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PubMed2000

Genes and genomes of Necator americanus and related hookworms.

The human hookworms (Necator americanus and Ancylostoma duodenale) infect over one billion people. The phylogenetic relationships of the human hookworms suggest independent acquisition of the human host. The hookworms probably have a haploid chromosome number n = 6, and an XX-XO sex determination mechanism is likely to be used. Genetic and molecular research on hookworms is in its infancy, but several important genes and gene products have already been identified. Of note are cathepsin genes, a family of secreted proteins known as Ancylostoma activation-associated proteins and a family of anticoagulants. The inception of an expressed sequence tag program on the human hookworm, N. americanus, promises to yield many new genes with novel functions in the biology of these important parasites.

Animals↗

PubMed2000

Hookworm burden reductions in BALB/c mice vaccinated with recombinant Ancylostoma secreted proteins (ASPs) from Ancylostoma duodenale, Ancylostoma caninum and Necator americanus.

Vaccination of mice with alum-precipitated recombinant Ancylostoma secreted protein-1 from the canine hookworm Ancylostoma caninum (Ac-ASP-1) results in protection against A. caninum larval challenge. Vaccine protection is manifested by host reductions in hookworm burden compared to control mice. The goal of this study was to determine whether ASP antigens cloned and expressed from different hookworm species will cross protect against A. caninum larval challenge. Cross-species protection against A. caninum challenge infections was observed with immunizations using recombinant ASP-1 from the human hookworms Ancylostoma duodenale and Necator americanus. However, the degree of protection was proportional to the extent of amino acid sequence homology between the ASP immunogen used for vaccination and the Ac-ASP-1 produced by the challenge larval strain. Vaccine protection was noted to decrease significantly as amino acid sequence homologies diverged 10% or more. It was also determined that Ac-ASP-2, a molecule cloned from A. caninum having 55% amino acid sequence homology to the C-terminus of Ac-ASP-1, did not elicit vaccine protection. These observations were partly reflected in the titer of antibodies that recognize Ac-ASP-1. The studies reported here will help to design immunogenic peptide vaccines based on the sequence divergence of hookworm ASPs.

Ancylostoma↗

PubMed1993

Hookworm infections and human iron metabolism.

Ancylostoma duodenale and Necator americanus are extremely common species of soil-transmitted helminth which flourish where poverty and malnutrition prevail. Hookworms contribute significantly to iron-deficiency anaemia, which remains one of the world's major nutritional problems, through the feeding activities of intestinal stages leading to chronic blood loss into the gut. In this article, a mathematical model is proposed to explain how human iron metabolism may respond to hookworm infection of varying intensity. The model draws attention to the importance of the regulation of stored iron levels in the process. The results from the model are presented for the effects of hookworm infection on the iron metabolism of a healthy adult male. Calculations are also presented in which the effects of hookworms on the iron metabolism of a non-pregnant woman are compared with those of a pregnant woman. Use of the model may help develop a better understanding of the pathology of hookworm disease.

Adolescent↗

PubMed2001

Mitigation of hookworm disease by immunization with soluble extracts of Ancylostoma ceylanicum.

Hookworms are a leading cause of anemia in developing countries, and a strategy aimed at reducing pathology caused by blood-feeding adult parasites would be a valuable addition to global control efforts. This article describes experiments designed to induce resistance to the major clinical sequelae (weight loss and anemia) of Ancylostoma ceylanicum hookworm infection in Syrian golden hamsters of the outbred LVG strain. Previously infected animals acquired long-lived resistance to weight loss and anemia caused by a secondary hookworm infection. Furthermore, transfer of pooled serum from twice-infected hamsters to animals undergoing a primary infection was associated with partial resistance to growth delay and anemia. Active vaccination of hamsters with soluble adult hookworm antigens emulsified in alum led to partial protection from hookworm-associated pathology in the absence of reductions in adult worm burden. This intriguing result may have important implications for human vaccine development.

Ancylostoma↗

PubMed1998

Epidemiological evidence for a differential effect of hookworm species, Ancylostoma duodenale or Necator americanus, on iron status of children.

BACKGROUND: The hookworms, Ancylostoma duodenale and Necator americanus, cause significant gastrointestinal blood loss. In clinical studies, greater blood losses have been reported with A. duodenale. However, there has been no evidence that endemic A. duodenale infection has greater impact than N. americanus infection on the iron status of populations. METHODS: In a sample of 525 school children in Pemba Island, Tanzania, we compared the degree of anaemia and iron deficiency associated with the two hookworm species at the individual and community (i.e. school) levels. Multiple regression was used to control for infection intensities and other child characteristics. RESULTS: In the 492 children with hookworm positive faecal cultures, haemoglobin and ferritin concentrations decreased with increasing proportions of A. duodenale. Among children with only N. americanus larvae, the prevalence of anaemia was 60.5 % and the prevalence of ferritin <12 microg/l was 33.1%, while in children with > or =50% A. duodenale larvae, the respective prevalences were 80.6% and 58.9%. When children were grouped by the prevalence of A. duodenale at the school level, children from high prevalence (> or =20%) schools had significantly worse iron deficiency and anaemia than children from low prevalence schools. CONCLUSIONS: The species of hookworm being transmitted in a community influences the burden of iron deficiency anaemia in the community, and should be considered in prioritizing and planning programmes for hookworm and anaemia control.

Ancylostomiasis↗

PubMed2002

Cleavage of hemoglobin by hookworm cathepsin D aspartic proteases and its potential contribution to host specificity.

Hookworms routinely reach the gut of nonpermissive hosts but fail to successfully feed, develop, and reproduce. To investigate the effects of host-parasite coevolution on the ability of hookworms to feed in nonpermissive hosts, we cloned and expressed aspartic proteases from canine and human hookworms. We show here that a cathepsin D-like protease from the canine hookworm Ancylosotoma caninum (Ac-APR-1) and the orthologous protease from the human hookworm Necator americanus (Na-APR-1) are expressed in the gut and probably exert their proteolytic activity extracellularly. Both proteases were detected immunologically and enzymatically in somatic extracts of adult worms. The two proteases were expressed in baculovirus, and both cleaved human and dog hemoglobin (Hb) in vitro. Each protease digested Hb from its permissive host between twofold (whole molecule) and sixfold (synthetic peptides) more efficiently than Hb from the nonpermissive host, despite the two proteases' having identical residues lining their active site clefts. Furthermore, both proteases cleaved Hb at numerous distinct sites and showed different substrate preferences. The findings suggest that the paradigm of matching the molecular structure of the food source within a host to the molecular structure of the catabolic proteases of the parasite is an important contributing factor for host-parasite compatibility and host species range.

Ancylostomatoidea↗

PubMed2004

Hookworm (Necator americanus) transmission in inland areas of sandy soils in KwaZulu-Natal, South Africa.

This study extended the association between hookworm transmission in KwaZulu-Natal and the sandy coastal plain by investigating the parasite's occurrence in isolated areas of sandy soils further inland. A school-based prevalence survey was carried out in selected inland sandy areas and in surrounding areas dominated by clay soils within a narrow altitudinal range of between 500 and 700 m to reduce the effect of altitude on climate-related factors (rainfall and temperature). Sandy areas situated on the coastal plain were included in the analysis for comparative purposes. Soil samples (0-50 mm depth) were collected from each locality to assess their nematode loadings and to analyse selected physical and chemical properties. Significant differences were found between the moderate prevalence of hookworm infection among children living in inland areas with sandy soils (17.3%) and the low prevalence in surrounding non-sandy areas (5.3%, P < 0.001), and between infection among children living in all inland areas (9.3%) and the high prevalence on the coastal plain (62.5%, P < 0.001). Amounts of fine and medium sand were highest in both the coastal plain soils and in inland sandy areas and these fractions showed a significant positive correlation with hookworm prevalence and nematode loadings. Clay, coarse sand and organic matter contents were highest in surrounding non-sandy soils and showed a significant negative correlation with the nematode variables. No statistically significant correlations were found with soil pH at study localities. We conclude that properties of inland sandy soils, particularly particle size distribution, correlate well with hookworm prevalence and nematode loadings and therefore provide a more suitable habitat for nematodes than surrounding non-sandy areas. These results suggest that particle size distribution of sand fractions, organic matter and clay content in the soil influence the survival of hookworm larvae and hence the parasite's transmission.

Adolescent↗

PubMed1992

Hyaluronidase from infective Ancylostoma hookworm larvae and its possible function as a virulence factor in tissue invasion and in cutaneous larva migrans.

During skin penetration, infective hookworm larvae encounter hyaluronic acid as they migrate between epidermal keratinocytes and through the ground substance of the dermis. A hyaluronidase would facilitate passage through the epidermis and dermis during larval invasion. Zoonotic hookworm larvae of the genus Ancylostoma were shown to contain a hyaluronidase activity that migrated on modified sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE) hyaluronic acid gels with an apparent Mr of 49,000. A second form with an Mr of 87,000 was also identified. The major etiologic agent of cutaneous larva migrans, A. braziliense, was shown to have the greatest enzyme activity, hydrolyzing up to 3.3 micrograms of hyaluronic acid per h per micrograms of total parasite protein at pH 6.0, whereas A. caninum and A. tubaeforme each had much less enzyme activity. The differences in enzyme activities between species correlated with differences in the intensities of the lytic zones at 49 and 87 kDa on SDS-PAGE hyaluronic acid gels. Hookworm hyaluronidase activity exhibited a broad pH optimum between 6.0 and 8.0 and did not hydrolyze chondroitin sulfate, two features that suggest that the hookworm enzyme is more like the invertebrate leech hyaluronidase than mammalian testicular or lysosomal hyaluronidase. Larvae of A. braziliense were shown to release hyaluronidase activity and degrade radiolabeled hyaluronic acid in vitro. Gold sodium thiomalate was identified as an enzyme inhibitor. The hyaluronidase is the second major virulence factor that we have identified from infective hookworm larvae.

Ancylostoma↗

PubMed2003

Oral transfer of adult Ancylostoma ceylanicum hookworms into permissive and nonpermissive host species.

Syrian hamsters become anemic and exhibit delayed growth following oral infection with third-stage Ancylostoma ceylanicum hookworm larvae. Here we describe experiments designed to determine the feasibility of adult worm transfer (AWT) between hosts, a technique that would facilitate the specific study of bloodfeeding hookworms in vivo without prior exposure of the host to larva-specific antigens, permit the ex vivo manipulation of adult parasites prior to reimplantation, and also allow for cross-species transfer of worms. Weanling hamsters given an oral AWT of 40 or 60 mixed-sex A. ceylanicum worms rapidly developed anemia; in the higher-dose group, hemoglobin levels declined from prechallenge levels by 44% within 4 days following AWT. Long-term survival of transferred worms was demonstrated by recovery of parasites from the intestines 42 days after AWT. AWT hamsters acquired humoral immune responses against soluble adult hookworm extracts and excretory-secretory products that were comparable in magnitude to those of animals given a typical infection with larvae. In AWT experiments employing the nonpermissive murine model, C57BL/6 mice given adult worms rapidly became anemic and lost weight in a manner similar to AWT hamsters. Infection of additional mouse strains demonstrated that while C57BL/10 and CD-1 mice also developed anemia following AWT, BALB/c mice were resistant. The technique of AWT to mice may further our understanding of hookworm pathogenesis by allowing the study of adult hookworm infections in a species with well-characterized genetics and an abundance of available reagents.

Administration, Oral↗

PubMed2004

Molecular characterization of Ancylostoma ceylanicum Kunitz-type serine protease inhibitor: evidence for a role in hookworm-associated growth delay.

Hookworm infection is a major cause of iron deficiency anemia and malnutrition in developing countries. The Ancylostoma ceylanicum Kunitz-type inhibitor (AceKI) is a 7.9-kDa broad-spectrum inhibitor of trypsin, chymotrypsin, and pancreatic elastase that has previously been isolated from adult hookworms. Site-directed mutagenesis of the predicted P1 inhibitory reactive site amino acid confirmed the role of Met(26) in mediating inhibition of the three target serine proteases. By using reverse transcription-PCR, it was demonstrated that the level of AceKI gene expression increased following activation of third-stage larvae with serum and that the highest level of expression was reached in the adult stage of the parasite. Immunohistochemistry studies performed with polyclonal immunoglobulin G raised against recombinant AceKI showed that the inhibitor localized to the subcuticle of the adult hookworm, suggesting that it has a potential in vivo role in neutralizing intestinal proteases at the surface of the parasite. Immunization with recombinant AceKI was shown to confer partial protection against hookworm-associated growth delay without a measurable effect on anemia. Taken together, the data suggest that AceKI plays a role in the pathogenesis of hookworm-associated malnutrition and growth delay, perhaps through inhibition of nutrient absorption in infected hosts.

Amino Acid Sequence↗

PubMed1999

Geographic distribution and epidemiology of Oesophagostomum bifurcum and hookworm infections in humans in Togo.

In contrast to the rest of the world, infections with Oesophagostomum bifurcum are commonly found in humans in northern Togo and Ghana. In addition, infections with hookworm are endemic in this region. In the present study, a detailed map of the geographic distribution of O. bifurcum and hookworm infections in northern Togo was made. There were a number of foci with high prevalence of infection with O. bifurcum. All the villages examined were infected with hookworm, and the distribution was quite patchy. Women were infected with O. bifurcum more often than men, while infections with hookworm were more prevalent in men than in women. The prevalence and intensity of infection with both parasites were clearly age-dependent. We estimate that more than a 100,000 people in Togo are infected with O. bifurcum and more than 230,000 are infected with hookworm.

Adolescent↗

PubMed1995

The changing epidemiology of human hookworm infection in Australia.

OBJECTIVE: To document the distribution of human hookworm infection in Australia. DESIGN: A retrospective study of published data and accessible archival records. METHODS: The core data were derived from the report of the Australian Hookworm Campaign, annual reports of various State health departments and the Queensland Institute of Medical Research, and unpublished files of the former Queensland Aboriginal Health Programme. FINDINGS: Today, hookworm infection is endemic only among Aboriginal communities of northern Western Australia and the Northern Territory, where Ancylostoma duodenale may be the sole species. Early this century, infection was also established in Queensland and northern New South Wales, in both Aboriginal and white communities, and Necator americanus predominated. The origin of these parasites is obscure. Sustained control programs seem to have eradicated hookworms from Queensland. CONCLUSIONS: Improved sanitation, hygiene and chemotherapy have eliminated hookworms from white populations of Australia. Continued anthelminthic campaigns will also clear the parasites from Aboriginal communities, but unless living conditions improve significantly infection will recur.

Ancylostomiasis↗

PubMed1999

Enzyme-linked immunoelectrotransfer blotting analysis of human serologic responses to infective hookworm larval antigen.

OBJECTIVE: To explore the possibility of using specific antigens for immunodiagnosis of hookworm disease in endemic area. METHOD: Infective third-stage larvae of the canine hookworm, Ancylostoma caninum (A. caninum), were prepared as the source of antigen. Enzyme-linked immunoelectrotransfer blotting (EITB) was employed as an immunodiagnostic method. RESULTS: Two immunodominant bands of hookworm antigens (42 kDa and 55 kDa) were recognized by the sera of hookworm-infected patients (serum dilution 1:200; antigen centrifuged at 36,000 r/m for 20 minutes, but not by sera from negative controls. CONCLUSION: The 42 kDa and 55 kDa A. caninum antigens might be the specific antigens that could be used for immunodiagnosis of hookworm disease in endemic area.

Ancylostoma↗

PubMed1990

[Effect of albendazole and pyrantel in treating intestinal helminthiasis and controlling the recurrence of hookworm infections].

A total of 720 human intestinal helminthic infections were divided into 4 groups and treated with albendazole 400mg/d x 3d, 400mg/d x 5d, pyrantel 1,500mg/d x 3d, or 1,500mg/d x 5d. Half a month after treatment, the negative rates of hookworm egg were 98.6, 98.6, 86.2 and 93.5%, those of ascaris egg were 96.5, 98.2, 92.9 and 96.3%, and those of whipworm egg were 86.4, 89.0, 68.9 and 67.0% respectively. Reduction rate of hookworm egg reached more than 98% in all the 4 groups. Six months after treatment, however, the positive rates of all the 4 groups rose again in varying degrees. The predominant species of hookworm infections was Necator americanus before the treatment and Ancylostoma duodenale after the treatment. It was demonstrated that the recurrence of hookworm infection resulted from A. duodenale infections, while a single dose of 400mg albendazole per day for 3 or 5 days showed good effect in controlling the recurrence of hookworm infections in a certain area.

Adolescent↗

PubMed1981

The relationship of hookworm infection, anaemia and iron status in a Papua New Guinea highland population and the response to treatment with iron and mebendazole.

In 345 apparently healthy Papua New Guinea male subjects, predominantly highlanders, 89% of whom were infected with hookworm (Necator americanus), there was a statistically significant inverse correlation of hookworm egg count with haemoglobin and serum ferritin level, but no significant correlation with serum albumin, folate or B12 values. A sub-group of 128 was chosen for a six-month study on the effect of treatment with the anthelmintic mebendazole and/or parenteral iron on haemoglobin and serum ferritin levels. Mebendazole-treated subjects remained worm-free and the hookworm egg counts of the controls decreased during the study period. Parenteral iron treatment had the expected effect of raising haemoglobin to a normal level. There was a statistically significant improvement in haemoglobin level in all treated groups but not in the control. Serum ferritin levels decreased significantly in all groups, but more in the control than in the treated groups, although treatment groups were not significantly different. Although probable inadequate uptake of iron by the subjects and blood donation by some subjects was apparently more detrimental to iron status than hookworm infection, the results of this study support the view that hookworm infection in this country contributes to lowered haemoglobin levels and iron status.

Anemia, Hypochromic↗

PubMed1993

[Field studies on different control schemes for hookworm infection].

Three different control schemes including selective chemotherapy, a combination of selective chemotherapy and management of fecal disposal and mass treatment with mebendazole-medicated salt were implemented for the control of hookworm infection in Luwo Town and Taiping Village, Wuming County, Guangxi Zhuang Autonomous Region during 1971-1990. Through ten years of selective chemotherapy, the results showed that the hookworm infection rate dropped from 42.6% to 3.9% and the infection intensity from 425.6 eggs per gram faeces to 1.6 eggs per gram faeces. After seven years of selective chemotherapy combined with management of fecal, the hookworm infection rate dropped from 58.7% to 10.9% and the infection intensity from 111.5 eggs per gram faeces to 8.5 eggs per gram faeces. However, 3 years after the termination of the combined treatment, the hookworm infection rate increased to 14.5%. As for the implementation of the third scheme, mebendazole-medicated salt was given at 100 mg daily for 30 consecutive days or at 50 mg daily for 30 consecutive days. Stool examination was made one year after the treatment, the results showed that the infection rate of the inhabitants dropped from 36% and 36% to 2% and 1.3%, respectively, and the infection intensity dropped from 110.3 and 246 eggs per gram faeces to 1.3 and 0.24 eggs per gram faeces. After three years, the infection rate of inhabitants remained under 2% and the infection intensity under 1 egg per gram faeces. It was concluded that mass treatment with mebendazole-medicated salt might be the most practical scheme for the control of hookworm infection.

Ancylostomiasis↗

PubMed1998

Ancylostoma duodenale is responsible for hookworm infections among pregnant women in the rural plains of Nepal.

Fecal specimens from 292 pregnant women (ages 15-40 yr) and 129 infants (ages 10-20 wk) were examined for helminth eggs by the Kato-Katz method and cultured for helminth larvae identification using a modified Harada Mori method. These specimens were collected from June 1995 through July 1996 in Sarlahi District in the southern rural plains of Nepal. Among pregnant women, the prevalence of helminth infection by the Kato-Katz method was 78.8%, 56.2%, and 7.9% for hookworm, Ascaris lumbricoides, and Trichuris trichiura, respectively. Using the modified Harada-Mori method, 66.1% and 2.0% of women's fecal cultures were positive for hookworm and Strongyloides stercoralis, respectively. All of the cultured hookworm larvae were identified as Ancylostoma duodenale. Among infants, 1 specimen was positive for hookworm and 1 for A. lumbricoides using the Kato-Katz method. The modified Harada Mori method detected no larvae in specimens from infants. There was 81.8% agreement between the 2 methods for the detection of hookworm infection. Ancylostoma duodenale is endemic in this study population and highly prevalent in pregnant women.

Adolescent↗

PubMed2005

Historic importance of some aspects of research by O. Wilford Olsen on hookworms (Uncinaria lucasi) in northern fur seals (Callorhinus ursinus) and Steller sea lions (Eumatopias jubatus) in 1951 on St. Paul Island, Alaska.

The purpose of this paper is to disseminate, more widely, certain historically important aspects of research by O. Wilford Olsen on hookworms (Uncinaria lucasi) in northern fur seals (Callorhinus ursinus) and Steller sea lions (Eumatopias jubatus) in 1951 on St. Paul Island, Alaska. Highly meaningful facets of the research were recorded only in a report with limited availability by Olsen in 1952. Portions of his research presented here include measurements of hookworm free-living infective third-stage larvae (L3) and adults, photographs of bursae, and the conclusion that the species of hookworms (U. lucasi) is probably the same in both pinniped hosts. This information is especially important because very little has been published on taxonomy of hookworms in E. jubatus.

Alaska↗