effective at 25–50 mg per kg.
Benzimidazoles Benzimidazole is a heterocyclic aromatic organic compound. This bicyclic compound may be viewed as fused rings of the aromatic compounds benzene and imidazole. It is a colorless solid. Preparation Benzimidazole is produced by condensation of o- ...
such as
mebendazole Mebendazole (MBZ), sold under the brand name Vermox among others, is a medication used to treat a number of parasitic worm infestations. This includes ascariasis, pinworm infection, hookworm infections, guinea worm infections, hydatid disease, ...
, parbendazole and
thiabendazole Tiabendazole (INN, BAN), also known as thiabendazole (AAN, USAN) or TBZ and the trade names Mintezol, Tresaderm, and Arbotect, is a preservative, an antifungal agent, and an antiparasitic agent. Uses Preservative Tiabendazole is used primar ...
are also highly effective.


Genome

A draft assembly of the genome of ''Ancylostoma ceylanicum'' has been sequenced and analyzed. It comprises 313 Mb, estimated to be 95% of the full genome, with transcriptomic data throughout infection showing expression of 30,738 predicted protein-coding genes. These include genes encoding three gene families upregulated during successive stages of infection: Activation-associated Secreted Protein Related (ASPRs), Secreted L4 Proteins (SL4Ps), and Secreted Clade V proteins (SCVPs). The genes of ''A. ceylanicum'' also include predicted targets for drugs and vaccines.


References


External links


Taxonomy at UniProt''A. ceylanicum'' genome at WormBase''A. ceylanicum'' genome at ParaSiteNemtode.net The Genome InstituteThe Genome Institute of Washington UniversityNEMBASE4Protein database at BerkeleyDisease informationSEM image at Visual Unlimited
{{Taxonbar, from=Q16974161 Parasitic nematodes of mammals Ancylostomatidae Nematodes described in 1911