Ancylostoma braziliense

Ge­o­graphic Range

An­cy­lostoma braziliense is the cat and dog hook­worm. En­demic in the south­ern United States, this species is pre­sent in a num­ber of sub­trop­i­cal re­gions around the world, in­clud­ing Cen­tral and South Amer­ica, and South­ern Asia. In South­ern Asia, the dis­tri­b­u­tion of An­cy­lostoma is con­fined to In­done­sia, Bor­neo, and Malaysia. The rea­son for this ge­o­graph­i­cal re­stric­tion is con­trary to an ex­pected wide­spread dis­tri­b­u­tion of the hook­worm due to the in­ter­min­gling of human pop­u­la­tions. Hu­mans and do­mes­tic an­i­mals have in­tro­duced species of An­cy­lostoma in other places around the world, such as Aus­tralia. (Richey, et al., 1996; Traub, et al., 2007)

Habi­tat

An­cy­lostoma braziliense eggs are passed into the en­vi­ron­ment through feces of cats and dogs. The eggs in­cu­bate on warm, moist soil. Adult hook­worms live in the small in­testines of many ver­te­brates. (Bal­four, et al., 2002; Richey, et al., 1996; Roberts and Janovy Jr, 2009)

Phys­i­cal De­scrip­tion

Hook­worms in the phy­lum Ne­ma­toda share a com­mon mor­phol­ogy as the an­te­rior end curves like a hook and the buc­cal cap­sule is scle­ro­tized and lined with teeth. Within the fam­ily An­cy­losto­mi­dae, the an­te­rior end is curved dor­sally. The ne­ma­tode body is cov­ered with a non-liv­ing cu­ti­cle shed by molt­ing. The width of the cu­tic­u­lar stri­a­tion pat­terns of A. braziliense is 3.45 µm.

Ne­ma­tode guts con­sist of a tube with an esoph­a­gus com­posed of the cor­pus, isth­mus, and bulb. Within the An­cy­losto­mi­dae, the esophogus is stout and mus­cu­lar. The buc­cal cav­ity (be­tween the mouth and esoph­a­gus) has a thick cu­tic­u­lar lin­ing which is large and heav­ily scle­ro­tized, termed a buc­cal cap­sule. Males of A. braziliense have a tu­ber­cu­lar process that is found at the buc­cal cap­sule of the mouth.

Male A. braziliense have two broad lat­eral lobes and a smaller dor­sal lobe with rays on the cop­u­la­tory bursa. The bur­sal rays dis­tin­guish dif­fer­ent species of An­cy­lostoma. In A. braziliense, the lat­eral bur­sal rays are sep­a­rated at the tips, and the po­si­tion of at­tach­ment of the ex­tern­odor­sal ray is unique in that it is closer to the be­gin­ning of the dor­sal trunk than in other species.

Fe­males are more dif­fi­cult to dis­tin­guish be­tween dif­fer­ent species, and usu­ally the teeth are the only di­ag­nos­tic tool that can be used. There is a prob­lem with this method, how­ever, be­cause for many species, the size and shape of the teeth do not ap­pear to be sig­nif­i­cantly unique. (Roberts and Janovy Jr, 2009; Traub, et al., 2007; Yoshida, 1971)

  • Sexual Dimorphism
  • female larger
  • Range length
    7.67 to 8.33 mm
    0.30 to 0.33 in

De­vel­op­ment

After the eggs are passed into the en­vi­ron­ment through the feces of cats or dogs, they hatch into first-stage ju­ve­niles. The first-stage ju­ve­nile feeds on soil bac­te­ria and feces for about a day. In the early stages of de­vel­op­ment, A. braziliense is non­in­fec­tious, and has a rhab­di­ti­form esoph­a­gus. The rhab­di­ti­form esoph­a­gus is char­ac­ter­is­tic of a feed­ing stage. Over a pe­riod of sev­eral days, the ne­ma­tode molts and reaches the third stage of de­vel­op­ment where be­comes an in­fec­tious fi­lar­i­form ju­ve­nile. The fi­lar­i­form esoph­a­gus is char­ac­ter­is­tic of non-feed­ing or liq­uid feed­ing stages, and the cor­pus is re­duced. At the third stage, the ju­ve­nile stops feed­ing be­cause it has ac­cu­mu­lated all the nu­tri­ents needed, and de­vel­op­ment pauses as the ju­ve­nile waits for a host. Once a proper de­fin­i­tive host has been found, the ju­ve­nile pen­e­trates the skin. At this stage, A. braziliense is found in the epi­der­mis, fol­li­cles, and glands of the skin, some­times ex­tend­ing to se­ba­ceous glands where they form coils. The ju­ve­nile mi­grates from the skin, and is car­ried to the heart and lungs by the blood­stream. In the lungs, the ju­ve­nile breaks into the alve­oli and is pro­pelled by cilia up the res­pi­ra­tory tract, where it is swal­lowed and ends up in the small in­tes­tine. The ju­ve­nile at­tachs to the in­testi­nal mu­cosa via the buc­cal cap­sule, and ma­tures to adult form by molt­ing twice. Once the last molt has fin­ished, the adult hook­worm is sex­u­ally ma­ture and ready to mate.

Al­though hu­mans can be­come in­fected, they are not true de­fin­i­tive hosts of A. braziliense. As a re­sult, the ju­ve­niles only mi­grate along the epi­der­mis, cre­at­ing le­sions. The ju­ve­nile even­tu­ally dies, never mak­ing it to the cir­cu­la­tory sys­tem or reach­ing adult­hood in the in­tes­tine. (Bal­four, et al., 2002; Brand and Haw­don, 2004; Bren­ner and Patel, 2003; Costa, et al., 2009; Nor­ris, 1971)

Re­pro­duc­tion

Males and fe­males mate using di­rect sperm trans­fer. Males have spicules that keep the fe­male re­pro­duc­tive open­ing gap­ing dur­ing sperm trans­fer. (Roberts and Janovy Jr, 2009)

Males and fe­males of A. braziliense have very dif­fer­ent roles dur­ing in­fec­tion and re­pro­duc­tion. The pri­mary role of fe­males is to lay eggs, and once a fe­male has suc­cess­fully in­fected a de­fin­i­tive host, it takes a min­i­mum of five weeks be­fore eggs are pro­duced. Males likely exist pri­mar­ily to fer­til­ize eggs, and pro­duce pro­tease in­hibitors. The role of pro­tease in­hibitors is still largely un­known, but thought that they are im­por­tant for par­a­sitism and sur­vival within the host.

Few stud­ies have been con­ducted on the re­pro­duc­tive genes of ne­ma­todes, but re­cent work has iden­ti­fied gen­der spe­cific genes in A. braziliense. Sci­en­tists in­ves­ti­gated sex spe­cific genes of un­known func­tion in A. braziliense, and com­pared them to sex spe­cific genes of known func­tion in other species. By look­ing for ho­mol­ogy, sci­en­tists dis­cov­ered that males of dif­fer­ent species also se­crete pro­tease in­hibitors, and that they some­how func­tion to en­sure sur­vival within the host. Sim­i­lar­i­ties among dif­fer­ent species of ne­ma­todes with known gene func­tion will help to de­fine the func­tion of pre­vi­ously un­known genes in A. braziliense. Such re­search also promises to bet­ter show the dif­fer­ent roles of males and fe­males in in­fec­tion, re­pro­duc­tion, and sur­vival within the host. (Costa, et al., 2009)

After eggs are laid, there is no parental in­vest­ment. (Costa, et al., 2009)

  • Parental Investment
  • no parental involvement
  • pre-hatching/birth
    • provisioning

Lifes­pan/Longevity

The longest lifes­pan is un­known, and the ex­pected lifes­pan varies de­pend­ing on the health of the host and the num­ber of adults caus­ing the in­fec­tion. (Bal­four, et al., 2002)

Be­hav­ior

The be­hav­ior of A. braziliense has not been widely stud­ied. How­ever, be­hav­ioral pat­terns of A. can­inum can be used as a way to gain a un­der­stand­ing of how in­fec­tive lar­vae lo­cate their hosts.

Tem­per­a­ture, light, and hu­mid­ity af­fect the be­hav­ioral pat­terns of A. can­inum, whereas car­bon diox­ide lev­els, light, and pH have no ef­fect. A tem­per­a­ture of ap­prox­i­mately 40 deg C at­tracts a ju­ve­nile, and it creeps to­wards the warmth in a snake-like way. Tem­per­a­tures above or below 40 deg C will not as ef­fec­tively stim­u­late host pen­e­tra­tion.

In ad­di­tion to warm tem­per­a­ture, host sur­face ex­tracts and serum also stim­u­late pen­e­tra­tion. Specif­i­cally, pen­e­tra­tion is stim­u­lated when ju­ve­niles of A. can­inum come into con­tact with the sur­face ex­tract of dogs, a de­fin­i­tive host. Sur­face ex­tracts and serum of other in­ter­me­di­ate hosts such as hu­mans, are not as ef­fec­tive in stim­u­lat­ing pen­e­tra­tion.

Lastly, A. can­inum and A. braziliense can­not sur­vive in­ges­tion by a de­fin­i­tive host, and as a re­sult, the ju­ve­niles do not es­tab­lish them­selves on pro­jec­tions like the ju­ve­niles of other ne­ma­tode species. In­stead, A. can­inum and A. braziliense pen­e­trate the skin of their hosts. Ex­cep­tions have been found among some species of An­cy­lostoma, how­ever, where the ju­ve­niles have sur­vived oral in­ges­tion by their de­fin­i­tive host. (Granzer and Haas, 1991)

Com­mu­ni­ca­tion and Per­cep­tion

Sen­sory or­gans on ne­ma­todes in­clude papil­lae, which are tac­tile re­cep­tors, and am­phids, which are chemore­cep­tors. Chem­i­cal cues are likely used to at­tract a mate. Sex­ual pheromones have been iden­ti­fied for at least 40 species of ne­ma­todes. Once a male and fe­male en­counter each other, tac­tile cues are used for re­pro­duc­tion.

Males of A. braziliense and A. can­inum can mon­i­tor the host-par­a­site re­la­tion­ship, but it is not known ex­actly how. An­cy­lostoma can­inum acts in­de­pen­dently and won't fol­low ju­ve­niles that have al­ready lo­cated and pen­e­trated a host. Re­cent stud­ies have shown that males se­crete pro­tease in­hibitors, which are thought to play an im­por­tant role in in­fec­tion and sur­vival within the host.

Sur­vival modes of A. braziliense have only re­cently been stud­ied. Pre­vi­ous re­search on other hook­worm species, such as Neca­tor amer­i­canus, has pro­vided the frame­work of how hook­worms evade the host im­mune sys­tem. Nec­tor amer­i­canus par­a­sitizes hu­mans, and it se­cretes prod­ucts that bind to human cells, in­hibit­ing a proper im­mune re­sponse. By tam­per­ing with the human im­mune sys­tem, N. amer­i­canus can en­sure its sur­vival. (Granzer and Haas, 1991; Roberts and Janovy Jr, 2009; Teix­eira-Car­valho, et al., 2008)

Food Habits

Adults feed on blood and in­testi­nal flu­ids in the gut of their host, lead­ing to iron de­fi­ciency and ane­mia for the host. The way in which hook­worms di­gest their blood meal is still being stud­ied, but it is known that adult hook­worms use a cas­cade of pro­teins to lyse red blood cells. A. braziliense ju­ve­niles do not feed on blood, rather they feed on dung and soil bac­te­ria. (Don, et al., 2007)

  • Animal Foods
  • blood
  • body fluids
  • Other Foods
  • dung
  • microbes

Pre­da­tion

There are no known preda­tors of A. braziliense. (Granzer and Haas, 1991)

Ecosys­tem Roles

An­cy­lostoma braziliense is an en­dopar­a­site of cats and dogs. Adult hook­worms eat blood from the in­tes­tine of cats and dogs, caus­ing ane­mia. (Don, et al., 2007)

Species Used as Host
  • Cat, Felis sylvestris
  • Dog, Canis lupis do­mes­ti­cus

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

A. braziliense has no pos­i­tive im­pact on hu­mans. (Bal­four, et al., 2002)

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Hu­mans are ac­ci­den­tal, dead-end hosts for hook­worms. In­fec­tious ju­ve­niles can pen­e­trate the epi­der­mis of hu­mans by tak­ing ad­van­tage of bro­ken skin or hair fol­li­cles. In­side the human host, the ju­ve­niles mi­grate through the epi­der­mis, com­monly called the creep­ing erup­tion. As the ju­ve­niles mi­grate, they cause ser­pingi­nous erup­tions on the skin, and they cause a tin­gling sen­sa­tion. The le­sions on the skin are typ­i­cally 3 mm wide, but in some cases the le­sions can ex­pand to reach 15 to 20 cm. The ju­ve­niles ad­vance through the epi­der­mis at a rate of a few mm per day. As the worms mi­grate, they fre­quently change di­rec­tion, and vesi­cles begin to form in their trail. The vesi­cles often be­come in­flamed, and as a re­sult the trails of the ju­ve­nile are vis­i­ble on the sur­face of the skin. The most com­mon site of in­fec­tion of hu­mans is the feet. Ad­di­tion­ally, it is quite com­mon for chil­dren to be­come in­fected on their but­tocks be­cause thin swim wear does not pro­vide a suf­fi­cient bar­rier to the in­fec­tious ju­ve­niles found in the warm, moist sand of trop­i­cal beaches.

Al­though the ju­ve­niles cre­ate un­com­fort­able le­sions on the skin, they die be­fore caus­ing any se­ri­ous harm be­cause hu­mans are dead-end hosts. Typ­i­cally, in­fected hu­mans do not need med­ical at­ten­tion be­cause the ju­ve­niles die after a few weeks or months. The most se­ri­ous neg­a­tive im­pact of A. braziliense on hu­mans comes from in­fect­ing their pets. In­fected cats and dogs re­quire ex­pen­sive med­ical at­ten­tion to stop the spread of the par­a­site. (Bal­four, et al., 2002; Roberts and Janovy Jr, 2009)

Con­ser­va­tion Sta­tus

As a par­a­site with only neg­a­tive af­fects on its hosts, A. braziliense has no con­ser­va­tion sta­tus.

Other Com­ments

The width of the cu­tic­u­lar stri­a­tion pat­terns of A. braziliense is 3.45 mi­crom­e­ters com­pared to 5.66 mi­crom­e­ters in A. cey­lan­icum. Males of A. braziliense have a tu­ber­cu­lar process that is found at the buc­cal cap­sule of the mouth, whereas this fea­ture is com­pletely ab­sent in other species, such as A. cey­lan­icum. (Traub, et al., 2007; Yoshida, 1971)

Con­trib­u­tors

Stephanie Chap­man (au­thor), Uni­ver­sity of Michi­gan-Ann Arbor, Heidi Liere (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor, John Marino (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor, Barry OCon­nor (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor, Renee Mul­crone (ed­i­tor), Spe­cial Pro­jects.

Glossary

Australian

Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.

World Map

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

agricultural

living in landscapes dominated by human agriculture.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

causes or carries domestic animal disease

either directly causes, or indirectly transmits, a disease to a domestic animal

chemical

uses smells or other chemicals to communicate

diapause

a period of time when growth or development is suspended in insects and other invertebrates, it can usually only be ended the appropriate environmental stimulus.

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

fertilization

union of egg and spermatozoan

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

infrared/heat

(as keyword in perception channel section) This animal has a special ability to detect heat from other organisms in its environment.

internal fertilization

fertilization takes place within the female's body

introduced

referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

parasite

an organism that obtains nutrients from other organisms in a harmful way that doesn't cause immediate death

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

Ref­er­ences

Bal­four, E., A. Zalka, R. La­zova. 2002. Cu­ta­neous larva mi­grans with parts of the larva in the epi­der­mis. Cutis, 69: 368-370. Ac­cessed April 16, 2011 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​12041816.

Brand, A., J. Haw­don. 2004. Phos­pho­inosi­tide-3-OH- ki­nase in­hibitor LY294002 pre­vents ati­va­tion of An­cy­lostoma can­inum and An­cy­lostoma cey­lan­icum third-stage in­fec­tive lar­vae. In­ter­na­tional Jour­nal for Par­a­sitol­ogy, 34: 909-914. Ac­cessed April 16, 2011 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​15217729.

Bren­ner, M., M. Patel. 2003. Cu­ta­neous larva mi­grans: The creep­ing erup­tion. Cutis, 72: 111-115. Ac­cessed April 16, 2011 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​12953933.

Car­valho, R., J. Araujo, F. Braga, S. Fer­reira, J. Araujo, A. Silva, L. Frassy, C. Alves. 2009. Bi­o­log­i­cal con­trol of An­clyos­tomis in dogs using the ne­ma­tode-trap­ping fun­gus Monacrospo­rium thau­ma­sium in south­east­ern Brazil. Vet­eri­nary Par­a­sitol­ogy, 165: 179-183.

Costa, A., A. Gomez- Ruiz, E. Ra­belo. 2008. Iden­ti­fi­ca­tion of gen­der-reg­u­lated genes in An­cy­lostoma braziliense by real-time RT-PCR. Vet­eri­nary Par­a­sitol­ogy, 153: 277-284. Ac­cessed April 16, 2011 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​18358613.

Costa, A., R. Gasser, S. Dias, E. Ra­belo. 2009. Male-en­riched tran­scrip­tion of genes en­cod­ing ASPs and Ku­niz-type pro­tease in­hibitors in An­cy­lostoma species. Mol­e­c­u­lar and Cel­lu­lar Probes, 23: 298-303. Ac­cessed April 16, 2011 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​19646525.

Don, T., Y. Oksov, S. Lustig­man, A. Loukas. 2007. Saposin-like pro­tein from the in­tes­tine of the blood-feed­ing hook­worm, An­cy­lostoma can­inum. Cam­bridge Jour­nals, 134: 427-436. Ac­cessed April 16, 2011 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​17109779.

Granzer, M., W. Haas. 1991. Host-find­ing and host recog­ni­tion of in­fec­tive An­cy­lostoma can­inum lar­vae. In­ter­na­tional Jour­nal for Par­a­sitol­ogy, 21 (4): 429-440. Ac­cessed April 16, 2011 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​1917283.

Labarthe, N., M. Lucia Ser­rao, A. Fer­reira, N. Almeida, J. Guer­rero. 2004. A sur­vey of gas­troin­testi­nal helminths in cats of the met­ro­pol­i­tan re­gion of Rio de Janeiro, Brazil. Vet­eri­nary Par­a­sitol­ogy, 123: 133-139. Ac­cessed Feb­ru­ary 05, 2010 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​1917283.

Nor­ris, D. 1971. The mi­gra­tory be­hav­ior of the in­fec­tive-stage lar­vae of An­cy­lostoma braziliense and An­cy­lostoma tubae­forme in ro­dent paratenic hosts. The Jour­nal of Par­a­sitol­ogy, 57 (5): 998-1009.

Richey, T., R. Gen­try, J. Fitz­patrick, A. Mor­gan. 1996. Per­sis­tant cu­ta­neous larva mi­grans due to An­cy­lostoma species. South­ern Med­ical Jour­nal, 89 (6): 609-611. Ac­cessed Feb­ru­ary 16, 2010 at http://​journals.​lww.​com/​smajournalonline/​Abstract/​1996/​06000/​Persistent_​Cutaneous_​Larva_​Migrans_​Due_​to.​10.​aspx.

Roberts, L., J. Janovy Jr. 2009. Foun­da­tions of Par­a­sitol­ogy. New York, NY: Mc­Graw- Hill.

Stothard, P., D. Pil­grim. 2003. Sex-de­ter­mi­na­tion gene and path­way evo­lu­tion in ne­ma­todes. Bioes­says, 25 (3): 221-231.

Teix­eira-Car­valho, A., R. Fu­ji­wara, E. Stemmy, D. Oive, J. Damsker, A. Loukas, R. Cor­rea-Oliveira, S. Con­stant, J. Bethony. 2008. Bind­ing of ex­creted and/or se­creted prod­ucts of adult hook­worms to human NK cells in Neca­tor amer­i­canus-in­fected in­di­vid­u­als from Brazil. Amer­i­can So­ci­ety For Mi­cro­bi­ol­ogy, 76 (12): 5810-5816.

Traub, R., R. Hobbs, P. Adams, J. Behnke, P. Har­ris, C.A Thomp­son. 2007. A case of mis­taken iden­tity-reap­praisal of the species of canid and felid hook­works (An­cy­lostoma) pre­sent in Aus­tralia and India. Par­a­sitol­ogy, 134: 113-119.

Williamson, A., S. Lustig­man, Y. Oksov, V. Deu­mic, J. Plieskatt, S. Mendez, B. Zhan, M. Bot­tazzi, P. Hotez, A. Loukas. 2006. An­cy­lostoma can­inum MTP-1, an astacin-like met­al­lo­pro­tease se­creted by in­fec­tive hook­worm lar­vae, is in­volved in tis­sue mi­gra­tion. In­fec­tion and Im­munol­ogy, 74 (2): 961-967. Ac­cessed April 16, 2011 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​16428741.

Yoshida, Y. 1971. Com­par­a­tive stud­ies on An­cy­lostoma braziliense and An­cy­lostoma cey­lan­icum. I. The adult stage. The Jour­nal of Par­a­sitol­ogy, 57 (5): 983-989. Ac­cessed April 16, 2011 at http://​www.​ncbi.​nlm.​nih.​gov/​pubmed/​4851100.