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At present, research is being done on the possibility that pigs are intermediate hosts for N. americanus. Research has shown that the approximately 15% of eggs ingested by a pig host will pass through in the feces.

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Harlen Hays, University of Michigan-Ann Arbor
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Associations

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These parasites are probably not preyed on directly, but are ingested from host to host. Larval mortality is high as most of the parasites do not reach appropriate hosts.

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Morphology

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As a nematode, Necator americanus has a cylindrical body, and a cuticle with three main outer layers made of collagen and other compounds, secreted by the epidermis. The cuticle layer protects the nematode so it can invade digestive tracts of animals.

Eggs range in size from 65-75 micrometers x 36-40 micrometers and are virtually indistinguishable from those of Ancylostoma duodenale, another common hookworm. Necator americanus has four larval stages. The first stage is referred to as rhabditiform larvae because the esophagus has a large bulb separated from the rest of the esophagus by a region called the isthmus. The third stage is referred to as filariform larvae because the esophagus has no bulb. Adult females range in size from 9 mm to 11 mm while the smaller males range in size from 7 mm to 9 mm. The mouth of the adults has two pair of cutting plates, one dorsal and the other ventral. The males of the species are characterized by fused spicules found on the bursa. The common name "hookworm" comes from the dorsal curve at the anterior end.

Range length: 7 to 11 mm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: female larger; sexes shaped differently

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Life Expectancy

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The adult worms can live up to 5 years.

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Habitat

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Adult N. americanus are found exclusively in tropical and temperate regions. Eggs require a moist, warm and shaded environment to hatch. Optimal temperatures for juveniles to mature are from 23 to 30 deg C. Eggs and juveniles die below freezing or with soil dessication. Heavy rains and warmer temperatures appear to have a high positive correlation with the rate of transmission. Necator americanus appears to prefer male hosts to female hosts. However, this may be due to the division of labor in areas of high infestation. Soil type also plays a major part in the habitat for these worms. Ideal soil conditions are where the water drains but not too quickly. The pore size of the soil particle is a major factor.

Habitat Regions: temperate ; tropical

Terrestrial Biomes: savanna or grassland ; forest ; rainforest ; scrub forest

Aquatic Biomes: lakes and ponds; temporary pools

Wetlands: marsh ; swamp

Other Habitat Features: urban ; suburban ; agricultural

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Distribution

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Necator americanus is found in Africa, Asia, and Europe but is predominately found in the Americas and in Australia. In the United States, the largest concentration is found in the southern and southwestern United States. In the rest of the world, N. americanus is found in tropical climates.

Biogeographic Regions: nearctic ; palearctic ; oriental ; ethiopian ; neotropical ; australian

Other Geographic Terms: cosmopolitan

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Trophic Strategy

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Adult N. americanus feed from the blood of their hosts. The worm will attach itself to the intestinal wall and use its cutting plates to cause bleeding. The worm feeds from this blood, possibly causing anemia to the host. Necator americanus does not permanently attach itself to the wall. This allows movement to new sites for feeding and reproduction within the host. Previous sites continue to bleed, adding the host's blood loss.

Animal Foods: blood; body fluids

Primary Diet: carnivore (Sanguivore , Eats body fluids)

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Associations

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Necator americanus mainly infects humans and appears to prefer male hosts to female hosts. However, this may be due to the division of labor in areas of high infestation.

Ecosystem Impact: parasite

Species Used as Host:

  • humans, Homo sapiens
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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Benefits

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None

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Benefits

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As a strict parasite of humans, N. americanus has played a major role in the development of the New and Old worlds. Large numbers of worms cause what is known as "hookworm disease".

Nutrition and blood loss are the major contributors to ill effects from the interaction between host and parasite. The severity of the disease is directly related to the number of worms in the host's body. Each worm is capable of ingesting .03 ml of blood per day. When small amounts of worms have infected a host, the patient will be asymptomatic. Once infections reach 25 to 100 worms, the patient will experience light symptoms that include fatigue, slight weight loss, and possible headaches. Once infestations reach between 100-500 the patient will experience fatigue, iron deficiency leading to anemia, loss of appetite and abdominal pains. Should the infestation reach over 500, the patient will experience anemia and, depending on the diet of the person, possibly death. Children are at greater risks for lifelong damage and death due to their smaller size and greater need for nutrition. Generally infestation does not lead to death, but many cause permanent damage. The mortality rate for N. americanus is around .005% while the morbidity rate is 12%.

Treatment of N. americanus infections is simple, but resistance has been detected to usual treatments. Preventive measures include proper waste disposal, clean water, and proper footwear. Since the larvae need direct skin contact to invade the host, protective clothing is a key to maintaining low incidence rates.

Negative Impacts: injures humans (causes disease in humans )

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Life Cycle

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Necator americanus adults are obligate internal parasites of humans. Both the first and second stage rhabditiform larvae are free-living. Eggs are passed out through the feces of humans. These eggs will hatch within 2 days, and the first rhabditiform larva emerges. This larva will molt twice within 10 days to become a third stage filariform larva. The filariform larva will come into contact with the host's skin, and burrow into it. Travelling through the circulatory system to the lungs, the third stage larva will either be coughed up and swallowed or will migrate up the bronchi to the throat area. Once swallowed, the larva will make its way to the intestine. Upon reaching the intestine, the larva will molt twice and an adult will emerge. This adult will use its hooks to attach itself to the inner wall of the intestine and will cause hemorrhaging. The worm will feed from this blood. Mating occurs in the intestine. The eggs are the passed out with the feces.

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Conservation Status

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None

US Federal List: no special status

CITES: no special status

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Behavior

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Nematodes within the Secernentea have phasmids, which are unicellular glands. Phasmids likely function as chemoreceptors. Females may produce pheromones to attract males.

Nematodes in general have papillae, setae and amphids as the main sense organs. Setae detect motion (mechanoreceptors), while amphids detect chemicals (chemoreceptors).

Communication Channels: tactile ; chemical

Other Communication Modes: pheromones

Perception Channels: tactile ; chemical

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Hays, H. 2001. "Necator americanus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Necator_americanus.html
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Reproduction

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Sexual maturity is reached at the final molt. Egg production in females occurs five weeks or more after the female matures. Mating occurs in the intestine of the host.

Males are required to find females and inject their sperm into the females. Females may produce a pheromone to attract males. The male coils around a female with his curved area over the female genital pore. The gubernaculum, made of cuticle tissue, guides spicules which extend through the cloaca and anus. Males use spicules to hold open the genital opening on the female to allow transfer of sperm, which are amoeboid-like and lack flagella. The fertilized females then lay eggs in the surrounding areas. Females are capable of producing 10,000 eggs per day. The eggs are the passed out with the feces.

Key Reproductive Features: gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); oviparous

There is no parental investment beyond egg laying.

Parental Investment: pre-fertilization (Provisioning)

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Brief Summary

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The human hookworms include the nematode species Necator americanus and Ancylostoma duodenale. A larger group of hookworm species normally infecting non-human animals can invade and parasitize humans or can penetrate the human skin--causing cutaneous larva migrans--but do not develop any further. Hookworm is the second most common human helminthic infection (after ascariasis) and among the most common chronic infections in the world, affecting around three quarters of a billion people in the tropics and subtropics, particularly in China and sub-Saharan Africa (deSilva et al. 2003). Hookworm species are worldwide in distribution, mostly in areas with moist, warm climates. Both N. americanus and A. duodenale are found in Africa, Asia, and the Americas. Necator americanus predominates in the Americas and Australia, while only A. duodenale is found in the Middle East, North Africa, and southern Europe.

Hookworms are transmiited to humans via contact with contaminated soil. Eggs are passed in the stool and under favorable conditions (moisture, warmth, shade), larvae hatch in 1 to 2 days.The released rhabditiform larvae grow in the feces and/or the soil and after 5 to 10 days (and two molts) they become filariform (third-stage) larvae that are infective. These infective larvae can survive 3 to 4 weeks in favorable environmental conditions. On contact with the human host, the larvae penetrate the skin and are carried through the blood vessels to the heart and then to the lungs.They penetrate into the pulmonary alveoli, ascend the bronchial tree to the pharynx, and are swallowed. The larvae reach the small intestine, where they reside and mature into adults. Adult worms live in the lumen of the small intestine, where they attach to the intestinal wall with resultant blood loss by the host. Most adult worms are eliminated in 1 to 2 years, but longevity may reach several years. Some A. duodenale larvae, following penetration of the host skin, can become dormant (in the intestine or muscle). In addition, infection by A. duodenale probably also occurs by the oral and transmammary route. Necator americanus, however, requires a transpulmonary migration phase. Iron deficiency anemia (caused by blood loss at the site of intestinal attachment of the adult worms) is the most common symptom of hookworm infection, and can be accompanied by cardiac complications. Gastrointestinal and nutritional/metabolic symptoms can also occur. In addition, local skin manifestations ('ground itch') can occur during penetration by the filariform (L3) larvae, and respiratory symptoms can be observed during pulmonary migration of the larvae.

(Primary source: Centers for Disease Control Parasites and Health Website)

Loukas et al. (2006) discussed the need for and prospects of developing a human hookworm vaccine. Hotez et al. (2004) provide a broad review of issues related to human hookworm infection.

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Necator americanus

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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 of human hosts.[1] Necatoriasis—a type of helminthiasis—is the term for the condition of being host to an infestation of a species of Necator. Since N. americanus and Ancylostoma duodenale (also known as Old World hookworm) are the two species of hookworms that most commonly infest humans, they are usually dealt with under the collective heading of "hookworm infection". They differ most obviously in geographical distribution, structure of mouthparts, and relative size.[2]

Necator americanus has been proposed as an alternative to Trichuris suis in helminthic therapy.[3]

Morphology

This parasite has two dorsal and two ventral cutting plates around the anterior margin of the buccal capsule. It also has a pair of subdorsal and a pair of subventral teeth located close to the rear. Males are usually 7–9 mm long, whereas females are about 9–11 mm long. The typical lifespan of these parasites is 3–5 years. They can produce between 5,000 and 10,000 eggs per day.[4]

Habitat

N. americanus is primarily found in tropical and temperate areas. This parasite thrives in warmer climates because to hatch, the eggs require a moist, warm, and shaded environment. The thin, smooth shells of this species cause the eggs and juveniles to die in freezing temperatures or with soil desiccation. Therefore, the type of soil where the parasite resides is also very important for their ideal living conditions. Ideal soil conditions tend to be in areas where water is able to drain at a standard pace, and the size of the soil particles is neither too large nor too small. That way, the degree of dampness, as well as the openings in the soil, allow the parasites to burrow to the surface and attach to the skin of their next host. High transmission rates seem to be congruent with the heavy rains and warm temperatures that are characteristic of tropical climates. One anomaly of this species is that it appears to prefer male hosts to female hosts, likely because of the division of labor in regions of its prevalence.[5]

Lifecycle

Life cycle of N. americanus inside and outside of the human body

This worm starts out as an unembryonated egg in the soil. After 24–48 hours under favorable conditions, the eggs become embryonated and hatch. This first juvenile stage 1 is known as 'rhabditiform'. The rhabditiform larvae grow and molt in the soil, transforming into a juvenile stage 2. The juvenile stage 2 molts once more until reaching the juvenile 3 stage, which is also called 'filariform'; this is also the infective form. The transformation from rhabditiform to the filariform usually takes 5–10 days.[6] This larval form is able to penetrate human skin, travel through the blood vessels and heart, and reach the lungs. Once there, it burrows through the pulmonary alveoli and travels up the trachea, where it is swallowed and carried to the small intestine. There, it attaches to the intestinal wall, and matures into an adult and begins reproduction. Adults live in the lumen of the intestinal wall, where they cause blood loss to the host. The eggs produced by the adults end up on the soil after leaving the body through the feces; female hookworms produce up to 30,000 eggs per day.[7][8] On average, most adult worms are eliminated in 1–2 years. The N. americanus lifecycle only differs slightly from that of A. duodenale. N. americanus has no development arrest in immune hosts and it must migrate through the lungs.

Pathogenesis and symptoms

The pathology of N. americanus is divided into two stages – larvae and adults. The larvae penetrate the uninfected skin and travel through various organs, including the respiratory tract and lymph nodes. Once in the lymph nodes, the larvae start entering the blood, lungs, and intestines. Some larvae cannot readily enter the dermis and remain trapped in the skin, causing skin irritation and cutaneous larva migrans. Other symptoms include excessive coughing and dyspnea (short of breath) during larval migration. Once attached to the intestinal wall, N. americanus resides and matures into adults, penetrates blood vessels, and sucks blood. The incubation process of the larvae begins once entered into the small intestine; therefore, symptoms may not arise for up to 40 days, but this is variable from person to person.[9] Blood loss from sites of intestinal attachment may cause iron-deficiency anemia and protein loss.[8] One individual N. americanus can cause 30 μl of blood loss per day.[10] Iron-deficiency anemia can cause intellectual disability and growth insufficiency in children. Further, infected patients experience abdominal pain (exacerbated by meals) with diarrhea, bloating, and nausea.[11]

Epidemiology

In the United States, 95% of human hookworm cases are caused by N. americanus, primarily in young school children in economically deprived rural areas. Historically, there have been high rates of infection among children in the American South.[12] Juveniles cannot survive freezing temperatures, so the highest prevalence occurs in areas with warmer temperatures and greater rainfall. The greatest incidence of infections occurs in Asia and sub-Saharan Africa, especially in poverty-stricken areas with poor sanitation.[8] A. duodenale infections occur at a lesser rate and are seen primarily in Europe and the Mediterranean.[13]

Genome

A draft assembly of the genome of Necator americanus has been sequenced and analyzed.[14] It comprises 244 Mbp with 19,151 predicted protein-coding genes; these include genes whose products mediate the hookworm's invasion of the human host, genes involved in blood feeding and development, genes encoding proteins that represent new potential drug targets against hookworms, and expanded gene families encoding likely immunomodulator proteins, whose products may be beneficial in treating inflammatory diseases and asthma.[15]

Diagnostics

The most common method for diagnosing N. americanus is through identification of eggs in a fecal sample using a microscope. N. americanus eggs have a thin shell and are oval shaped, measuring roughly 56–74 by 36–40 μm.[16]

Treatments

The most common treatment for N. americanus infection are benzimidazoles, specifically albendazole and mebendazole. Benzimidazoles kill adult worms by binding to the nematode’s β-tubulin and subsequently inhibiting microtubule polymerization within the parasite.[17] The efficacy of single-dose treatments for hookworm infections were: 72% for albendazole, 15% for mebendazole, and 31% for pyrantel pamoate.[18] A current concern with this parasite is the increase in drug resistance, such as benzimidazoles and mebendazoles.[19]

Pregnant women should not be treated within their first trimester.[9]

During the 1940s, the treatment of choice was tetrachloroethylene,[20] given as 3 to 4 cc in the fasting state, followed by 30 to 45 g of sodium sulfate. Tetrachloroethylene was reported to have a cure rate of 80 percent for Necator infections, but 25 percent in Ancylostoma infections, and often produced mild intoxication in the patient.

Prevention and control

Infection and transmission of others can be prevented by not defecating outdoors or using human feces as fertilizer.[21] This parasite is not transmittable directly from person to person.[9] Pigs may be an intermediate host for N. americanus.[1]

Economic burden

N. americanus causes hookworm diseases, which are associated with blood loss and anemia. Patients who are infected with around 25 to 100 worms experience symptoms such as fatigue, weight loss, and slight headaches. As the infestation number reaches 100 to 500 worms, the patient experiences extreme fatigue, iron deficiency, and abdominal pain. The symptoms worsen and result in possible death when the infestation reaches over 500 hookworms. Children and pregnant women affected by N. americanus are at greater risk due to anemia and the greater need for dietary iron and protein. The demand is high for an improvement of sanitation to reduce fecal contamination in regions with high prevalence of N. americanus infections. The current control strategies include a combination of mass drug administration for children at age 4–6 years to prevent or eliminate N. americanus infections.[22]

References

  1. ^ a b "Necator americanus". Animal Diversity Web. Archived from the original on 2018-04-18. Retrieved 2018-04-21.
  2. ^ Georgiev VS (May 2000). "Necatoriasis: treatment and developmental therapeutics". Expert Opinion on Investigational Drugs. 9 (5): 1065–78. doi:10.1517/13543784.9.5.1065. PMID 11060728. S2CID 8040066. Archived from the original on 2018-08-20. Retrieved 2018-08-20.
  3. ^ Croese J, O'Neil J, Masson J, et al. (January 2006). "A proof of concept study establishing Necator americanus in Crohn's patients and reservoir donors". Gut. 55 (1): 136–7. doi:10.1136/gut.2005.079129. PMC 1856386. PMID 16344586.
  4. ^ Roberts, Janovy & Schmidt 2010
  5. ^ "Necator americanus". Animal Diversity Web. Archived from the original on 2018-04-18. Retrieved 2018-04-21.
  6. ^ "Hookworm (Intestinal) | CDC-DPDx". September 17, 2019. Archived from the original on 2009-12-15. Retrieved 2021-11-11..
  7. ^ "Hookworm disease". Encyclopædia Britannica. 2009. Archived from the original on 2021-11-11. Retrieved 2021-11-11.
  8. ^ a b c Hotez, Peter J; Bethony, Jeff; Bottazzi, Maria Elena; Brooker, Simon; Buss, Paulo (2005). "Hookworm: "The Great Infection of Mankind"". PLOS Medicine. 2 (3): e67. doi:10.1371/journal.pmed.0020067. ISSN 1549-1676. PMC 1069663. PMID 15783256.
  9. ^ a b c "Necator americanus". Archived from the original on 2018-04-19. Retrieved 2018-04-21.
  10. ^ St. Georgiev, Vassil (1999). "Parasitic Infections. Treatment and Developmental Therapeutics. 1. Necatoriasis". Current Pharmaceutical Design. Current Pharmaceutical Design. Vol. 7. Bentham Science. pp. 545–54. ISSN 1381-6128.
  11. ^ Phosuk I, Intapan PM, Thanchomnang T, Sanpool O, Janwan P, Laummaunwai P, Aamnart W, Morakote N, Maleewong W (2013). "Molecular detection of Ancylostoma duodenale, Ancylostoma ceylanicum, and Necator americanus in humans in northeastern and southern Thailand". Korean Journal of Parasitology. 51 (6): 747–49. doi:10.3347/kjp.2013.51.6.747. PMC 3916468. PMID 24516284.
  12. ^ Bleakley, Hoyt (2007). "Disease and Development: Evidence from Hookworm Eradication in the American South". The Quarterly Journal of Economics. 122 (1): 73–117. doi:10.1162/qjec.121.1.73. ISSN 0033-5533. PMC 3800113. PMID 24146438.
  13. ^ John, D.T.; Petri, W.A. (2013). "8 The Intestinal Nematodes: Necator americanus". Markell and Voge's Medical Parasitology (9th ed.). Elsevier Health Sciences. pp. 251–52. ISBN 978-0-323-27764-8. (N. americanus) is the native hookworm of Africa south of the Sahara and the only one found in parts of India. Ancylostoma and Necator occur together elsewhere in India, in much of China, in southeast Asia, in the islands of the South and Southwest Pacific, and in parts of Australia. ..., it is probable that the geographic boundaries between the two genera of human hookworms will disappear.
  14. ^ Tang YT, Gao X, Rosa BA, Abubucker S, Hallsworth-Pepin K, Martin J, Tyagi R, Heizer E, Zhang X, Bhonagiri-Palsikar V, Minx P, Warren WC, Wang Q, Zhan B, Hotez PJ, Sternberg PW, Dougall A, Gaze ST, Mulvenna J, Sotillo J, Ranganathan S, Rabelo EM, Wilson RK, Felgner PL, Bethony J, Hawdon JM, Gasser RB, Loukas A, Mitreva M (Mar 2014). "Genome of the human hookworm Necator americanus". Nature Genetics. 46 (3): 261–69. doi:10.1038/ng.2875. PMC 3978129. PMID 24441737.
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Necator americanus: Brief Summary

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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 of human hosts. Necatoriasis—a type of helminthiasis—is the term for the condition of being host to an infestation of a species of Necator. Since N. americanus and Ancylostoma duodenale (also known as Old World hookworm) are the two species of hookworms that most commonly infest humans, they are usually dealt with under the collective heading of "hookworm infection". They differ most obviously in geographical distribution, structure of mouthparts, and relative size.

Necator americanus has been proposed as an alternative to Trichuris suis in helminthic therapy.

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